Tick and Tickborne Disease

Ticks and Tick-borne Diseases
Geographical Distribution and Control Strategies

in the Euro-Asia Region
This page intentionally left blank
Ticks and Tick-borne Diseases

Geographical Distribution and Control Strategies
in the Euro-Asia Region
Edited by
Mo Salman and Jordi Tarrs-Call

Authors:
Agustn Estrada-Pea, Rbert Farkas, Thomas G.T. Jaenson,

Frank Koenen, Maxime Madder, Ilaria Pascucci, Mo Salman,
Rita de Sousa and Alan R. Walker
CABI is a trading name of CAB International

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Library of Congress Cataloging-in-Publication Data
Salman, Mowafak Dauod.
Ticks and tick-borne diseases : geographical distribution and control strategies in the Euro-Asia region / edited by Mo Salman and Jordi Tarrs-Call;
authors, Agustn Estrada-Pea [et al.].
p. ; cm.
Includes bibliographical references and index.
ISBN 978-1-84593-853-6 (alk. paper)
I. Tarrs-Call, Jordi. II. Estrada-Pea, Agustn. III. Title.
[DNLM: 1. Tick-Borne Diseases--epidemiology--Asia. 2. Tick-Borne Diseases-epidemiology--Europe. 3. Tick-Borne Diseases--prevention & control--Asia.
4. Tick-Borne Diseases--prevention & control--Europe. 5. Ticks--pathogenicity-Asia. 6. Ticks--pathogenicity--Europe. 7. Topography, Medical--Asia. 8. Topography, Medical--Europe. WC 600]
614.5'263094--dc23
2012024123
ISBN-13: 978 1 84593 853 6
Commissioning editor: Sarah Hulbert
Editorial assistant: Alexandra Lainsbury
Production editor: Shankari Wilford
Typeset by SPi, Pondicherry, India
Printed and bound in the UK by the MPG Books Group
Contents
Contributors
ix
Foreword
Jeremy Gray
xi
Acknowledgements
Abbreviations
Introduction
Significance of Tick-borne Diseases
Geographic Scope
Methods Used for the Systematic Review of the Literature
Identification Methods for Tick Species, Including the Most Recent
Advanced Techniques and their Limitations
1
Emerging (Re-emerging) Tick-borne Infections and the

Dissemination of Ticks
Mo Salman and Agustn Estrada-Pea
Basic Biology and Geographical Distribution of Tick Species Involved
in the Transmission of Animal Pathogens, Including Zoonoses
Rbert Farkas, Agustn Estrada-Pea, Thomas G.T. Jaenson, Ilaria Pascucci
and Maxime Madder
2.1 Soft Ticks (Argasidae)
2.2 Hard Ticks (Ixodidae)
Factors Influencing the Spread and Distribution of Ticks
Maxime Madder and Ilaria Pascucci
3.1 Climate Changes
3.2 Changes in Land Use
3.3 Movement of Animals
xiii
xv
xvii
xvii
xviii
xviii
xxi
6
11
27
27
28
29
vi
Contents
Tick-borne Infections (Including Zoonoses) in Europe

and the Mediterranean Basin
Frank Koenen, Ilaria Pascucci, Thomas G.T. Jaenson, Maxime Madder,
Rita de Sousa, Agustn Estrada-Pea, Rbert Farkas and Mo Salman
4.1 African Swine Fever
4.2 Crimean-Congo Haemorrhagic Fever
4.3 Tick-borne Encephalitis Group
4.4 Anaplasmoses
4.5 Ehrlichioses
4.6 Rickettsioses
4.7 Lyme Borrelioses
4.8 Recurrent (Relapsing) Fever
4.9 Piroplasmoses
4.10 Hepatozoonosis
4.11 Other Potentially Tick-borne Infections
Maps of Reported Occurrence of Tick-borne Pathogens
Agustn Estrada-Pea, Frank Koenen, Ilaria Pascucci, Thomas G.T. Jaenson,
Maxime Madder, Rita de Sousa and Rbert Farkas
5.2 Crimean-Congo Haemorrhagic Fever
5.3 Tick-borne Encephalitis Group Viruses
5.4 Anaplasmoses and Ehrlichioses
5.5 Rickettsioses
5.6 Borrelioses
5.7 Babesioses
5.8 Theilerioses
5.9 Equine Piroplasmoses
5.10 Bartonelloses
5.11 Tularaemia
5.12 Q fever
Maps of Reported Occurrence of Ticks
Agustn Estrada-Pea, Thomas G.T. Jaenson, Rbert Farkas and Ilaria Pascucci
6.1 Soft Ticks (Argasidae), Genus Ornithodoros
6.2 Hard Ticks (Ixodidae)
Surveillance and Control Measures
Rbert Farkas, Mo Salman and Agustn Estrada-Pea
7.1 Introduction to Surveillance and its Aim
7.2 Control Options for Hard Ticks
7.3 Control Possibilities for Soft Ticks
Glossary
33
33
36
37
39
43
44
51
52
53
56
57
76
77
77
78
80
80
81
82
83
84
85
86
86
89
89
90
98
98
104
110
115
Appendices
Appendix 1 Tables of Geographic Distribution Data for Ticks and
Tick-borne Diseases
Table A1.1. Geographic distribution data for African swine fever virus (ASFV)
Table A1.2. Geographic distribution data for Crimean-Congo
haemorrhagic fever virus (CCHFV)
117
117
118
Contents
Table A1.3. Geographic distribution data for the tick-borne encephalitis

(TBE) group
Table A1.4. Geographic distribution data for Anaplasma spp. and Ehrlichia spp.
now included in Anaplasma spp.
Table A1.5. Geographic distribution data for Rickettsia spp.
Table A1.6. Geographic distribution data for Borrelia spp.
Table A1.7. Geographic distribution data for Babesia spp.
Table A1.8. Geographic distribution data for Theileria spp.
Table A1.9. Geographic distribution data for equine piroplasmoses
Table A1.10. Geographic distribution data for Bartonella spp.
Table A1.11. Geographic distribution data for Francisella tularensis
Table A1.12. Geographic distribution data for Coxiella burnetii
Table A1.13. Geographic distribution data for Ornithodoros spp.
Table A1.14. Geographic distribution data for Ixodes ricinus
Table A1.15. Geographic distribution data for Haemaphysalis punctata
Table A1.16. Geographic distribution data for Haemaphysalis concinna
Table A1.17. Geographic distribution data for Haemaphysalis inermis
Table A1.18. Geographic distribution data for Hyalomma marginatum
Table A1.19. Geographic distribution data for Rhipicephalus sanguineus
group (Rh. sanguineus and Rh. turanicus)
Table A1.20. Geographic distribution data for Rhipicephalus bursa
Table A1.21. Geographic distribution data for Rhipicephalus (Boophilus) annulatus
Table A1.22. Geographic distribution data for Dermacentor marginatus
Table A1.23. Geographic distribution data for Dermacentor reticulatus
Table A1.24. Data from serological surveillance of Crimean-Congo
haemorrhagic fever (CCHF) in domestic animals
Table A1.25. Geographic distribution data for Rickettsia spp.
vii
120
125
134
139
159
166
168
169
170
171
174
176
201
205
205
206
213
237
244
246
252
255
258
Appendix 2 List (By ID Number) of References from which Geographic Data

Were Extracted (hosted on http://www.cabi.org/resources/tatbd).
272
Appendix 3 List of Scientific Papers That Could Not Be Retrieved

(hosted on http://www.cabi.org/resources/tatbd).
272
Appendix 4 Keys for Genera and Species of Ticks in the Mediterranean Basin
273
Index
289
The colour plate section can be found following p. 8
Contributors
Agustn Estrada-Pea, Professor of Parasitology, Department of Animal Pathology

(Parasitology), Faculty of Veterinary Medicine, University of Zaragoza, Miguel Servet 177,
50013-Zaragoza, Spain. E-mail: AEstrada@unizar.es
Rbert Farkas, Professor in Parasitology, Department of Parasitology and Zoology, Faculty of
Veterinary Science, Szent Istvn University, H-1078 Budapest VII, Istvn u. 2, Hungary.
E-mail: Farkas.Robert@aotk.szie.hu
Thomas G.T. Jaenson, Professor in Medical Entomology, Medical Entomology Unit,
Department of Systematic Biology, Evolutionary Biology Centre, Uppsala University,
Norbyvgen 18d, SE-752 36 Uppsala, Sweden. E-mail: Thomas.Jaenson@ebc.uu.se
Frank Koenen, Operational Director, Interactions and Surveillance, Veterinary and
Agrochemical Research Centre (Centrum voor Onderzoek in Diergeneeskunde en
Agrochemie-Centre dEtude et de Recherches Vtrinaires et Agrochimiques) (CODACERVA), Groeselenberg 99, B-1180 Brussels, Belgium. E-mail: Frkoe@var.fgov.be
Maxime Madder, Senior Researcher, Biologist, PhD, Veterinary Entomology Group,
Department of Biomedical Sciences, Institute of Tropical Medicine, Nationalestraat
155B-2000 Antwerp, Belgium, and Extra-ordinary Lecturer, Department of Veterinary
Tropical Diseases, Faculty of Veterinary Science, University of Pretoria, Onderstepoort
0110, South Africa. E-mail: Mmadder@itg.be
Ilaria Pascucci, DMV, PhD, Istituto Zooprofilattico Sperimentale dellAbruzzo e del Molise
G. Caporale, Campo Boario, I-64100 Teramo, Italy. E-mail: I.Pascucci@izs.it
Mo Salman, Professor of Veterinary Epidemiology, Campus Stop 1644, Animal Population Health Institute, College of Veterinary Medicine and Biomedical Sciences,
Colorado State University, Fort Collins, CO 80523-1644, USA. E-mail: M.D.Salman@
ColoState.edu
Rita de Sousa, Centro de Estudos de Vectores e Doenas Infecciosas Doutor Francisco
Cambournac (Center for the Study of Vectors and Infectious Disease Research Doutor
Francisco Cambournac) (CEVDI), Instituto Nacional de Sade Doutor Ricardo Jorge, IP
(National Institute of Health Doutor Ricardo Jorge, IP) (INSA), Av. da Liberdade 5, 2965-575
guas de Moura, Portugal. E-mail: Rita.Sousa@insa.min-saude.pt
ix
Contributors
Jordi Tarrs-Call, DMV, PhD, Scientific Officer of the FEED Unit (formerly Scientific Officer in
the Animal Health and Animal Welfare Unit), European Food Safety Agency (EFSA), Via
Carlo Magno 1/A, I-43126 Parma, Italy. E-mail: Jordi.Tarres-Call@efsa.europa.eu1
Alan R.Walker, Royal (Dick) School of Veterinary Studies, University of Edinburgh, Edinburgh,
EH25 9RG, Scotland, UK. E-mail: Alan.R.Walker@ed.ac.uk
The editor Jordi Tarrs-Call is employed by the European Food Safety Authority (EFSA). The positions
and opinions presented in this monograph are those of the authors and editors alone and are not intended to
represent the views or scientific works of EFSA.
Foreword
Jeremy Gray
Emeritus Professor of Animal Parasitology, University
College Dublin, Republic of Ireland
Ticks are the most important ectoparasites worldwide in veterinary science, both as parasites
in their own right and also as vectors of many different pathogens that cause diseases in livestock and companion animals such as anaplasmosis, babesiosis, cowdriosis, ehrlichiosis and
theileriosis. Although few tick species are adapted to parasitize mankind, they are also of
great significance in human medicine as vectors of zoonotic pathogens, notably in the temperate northern hemisphere, where Lyme borreliosis, caused by the spirochaete species complex,
Borrelia burgdorferi sensu lato, is now acknowledged as the most prevalent vector-borne disease. Despite long-standing knowledge of the subject, relatively few books have been devoted
to ticks and tick-borne diseases, with the two-volume 1992/3 publication Biology of Ticks by
D.E. Sonenshine constituting the most important recent contribution, and a new multi-author
version currently in preparation. Several other books such as A. Bowman and P. Nuttalls
Ticks: Biology, Disease and Control and Tick-borne Diseases of Humans by J.L. Goodman et al.
address current hot topics, and there are several published proceedings of conferences and,
reflecting growing public interest, a plethora of popular books. However, none of these publications are concerned with the very important subject of the geographical distribution of
tick-borne pathogens and their vectors. This topic has acquired special significance in light of
the predicted effects of climate and environmental change brought about by global warming
and other anthropogenic factors, especially in the context of remote sensing and risk mapping. For such information pertaining to Europe and the Mediterranean, tick biologists have
had to rely in the past on texts such as Ticks of North-west Europe by P.D. Hillyard and Ticks of
the Domestic Animals in the Mediterranean Region by A. Estrada-Pea et al., though both are
quite short publications and are primarily concerned with taxonomy and identification rather
than distribution. Not since the outdated and relatively limited monographs compiled by the
great Harry Hoogstraal and co-workers in the 1950s has such a comprehensive record of tick
distribution been published as is presented in this monograph, and to my knowledge no such
attempt to record the distribution of tick-borne diseases in these particular geographical
regions has been attempted before.
This monograph comes in two parts: first is the main text with descriptions and accompanying maps, applying to both the ticks and to the diseases they transmit, and second is an
appendix detailing the sources of the data and other reference material. The main text starts
with an introduction that focuses on the significance of tick-borne diseases, the geographic
scope of the work, the methods used for reviewing the literature and a brief account of tick
xi
xii
Jeremy Grey
identification methods. Information on the literature was extracted from PubMed and the ISI
Web of Knowledge using several databases including those of CABI, Current Contents, Food
Science and Technology Abstracts, Journal Citation Reports, MedLine and Web of Science. This
resulted in the citation of more than 1400 papers concerning the distribution of ticks and tickborne diseases, with many others addressing other topics, and the vast majority published in
English.
The first chapter consists of an account of the factors behind the emergence and re-emergence
of tick-borne diseases, highlighting the theme of environmental and climatic change and also
the renewed interest in ticks and the diseases they transmit, which has been stimulated by an
increased awareness of tick-borne zoonoses. Chapter 2 describes the basic biology of a total of
25 important tick species endemic to part or all of the geographical region under consideration,
and also includes short accounts of their life cycles, geographical distributions, and significance
as vectors. It contains original photographs of examples of the vast majority of the tick species
considered in the publication, some of which have evidently been collected from host animals.
In addition there are comments on some non-endemic species such as Amblyomma (Am.) variegatum and Am. hebraeum, which are of immense significance south of the Sahara and whose distribution may change as a result of accidental introduction, as in the Caribbean. In Chapter 3 the
factors responsible for the spread and distribution of ticks are considered, which include climate, land use, animal movement both wild and domestic, and the importation of exotic
vertebrates. Tick-borne infections are reviewed in Chapter 4, and within each section of this
chapter topics as varied as molecular taxonomy, pathology and epidemiology are covered.
Approximately 29 bacterial species, 12 protozoan and three viral species are considered at varying levels of detail depending on their significance, and these descriptions provide useful
thumbnail sketches of the species biology to put alongside their distributions. The geographical
distribution of tick-borne pathogens is the focus of Chapter 5, in the form of maps with accompanying qualifying and illustrative comments; Chapter 6 addresses the distributions of the vector ticks in the same way. In both cases, the longitude and latitude coordinates were utilized; if
these were not available, the nomenclature of territorial units for statistics (NUTS) was used for
EU countries, while for non-EU countries the name of the administrative area given in the publication was used. Chapter 7 addresses the surveillance and control of ticks and tick-borne
diseases. It includes a brief description of tick sampling methods, an introduction to the principles of surveillance and monitoring, and control options for both ixodids and argasids.
The text of the book is supported by an extremely extensive appendix. Nearly all of this
(as Appendix 1) consists of tables of geographic data for individual tick-borne diseases and
tick species that have been derived from the very numerous bibliographic records (Appendices
2 and 3; available at http://www.cabi.org/vetmedresource/) from which the distribution data
were extracted. Finally, Appendix 4 presents illustrated identification keys for the genera and
species of ticks in the Mediterranean basin. Most of the focus is on identification to genus level
but there are notes on selected species accompanying the illustrations. All the important tick
species that occur in northern Europe and that are mentioned in the main text are also included
here, with the exception of Ixodes persulcatus, which is primarily a tick of temperate Asia.
There is no question that this publication will prove to be extremely useful for all those
working with ticks and tick-borne diseases in Europe and around the shores of the Mediterranean.
It fills an obvious gap in the tick literature and will complement the more specialized content of
other recent publications, including the forthcoming new edition of Biology of Ticks. It will be a
useful addition to the library of every European university with a biology department, and it
should also find a place in all institutions in this geographical region that are concerned with
the surveillance and monitoring of vector-borne diseases, which are likely to become increasingly relevant to those concerned with the determination of public health policies and the
implementation of control measures.
Acknowledgements
The initial mandate for this work was conducted under the auspices of the Animal Health and
Animal Welfare Panel of European Food Safety Authority (EFSA). The editors and the
contributors of this monograph would like to express their appreciation for the full support of
EFSA for producing this monograph.
The authors acknowledge the valuable assistance of Drs Milen Georgiev and Sofie
Dhollander (Scientific Officers of the Animal Health and Animal Welfare Unit of EFSA); and
of Dr Nely Snchez (Veterinary Parasitologist of the Department of Animal Pathology
Parasitology of the University of Zaragoza) during the systematic literature review.
Thomas Jaensons contributions were related to his research on ticks and tick-borne infections which is funded by Carl Tryggers Stiftelse (Norwegian Foundation) and Magnus
Bergvalls Stiftelse.
The data set produced for this monograph was a joint effort made possible in part by the
International Consortium of Ticks and Tick-Borne Diseases, funded by the European Union
and led by Frans Jongejan (University of Utrecht, The Netherlands), and EFSA.
Parts of the data set were compiled with records curated in the collection of the Institute
of Parasitology of the Czech Republic, with the help of Frantsek Dusbbek and the support of
the European Concerted Action on Lyme Borreliosis.
We express our thanks to the people in these concerted actions who contributed with
expert assistance to the compilation: Danielle de Meneghi, Peter Zemman and Milan Daniel.
We also express our thanks to Ali Bouattour, Laurence Vial, Miguel A. Habela, Ramn Juste,
Ana Garca-Prez, Joaquim Castell, Margarida Santos Silva and Sofia Nncio for providing
advice and lists of records in the collections of ticks under their charge. Byron Papadopoulos
contributed with records in Macedonia. Lise Gern contributed with much help on data on tick
distribution in Switzerland. John Scharlemann provided a list with records of Ixodes ricinus in
the UK. Zati Vatansever compiled unreported reports of ticks in Turkey and produced the list
of species for the country as included in the final data set. Jean-Louis Camicas and Ali Bouattour
examined earlier maps of the species reported in the Mediterranean region. We want to extend
our gratitude to collectors and systematists who made the creation of this data set possible.
xiii
Abbreviations
A.: Anaplasma
AF: Astrakhan fever
Am.: Amblyomma
Ar.: Argas
ASF(V): African swine fever (virus)
B.: Borrelia or Babesia
Bo.: Boophilus
CCHF(V): Crimean-Congo haemorrhagic fever (virus)
D.: Dermacentor
DEBONEL: Dermacentor-borne necrosis erythema lymphadenopathy
DEET: diethyl toluamide
E.: Ehrlichia
EPNs: Entomopathogenic nematodes
H.: Hepatozoon
Ha.: Haemaphysalis
Hy.: Hyalomma
I.: Ixodes
ISF: Israeli spotted fever
ICTTD: Integrated Consortium on Ticks and Tick-borne Diseases (European project)
LAR: Lymphangitis-associated rickettsiosis
LI(V): Louping ill (virus)
MSF: Mediterranean spotted fever
NUTS: Nomenclature for territorial units for statistics
O.: Ornithodoros
R.: Rickettsia
RF: Russian Federation
Rh.: Rhipicephalus
T.: Theileria
TBD: Tick-borne disease
TBE(V): Tick-borne encephalitis (virus)
TCC: Transcaucasian Countries
TIBOLA: Tick-borne lymphadenopathy
xv
Introduction
Mo Salman1 and Jordi Tarrs-Call2

Colorado State University, USA; and 2European Food Safety
Agency (EFSA), Parma, Italy
Significance of Tick-borne Diseases

The aim of this monograph is to provide a general overview of the geographical distribution
of various tick species which have proven their involvement in the transmission of the pathogens causing animal diseases and zoonoses in Europe, the Middle East and the Mediterranean
basin. As blood-feeding parasites, ticks are able to transmit to their hosts a wide variety of
pathogens which may cause tick-borne infections and tick-borne diseases (TBDs) that affect
wild and domestic animals, including companion animals. The transmission of pathogens
among ticks may occur transovarially, i.e. the pathogen is transmitted via the eggs from females
to their offspring, trans-stadially from larva to nymph and/or from nymph to adult, and venereally during copulation from male to female tick. Vectorial competence is the overall ability of
a vector tick species to transmit a pathogen to a range of receptive vertebrate hosts in a given
location at a specific time. The TBDs usually are geographically distributed within the range of
their vectors. Some of these infections/diseases can give clinical signs which can be severe (in
the acute phase) but can also present as subclinical forms (mainly in endemic areas) in animals or
humans. Furthermore, co-infection with different pathogens can occur in the same vertebrate
animal when the same tick species transmits more than one pathogen (e.g. Hepatozoon canis,
Ehrlichia canis and Anaplasma platys transmitted by Rhipicephalus sanguineus), or when two or
more tick species infest an animal or human at the same time (Belongia, 2002; Stanczak et al.,
2002; Bremer et al., 2005; Halos et al., 2005; Swanson et al., 2006). TBD co-infections by ticks are
frequent in companion animals living in endemic areas and this may often impair an appropriate aetiological diagnosis (EFSA, 2007).
Several recent reports have indicated the wider spread of specific tick species as vectors in
geographic areas that were not considered previously infested with these tick species (e.g.
Jskelainen et al., 2006, for Ixodes persulcatus; Nijhof et al., 2007, for Dermacentor reticulatus; Gray
et al., 2009, Jaenson et al., 2012, for I. ricinus and D. reticulatus). Furthermore, other reports have
shown that the host range of some tick species was wider than previously known. There are
limited data on vector competence for many tick species and a lack of information on the effect
of environmental factors, including climatic factors, on the transmission of pathogens.
The taxonomy of tick-borne pathogens is also evolving and subject to change. Based on
recent molecular studies, the taxonomy of certain tick species has been adapted accordingly.
This monograph uses the taxonomy proposed by Uilenberg (2006).
CAB International 2013. Ticks and Tick-borne Diseases (eds M. Salman and J. Tarrs-Call)
xvii
xviii
Geographic Scope
The list of countries that are covered in this monograph includes the European countries
(Albania, Andorra, Austria, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Cyprus, the
Czech Republic, Denmark, Estonia, Faroe Islands, Finland (including land Islands), France,
Germany, Greece, Hungary, Ireland, Italy, Kosovo, Latvia, Liechtenstein, Lithuania,
Luxembourg, Macedonia, Malta, Montenegro, The Netherlands, Norway, Poland, Portugal,
Romania, San Marino, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, the UK). In
addition, the following countries are also included: Algeria, Egypt, Israel, Jordan, Lebanon,
Libya, Morocco, Syria, Tunisia and Turkey, and the geographic areas covering Palestine and
the Western Sahara. These neighbouring countries were included owing to their proximity to
Europe and the potential for the spread of ticks from them. The eastern border of Europe
(Armenia, Belarus, Georgia, Moldova and Ukraine, and a buffer zone of some 600 km in
European Russia) is also included to account for the important tick species recorded in that
area which are of potential interest for Europe under future scenarios.
Methods Used for the Systematic

Review of the Literature
A systematic literature review was carried out based on the general principles of the systematic
review methodology (CRD, 2009; Higgins and Green, 2009) with the aim of gathering information about the geographical distribution of tick species and tick-borne pathogens. The literature review was based on scientific papers published from January 2000 to March 2010 (some
of the most recent data published or generated after this monograph was initiated are not
present in the maps).These papers were retrieved from the citation indexing service of ISI Web
of KnowledgeSM (Thomson Reuters) and from PubMed (US National Library of Medicine).
The active databases during the search in the ISI Web of Knowledge were:
Web of Science
Current Contents Connect
CAB Direct
Food Science and Technology Abstracts
MEDLINE
Journal Citation Reports
Additional publications within which the topics were considered relevant to the review,
were submitted by the other contributors to this monograph. Two different sets of search
strings were used one for ticks and another for tick-borne pathogens. These strings were
applied to the title and the abstract (Table I.1) of the publications. No language restrictions
were set. The geographical limitation was applied as per the rustication to countries and
regions listed above. Published validated data from the European Union Project Integrated
Consortium on Ticks and Tick-borne Diseases (ICTTD-3), collected by one of the contributors
to this monograph, were also considered. All data derived from these sources were collated in
a relational database (using FileMaker software, see http://www.filemaker.com), from which
the maps were created.
Duplicate references were deleted automatically by means of a reference management
systemi. References were then checked manually and duplicates removed. The search in ISI
Web of Knowledge and in PubMed produced a list of 2197 references.
The title and abstract of each reference were screened following the criteria listed in Table I.2.
After the first screening, 1222 references were considered relevant, 309 doubtful, and 666
non-relevant. The doubtful references (title and abstract) were further revised by two contributors to this monograph who still found ten relevant references. By checking the full text,
Introduction
xix
Table I.1. The search strings for publications on ticks and tick-borne pathogens.
Search terms related either to ticks or
tick-borne pathogens
Tick
Argas OR Ornithodoros
OR Dermacentor OR
Haemaphysalis OR Hyalomma
OR Ixodes OR Rhipicephalus
OR Boophilus
Tick-borne African Swine Fever virus OR ASF

pathogen virus OR ASFV OR Anaplasma
OR A phagocytophilum OR
Ehrlichia phagocytophila OR
Babesia OR Crimean Congo
Haemorrhagic Fever virus
OR CCHF virus OR CCHFV
OR Hepatozoon OR Lyme
disease agent OR Borrelia OR
B burgdorferi OR B garinii OR
B spielmanii OR B lusitaniae
OR Rickettsia OR R conorii
OR Ehrlichia canis OR Borrelia
hispanica OR B hispanica OR
Theileria OR T parva OR T hirci
OR T ovis OR T lestoquardi OR
Tick borne encephalitis virus
OR Louping ill virus OR TBE
virus OR TBEV OR tick borne
avivirus OR TBEF OR TBEFV
OR TBE group OR Francisella
OR F tularensis OR Bartonella
OR Q fever OR Coxiella OR
African horse sickness virus OR
AHSV OR tick borne orbivirus
AND
AND (search terms related to the

geographic region)
Aland OR Albania OR Andorra OR

Austria OR Belgium OR Bosnia
and Herzegovina OR Bulgaria
OR Croatia OR Cyprus OR
Czech Republic OR Denmark OR
Germany OR Spain OR Estonia
Distribution OR
OR Faroe Islands OR Finland OR
presence OR
France OR Greece OR Hungary
occurrence OR
OR Ireland OR Italy OR Kosovo
reported
OR Latvia OR Liechtenstein
OR Lithuania OR Luxembourg
OR Macedonia OR Malta OR
Montenegro OR The Netherlands
OR Norway OR Poland OR
Portugal OR Romania OR San
Marino OR Serbia OR Slovakia
OR Slovenia OR Switzerland OR
Sweden OR United Kingdom OR
Algeria OR Egypt OR Israel OR
Jordan OR Lebanon OR Libya OR
Morocco OR Syria OR Tunisia OR
Turkey OR Palestine OR Western
Sahara OR Armenia OR Belarus
OR Georgia OR Moldova OR
Ukraine OR Russia OR USSR
Table I.2. Criteria used in the rst screening for relevance of the selected references.
Criterion
Included
Excluded
Concerns occurrence of a tick species or a tick-borne

pathogen in the area considered
Concerns a tick species with proven involvement in
transmitting pathogens of animal diseases or zoonoses
Contains geographic information on the distribution
of the tick species or the tick-borne pathogen
Yes
No
Yes
No
Yes
No
43 other initially doubtful references were considered as relevant. This produced a total of 1275
relevant references and 822 non-relevant references. The full articles of the relevant references
were then retrieved, but some could not be found. There were 125 scientific articles (six in
English language and 119 in other languages) that were missed.
The second screening of articles was performed in parallel with the data extraction
by two experienced veterinary parasitologists of the Veterinary School of the University
of Zaragoza (Spain), one of whom is a contributor to this monograph. All contributors were
xx
consulted when doubts arose. Apart from the three criteria considered in the first screening, the contributors were asked to also consider the secondary criteria listed in Table
I.3.
This second screening resulted in a total of 637 scientific papers considered appropriate to
be included in the review. These are listed in Appendix 2, sorted by the identification number
of the reference.
Data were extracted to an Excel spreadsheet containing the following fields:
Tick genus and species, recorded as in the original paper.

Location of the tick species: based on the nomenclature of territorial units for statistics
(NUTS) for the EU (European Union) countries. For non-EU countries, locations were
recorded by the name provided in the original report, at the equivalent level of geographical precision. The coordinates (latitude/longitude) were recorded if given in the
article.
Whether molecular techniques were used to identify the tick species (yes/no).
The source of the tick specimen: free living (questing), livestock, pet, human, wildlife (taxonomic order of the host).
The corresponding bibliographic reference.
The tick-borne pathogen genus and species (as it appeared in the original work).
Location of the pathogen: NUTS for European countries. For the countries that do not
have the NUTS, the name of the administrative region at the level of precision provided in
the scientific paper. Geographical coordinates if provided.
Diagnostic/identification method of the pathogen: isolation, molecular, serology.
Table I.3. Criteria used in the second screening for relevance.

Criterion
Language publication
Included
Excluded
English abstract and text Abstract not available

available in English,
or text not in English,
French, German, Italian, French, German,
Spanish, Hungarian,
Italian, Spanish,
Bulgarian or Dutch
Hungarian, Bulgarian
or Dutch
Original work (not a review document)
Yes
No
Contains geographic information on the distribution Yes
No
of the tick species or the tick-borne pathogen
Does not specify a concrete geographic location.
No
Yes
Rather refers it to the entire country
Concerns a prompt importation of a tick species that No
Yes
does not reach sufcient epidemiological threshold
for its establishment in the specied location
Case reports of human infections that, in order to
No
Yes
protect the privacy of personal data, provide the
address of a reference hospital instead of the
residence of the infected patient
Tick identication is unequivocal and appropriate
Yes
No
The diagnostic method for the tick-borne pathogen Yes
No
is appropriate (serology, isolation, biological
methods such as PCR)
Introduction
xxi
Source of the sample (pathogen): livestock, pet, human, wildlife (taxonomic order of the
host), tick.
The corresponding bibliographic reference.
A more detailed description of the systematic literature review process is given in

EFSA (2010).
Identification Methods for Tick Species, Including the Most

Recent Advanced Techniques and their Limitations
Identification of ticks can be based on different criteria. Morphology can be the only criterion
in some occasions but, in a more holistic approach, identification may be based on host species;
predilection sites, geographical occurrence, seasonality, and other criteria. Morphological
identification is based on: presence/absence of eyes; the presence of anal plates; ornamentation on the legs and scutum; shape, size and the patterns of the scutum; and other unique
characteristics. Figure I.1 describes these morphological terms, and Tables I.4 and I.5 show
how these morphological characteristics are present in hard and soft ticks, respectively.
The taxonomy of ticks is not exempt from complications. There are problems of morphological identification and sympatric tick speciation. Identification of tick species requires extensive experience with the subject over many species. Molecular biology has reshuffled many
names at both species and genus level, although it must be stressed that the conventional
identification was in many cases able to identify the different tick entities. For instance, in the
case of the changes of Boophilus microplus to Rhipicephalus (Rh.) microplus, and of Hyalomma
(Hy.) marginatum marginatum to Hy. marginatum, it was only the name that changed, not the
species identification. In other situations, the identification changed owing to tick physical
appearance and not because of molecular characteristics, such as the change of Haemaphysalis
(Ha.) leachi to Ha. elliptica in South Africa.
The colour plate section contains photographs of some hard and soft tick species involved
in the transmission of animal diseases and zoonoses, and Appendix 4 contains identification
keys for the morphological identification of hard and soft ticks in the Mediterranean basin.
Limitations
This monograph addresses data on the distribution of ticks and TBDs in Europe and the
Mediterranean basin that were retrieved from existing literature and technical reports. The
reports findings, however, are limited as a result of the following issues:
The presence of pathogens was based on antigen detection assays without consideration
of serological evidence.
Cases were only reported from the literature without consideration of other sources that
may be available through other means of searching.
Literature was retrieved mainly in the English language, with only limited inclusion of
other languages.
Although intended to be comprehensive in the literature search strings, the search may
have missed some publications.
The maps have not been updated since March 2010, so some of the most recent data published or generated after the completion of this monograph are not included.
xxii
External structure of adult argasid ticks (the example is Ornithodoros)
claws
mouthparts
four pairs of legs

genital
aperture
mammillated texture
spiracle
anus
DORSAL VIEW
VENTRAL VIEW
External structure of adult ixodid ticks (the example is Hyalomma)
four pairs
of legs
conscutum
palp
scutum
basis capituli
alloscutum
festoons
MALE, DORSAL VIEW
FEMALE, DORSAL VIEW
pulvillus
hypostome
claw
coxa I
genital aperture
genital aperture
coxa IV
spiracle
anal groove
anus
ventral plates
FEMALE, VENTRAL VIEW
MALE, VENTRAL VIEW
Fig. I.1. Main anatomical characteristics and terms used to describe the morphology of adult hard ticks,
from Walker et al., 2003.
Note
i
EndNote X1.0.1 [Bld 2682], Thomson. http://www.endnote.com
Table I.4. Generic distinguishing morphological features of different genera of hard ticks.
Feature
Hyalomma
Size (unfed Large

adults,
(0.5
total
0.8 cm)
length)
Mouthparts Long
Basis
capituli
Ornate
Caudal
process
Dermacentor Haemaphysalis Amblyomma
Aponomma *
Rhipicentor * (Amblyomma) Margaropus*
Medium
Small
Small
Medium-large Small
Very large
Medium
(0.30.6 cm) (0.10.3 cm) (0.20.3 cm) (0.40.9 cm) (0.10.3 cm)
(0.51.0 cm) (0.3
0.5 cm)
Large
(0.5
0.6 cm)
Small
(0.1
0.3 cm)
Short to
medium
Hexagonal
Long
Very short
Very short
Long
Medium
Short
Very long
Medium
Hexagonal
Rectangular
Rectangular
Yes
No
Yes
No
Some species No
Yes
Yes
No
No
Yes
No
Yes
No
Yes
No
No
No
Posterior,
chaliceshaped
No
Absent
No
Yes
Yes
Yes
No, except
for 4
species
Yes
Yes
Yes
With ventral
auriculae
No
Yes
No
Yes
No
No
No
Yes
Yes
No
No
Yes
No
Yes
No
No
No
No
No
Yes
Yes
None, or
very small
No
Posterior,
chaliceshaped
No
Posterior,
chaliceshaped
Present
in some
species
Bid
Posterior,
Anterior,
Posterior,
vertical line
semicircular
chaliceor U-shaped shaped
Yes, on some No
No
Posterior,
chaliceshaped
No
Posterior,
Posterior,
semicircular chaliceshaped
No
No
No
Coxae I
Coxae IV
Bid
Legs
Banded
Hexagonal
Large, bid
Very large
Grouped
anteriorly
Banded
Yes, pilose
posterior
margin
Large, bid
Very large,
with spurs
Banded
Banded,
enlarged
segments
in males
xxiii
(*) These genera are currently not present in European member states, but they may be imported in the future.
Introduction
Eyes
Festoons
Adanal
plates
Sub-anal
plates
Anal
groove
Rhipicephalus
Rhipicephalus (Boophilus)
Ixodes
xxiv
Table I.5. Generic distinguishing morphological features of different genera of soft ticks.
Feature
Argas
Ornithodoros
Body
Presence of suture (differentiated tegumental
tissue between dorsal and ventral surface)
Presence of cuticle extensions rounded hook-like
cones shaped
Eyes
Tegument
Discs of tegument
Flattened
No
Thick
Yes
No
Yes
No
Granular
Usually in radial rows
Yes
Squamous
Not in radial rows
References
Belongia E.A. (2002) Epidemiology and impact of coinfections acquired from Ixodes ticks. Vector-Borne and
Zoonotic Diseases 2, 265273.
Bremer, W.G., Schaefer, J.J., Wagner, E.R., Ewing, S.A., Rikihisa, Y., Needham, G.R., Jittapalapong, S., Moore,
D.L. and Stich R.W. (2005) Transstadial and intrastadial experimental transmission of Ehrlichia canis
by male Rhipicephalus sanguineus. Veterinary Parasitology 131, 95105.
CRD (2009) Systematic Reviews, CRDs Guidance for Undertaking Reviews in Health Care. January 2009,
Centre for Reviews and Dissemination, University of York, UK.
EFSA (European Food Safety Authority) (2007) Opinion of the Scientific Panel on AHAW on a request from the
European Commission on the risk of tick introduction into the UK, Ireland, and Malta as a consequence
of abandoning the national rule. EFSA Journal 469, 1102.
EFSA Panel on Animal Health and Welfare (AHAW) (2010) Scientific opinion on the role of tick vectors in
the epidemiology of Crimean Congo hemorrhagic fever and African swine fever in Eurasia. EFSA
Journal 8(8): 1703.
Gray, J.S., Dautel, H., Estrada-Pea, A., Kahl, A. and Lindgren, E. (2009) Effects of climate change on ticks and
tick-borne diseases in Europe. Interdisciplinary Perspectives on Infectious Diseases 2009, 112, Article
ID 593232.
Halos, L., Jamal, T., Maillard, R., Beugnet, F., Menach, A.L., Boulouis, H.J. and Vayssier-Taussat, M.
(2005) Evidence of Bartonella sp. in questing adult and nymphal Ixodes ricinus ticks from France and
co-infection with Borrelia burgdorferi sensu lato and Babesia sp. Veterinary Research 36, 7987.
Higgins, J.P.T. and Green, S. (eds) (2009) Cochrane Handbook for Systematic Reviews of Interventions Version
5.0.2 (updated September 2009). The Cochrane Collaboration, Oxford, UK.
Jskelinen, A.E., Tikkakoski, T., Uzctegui, N.Y., Alekseev, A.N., Vaheri, A. and Vapalahti, O. (2006) Siberian
subtype tickborne encephalitis virus, Finland. Emerging Infectious Diseases 12, 10. Available at: http://
dx.doi.org/10.3201/eid1210.060320 (accessed 22 June 2012).
Jaenson, T.G.T., Jaenson, D.G.E., Eisen, L., Petersson, E. and Lindgren, E. (2012) Changes in the geographical
distribution and abundance of the tick Ixodes ricinus during the past 30 years in Sweden. Parasites
and Vectors 5, 8, doi:10.1186/1756-3305-5-8. Available at: http://www.parasitesandvectors.com/
content/5/1/8 (accessed 22 June 2012).
Nijhof, A.M., Bodaan, C., Postigo, M., Nieuwenhuijs, H., Opsteegh, M., Franssen, L., Jebbink, F. and Jongejan,
F. (2007) Ticks and associated pathogens collected from domestic animals in the Netherlands. Vectorborne and Zoonotic Diseases 7, 585595.
Stanczak, J., Racewicz, M., Kruminis-Lozowska, W. and Kubica-Biernat, B. (2002) Coinfection of Ixodes
ricinus (Acari: Ixodidae) in northern Poland with the agents of Lyme borreliosis (LB) and human granulocytic ehrlichiosis (HGE). International Journal of Medical Microbiology 291(Supplement 33), 198201.
Swanson S.J., Neitzel, D., Reed K.D. and Belongia, E.A. (2006) Coinfections acquired from Ixodes ticks.
Clinical Microbiological Reviews 19, 708727.
Uilenberg, G. (2006). Babesia a historical overview. Veterinary Parasitology 138, 310.
Walker, A.R., Bouattour, A., Camicas, J.-L., Estrada Pea, A., Horak, I.G., Latif, A.A., Pegram, R.G. and Preston,
P.M. (2003) Ticks of Domestic Animals in Africa: A Guide to Identification of Species. Bioscience Reports,
The University of Edinburgh, Edinburgh, UK.
Emerging (Re-emerging) Tick-borne

Infections and the Dissemination of Ticks
Mo Salman1 and Agustn Estrada-Pea2

Colorado State University, USA; and 2University of Zaragoza, Spain
Ticks are arthropods and therefore they are

susceptible to changes in climate. All stages
of the life cycle of ticks are associated with
vegetation, so small changes in the prevailing
environment may have important consequences for the overall impact of ticks and
tick-borne infections in surrounding areas.
Nevertheless, this issue is not as simple as it
appears. On completion of their life cycles,
ticks are also sensitive to the availability of
hosts and to several other factors related to
the surrounding landscape. For instance, the
fragmentation of a forest into smaller patches
of vegetation surrounded by a matrix of nonhabitat landscape may deeply affect the
behaviour of the hosts, and this might have
profound consequences for the life cycle of
the ticks.
The different life stages of the tick are
sensitive to temperature, i.e. they occur faster
if a temperature near the optimum for that
stage is available in the very few centimetres
above the ground. While a higher temperature will result in a faster completion of a
given stage, it will probably also generate a
higher level of evaporation. This effect is
hazardous for tick survival, as ticks are sensitive to desiccation. A higher temperature is,
therefore, not always better for the overall
performance of the tick population. The effect
of temperature and desiccation are dependent upon the preferences of the species. For
example, limited variations in temperature

may have serious consequences for populations of ticks that colonize cold and wet
environments in the forests of northern
Europe, but are probably not so disastrous for
species living in Mediterranean conditions,
for which a high seasonal variability of climatic characteristics is the norm. Climatic
and other factors also have an interactive
effect on the survivability and spread of ticks
and their infections in host populations.
It is now evident that some tick species
are spreading in parts of the world that were
not previously considered to be favourable
areas for ticks. The reason for this spread is
not only due to the effects of climate change,
but also to subtle and continuous trends of
changes in the vegetation, landscape features
and social habits. These changes are leading
to new areas of contact between ticks, their
pathogens and the interface between animals
and humans. No unique pattern can explain
the spread of the ticks and of the pathogens
that they transmit, although in some parts of
their range, climate may account for their
changes in distribution. For example, Ixodes
ricinus has spread in parts of Scandinavia as
well as in the mountain ranges of the Czech
Republic (Daniel et al., 2003). These changes
have been attributed to climate because
permanent populations of I. ricinus are now
detected in northern areas of Sweden
(Talleklint and Jaenson, 1998) and colonize a

higher altitude range in the mountains.
The impact of this wide spread of I. ricinus
is related not only to parasitism by this tick
species of both humans and animals, but also
to its associated pathogens and to infection of
its hosts by these pathogens. I. ricinus is the
main vector in the Palaearctic region for the
prominent pathogen Borrelia burgdorferi sensu
lato (s.l.), the aetiological agent of Lyme disease. While there are no adequate figures
available on the importance of this disease in
the countries of the European Union (as it is
not a compulsorily reportable disease) it is
estimated to be the most widespread infection transmitted by ticks in Europe (Rizzoli
et al., 2011). A recent paper suggested that this
pathogen is present everywhere that the tick
vector is, and that the prevalence rates of
B. burgdorferi s.l. infection are directly dependent upon a combination of features of climate
and vegetation (Estrada-Pea et al., 2011).
Other studies have confirmed the probable
northward spread of the tick vector and of
the risk of contracting Lyme disease in
Scandinavia (Jaenson and Lindgren, 2011),
and that the northern limit of I. ricinus may be
predicted by some features of vegetation,
which are obviously dependent upon a combination of climate factors. Lyme disease is
also dependent upon a combination of reservoirs, such as rodents, birds and lizards,
which are necessary for the maintenance of
the pathogen population.
I. ricinus is also the only known vector in
Europe for tick-borne encephalitis virus
(TBEV), which causes tick-borne encephalitis
(TBE) in humans. Although the ecological
determinants of the spread of TBEV have
been ignored, the map of reported cases of the
disease has changed over the last few years.
However, while there is a direct relationship
between the presence of I. ricinus and cases of
Lyme disease (or the presence of the pathogen in the tick vector), the incidence of TBE is
not directly related to the abundance of tick
vectors. Other authors have speculated on
different theories to explain the presence of
a given disease reservoir as the shape of
the area where the infection is present or have
evaluated the importance of a given climate
pattern and its impact on the life cycle of ticks
(Rogers and Randolph, 2006). It is, however,

necessary to further evaluate how these factors may evolve, and how the areas where the
infection is predicted to exist associate with
the actual distribution of reported cases of the
disease (or the records of the pathogen).
A process-driven model for the life cycle of
the tick I. ricinus would also be of great importance in providing an additional tool to
understand the patterns of disease.
In the last few years, infection by the
Crimean-Congo haemorrhagic fever (CCHF)
virus (CCHFV) has gained much attention as
a result of several serious outbreaks. CCHF is
an important disease that is prevalent over
wide areas of Africa, Asia, the Near East and
parts of Europe (Ergnl, 2006). It has not
been reported in the Americas and Australia
because of the absence of the main vectors.
The pathogen is mainly transmitted by the
bite of ticks of the genus Hyalomma (and especially by Hyalomma marginatum), which are
most prevalent in a Mediterranean-type environment, as well as in the African steppes.
The disease was first diagnosed around the
middle 1940s in the Crimea and then in the
Congo. CCHF has not yet been diagnosed in
sites or countries outside its main distribution
area for 60 years. In the last few years though,
this disease has re-emerged in Turkey, with
more than 3000 cases, and the outbreak seems
to be independent of climatic causes. Studies
have hypothesized that the re-emergence of
the disease is based on a series of factors that
interact, mainly an abundance of ticks created
by social influences and an increased availability of hosts (Ergnl, 2006).
As for other tick-borne diseases, climate
trends have been commonly linked to clinical
cases of CCHF. Ticks are highly sensitive to
small changes in climate, and long-term, sustained and small differences in key climatic
factors may drive a serious change. A framework linking the response of the tick to subtle
climatic variations is, however, needed to
understand the actual significant factors.
An assessment of the effects of climate on the
presence of human CCHF clinical cases in
Turkey was carried out by Estrada-Pea et al.
(2009). The analysis included monthly data
for several climate factors. The study concluded that climate was not different between
Emerging Tick-borne Infections and Dissemination
sites with active foci of the infection and sites

where Hyalomma marginatum is common but
human cases are not reported. The study also
concluded that it is not possible to predict,
based solely on climate factors, where new
cases could appear in the reasonably near
future. Climate is not the sole factor driving
the virus amplification, even if climate in the
autumn/winter may have a strong regulating
role on the survival of tick populations
(Hoogstraal, 1979). It is noteworthy that
Hyalomma endemic areas with the mildest
autumns and winters in the Mediterranean
basin have historically been, and are currently, free of the infection, so the impact of a
warmer climate seems to be not the only factor determining the occurrence of the disease.
Efforts to link occurrence of the disease solely
to an array of climate conditions in sites where
Hyalomma ticks are absent (Randolph and
Rogers, 2007) may be unrealistic. Claims,
therefore, that a warmer climate influences
the presence or absence of the infection
(Purnak et al., 2007) are unsupported when
climate series are examined and compared.
Bold statements such as the probability of
hospitable climates not only extended to
Spain, Italy, and northern African countries
across the Mediterranean Sea, but even in
large parts of North America, smaller parts of
South America, and Australia (Maltezou and
Pappa, 2010) are unreliable without further
knowledge of the system. Even though climatic characteristics may be consistent with
sites where the disease has been reported,
adequate vectors for CCHFV are known to be
absent in the New World and Australia. The
role of the tick vectors needs therefore to be
carefully evaluated under both laboratory
and field conditions, and the lack of concordance between the groups of data clarified.
There is, however, evidence that a fragmented landscape, with many small patches
existing within a matrix of unsuitable tick
habitat, may lead to isolated populations of
both ticks and hosts, producing an amplification cycle whereby ticks feed on infected
hosts (Estrada-Pea et al., 2011). For CCHFV
eco-epidemiology, the degree of habitat
patchiness contributes to the increased contact rate among reservoir hosts, humans and
ticks. It also leads to the occurrence of isolated
amplification foci, with consequently higher

tick exposure to humans (Estrada-Pea et al.,
2009). Isolated fragments surrounded by a
matrix of agricultural land commonly have
the poorest diversity of mammals. Species
composition changes within these fragmented
and smaller isolated patches, and each species differs in its ability to cross the matrix
between fragments (e.g. Pires et al., 2002;
Lira et al., 2007; Forero-Medina and Vieira,
2009). Although Hyalomma ticks can move
horizontally, they require a vertebrate host to
move over significant distances. Because
these host populations are relatively isolated,
there are few local movements of hosts and
therefore new naive animals carrying uninfected ticks do not dilute the prevalence rates
in the isolated patch. However, the immune
response of such isolated populations against
the ticks and the pathogen may seriously
decrease the population of infected ticks.
These aspects of the disease have never been
studied for CCHF.
It is unlikely that the recent epidemic of
CCHF in Turkey is caused by a spreading
wave of infected ticks through the movement
of hosts, i.e. ticks infected with an imported
virus strain (or by a new, more infective
genetic variant of the virus) spreading from
central Anatolia to other regions of Turkey, as
was hypothesized by Karti et al. (2004).
Research on the epidemiology of the CCHF
infection in Turkey has focused on the region
of Anatolia in north-central Turkey, due to the
high incidence of the disease in this area. The
first clinical case of CCHF in the Thrace region
(in north-western Turkey) was noticed in 2007
(Midilli et al., 2009). Findings from a serosurvey on 193 human samples from three
localities showed that anti-CCHFV IgM and
IgG positivity rates were compatible with an
epidemiological situation in which the virus
has been present for a long time and is not the
result of a recent invasive event from the
main epidemic centre in Anatolia (Gargili
et al., 2011). The detection of CCHFV IgM
positivity and an elevated proportion of IgG
seropositivity indicated that the genetic variants Europe 1 (the clade in which viral strains
from Anatolia are included) and Europe 2
are very active in the region, and that high
IgG seroprevalence in humans indicates an
active immune protection (Gargili et al.,

2011). These figures support the idea that the
infection had been present in Turkey for a
long time before the epidemic reports.
Conditions have existed for continued cycles
of amplification, with the epidemic threshold only recently being reached. However,
these conditions are not the final step in a
chain of recent climate trends. Such an
epidemic threshold, whatever the factors
behind it, has been observed simultaneously
in distant regions of Turkey in only 24 years,
where it is not the result of a moving wave of
infected ticks feeding on hosts either. While
landscape patchiness, and thus isolated amplification systems, seems to be behind the proximal causes of the epidemic, the factors driving
the enzootic amplification cycle in Turkey

remain unknown. Land use practices are also
important; the agriculture in CCHF areas is
based on crop rotation and land fallowing,
and in a given village land is used for wheat
production during half of the year, while for
the remaining months it is used either for
chickpea/vetch production or is left fallow.
The following year the land use is rotated.
All this discussion highlights the variable
impact of different biotic and abiotic factors
in the delineation of the foci of pathogens
transmitted by ticks. The shifting nature of
these foci is far from being fully recognized,
and efforts are actively being made to evaluate the main forces that make up such an
intrinsically complex epidemiology.
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Emerging Tick-borne Infections and Dissemination
Purnak T., Selvi, N.A. and Altundag, K. (2007) Global warming may increase the incidence and geographic
range of Crimean-Congo hemorrhagic fever. Medical Hypotheses 68, 924925.
Randolph, S.E. and Rogers, D.J. (2007) Ecology of tick-borne disease and the role of climate. In: Ergonul, O.
and Whitehouse, C.A. (eds) Crimean-Congo Hemorrhagic Fever. A Global Perspective. Springer,
Dordrecht, The Netherlands, pp. 313.
Rizzoli, A., Hauffe, H.C., Carpi, G., Vourch, G.I., Neteler, M. and Ros, R. (2011) Lyme borreliosis in Europe.
Eurosurveillance 16(27), pii=19906.
Rogers, D. and Randolph, S.E. (2006) Climate change and vector-borne diseases. Advances in Parasitology
62, 345381.
Talleklint, L. and Jaenson, T.G.T. (1998) Increasing geographical distribution and density of Ixodes ricinus
(Acari: Ixodidae) in Central and Northern Sweden. Journal of Medical Entomology 35, 521526.
Basic Biology and Geographical

Distribution of Tick Species Involved
in the Transmission of Animal
Pathogens, Including Zoonoses*
Rbert Farkas,1 Agustn Estrada-Pea,2 Thomas G.T. Jaenson,3
Ilaria Pascucci4 and Maxime Madder5
1
Szent Istvn University, Budapest, Hungary; 2University of Zaragoza,
Spain; 3Uppsala University, Sweden; 4Istituto Zooprofilattico Sperimentale
dellAbruzzo e del Molise G. Caporale, Teramo, Italy; and 5Institute of Tropical
Medicine, Antwerp, Belgium and University of Pretoria, South Africa
The class Arachnida falls within the phylum

Arthropoda (arthropods) and encompasses
the following taxa: Acari (ticks and mites),
Araneae (spiders), Opiliones (harvestmen
or daddy-long-legs), Palpigradi (micro whip
scorpions), Amblypygi (tailless whip scorpions), Thelyphonida (whiptailed scorpions or uropygids), Ricinulei (ricinuleids),
Scorpionida (scorpions), Pseudoscorpiones
(book scorpions) and Solifugae (camel
spiders, sun spiders or solpugids). The
Ixodida (ticks) is one of several taxa that
constitute the subclass Acari (mites). Hence,
ticks are mites but not all mites are ticks
(Woolley, 1988).
There are three families of ticks comprising a total of at least 907 species of ticks:
(i) Nutalliellidae, with a single species,
Nutalliella namaqua; (ii) Argasidae, soft-bodied
or soft ticks, with 186 species (Section 2.1
below); and (iii) Ixodidae, hard-bodied or
hard ticks, with 720 species considered to
have valid genus and species names (Section

2.2 below) (Barker and Murrell, 2008).
Ticks can be easily distinguished from
adult insects because the ticks body is not
divided into three sections; rather there is strong
fusion of the body segments giving the tick
body a sac-like appearance. In common with
most arachnids, the adult tick and the nymph
have eight legs, whereas the larva has six legs
like adult insects. The tick has no distinct head,
but the mouthparts and the basis capituli form
the capitulum. All nymphal and adult ticks
have a pair of spiracles located latero-ventrally
on the abdomen. Depending on the species, a
pair of simple eyes may be present.
2.1
Soft Ticks (Argasidae)
Argasid ticks, known as soft ticks, include

about 180 species of four genera: Argas,
Ornithodoros, Otobius and Carios (Klompen
* Rbert Farkas wrote Sections 2.1 (Introduction), 2.1.1, 2.2.2, 2.2.6; Agustn Estrada-Pea, Sections 2.1.2,
2.2.5; Thomas G.T. Jaenson, introduction of Chapter 2 Sections 2.2 (Introduction), 2.2.1; Ilaria Pascucci, Sections
2.2.3, 2.2.4; and Maxime Madder supplied the colour plates of different tick species that are cited in the text.
Biology and Distribution of Tick Vectors
and Olivier, 1993). The Argasidae attain their

greatest abundance in dry regions, and when
they extend into humid areas they tend to
select drier niches where hard ticks would
not survive. This behaviour is related to their
leathery cuticle, which is effective in preventing water loss. A variety of argasids
occur throughout most of the tropics and
subtropics of the world. Fewer species live in
temperate areas and very few inhabit arctic
climates. Soft ticks often live for many years.
In contrast to hard ticks, they live in
close proximity to their hosts: in the nests of
birds, pigeon lofts, animal burrows or dens,
and caves.
The body of an argasid tick has an
unsclerotized, leathery cuticle with a textured surface that may be marked with
grooves or folds. The many small integumental folds usually have a button-like
appearance. The ticks have a flattened body
margin. In nymphs and adults, the gnathosoma (mouth and feeding parts) is not visible from the dorsal view, being located
ventrally. There is little sexual dimorphism.
The life cycles vary greatly between the
Ixodidae and the Argasidae. In the argasids,
mating usually occurs off the host. Females
take repeated small blood meals and lay
small batches of eggs (50500) in places
where they seek shelter (e.g. cracks and
crevices) after each feeding. As many as six
gonotrophic cycles have been reported in
some species. The six-legged larvae hatch in
14 weeks and, after seeking and attaching
to a host, feed for 38 days. They then drop
off and moult to the first nymphal stage.
Most argasids have two or more (six or even
seven in some species) nymphal stages in
their life cycle, each of which must have a
blood meal once before moulting to the next
stage. These nymphal stages occur off the
host in cracks or crevices, or beneath debris
in or near the nest and burrows. In contrast
to the slow-feeding hard ticks, the feeding of
nymphs and adults usually lasts 1 hour or
less. They are also highly resistant to starvation, which can extend their longevity even
further (Hoogstraal, 1985).
Argasid ticks have a considerable economic and medical impact in many parts of
the world. However, at the present time they
are apparently of less importance in Europe

than are the ixodid ticks. It should be noted
that argasid ticks in general are xerophilic
arthropods. Although in localities of
extremely low relative humidity argasids
may seek a somewhat more humid microhabitat, the cracks and crevices that they use
seldom have a significantly high relative
humidity; within this range, individual species have varying degrees of tolerance to
humidity. The soft ticks are vectors of a large
number of poorly known viruses (Hoogstraal,
1985) which seem of little economic interest
for domestic animals. A few studies exist
about those pathogens. However, perhaps
the most prominent pathogen transmitted
by a soft tick in the Mediterranean basin is
the African swine fever (ASF) virus (ASFV).
This virus has produced several epidemics
in Spain and Portugal, and is transmitted by
Ornithodoros erraticus (EFSA Panel on Health
and Welfare, 2010a).
2.1.1
Genus Argas
Argas species are about 510 mm long and

36 mm wide. In general, they are oval shaped
and dorsoventrally flattened. The cuticle is
leathery and wrinkled with definite margins
that can be seen even when the tick is
engorged. The unfed adults are yellowish,
but after a blood meal they appear slate grey.
Eyes, when present, occur on folds lateral to
the coxae. The mouthparts are on the ventral
surface of the body and small. The spiracular
plates, located between the coxae of legs III
and IV, are relatively small and inconspicuous. Members of this genus are most commonly associated with birds and bats,
although some species are found on reptiles
and, to a lesser extent, on insectivorous mammals other than bats. These soft ticks are nocturnal and endophilic, and they mate in the
environment. The females lay batches of
20100 eggs in hidden places after each blood
meal. The eggs hatch in about 23 weeks.
Argas larvae attach to hosts and feed for about
a week; they then drop to the ground to moult
into a nymph. There are usually two nymphal
stages. Like adults, each eight-legged nymph
Rbert Farkas et al.
instar is nocturnal; they feed at night and

moult in cracks and crevices to the next nymphal stage and, finally, to the adult male or
female. The life cycle from eggs to adults
takes about a month or longer. The adults can
live up to 2 years or more without a blood
meal. The larvae and nymphs may survive
for a few months. The larvae and first
nymphal stage are most active in early summer, the other nymphal stages in midsummer, and the adults in late summer and
autumn. They overwinter in the adult stage
or as eggs. Three species of Argas have been
commonly reported in Europe, namely Argas
(Ar.) persicus, Ar. reflexus and Ar. polonicus.
Argas persicus
The fowl or chicken tick, Ar. persicus, has
spread widely in most parts of the world.
These ticks construct large, numerous, and
fairly permanent nests in the vicinity of
human activity. All stages congregate on
walls, in crevices or between the boards of
poultry houses. Populations of the species
can reach enormous numbers in poultry
houses, causing irritation, loss of egg production, even high mortality because of anaemia.
It is reported that larvae of Ar. persicus can
cause fatal paralysis in young chickens.
Occasionally, they parasitize small mammals
and reptiles, but seldom humans. Adults feed
once per month for about 2 h, but can survive
for up to 45 years unfed. Ar. persicus may be
the vector of the rickettsial agent Aegyptianella
pullorum, and it is a vector of Borrelia anserina,
the agent of fowl spirochaetosis in the
Mediterranean region. The colour plate section contains two images (dorsal and ventral)
of an adult male of Ar. persicus (Plate 1a,b).
Argas reflexus
Ar. reflexus, the European pigeon tick, originates from the Mediterranean/Near East and
can be distinguished from Ar. persicus by its
body margin. The host of Ar. reflexus is the
wild rock pigeon, Columba livia, and its various domesticated forms, as well as feral town
pigeons, C. livia var. domestica (Hoogstraal,
1956). Ar. reflexus lives in several European
countries up to about 55N (Dautel et al., 1991).
In central Europe, Ar. reflexus occurs only in

or at buildings serving as pigeon breeding
sites where there are microclimatically elevated temperatures, especially in summer
and autumn but not necessarily in winter.
Such conditions are necessary for the completion of the embryonic development. The
life cycle appears to be much like that of Ar.
persicus. All nymphal stages as well as the
adults are nocturnal feeders taking blood
from sleeping pigeons. Feeding is for up to
12 h and, therefore, post-larval stages might
not have the opportunity to be transported
to distant habitats. Heavy infestations of
pigeons may cause death as a result of severe
anaemia.
Ar. reflexus has a relatively long minimum generation time of 24 years (Dautel
and Knlle, 1997). Hiding places of these soft
ticks are easily found in the cracks and crevices of pigeon cotes. German scientists (Dautel
et al., 1994) investigated the seasonal activities of this soft tick species in Berlin. They
reported that activities of all post-embryonic
stages peaked from March through to early
June, irrespective of whether or not hosts
were available to the ticks. The immature
stages displayed another peak of activity in
late summer/autumn. The only survey thus
far for pigeon ticks was also performed in the
city of Berlin, where more than 200 infested
buildings were discovered between 1989 and
1998. Most of the infestations were found in
older buildings constructed before 1918
(Dautel et al., 1999). Domestic chickens are
apparently considerably less liable to be
attacked by Ar. reflexus than are pigeons. In
the laboratory, any available mammal may
serve as host. Although the European pigeon
tick is nearly always associated with pigeons,
the exigencies of its domestic existence drive
it to attack humans, possibly more frequently
than does Ar. persicus (Rolla et al., 2004;
Spiewak et al., 2006). There have been a
number of reports describing human cases of
a specific anti-Ar. reflexus allergy (Chappard
et al., 1996; Veraldi, 1996; Kleine-Tebbe et al.,
2006). Allergy to Ar. reflexus has to be considered in patients having allergy symptoms
and living in buildings where pigeons have
their nests. Severe systemic anaphylactic
reactions (urticaria, angiooedema, dyspnoea,
cardiovascular dysregulation, unconsciousness) were reported in 12 of 148 (8%) humans

after bites of the soft tick Ar. reflexus and 146
of 148 (99%) had local reactions (Kliene-Tebbe
et al., 2006). Man is frequently attacked, especially in the vicinity of long unoccupied
pigeon sites. The pigeon tick can withstand
long periods of starvation; it may remain
unfed in or near pigeon houses for many
months, or even for several years. This species can also be the vector of B. anserina. There
is no evidence that Ar. reflexus transmits pathogens to humans in central Europe. The colour plate section contains two photographs
(dorsal and ventral) of an adult male of
Ar. reflexus (Plate 2a,b).
Argas polonicus
Ar. polonicus is a poorly known species that
seems to be restricted to domestic and wild
pigeons. It has been largely confused with a
related species, Ar. vulgaris, and it differs distinctly from Ar. reflexus (Siuda et al., 1979).
Phenotypic (involving the use of cuticular
hydrocarbons) and genetic analysis also
showed that the two species are different
(Estrada-Pea and Dusbabek, 1992).
2.1.2
Genus Ornithodoros
The soft tick species within the genus

Ornithodoros are able to transmit ASFV. All
Ornithodoros species tested up to the present
have been proved to be competent vectors
for ASFV (Table 2.1). Some of these belong
to the taxonomic group of O. erraticus, a
group of species distributed mainly in the
Mediterranean basin and Middle East, including Transcaucasia and parts of Russia
(Morel, 1969). Snchez-Botija (1963) demonstrated that O. erraticus found in pig sites
can be a vector of ASFV in Europe, while
Table 2.1. Laboratory and eld studies on Ornithodoros species to determine their vectorial capacity for
African swine fever virus (ASFV) (EFSA, 2010a,b).
Virus presence
(max. period
post infection)
Species
Region
O. coriaceus
North
502 days
America
O. turicata
North
America
O. puertoricensis North
America
O. parkeri
North
America
O. savignyi
Africa
O. moubata
Africa
O. erraticus/
Europe
O. marocanus
Virus transmissiona
Transstadialb Transovarial Sexual To pigs
References
+ (4)
ND
23 days
NDd
ND
ND
+ 502 dpic Hess et al., 1987;

Endris and
Hess, 1994
+ 23 dpi
239 days
ND
ND
+ 239 dpi
70 days
+ (1)
ND
ND
106 days
+ (1)
ND
3 years
+ (5)
5 years
+ (5)
+ 106 dpi
Mellor and
Wilkinson,
1985
+ 3 years Kleiboeker et al.,
1998, 1999;
Plowright,
1977; Plowright et al.,
1970a,b, 1974,
1994; Rennie
et al., 2001
+ 588 dpi Hess et al.,
1989; Endris
et al., 1992;
Boinas, 1994
a
Only the maximum lengths of time for virus presence and transmission to pigs, or horizontal or vertical transmission
among tick populations, are considered; b number of moults; cdays post infection; dND, no data.
10
Rbert Farkas et al.
Plowright et al. (1969a,b) demonstrated that

O. moubata/porcinus ticks associated with
warthogs were also involved in virus transmission in Africa. The group of species O.
moubata/porcinus are not found in Europe or
in northern Africa. O. erraticus is accepted as a
reservoir for ASFV with a well-established
role in the maintenance of the infection in
nature (Plowright et al., 1970a,b). Replication
of ASFV in this tick species has been demonstrated, and this allows the species to remain
infectious for months and up to 5 years, posing a threat of transmitting the virus each
time the ticks feed on pigs (Boinas, 1994).
The adults and nymphal stages 4 and 5 of
O. erraticus remain infected for longer than
the larvae up to nymph 3 (Boinas, 1994; Basto
et al., 2006). It has also been reported that the
prevalence of infected O. erraticus and the
average titres of ASFV per tick decreased over
time when pigs were not present (Boinas,
1994). The maintenance of virus infection in
the tick can constitute risk if the ticks are able
to feed on pigs for periods of 40 weeks after
an ASFV infective meal (Basto et al., 2006).
More recently, Vial et al. (2007) demonstrated
ASFV DNA in O. sonrai collected from the
field in West Africa. O. sonrai, although a
different species, is morphologically undistinguishable from O. erraticus. The two species
may copulate but the progeny are not fertile.
At present, as there is not a solid ground test
to determine which of these two tick species
is which, any overlapping distribution in
some areas is ignored, as well as the basic biological mechanisms of separation of the two
species. It is probably that the main mechanism of isolation of the species leading
to reproductive isolation is the simple
geographical partitioning of the populations
(Vial et al., 2007).
Some authors consider ASFV and the
Ornithodoros tick as co-evolving organisms.
In fact, there are noticeable telomeric similarities in the genomes of ASFV and Borrelia,
with the latter sharing the same Ornithodoros
tick host in Africa and considered to be an
original pathogen of soft ticks, suggesting
that ASFV is also a primary organism of
Ornithodoros ticks and co-adapted to its
tick hosts (Hinnebusch and Barbour, 1991).
This hypothesis could explain discrepancies concerning infection success rates

that have been noticeable between several
past surveys.
For example, De Tray (1963) reported a
consistent establishment of the virus isolate
Uganda in specimens of the O. moubata
group (34/35 were infected), whereas another
isolate, Tengani, only caused persistent
infection in a small proportion of ticks (2/46
were infected). More recently, Kleiboeker
et al. (1999) compared oral and intra-haemocoelic experimental infections of Ornithodoros
ticks collected from warthog burrows in
Kruger National Park and the Northern
Transvaal region of South Africa, as well as
infections of ticks from Masai Mara Reserve
in Kenya, by three different viruses from
South Africa, Malawi and Zimbabwe; all the
viruses were originally isolated from ticks
collected from the field. The oral infection
conducted with the isolate from Malawi was
self-limiting (with a decline of virus titres
and the number of ticks containing virus),
while the other infections persisted. According
to Kleiboeker et al. (1999), the cytopathology
caused by the Malawi strain in infected ticks
suggested the non-adaptation of the isolate to
express specific genes that allow the production of large quantities of progeny virus without damaging the host cell. The reason why
this virus was originally isolated from ticks
could be the large opportunity for those
ticks to feed on infected pigs with high viraemic titres during an ASF outbreak and the
leakage of midgut contents into the haemocoel without tick mortality, instead of real
adaptation of this virus isolate to the tick host
(Kleiboeker et al., 1999).
Dixon and Wilkinson (1988) analysed
genomes of ASFV isolates collected over a
2 year period from ticks inhabiting warthog
burrows in four regions of Zambia and
observed additional sequences in the region
close to the left-end terminus of the genome,
which are not observed in domestic pig isolates. The author concluded that virus replication in ticks and warthogs may require
additional host-specific genes not necessary
for multiplication in domestic pigs and that
the introduction of virus from tick/warthog
sources into domestic pig populations

would remove the selection pressure for
maintaining these genes (Dixon and
Wilkinson, 1988). More recently, Alfonso
et al. (2004) and Burrage et al. (2004) demonstrated that the ASFV multigene family in
the left variable region of the genome
encodes a host range determinant required
for efficient replication and generalization
of infection in pigs and ticks. However, no
more information is yet available on specific determinants for tick/warthog hosts,
as it was previously suggested. In addition,
it is unknown if ASFV is able to come back
from the domestic to the sylvatic cycle,
although some authors suggest that recombination processes during co-infections in
ticks may exist (Plowright, 1977; Dixon and
Wilkinson, 1988).
Identification
Ticks in the family Argasidae are easily separated from those in the family Ixodidae
because of the presence of the mouthparts
(capitulum) in a ventral position, instead of
in the terminal position as in members of the
family Ixodidae, and by the absence of a dorsal scutum in the adults. Furthermore, the
scutum (or conscutum, in males) is very
prominent in the specimens of the family
Ixodidae. The species in the genus
Ornithodoros share the features of the family,
as well as the absence of a lateral suture,
which separate species in the genus Argas
from species in the genus Ornithodoros. Other
than this feature, the mamillate structures on
the dorsal surface of the genera Argas and
Ornithodoros are different.
The identification of species of
Ornithodoros is difficult because most of the
species are not morphologically different.
Wallis and Miller (1983) conducted electrophoretic enzyme analysis to differentiate
between some of the species. In Eurasia, the
adults of most species of Ornithodoros have a
similar morphology, rendering the identification on that stage difficult, and the larvae
are the most useful stages to provide an
adequate determination. According to the
morphological features outlined by Filippova
11
(1966), ticks in the O. erraticus complex can be

separated by the setal dorsal complement in
the larvae, which is reduced to only two pairs
of dorsal setae. Currently, however, there is
no accurate way to separate the different species in the complex. Furthermore, there are
no adequate DNA data for the species found
in the wide distribution range of the complex, and there is strong evidence pointing to
a greater diversity than has currently been
reported. The colour plate section contains
two photographs (dorsal and ventral) of
an adult (sex undetermined) of O. erraticus
(Plate 3a,b).
2.2
Hard Ticks (Ixodidae)
Several physiologicalbehavioural and morphological characteristics distinguish the

hard ticks, Ixodidae from the soft ticks,
Argasidae. The life cycle of the hard ticks has
only one nymphal stage. A single but comparatively large blood meal is ingested by the
larva, the nymph and the adult female. The
ixodid female, in particular, is capable of
imbibing a very large amount of blood so that
the abdomen becomes enormously distended.
So in general, the ixodid ticks, except the
males in some species, stay attached to the
host for a period of several days. The morphology of ixodids has the following characteristics: (i) an anteriorly located capitulum
visible from above; (ii) behind the capitulum
a hard shield, the scutum; (iii) distinct sexual
dimorphism in that the dorsum of the adult
ixodid male is covered completely by the scutum, but the larva, nymph and adult female
have a scutum that only covers the anterior
part of the dorsal abdomen.
According to Barker and Murrell (2008),
the Ixodidae encompass the following genera:
Amblyomma, Anomalohimalaya, Bothriocroton,
Cornupalpatum, Cosmiomma, Dermacentor, Haemaphysalis, Hyalomma, Ixodes, Margaropus,
Nosomma, Rhipicentor and Rhipicephalus.
It may be noted that five species previously
in the genus Boophilus have been incorporated into the genus Rhipicephalus (subgenus
Boophilus).
12
Rbert Farkas et al.
2.2.1
Genus Ixodes
This is the largest genus of the family

Ixodidae, the hard ticks. The ticks in this
genus are so-called prostriate hard ticks,
which are characterized by the anal groove
encircling the anus anteriorly. The genus is
present on all continents, including Antarctica.
The species of most medical and veterinary
importance are I. scapularis and I. pacificus in
North America, I. ricinus in Europe, western
Asia and the Mediterranean coast of North
Africa, and I. persulcatus in north-eastern
Europe and northern Asia. In Eurasia, in view
of their roles as vectors of zoonotic pathogens among the natural host animals, I. hexagonus (present in Europe only), I. caledonicus,
I. frontalis, I. trianguliceps and I. uriae may also
be considered as medically and epidemiologically important tick species. Among
all Ixodes species, approximately 20% parasitize birds and 80% parasitize mammals
(Kolonin, 2009).
The host-seeking behaviour of this group
of ticks can be classified as endophilic or
exophilic. Endophilic ticks remain hidden
close to their hosts nest or burrows and only
attach to the host when it arrives, whereas
exophilic ticks seek hosts by climbing up the
vegetation waiting for them to pass. This
host-seeking behaviour of exophilic Ixodes
spp. should be distinguished from that of
hunting ticks, e.g. Hyalomma spp., which
actively run towards potential hosts. Because
of the difference in host-finding behaviour,
only exophilic species can be collected by
flagging or dragging. In the genus Ixodes,
exophilic species such as I. ricinus usually
mate on the host. Endophilic species such as
I. hexagonus normally mate in the hosts refuge (Hillyard, 1996).
Ixodes ricinus
In Europe, the most well-known tick species
is I. ricinus; this is often called the common
tick, castor bean tick or sheep tick. It is the
most commonly encountered tick species
in most parts of central, western and northern Europe. The distribution of this species
ranges from Ireland, Britain and Scandinavia,
Finland and western Russia across continental
Europe southwards to the Mediterranean

area into northern Africa and eastwards
to Iran.
I. ricinus is a three-host tick with a very
wide host range (telotropic host behaviour),
which includes lizards, many species of
birds and small, medium-sized and large
mammals. This tick species will readily
attach to humans. Jaenson et al. (1994) listed
two lizard species, 56 bird species and 29
mammal species as hosts recorded for
I. ricinus in Sweden alone. The colour plate
section contains photographs (dorsal and
ventral views) of adult male (Plate 4a,b)
and female (Plate 4c,d) specimens of
I. ricinus.
In Europe, I. ricinus is the vector of the
agents of several zoonoses, including Lyme
borreliosis (at least eight Borrelia genospecies
have been recorded from I. ricinus; Rudenko
et al., 2011), TBE, anaplasmosis (granulocytic
ehrlichiosis, tick-bite fever) due to Anaplasma
phagocytophilum and Rickettsia helvetica. I. ricinus also transmits Babesia divergens (the causative agent of babesiosis in cattle and a
life-threatening infection in splenectomized
humans), B. microti and B. venatorum, tularaemia due to Francisella tularensis and the louping ill virus (LIV).
Ixodes persulcatus
The taiga tick is an extremely important
vector of the TBE virus, Lyme borreliosis spirochetes and other pathogens in Finland, the
Baltic States and Russia, and also eastwards
to Japan. In the Baltic region and northwestern Russia, I. persulcatus overlaps with
the range of I. ricinus and replaces I. ricinus
eastwards through Siberia. I. persulcatus transmits, in general, the same pathogens as
I. ricinus, i.e. the TBE virus, Lyme borreliosis
spirochetes, A. phagocytophilum and B. divergens. Both tick species are occasionally also
vectors, and possibly long-term reservoirs, of
F. tularensis. In I. ricinus, it is the nymphal
stage which is the most important one for
transmission of pathogens to humans. In contrast, the nymphs of I. persulcatus rarely bite
humans; therefore it is the adult females of
this species which play this main role of transmitting pathogens to humans.
Ixodes hexagonus
This tick is better known as the hedgehog tick
and ranges through western, central and
southern Europe, including Ireland, the UK,
Norway, Sweden and Poland, to Portugal,
Spain, Italy, Greece, Romania and Ukraine
(Kolonin, 2009). It can easily be mistaken for
I. ricinus, but there is a pronounced hexagonal shield in female ticks. The main hosts
are carnivorous mammals (Canidae, Felidae
and Mustelidae), and the hedgehog, i.e.
mammals with a permanent dwelling
(Arthur, 1963; Jaenson et al., 1994; Kolonin,
2009). It is rarely found on sheep, cattle,
horses and birds (Hillyard, 1996). This tick
bites man occasionally and was a pest in
underground shelters in the London area
during wartime (Browning, 1944). I. hexagonus is a competent vector of Borrelia burgdorferi s.l. and transfers the spirochaete both
trans-stadially and transovarially (Gern et al.,
1991); it is also possibly a competent vector of
TBEV and Babesia microti. The species has an
endophilic feeding behaviour and remains
close to the shelter of its hosts. Although relatively abundant, it is never caught by dragging or flagging. The colour plate section
contains photographs (dorsal and ventral
views) of adult male (Plate 5a,b) and female
(Plate 5c,d) specimens of I. hexagonus.
Ixodes canisuga
I. canisuga Johnston 1849 (= I. crenulatus Koch
1844; according to Kolonin, 2009) is often confused with I. hexagonus, which is usually more
commonly encountered. I. canisuga is distributed from western, northern and southern
Europe through Russia into Iran, Afghanistan,
India and China (Hillyard, 1996; Kolonin,
2009). All stages are parasitic on rodents and
carnivores, including badgers, foxes, mustelids, dogs and cats, i.e. hosts that regularly
return to a burrow or lair (Hillyard, 1996).
The colour plate section contains photographs
(dorsal and ventral views) of an adult female
of I. canisuga (Plate 6a,b).
Ixodes gibbosus
The main hosts of the larvae, nymphs and
adults of I. gibbosus are sheep and goats, but
13
the larvae and nymphs also feed on small

mammals and birds. The species is distributed from former Yugoslavia through Turkey
to Israel (Kolonin, 2009).
Ixodes uriae
This is the seabird tick, which inhabits islands
and mainland coasts in the subarctic and
temperate regions of the northern and southern hemispheres. I. uriae parasitizes colonynesting marine birds. This tick will
occasionally bite people visiting seabird
colonies. It is a vector of the human Lyme
disease spirochaete Borrelia garinii (Olsen
et al., 1993). The colour plate section contains
photographs (dorsal and ventral views) of
an adult male (Plate 7a,b) and female
(Plate 7c,d) of I. uriae.
Ixodes frontalis
Several arboviruses (Bahig, Kemerovo, TBEV)
have been isolated from the bird-parasitizing
tick I. frontalis, which is an ornithophagous
ectoparasite that does not bite humans.
However, it is presumably indirectly of enzootic
and epidemiological importance for TBEV.
Ixodes trianguliceps
I. trianguliceps is a nidicolous ectoparasite of
small mammals. It is distributed throughout
most of Europe eastwards through Ukraine to
Georgia, Armenia and Azerbaijan (Kolonin,
2009). Although this tick does not bite humans
it may indirectly be of medical and veterinary
importance in view of its presumed role as a
maintenance vector of TBEV, Borrelia afzelii
and Babesia microti, among its natural small
mammalian hosts (rodents and insectivores).
2.2.2
Genus Haemaphysalis
Few of the about 160 species of Haemaphysalis

parasitize livestock, but those that do are economically important in Eurasia, where they
inhabit humid, well-vegetated habitats. The
size of unfed adults ranges from 2.0 to 4.0 mm.
The genus is characterized by short mouthparts and a large rear-facing spur on trochanter
14
Rbert Farkas et al.
I. The capitulum is short and the palps are

wide. Ornamentation and eyes are absent but
festoons, usually 11 in number, are present on
the posterior margin. Coxa I lacks external
spurs. The anal groove passes behind the
anus. Nymphs of Haemaphysalis possess festoons but lack ornamentation and eyes
(Hillyard, 1996). Species of the genus Haemaphysalis are three-host ticks, with the larvae
and nymphs feeding on small mammals and
birds. Adults infest larger domestic and wild
mammals and, importantly, livestock; they are
especially adapted to domestic cattle and, to a
lesser extent, to sheep and goats.
Haemaphysalis punctata
The palps of the females and males are short
and wide, projecting laterally beyond the
basis capituli. The basis capituli of females is
rectangular, about twice as broad as long, but
it is only one and a half times broad as long in
males. The scutum of males is longer than it
is broad; in females it is about as broad.
Unlike the female, the basis capituli of
nymphs, in dorsal view, has sides with sharp
lateral points (Hillyard, 1996). The colour
plate section contains photographs (dorsal
and ventral views) of an adult male (Plate 8a, b)
and an adult female of Haemaphysalis (Ha.)
punctata (Plate 8c,d). Ha. punctata has the largest distribution range in Eurasia. It is also
known as the red sheep tick and is found in a
very wide variety of habitats, providing that
a high degree of humidity is present. This
tick has many similarities with I. ricinus in its
distribution, but is distinct in its physical
characters from that species. This species is
widely distributed throughout Europe, and it
occurs in cold and wet sites in most
Mediterranean countries (Estrada-Pea et al.,
2004). Ha. punctata is one of the two species of
the genus Haemaphysalis that are likely to be
found on livestock in North Africa, and it
needs to be distinguished from Ha. sulcata
(Bouattour et al., 1999). It is associated with
wild and domestic ruminants (Encinas
Grandes, 1986; Manilla, 1998a; Cringoli et al.,
2002). It is frequently found where sheep,
goats and cattle feed in certain open forests
and shrubby pastures in the greater part of
Europe, including southern Scandinavia and
Britain. The species is also found on horses,

deer and hares, and occasionally it attaches to
humans (Hillyard, 1996). Engorgement on
the host may take 630 days to complete and
each female lays 30005000 eggs. Immature
ticks infest birds (e.g. crows, magpies, pheasants, sparrows, blackbirds), hedgehogs,
rodents and lizards. In humid climates, adults
may be found feeding from October to March.
Under favourable conditions the life cycle
can be completed within 1 year but it usually
takes 3 years.
Ha. punctata is a vector of many Babesia
species (B. major, B. bigemina, B. bovis, B. motasi) and Theileria species (T. ovis, T. mutans,
T. recondite). It is also able to transmit F. tularensis and Coxiella burnetii. Specimens of this
species have been found to be infected by
tick-borne encephalitis virus (TBEV) and
Bhanja virus. The tick is also suspected of
being able to transmit Crimean-Congo
haemorrhagic fever virus (CCHFV), Listeria
monocytogenes and Brucella abortus. The bite
of Ha. punctata may cause tick paralysis
(Hillyard, 1996).
Haemaphysalis concinna
Ha. concinna is widely distributed in Europe
(Walter et al., 1986; Srter et al., 2003). The species is restricted to temperate broadleaved
and mixed forests where the environment is
relatively unaltered, and the distribution is of
relict type (Hoogstraal, 1972). The most
important hosts for adult Ha. concinna are roe
deer, but the ticks may occur on cattle, horses,
dogs, cats and hares. The sexes are usually
found attached close together and copulation
occurs on the host in April. Males can be more
numerous than females. The larvae and
nymphs feed on deer, hedgehogs, rodents,
reptiles and birds. Nymphs have also been
removed from humans (Hillyard, 1996). The
larvae and nymphs may be found in many
months of the year; they are most numerous
on small mammals in June and July. In
Hungary, Ha. concinna showed peak activity
in May (Hornok, 2009). Ha. concinna is known
to be a carrier of F. tularensis. The colour plate
section contains photographs (dorsal and
ventral views) of an adult male of Ha. concinna
(Plate 9a,b).
Haemaphysalis inermis
Ha. inermis, known as the winter tick, is a
rather small tick. The capitulum of females
differs from the normal form in the genus.
The palps are relatively long and narrow,
and when folded they do not project laterally beyond the relatively wide basis capituli. The scutum is broader than it is long.
All coxae have a small spur. In males, the
palps lack lateral projections and there are
11 well-marked festoons on the posterior
margin (Hillyard, 1996). The colour plate
section contains a photograph of an adult
female (ventral view, Plate 10) of Ha.
inermis.
This tick is present in some foci within
many European countries (e.g. France,
Poland, the Mediterranean region and the
Balkan Peninsula). Ha. inermis is winter
active and inhabits deciduous and mixed
forests, grasslands and mountain slopes. The
species is primarily associated with deer,
with the immature stages found on small
mammals, rodents and insectivores. The
adults also feed on large and small domestic
and wild mammals (e.g. cattle, horses, sheep,
fox and hedgehogs). Humans may also be
attacked (Hillyard, 1996). Copulation takes
place on the host but the sexes do not attach
alongside each other. The species has peak
activity in November and December in
Hungary (Hornok, 2009). The immature
stages are rarely found in nature, primarily
because they feed and drop off the host
within hours an exceptionally short period
for engorgement. However, the nymphs
rapid feeding contrasts with the very long
time needed to develop to the next stage;
nymphs may take a year or more before
moulting to the adult stage. Ha. inermis has
been noted as causing tick paralysis. In
Poland it is suspected of carrying the virus
that causes tick-borne encephalitis (TBE)
(Hillyard, 1996).
Haemaphysalis sulcata
Ha. sulcata is one of the two species of the
genus Haemaphysalis that are likely to be
found on livestock in a wide range of climates
in the Mediterranean region; it needs to be
15
distinguished from Ha. punctata. In Europe,

this tick species has been detected in several
countries (Cyprus, Greece, Romania, former
Yugoslavia, Hungary, Italy, France and Spain),
always under conditions of steppe vegetation. In North Africa, the species occurs in
Morocco, Algeria, Tunisia and Libya, as well
as in parts of Turkey (Estrada-Pea et al.,
2004). Adults of the species feed on ungulates.
Small ruminants, particularly sheep and
goats, are the most common hosts. The preferred feeding sites for the ticks are the neck,
shoulder and back. Adults are active in the
autumn and winter between October and
March. Immature stages feed on reptiles such
as lizards, but the range of hosts of larvae and
nymphs is similar to that of Ha. punctata.
Larvae and nymphs feed on reptiles in the
spring and summer (April and July). Probably
because of misidentification in the past, the
role in the transmission of pathogens by this
species is not yet clearly defined. It is suspected of transmitting T. annulata and
Anaplasma ovis. The Bhanja virus has also
been isolated in Ha. sulcata during a survey
performed in Bulgaria (Pavlov et al., 1978).
Other species of Haemaphysalis are potentially important in Europe and the
Mediterranean region. For example, examination of animals in El Arich city and
El Hassanah centre in Egypt revealed Ha.
parva on sheep, Ha. sulcata on goats, and Hy.
lusitanicum on cattle (Mazyad and Khalaf,
2002). Ha. franchinii were also identified in
small numbers in Tunisia (Bouattour et al.,
1999). Another important species is the rabbit
tick, Ha. leporispalustris.
2.2.3
Genus Hyalomma
Hyalomma species are large ticks with pale

rings on most segments of their legs. They also
have long mouthparts which project to the
anterior of the body and, in males, anal plate
(Manilla, 1998b; Estrada-Pea et al., 2004).
The hard tick genus Hyalomma is a complex of a few species that exhibit an almost
endless variation of morphology. Its original
centre of dispersal was probably Iran or
southern Russia. Genetic instability may in
16
Rbert Farkas et al.
part account for the wide morphological

differences found in many specimens
(Hoogstraal, 1956).
For identification purposes, there is a
need to consider only the differential features
not subject to change on an individual basis;
these are mainly seen in replete ticks. To
date, the most reliable diagnostic features are
actually scanty, especially for the females.
Adult Hyalomma ticks share the following common features: a basis capituli that is
usually rectangular, clearly wider than long; a
long hypostome with a rounded tip and 3/3
denticles; long palps with the 2nd article
longer than the 3rd; shiny eyes with a surrounding orbit (a circular groove surrounding
the eye), which are often convex and placed
in the confluence between the anterior and
the posterior edges of the scutum in females
and in the corresponding place in males; a
dark scutum rich in punctations and grooves;
festoons that are more evident in unfed ticks;
long, strong and banded legs with two large
spurs on the first pair of coxae; and adanal,
subanal and accessory plates in the male.
In most cases, it is not possible to identify
immature stages to the species level using
only morphology (Manilla, 1998b).
Hyalomma ticks are often the most abundant tick parasites of livestock, including
camels, in warm, arid and semiarid, generally
harsh, lowland and middle altitude biotopes,
and in biotopes with long dry seasons, from
central and south-west Asia to southern
Europe and southern Africa. Extraordinary
survival factors play a large part in permitting these ticks to exist and even thrive where
few or no others live.
Many Hyalomma species have been
redescribed recently based on morphological characteristics (Apanaskevich, 2003;
Apanaskevich and Horak, 2005). However,
these characteristics may vary with populations, host and other environmental factors,
making the ticks difficult to identify. Hyalomma
ticks are mainly two- and three-host species
(Matthysse and Colbo, 1987).
Hyalomma marginatum
Hyalomma marginatum is a large tick that can
reach 18 mm length in females and 6 mm
in males. Its dorsal shield is wider than long,

brown in colour and rich in punctations of
different size. The legs have white-yellowish
rings, and the body of the replete male has
subanal triangular plates that can be visible
beyond the posterior edge (Manilla, 1998b).
(dorsal and ventral views) of adult male (Plate
11a,b) and female (Plate 11c,d) specimens of
Hy. marginatum.
This species belongs to a hard tick species complex occurring in southern Europe,
southern Asia and most of Africa. It is a
characteristic species of steppe, savannah
and lightly wooded hill and valley biotopes
with fairly low humidity. In particular,
Hy. marginatum inhabits the Mediterranean
climatic zone, as well as steppe and foothill
landscapes
(south-western
Palaearctic
Region), from the Caspian Sea to southern
Ukraine and Bulgaria, and spreads westward to Spain, Portugal, Morocco and
Algeria (Manilla,1998). Sporadic reports of
the species in the humid biotopes of northern Europe can be explained as introductions by migratory birds that are
frequently included in the host range of the
immature stages. Hy. marginatum is a twohost tick and the immature stages stay for a
long time on the host (1226 days) which
enables them to be passively transported by
migrating birds over hundreds or even
thousands of kilometres (Rechav et al., 1987;
Manilla, 1998b).
Hy. marginatum is the main vector of
CCHFV in Eurasia. The virus can also be
transmitted transovarially with variable
efficiency depending upon the viral strain
(Zeller et al., 1997). The ticks are also a vector of Babesia caballi and T. equi, and of
T. annulata (Viseras and Garca-Fernndez,
1999). T. annulata is the agent of tropical
theileriosis that occurs in northern Africa,
including the sub-Saharan territories, Sudan
and Eritrea, southern Europe, the Near and
Middle East, Central Asia, India and northern China. Its distribution reaches the Far
East, where it overlaps with theileriosis
caused by T. sergenti infection (Pipano and
Shkap, 2004). Hy. marginatum is also
involved in the maintenance of the enzootic
cycle of C. burnetii.
Hyalomma anatolicum and Hyalomma

excavatum
The study of the ecology, development and
epizootic role of these two similar species is
very difficult. Their taxonomic status is not
even uniformly agreed on, and their occurrence within the same geographical area,
together with the presence of hybrid forms,
morphological variations and impossibility
of distinguishing the larvae and nymphs,
complicate this work. Ecological conditions
apparently determine which form will
develop in a given location (Liebish et al.,
1989; Apanaskevich, 2003).
A recent study that compared the morphology of each stage of the two species
using several specimens belonging to
different collections allowed researchers to
definitely separate the two specific identities of Hy. anatolicum and Hy. excavatum
(Apanaskevic and Horak, 2005).
Hy. anatolicum is common in the southern Mediterranean area and
also in eastern Africa (Sudan, Somalia and
Ethiopia) and central Asia (Ahmed et al., 2007;
Rasulov, 2007). This is a small tick; the length
and breadth of the conscutum is usually less
than 4 mm and 2.3 mm in the male, and 2 mm
and 1.9 mm in the female, respectively. The
scutum, gnathosoma and coxae are light yellowish- or reddish-brown. Ivory coloured
marbling is absent on the scutum, with large
and sparse punctations in the caudal field.
The species transmits a variety of protozoan parasites trans-stadially and transovarially and is a threat to animal improvement
programmes. It transmits the protozoans
T. annulata (which causes tropical theileriosis
in cattle) T. lestoquardi (which causes malignant
ovine theileriosis), B. caballi and T. equi (which
cause equine piroplasmosis). Hy. anatolicum is
also a vector of CCHFV to humans.
HYALOMMA ANATOLICUM
HYALOMMA EXCAVATUM
Hy. excavatum is larger
than Hy. anatolicum. The scutum, gnathosoma
and coxae are dark red- or black-brown. Ivory
coloured enamelling is diffuse on the dorsal
aspects of the leg segments. In the female, the
cervical and lateral scutal grooves are deeper.
17
(dorsal and ventral views) of adult male

(Plates 12a,b) and female (Plates 12c,d) specimens of Hy. excavatum.
The species is adapted to the Mediterranean and steppe climatic regions of North
Africa, and to steppe climatic regions elsewhere in its wide range. Hy. excavatum is often
less commonly found on livestock than
Hy. anatolicum, but it has a wider geographical distribution and is found in much of North
Africa from Mauritania to Egypt, and also
into Sudan, Ethiopia and Eritrea. It is not
found commonly south of the Sahara, but it is
reported eastwards through to Iran and
Turkmenistan.
The biological differences between the
two species influence their vector competences. Hy. excavatum is questioned as a vector of Theileria, because its vectorial capacity
is affected by its feeding habits on rodents
(Manila, 1998b, Walker et al., 2003).
Hyalomma scupense
Two years ago, and despite some morphological differences, the Hyalomma subspecies
that were known as Hy. detritum detritum and
Hy. detritum scupense were brought together
to comprise a unique species renamed as
Hy. scupense (Guglielmone et al., 2009).
Hy. scupense is a two-host tick species
that feeds on large domestic ungulates.
Compared with other Hyalomma ticks, it is
small, less punctated and does not have pale
rings on its legs. The former subspecies Hy. d.
detritum and Hy. d. scupense are different in
their body size (Hy. d. detritum is generally
smaller than Hy. d. scupense), the length of the
legs, and the thickness of the tails of the spiracle plates, which are narrower in Hy. d. detritum than in Hy. d. scupense (Manilla, 1998b;
Walker et al., 2003). The colour plate section
(Plate 13c,d) specimens of Hy. scupense.
This Hyalomma tick spreads in central
Asia from northern India to the Middle East,
and is also found in the southern Balkans,
southern Europe and along the Mediterranean
coast of Africa as far as Algeria and Morocco
in the west. It is present in sub-Saharan Africa
in north-central Sudan as well, perhaps
18
Rbert Farkas et al.
having invaded from the Red Sea coast or via

the Nile river valley (Manilla, 1998b).
Hy. scupense is able to transmit T. annulata to cattle and T. equi to horses, mules
and donkeys. The species is also a vector of
C. burnetii.
2.2.4
Genus Amblyomma
Amblyomma species are large ticks with long

mouthparts and pale rings on their legs. They
usually have ornate eyes and coxal spurs.
Like all Ixodidae, they are protected by a rigid
shield that completely covers the dorsal body
surface in the male; female shields are smaller,
enabling significant distension of the abdominal wall during blood feeding.
In some species, the shield is richly coloured. The long mouthparts are clearly visible
on the dorsal view of the body, and these
make Amblyomma ticks especially difficult to
remove. Manual removal frequently causes
serious wounds which may become secondarily infected by bacteria or screw worms
(Walker et al., 2003).
According to the Ixodidae evolutionary
hypothesis formulated by Hoogstraal and
Kim (1985), the tick genera belonging to the
metastriate group originated from a group of
ticks that resemble the present subfamily
Amblyomminae. These ticks reached their
greatest evolutionary success by simultaneously adapting themselves to the dinosaurs
during the geological time from the end of the
Palaeozoic to the end of the Cretaceous
period. Nowadays, adults and immature
stages of 37 of the around 102 known
Amblyomma species parasitize reptiles.
Moreover, reptiles, together with small mammals and birds, are often included in the host
range of immature Amblyomma ticks that feed
on mammals when they moult to become
adults (Manilla, 1998).
The genus Amblyomma is generally confined to the tropics and subtropics. Amblyomma
(Am.) variegatum and Am. hebraeum are the
most important species of the genus in Africa.
They occur in different geographical areas,
although they are adapted to the same type of
habitat. Their presence is affected by climate
and the surrounding environment according
to their developmental stage. Even though

individuals of both species are active throughout the year, in the African ranges of distribution a certain seasonality is reported for both
species. Generally, adults are more numerous
during the rainy season, while larvae and
nymphs start to be more active at the end
of the rainy season. Am. variegatum and
Am. hebraeum are morphologically similar.
Am. variegatum adults have long mouthparts
and banded legs like Am. hebraeum, but the
colour pattern on the scutum is different and
they have beady eyes. In addition, the festoons of the males are dark brown in colour.
Am. variegatum (the tropical bont tick,
TBT) is the most widespread species. This
tick originates from Africa and has spread
widely south of the Sahara desert. It was first
introduced into the Caribbean in around
1828, following the importation of cattle
from Senegal, and later spread to the rest
of the West Indies (Uilenberg et al., 1984,
Pegram et al., 2005). Am. variegatum was
similarly introduced from Africa into the
Arabian Peninsula and Mauritius (Norval
and Horak, 2004). It is seldom reported in the
Mediterranean, but was probably introduced
there by migratory birds (Albanese et al.,
1971, Papadopolous et al., 1996).
Am. hebraeum is exclusively a southern
African tick. It occurs in South Africa and in
southern Mozambique. Its distribution also
extends inland through the lowveld areas of
Swaziland and eastern Botswana, and to
southern and western Zimbabwe. The distribution of Am. hebraeum is affected by interspecific competition with Am. variegatum.
Both Am. variegatum and Am. hebraeum
are the main vectors of Ehrlichia ruminantum,
causing heartwater, a disease that is one of
the main constraints to livestock production
in sub-Saharan Africa. These ticks also transmit T. velifera and T. mutans to cattle. In the
tropics, Am. variegatum is also strictly associated with dermatophilosis and the transmission of African tick bite fever (caused by
R. africae) (Norval and Horak, 2004).
In America, Am. maculatum and Am.
americanum are two of the main species of
Amblyomma ticks. Am. maculatum is an important pest of livestock and is present in the
Gulf coast area. It has been experimentally
demonstrated to be a good vector for the

agents transmitting heartwater (Uilenberg,
1982). Am. americanum, the lone-star tick, is
abundant in the southern USA in Texas and
Missouri and up the Atlantic Coast, ranging
northward into New Jersey. It is also a notorious pest in Mexico as well as in Central and
South America. Its distinctive feature is the
silver spot on the scutum (the lone star).
Am. americanum is the vector of tularaemia,
Rocky Mountain spotted fever, Q fever, and
Lyme disease, and it may cause tick paralysis
in humans and dogs. It is also a vector of
monocytic ehrlichiosis caused by E. chaffeensis
and E. ewingii in humans and dogs.
2.2.5
Genus Rhipicephalus
The genus Rhipicephalus is one of the largest

of the family Ixodidae. Species in this genus
have some features in common: a short
hypostome and palps, a usually hexagonal
basis capituli and the presence of eyes and
festoons and anal plate in the males. With
the exception of few species, they are inornate, hence the common name of the brown
ticks. This is mainly an African genus
(Walker et al., 2000).
Adults of most species parasitize wild
and domestic artiodactyls (swine and ruminants), perissodactyls (horses) or carnivores.
Immature stages feed mostly on smaller
mammals; however, of those that parasitize
rodents and of those that parasitize artiodactyls, a few feed on the same host as the adults.
The life cycle is typically three host, but in the
Mediterranean climatic zone (long, warm
summer with low rainfall), Rhipicephalus (Rh.)
bursa is a two-host species (Merck, 2010).
Many Rhipicephalus species have long
been difficult to identify or have been incorrectly identified. Current concepts of tick
phylogeny, taxonomy and nomenclature are
being revised and expanded based on molecular analyses. This ongoing work is likely to
expand and alter the current understanding
of the phylogeny and evolution of the subfamily Rhipicephalinae. On the basis of those
studies, the genus Boophilus has recently been
included in the genus Rhipicephalus, but even
19
though Boophilus species now belong to the

same genus as Rhipicephalus species, the morphology and biology of the Boophilus species
are significantly different.
The genus Rhipicephalus comprises 79
species, including the five species that were
formerly in the genus Boophilus. Probably the
species of Rhipicephalus with the largest geographical range is Rh. sanguineus, the kennel
tick or brown dog tick, which has travelled
worldwide with domestic dogs. It is now
established in buildings as far north as
Canada and Scandinavia, and as far south as
Australia. It can be found almost worldwide,
mainly within latitudes 35S and 50N
(Dantas-Torres, 2008). Although this species
feeds primarily on dogs, it can be found on a
diverse range of wild and domestic animals,
including humans (Dantas-Torres, 2008).
However, in urban situations everywhere
within the range of distribution, dogs are virtually the only hosts of the immature stages
and the adults. Rh. sanguineus is active
throughout the year in the tropics and subtropics, but only from spring to autumn in
temperate zones. Newly active adults and
nymphs are frequently seen climbing walls
from floor-level cracks (Merck, 2010).
Rhipicephalus ticks are recognized worldwide
as vectors of many important disease agents
of animals and humans diseases such as:
Mediterranean spotted fever and other rickettsioses, Q fever, monocytic ehrlichiosis,
canine babesiosis, hepatozoonosis, Nairobi
sheep disease and East coast fever.
Rhipicephalus sanguineus, Rhipicephalus
camicasi and Rhipicephalus turanicus
Rh. sanguineus is universally known as the
kennel tick (Manilla 1998c; Walker et al., 2000;
Estrada-Pea et al., 2004). It is a medium sized,
pale yellowish-brown or reddish-brown tick.
The Rh. sanguineus group comprises several
tick species, and the biosystematic status of
the majority of them has been confused
(Walker et al., 2000; Estrada-Pea et al., 2004).
Two species of the group that closely resemble
Rh. sanguineus are Rh. camicasi and Rh. turanicus. Females of Rh. sanguineus are differentiated from those of Rh. camicasi and Rh. turanicus
by the genital aperture, which usually has a
20
Rbert Farkas et al.
broad V shape in Rh. sanguineus but a narrow

U shape in Rh. camicasi and Rh. turanicus. Both
sexes of Rh. sanguineus have spiracle plates
with tails which are narrow, and less than the
width of the adjacent festoon. In Rh. turanicus,
these tails are broad. Males of Rh. sanguineus
do not have a depression of the cervical fields
compared with the small depression that there
is in Rh. camicasi (Walker et al., 2000; EstradaPea et al., 2004). The adults may be separated
by a combination of adanal and spiracular
plate characteristics, the dorsal punctation
pattern (males), and by the feature of the genital aperture (females). To study the morphology of the genital aperture, it is necessary to
dissect and mount it, and allow adequate
cleaning to remove the chitinized structures.
(dorsal and ventral views) of adult male
(Plate 14a,b) and female (Plate 14c,d) specimens of Rh. sanguineus. There are also photographs (dorsal and ventral views) of adult
male (Plate 15a,b) and female (Plate 15c,d)
specimens of Rh. turanicus.
Rhipicephalus bursa
Adults of Rh. bursa are large ticks. Males have
very conspicuous anal plates, almost in a regular triangular shape, with a projection easily
seen in the median portion and protruding to
the anus (medial). Both males and females
have a noticeable large dorsal punctation pattern, deeper than those observed in other species of the genus collected in the Mediterranean
region. The colour plate section contains photographs (dorsal and ventral views) of adult
male (Plate 16a,b) and female (Plate 16c,d)
specimens of Rh. bursa.
Rhipicephalus annulatus ( previously
known as Boophilus)
Rhiphicephalus (Boophilus) ticks are unique in
that their entire life cycle from the larva to the
engorged, mated adult is confined to a single
host. Females drop to the ground to oviposit.
This single-host type of life cycle has numerous biological advantages. It also allows for
particularly easy control by dipping infested
animals. The boophilid type of life cycle
eliminates danger-ridden periods between
two or three different kinds of hosts, possibly

in inhospitable areas and for indefinite periods. The predilection of these ticks for large
domestic animals particularly favours widespread dispersal and survival, not only within
a continent but also from continent to continent on imported hosts. Cattle are the chief
hosts throughout the world; horses, other
domestic stock, and wild antelopes and deer
are less frequently attacked. Other wild
animals are not commonly infested. The
veterinary importance of these ticks is
considerable and they are vectors of important pathogens of livestock. The only species present in the Mediterranean basin is
Rh. annulatus. This is an important pest of
domestic cattle in every kind of Mediterranean
environment. Most interesting is that this tick
is well adapted to xerophilic areas with periodic rains. Temperature seems to be the only
restrictive factor in the dissemination of the
tick towards northern latitudes. Hence it is
widespread at scattered points across the
Mediterranean basin. Rh. annulatus is the
vector of some prominent pathogens, such
as several species of Babesia, most importantly
B. bovis, which is a potentially dangerous
pathogen of cattle. The colour plate section
(Plate 17c,d) specimens of Rh. (Bo.) annulatus.
2.2.6
Genus Dermacentor
The genus Dermacentor is small, with about 30

species of hard ticks, most of which are found
in the New World. They are medium sized to
large ticks, usually with ornate patterning,
and resemble those of Hyalomma. Most species of Dermacentor are three-host ticks, but a
few are one-host ticks. Two species, D. marginatus and D. reticulatus, are widely distributed in Europe (Hillyard, 1996).
Dermacentor marginatus
D. marginatus, also known as the ornate sheep
tick, has an ornate scutum. It is distinguished
from D. reticulatus by its palp which lacks a
prominent rear-facing spur. The length of
unfed adults is 4.65.8 mm; an engorged
female measures up to 1.5 cm. The unfed

nymph is 1.41.8 mm in length.
This species is one of the two representatives of the genus Dermacentor that infest
domestic animals in the Mediterranean region.
D. marginatus occurs widely in Europe, but
has a restricted distribution in North Africa
(Nosek, 1972). In the European part of its distribution, this tick species is restricted to areas
with dense bush and tree cover. It is common
under oak and pine vegetation. D. marginatus
has more thermophilic requirements than
other species that can occur together, such as
I. ricinus and Ha. punctata. In North Africa,
D. marginatus is restricted to the same type of
habitat as I. ricinus. Thus it is found in the
cooler and more humid parts of the
Mediterranean climatic region associated with
the Atlas Mountains, and is restricted to small
areas of Morocco, Algeria and Tunisia (EstradaPea et al., 2004). The ticks inhabit pastures,
temperate forest and grassland, having a preference for xerophilic vegetation.
The activity of the species varies considerably according to region. Adults are active
during the spring, early summer and autumn.
In a study in Hungary, D. marginatus was
found to be most numerous in February and
March (Hornok, 2009). The larvae appear in
June and the nymphs in July. A generation
from egg to adult usually develops within a
year. Adults feed on large mammals such as
domesticated and wild ruminants, but also
on dogs, hares, hedgehogs and humans. Wild
boars (Sus scropha) are also important wild
hosts. The larvae and nymphs feed mostly on
small mammals such as rodents and birds
(Nosek, 1972; Hillyard, 1996).
D. marginatus is a vector of C. burnetii,
R. conori, R. slovaca and R. sibirica sibirica.
F. tularensis may also be transmitted by this
tick species (Nosek, 1972; Hillyard, 1996;
Raoult et al., 2002). The colour plate section
(Plate 18c,d) specimens of D. marginatus.
Dermacentor reticulatus
D. reticulatus (syn. D. pictus) is also known as
the ornate dog tick, the marsh tick or the
ornate cow tick. This metastriate tick species
21
has been confused with others in the genus

because it has considerable morphological
variability (Estrada-Pea and Estrada-Pea,
1991a,b; Zahler and Gothe, 1997). The length
of unfed adults is 3.84.8 mm; an engorged
female measures up to 1.0 cm. The unfed
nymph is 1.41.8 mm in length. The colour
plate section contains photographs (dorsal
and ventral views) of adult male (Plate 19a,b)
and female (Plate 19c,d) specimens of
D. reticulatus.
The distribution area of the species
extends from France and south-western
England in the west to Central Asia in the east
(Gilot et al., 1989). D. reticulatus is recorded in
almost all European countries but formerly
was relatively rare in cold continental
climates. In western and Central Europe, it
does not occur north of 5354N latitude, for
example in Scandinavia, nor in the
Mediterranean climate zone. In East Europe,
however, it may occur as far north as
St Petersburg (60N). Within this large distribution area, the occurrence of this species is
highly focal (Dautel et al., 2006). In the last
two decades, several European countries
have noticed changes in the distribution of
D. reticulatus (Szymanski, 1987; Zahler and
Gothe, 2001; Srter et al., 2005; Dautel et al.,
2006; Bullova et al., 2009). The spread of
Dermacentor ticks is thought to be linked to
importation (Cochez et al., 2011) and climate
changes, notably the shortening of the winter
period and increasing minimum temperatures. The ticks are absent from Northern
Africa (Estrada-Pea et al., 2004). Most important for the survival and activity of D. reticulatus are foci of adequate microclimate
(Estrada-Pea, 2008). Therefore, the tick is
abundant in open, unused habitats both in
the middle and on the periphery of villages,
in unused gardens, on abandoned prairies, as
well as in grazing pastures and woodlands.
D. reticulatus is absent from mountain regions,
but very abundant in low-altitude hills.
Favourable landscapes for the survival of the
species occur where different forms of cultivation overlap and urbanization modifies
landscape structures, such as is found in periurban colonies.
The tick has a 1 or 2 year life cycle depending on the environmental conditions. It prefers
22
Rbert Farkas et al.
cold regions with an adequate amount of air

relative humidity, so it is absent in the
Mediterranean region where the relative
humidity is not sufficient for its survival. The
main activity period for adults is the spring,
with a secondary peak in autumn, but it varies
considerably according to region (Szymanski,
1987). For example, D. reticulatus was most
numerous in September and October in
Hungary (Hornok, 2009), and high numbers
of questing adults were found in Belgium during winter when temperatures were just above
0C (Madder, unpublished data). Adults parasitize larger domestic and wild mammals, cattle, horses, sheep, goats and pigs, but only
occasionally bite humans. D. reticulatus is one
of the most common tick species on dogs and
wild canids wherever it exists (Gilot et al.,
1989; Fldvri and Farkas, 2005). Copulation
takes place on the host. The female feeds for
915 days before dropping to the ground to
lay approximately 4000 eggs in sheltered

spots. The larvae hatch from the eggs after 23
weeks and will feed for approximately 2 days
on the host, then drop to the ground and
moult to the nymphal stage. This stage feeds
over a period of several days, drops off and
moults to become an adult. The larvae and
nymphs feed on a variety of small mammals,
such as small rodents and carnivores, and
occasionally birds (Nosek 1972; Hillyard,
1996). The immature stages are generally
active from midsummer to late autumn.
D. reticulatus is known to be a vector of
Babesia spp. (B. canis, B. divergens, B. ovis,
B. caballi), T. equi, A. ovis, R. conori and
R. raoultii (formerly Rickettsia sp. strain RpA4)
(Nosek 1972; Martinod and Gilot, 1991;
Hillyard, 1996; Zahler and Gothe, 2001; Dautel
et al., 2006). In Portugal, recent studies have
confirmed the presence of F. tularensis in
D. reticulatus (Lopes de Carvalho et al., 2011).
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3
1
Factors Influencing the Spread

and Distribution of Ticks
Maxime Madder1 and Ilaria Pascucci2

Institute of Tropical Medicine, Antwerp, Belgium and University of
Pretoria, South Africa; and 2Istituto Zooprofilattico Sperimentale
dellAbruzzo e del Molise G. Caporale, Teramo, Italy
3.1
Climate Changes
Long-time series of historical data related to

climatic factors and the geographical distribution of ticks and tick-borne pathogens are
required to determine the impact of climate
changes on the spread of these vectors and
the pathogens they carry. To our knowledge,
few such attempts have been made, mainly
because of the lack of data collection over a
sufficiently long period of time to allow for
comparison in relation to existing climatic
data. One of the most recent and long-term
studies has been described by Jaenson et al.
(2012). In this article it was shown that deer
abundance and climate change seem to be the
most important factors explaining the spread
of Ixodes ricinus northwards in Sweden during the last 30 years. It is suggested that the
milder climate has made it possible for roe
deer to expand northwards and into northern
Sweden. As the roe deer is, generally, the most
important host of adults of I. ricinus in Sweden
the expansion of this tick host has favoured
the subsequent expansion of the tick into the
same northern area (Jaenson et al., 2012).
It is possible, however, to affirm that
despite some evidence it is difficult to implicate climate change as the main cause for an
increase in tick-borne diseases. In order to
predict future tick-borne scenarios, climate
change models are required that take into
account the dynamic biological processes

involved in vector abundance and pathogen
transmission which affect the complex ecology and epidemiology of tick-borne diseases
such as Crimean-Congo haemorrhagic fever
(CCHF), tick-borne encephalitis (TBE) and
Lyme borreliosis. According to Gray et al.
(2009), climate changes have influenced the
survival of ticks (direct effect). Empirical
proofs have been obtained that climate may
be in part responsible for shaping the prevalence rates of Borrelia burgdorferi s.l. in questing nymphal I. ricinus (Estrada-Pea et al.,
2011). However, studies on the dynamics of
CCHF cases in Turkey did not show any influence of climate when sites where the disease
had been reported were compared with sites
where the disease was absent (Estrada-Pea
et al., 2010).
Changes in climate and the length of the
different seasons directly affect tick survival,
activity and development, but there is limited
evidence that rising temperature results in a
greater abundance of ticks simply by increasing rates of development. Effects of the climate on the geographical distribution of a
tick species are generally more evident along
the border of its distribution, affecting the
geographic range potentially suitable for the
tick. A study by Jaenson and Lindgren (2011)
has shown that a warmer climate in Northern
Europe has extended the vegetation period
27
28
which has allowed ticks to be active for a

longer time during the year; this seems to
have enlarged the geographic range available
for the tick. Then again, changes in development rates make tick cohorts available to different diapause windows (largely determined
by day length), thus changing the patterns of
seasonal activity and significantly altering
generation length. Probably the most interesting tick species to work on and to explore the
diversity of seasonal patterns is I. ricinus, the
most abundant tick in Europe and an important pest of both animals and humans.
However, a process-driven model, which is
necessary to study such diverse seasonality
patterns, is still not available.
The indirect effects of climate change
are more complicated to evaluate. Climate
changes can indirectly alter survival of the
ticks by modifying the type of vegetation and,
as mentioned above, the host abundance. For
instance, climate will affect the vegetation,
which in many ways may influence the
number of infected ticks. In addition, dense
humid vegetation will, for some tick species,
be beneficial to tick survival and could lead to
an increase in tick numbers as well. Such vegetation may also benefit pathogen-reservoir
hosts such as rodents and shrews, and
increase the survival and abundance of tickmaintenance hosts such as deer (Jaenson et al.,
2012). All these factors may boost the density
of infected ticks, but the direct impact on animal and human infection rates is unknown
and has not yet been evaluated for any of the
many pathogens that the ticks can transmit.
The influence of mast years on rodent
abundance and consequently on tick density
is one example of how vegetation will indirectly affect the density of Borrelia-infected
Ixodes ticks (Ogden et al., 2005). After mast
years, rodent populations significantly
increase as a result of an extended breeding
season and higher survival during winter.
This is due to an increase in food supply and
consequently in body fat. The higher host
population density affects the tick density in
the following year and possibly also the risk
of disease transmission (Tack et al., 2010).
Climate change may also influence the
risk of infection by affecting human activities;
for example, the long-term use of land, e.g. for
agriculture or tourism, could be affected by

climate change. Weather patterns also have
an effect by influencing short-term human
behaviour so that tick-bite risk may be linked
to the seasonality of certain human activities
such as hiking, hunting, and berry and mushroom picking. Generally climate effects are
more easily noticeable close to the geographical distribution limits of both vector and
pathogen, but the magnitude of the effects of
climate change in an endemic area is the result
of the interaction of many other parameters
associated with socio-economics, human
migration and settlement, ecosystems and
biodiversity, bird migrations, land use and
land cover, human culture and behaviour,
and immunity in the human and animal populations (Gray et al., 2009).
3.2
Changes in Land Use
Changes in land use often affect the hosttick

relationship and consequently increase the
transmission of tick-borne pathogens. For
instance, during World War II, after the occupation of Crimea (19411944), normal agricultural activities were disrupted and the
common sport of hunting European hares
was abandoned. When Soviet troops reoccupied the hilly Crimean steppes in 1944, the
hares had become excessively abundant and
neglected pastures were overgrown with
weeds. European hares are a reservoir of
CCHF virus (CCHFV) and one of the primary hosts of immature Hyalomma, and with
the expansion of the hare population came the
first documented CCHF outbreak of the
modern era (Ergnl, 2006). A similar explanation was suggested for an outbreak in
Turkey as hunting and pasturing had been
abandoned in the fields in the affected region
from between 1995 and 2001 because of terrorist activities (Ergnl, 2006). In this period,
the numbers of small mammals (e.g. hares)
and wild animals (e.g. wild boars) increased,
as did the population of Hyalomma ticks.
When the fields became available again for
hunting and pasturing, cattle, sheep and
humans were exposed to virus-carrying ticks
(Ergnl, 2006).
Factors Influencing Tick Spread and Distribution
3.3
Movement of Animals
3.3.1
Migratory birds
The introduction and spread of tick species

by migratory birds are documented for those
tick species which are ornithophilic in the larval and nymphal stages. Hyalomma (Hy.) marginatum and Hy. rufipes, and to a lesser extent
some species of Amblyomma, belong to this
group of ticks. Their potential spread northwards and the establishment of permanent
populations in the north are therefore of great
importance, particularly as the immature
stages of Hyalomma are frequently found on
migratory birds flying northward to temperate Europe (e.g. Jaenson et al., 1994; Molin
et al., 2011). The dispersal of ticks by birds
may be restricted to a short distance during
local post-breeding flights or over extremely
long distances during migration flights. In
studies conducted on birds migrating through
Egypt between 1955 and 1973, it was discovered that birds migrating from Eurasia to
Africa carried tick species that were characteristic of the fauna of Europe and Asia
(Hoogstraal, 1963). As another example, the
introduction of Am. variegatum in southern
Italy and Greece by migratory birds has been
sporadically reported (Albanese et al., 1971;
Papadopoulus et al., 1996).
Hy. rufipes is the most common tick found
on birds migrating northwards from subSaharan Africa to Eurasia in the spring
(Hoogstraal, 1963; Linthicum and Bailey,
1994). The risk of diffusion of CCHFV into
areas where the virus does not occur is linked
to the possibility of infected Hyalomma ticks
reaching and becoming established in such
uninfected areas. This probability is greater
for ticks associated with migratory birds
(Manilla, 1998a). Hoogstraal (1979) reported
that many bird species are responsible for the
intracontinental and intercontinental dissemination of ticks associated with CCHFV.
While this fact is not the most important
factor in the introduction of CCHFV to
Europe, it has nevertheless been hypothesized
that the outbreak of CCHF in Turkey in 2002
could have been caused by birds carrying
CCHFV-infected ticks from the Balkans (Karti
et al., 2004). The occurrence of this disease in
29
Turkey is under discussion, taking into consideration the available epidemiological data.
It currently seems impossible that migrating
birds from the Balkans could have introduced
the disease into Turkey and have it spread in
an interval of weeks. The current opinion is
that the virus has been there many years and
that some change/s favoured its contact with
humans so that disease incidence increased
(Ozdarendeli et al., 2010).
3.3.2
Importation of exotic and

wildlife species
There are many examples of exotic ticks that

have been introduced into countries by the
importation of exotic animals. Some reptileassociated African Amblyomma tick species
were found on the American mainland,
mainly as a result of the importation of tickinfested reptiles from Africa (i.e. infested by
Am. marmoreum or Am. sparsum). Also, in
Poland, Am. latum and Am. transversale were
found on pythons and Am. exornatum,
Am. flavomaculatum, Am. latum and Am. nuttalli on monitor lizards, both reptiles imported
from Africa (Nowak, 2010). In addition, in
the Netherlands, Am. flavomaculatum was collected from an African savannah monitor lizard (Nijhof et al., 2007), and other tick species
such as Am. marmoreum, Am. dissimile and
Hy. rufipes have occasionally been collected
from imported tortoises and snakes (Bronswijk
et al., 1979; Jongejan, 2001). Similarly, a constant threat for the introduction of heartwater
into the American mainland exists mainly by
the importation of livestock from the
Caribbean Islands infested with Am. variegatum ticks, but also through the introduction of
lightly infested wild ungulates and tickinfested reptiles from Africa, as mentioned
above (Burridge et al., 2002).
3.3.3
Movement of domestic animals
Ticks associated with pets are able to reach

and establish themselves in areas far from
their origins. One of the most typical examples is Rhipicephalus (Rh.) sanguineus, universally known as the kennel tick (Manilla,
30
1998b; Walker et al., 2000; Estrada-Pea et al.,

2004). This is a monotropic three-host tick
strictly associated with the domestic dog.
Rh. sanguineus is probably one of the most
widely distributed tick species in the world.
Travellers with dogs have brought this important vector species permanently into the
latitudes approximately between 50N and
30S (Walker et al., 2000). Furthermore,
Dermacentor reticulatus (the ornate dog tick,
marsh tick or cattle tick) may have been introduced into north-western Europe in a similar
way, in addition to the routes described
previously.
D. reticulatus is a tick species that appears
to be expanding its range in north-western
Europe. A study performed in Germany by
Dautel et al. (2006) in 2003 and 2004 showed
that the distribution and abundance of
D. reticulatus has increased on deer and vegetation. Populations have also been found over
the last few years in the Netherlands and
Belgium (Nijhof et al., 2007; Cochez et al., 2011).
In the Netherlands, several populations have
been found in freshwater tidal marshes,
mainly in the south of the country; these were
most likely established after the introduction
of southern European cattle breeds. Ponies
were exported to Belgium to manage vegetation, and these ponies then introduced this
tick species to that country. In Belgium,
D. reticulatus has been found on tick (flag or
drag) cloth dragged in different nature
reserves and peri-urban areas, and on dogs.
Most of these areas are frequented by dogs,
possibly explaining further spread of D. reticulatus, as no large mammals are present there.
The introduction of Am. variegatum into
the Caribbean area is an example of how the
movement of domestic animals can spread
exotic ticks. The initial introduction of Am.
variegatum into Guadeloupe and probably
also into most of the West Indies islands was
due to the importation of infested NDama
cattle in the 1800s, or to tick-infested litter
used during cattle transportation by ships
from Senegal to the West Indies (Barr et al.,
1987), or from East Africa to the Indian Ocean
islands. Cattle transportation between islands
also explains the introduction of the tick at
least into Marie-Galante, Antigua and
Martinique. But from the late 1960s, the rapid
increase in the number of infested islands in

the Caribbean area was linked to the presence
and spread of the cattle egret in the West
Indies. This cattle-associated African bird
species was also introduced into the Caribbean
area (Corn et al., 1993; Barr et al., 1995). The
reinfestation of some countries, for example
Puerto Rico, after a successful eradication
campaign might also be due to the movement
of this bird (Bokma and Shaw, 1993). Am. variegatum is thus the only African vector of cowdriosis that has established itself successfully
outside the continent (Walker and Olwage,
1987). It remains a threat in areas where climatic conditions are suitable for its establishment, as it has already succeeded in colonizing
new territories through the movement of
migratory birds which cannot be controlled.
Rh. (Boophilus) microplus, the pantropical
blue tick, is without a doubt one of the most
important vectors of livestock pathogens
in the world, mainly because of its large
geographical distribution and its ability to
transmit both Asiatic and African redwater
babesiosis caused by Babesia bovis and
B. bigemina (Coetzer and Tustin, 2004).
This tick species originates from Asia,
but during the latter half of the 19th century
it, and the babesiae it transmits, was spread by
cattle transported to Australia, Madagascar,
South Africa, Latin America, Mexico and the
USA (Hoogstraal, 1956; Temeyer et al., 2004).
Rh. microplus was eradicated from the USA in
1943 through the costly Cattle Fever Tick
Eradication Program which started in 1906.
The most resistant strain studied to date
has been shown to survive the dipping procedure used at the USAMexico border as
part of an importation barrier to prevent
the re-entry of Rh. microplus into the USA
(Temeyer et al., 2004).
Until recently, Rh. microplus was not
present in West Africa, but this tick species has
unfortunately been introduced into the Ivory
Coast (Madder et al., 2007, 2011) and Benin
(Madder et al., 2012) as a result of uncontrolled
cattle imports for cattle improvement programmes. Because of the high adaptability of
ticks and, more specifically, of Rh. microplus,
the risk of the introduction of this species into
the Mediterranean area and the establishment
of local populations seems possible.
Factors Influencing Tick Spread and Distribution
31
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climate on geographic range and seasonality of the tick Ixodes scapularis. International Journal for
Ozdarendeli, A., Canakoglu, N., Berber, E., Aydin, K., Tonbak, S., Ertek, M., Buzgan, T., Bolat, Y., Aktas, M.
and Kalkan, A. (2010) The complete genome analysis of Crimean-Congo hemorrhagic fever virus isolated in Turkey. Virus Research 147, 288293.
Papadopoulus B., Morel P.C. and Aeschliman A. (1996) Ticks of domestic animals in the Macedonia region of
Greece. Veterinary Parasitology 63, 2540.
Tack, W., Madder, M. and Verheyen, K. (2010) De impact van mastjaren op knaagdieren, teken en de ziekte
van Lyme. Antenna 4(3), 2931.
Temeyer, K.B., Pound, J.M., Miller, J.A., Chen, A.C., Pruett, J.H. Jr, Guerrero, F., Davey, R.B., Untalan, P.M.,
Lohmeyer, K.H., Li, A.Y., Miller, R. and George, J.E. (2004) Organophosphate resistance in Mexican
strains of Boophilus microplus: a major threat to the U.S. cattle industry. Southern Association of
Agricultural Scientists Bulletin of Biochemistry and Biotechnology 17, 4351.
Walker, J.B. and Olwage, A. (1987) The tick vectors of Cowdria ruminantium (Ixodoidea, Ixodidae, genus
Amblyomma) and their distribution. Onderstepoort Journal of Veterinary Research 54, 353379.
Walker, J.B., Keirans, J.E. and Horak, I.G. (2000) The Genus Rhipicephalus (Acari, Ixodidae): A Guide to the
Brown Ticks of the World. Cambridge University Press, Cambridge, UK.
Tick-borne Infections
(Including Zoonoses) in Europe
and the Mediterranean Basin*
Frank Koenen,1 Ilaria Pascucci,2 Thomas G.T. Jaenson,3 Maxime Madder,4
Rita de Sousa,5 Agustn Estrada-Pea,6 Rbert Farkas7 and Mo Salman8
1
Veterinary and Agrochemical Research Centre, CODA-CERVA, Brussels, Belgium;
2
Istituto Zooprofilattico Sperimentale dellAbruzzo e del Molise G. Caporale, Teramo,
Italy; 3Uppsala University, Sweden; 4Institute of Tropical Medicine, Antwerp, Belgium
and University of Pretoria, South Africa; 5National Institute of Health Doutor Ricardo
Jorge, IP (INSA), guas de Moura, Portugal; 6University of Zaragoza, Spain; 7Szent
Istvn University, Budapest, Hungary; and 8Colorado State University, USA

African swine fever (ASF) is a viral swine disease caused by the African swine fever virus
(ASFV), an icosahedral complex DNA virus that
is a unique member of the Asfarviridae family.
It affects only porcine species, those of all breeds
and ages. The disease was first described in
Kenya by Montgomery in 1921 when the virus
spread from infected warthogs (Phacochoerus
aethiopicus) to domestic pigs (Sus scrofa) and
resulted in a 100% case fatality rate. The disease
is currently present in Africa, mainly in countries located south of the Sahara, and in most
cases it is endemic. In Europe, ASF is still
endemic in Sardinia. More recently, in 2007,
ASFV spread to the Transcaucasian countries (TCC) and the Russian Federation (RF)
(Snchez-Vizcano, 2006).
Pigs are the only domestic animal species that are naturally infected by ASFV. Wild
boar have also been identified as susceptible

to ASFV infection, with clinical signs and
case fatality rates similar to those observed
in domestic pigs in Spain, Portugal and
Sardinia (Italy) and, experimentally, in feral
pigs in Florida (Snchez-Vizcano, 2006).
Wild boar and feral pigs can transmit the
virus directly to domestic swine. In Africa, it
has been observed that ASFV induces an
inapparent infection in three species of wild
pigs (warthogs, bush pigs and red river
hogs), while the role played by the giant forest hog has not yet been clarified (Jori and
Bastos, 2009).
At present, no treatment or effective
vaccine against ASFV is available. Since
1963, when the first live attenuated vaccine
was used in Portugal, many efforts have
been made in this area, but with unsatisfactory results. As no vaccine for ASFV currently exists, the prevention of this disease in
* Frank Koenen wrote Section 4.1; Ilaria Pascucci, Section 4.2; Thomas G.T. Jaenson, Sections 4.3, 4.7,
4.11.2; Maxime Madder, Sections 4.4, 4.5; Rita de Sousa, Section 4.6; Agustn Estrada-Pea, Section 4.8;
Rbert Farkas, Sections 4.9, 4.10, 4.11.3, 4.11.4; Mo Salman, Section 4.11.1.
33
34
Frank Koenen et al.
disease-free areas depends on preventing

the introduction of the virus (SnchezVizcano, 2006).
ASFV is spread among domestic pigs
via the oralnasal route. However, it has
also been demonstrated that the virus can
be infectious by a number of other routes,
including tick bites, experimental inoculation via cutaneous scarification, and by the
intramuscular, intravenous, subcutaneous
and intraperitoneal routes. The infection
usually commences in the monocytes
and macrophages of the tonsils and
mandibular lymph nodes. From there it
spreads through the draining lymph nodes
and blood to the target organs (lymph
nodes, bone marrow, spleen, lung, liver
and kidney) which are the principal sites of
secondary replication.
The clinical signs of ASF can resemble a
variety of other swine haemorrhagic diseases
and can easily be confused with classical
swine fever (hog cholera) and erysipelas.
Laboratory tests are necessary to confirm the
diagnosis. ASF can also present different clinical signs that depend primarily upon the
virulence of the virus, infectious dose and
mode of infection, with a range of clinical
forms varying from acute to subclinical and
chronic.
Some species of soft ticks have proved to
be ASFV reservoirs and vectors, such as
Ornithodoros moubata and O. porcinus in Africa
and O. erraticus in the Iberian Peninsula
(Spain and Portugal). In O. moubata, transovarial and trans-stadial ASFV transmission
have been described; in O. erraticus, only
trans-stadial transmission has been demonstrated. Other soft tick species that are widely
distributed in North and South America have
been identified as harbouring and transmitting ASFV, and in the experimental setting,
O. savignyi, present in Africa, has been
shown to transmit ASFV to domestic pigs (see
Table 2.1).
In Africa, ASFV is maintained by a cycle
of infection between wild pigs and soft ticks.
In some of these wild pigs, ASFV infection is
characterized by low levels of virus in the tissues and low or undetectable levels of viraemia; however, these levels of virus are
sufficient to infect soft tick vectors and for
tick transmission of the virus to domestic

pigs. This cycle of the virus makes ASF very
difficult to eradicate in Africa. In Sardinia,
where ASF is still present, wild boars are as
susceptible as domestic pigs. No ticks from
the O. erraticus complex have been found in
Sardinia.
Experience of past outbreaks of ASF
outside Africa has shown that the introduction of ASFV into a non-infected pig population within a free region is most often linked
to entry through international ports or
airports. The infected material is typically
garbage containing uncooked pork that is
used for pig feeding (Snchez-Vizcano,
2006). Once ASFV is established in domestic
swine, infected animals are the most important source of virus dissemination to susceptible pigs. In Europe, ASFV was introduced
for the first time in 1957 into Portugal
through waste from international flights.
Although this first outbreak was rapidly
eradicated, in 1960 the virus entered Europe
again in Lisbon (Portugal) and spread
through the rest of Portugal and Spain,
where ASFV remained endemic until 1995.
During this period, some other outbreaks
occurred in other European countries, affecting Andorra (1975), Belgium (1985), France
(1964, 1967 and 1974), Malta (1978), the
Netherlands (1986) and Italy (1967, 1969 and
1993), including the island of Sardinia, where
ASF has remained endemic since 1978. All of
these virus introductions were also linked to
swill feeding.
In Europe, several epidemiological
paths are known to be able to maintain ASFV
in domestic pig populations and this complicates the control of the disease. The main
routes of transmission are: swill, domestic
pig and wild boar interactions, and pigtick
interactions (Snchez-Vizcano, 2006). In
2010, a scientific opinion issued by the EFSA
(European Food Safety Authority) Panel on
Animal Health and Animal Welfare (2010a)
on ASF contained an assessment of the risk
of introduction of this virus into the European
Union (EU), especially from the Caucasus
(Wieland et al., 2011). Another scientific
report from the EFSA Panel on Animal
Health and Animal Welfare (2010c) discusses
the role of tick vectors in the epidemiology
Tick-borne Infections in Europe and Mediterranean Basin
of Crimean-Congo haemorrhagic fever

(CCHF) and ASF in Eurasia, and contains
geographic distribution maps of the tick vector and ASFV.
4.1.1 Virustick interaction

Several factors can influence the vector competence of soft tick species for ASFV. Some
authors consider ASFV and the Ornithodoros
tick as co-evolving organisms. Actually,
noticeable telomeric similarities in the
genomes of ASFV and Borrelia (the latter
shares the same Ornithodoros tick host in
Africa and is considered to be an original
pathogen of soft ticks) suggest that ASFV is
also a primary organism of Ornithodoros ticks
and that it coadapts to its tick hosts
(Hinnebusch and Barbour, 1991). This
hypothesis could explain the noticeable
discrepancies concerning infection success
rates that have been reported in several past
surveys. For example, De Tray (1963)
reported consistent establishment of the
virus isolate Uganda in specimens of the
O. moubata group (34/35 were infected)
whereas another isolate, Tengani, caused
persistent infection only in a small proportion of ticks (2/46 were infected). More
recently, Kleiboeker et al. (1999) compared
oral and intra-haemocoelic experimental
infections of Ornithodoros ticks collected
from warthog burrows in Kruger National
Park and the Northern Transvaal region of
South Africa, as well as infections of ticks
from Masai Mara Reserve in Kenya. These
researchers used three different viruses from
South Africa, Malawi and Zimbabwe, all
originally isolated from ticks collected in the
field. The oral infection using the isolate
from Malawi was self-limiting (decline of
virus titres and number of ticks containing
virus), while the others persisted. According
to Kleiboeker et al. (1999), the cytopathology
caused by the Malawi strain in infected ticks
suggested the non-adaptation of the isolate
to express specific genes that allow the production of large quantities of progeny virus
without damaging the host cell. The reason
why this virus was originally isolated from
35
ticks could be the large opportunity for those

ticks to feed on infected pigs with high viraemic titres during an ASF outbreak and the
leakage of midgut contents into the haemocoel without tick mortality, instead of real
adaptation of this virus isolate to the tick
host (Kleiboeker et al., 1999). Dixon and
Wilkinson (1988) suggested that virus replication in ticks and warthogs may require
additional host-specific genes that are not
necessary for multiplication in domestic
pigs, and that the introduction of virus from
tick/warthog sources into domestic pig populations would remove the selection pressure for maintaining these genes. However,
no more information is currently available
on specific determinants for tick/warthog
hosts, as was previously suggested. In addition, it is unknown whether ASFV is able to
come back from the domestic to the sylvatic
cycle, although some authors suggest that
recombination processes during co-infections
in ticks may exist (Plowright, 1977; Dixon
and Wilkinson, 1988).
4.1.2
Genetic diversification of ASFV
By sequencing the C-terminal end of the p72

gene, Bastos et al. (2003) and Lubisi et al.
(2005) observed higher genetic variations in
genotypes directly isolated from Ornithodoros
ticks and warthogs, or in genotypes circulating in East and southern Africa where the
sylvatic cycle plays a crucial part in the epidemiology of ASF. Some other genotypes
were only found in domestic pigs and presented low genetic divergence (Lubisi et al.,
2005). In Madagascar, using concatenated
sequences of the p22 and p32 genes, Michaud
(Michaud et al., 2007) detected relatively
high genetic divergence between Malagasy
virus isolates collected on domestic pigs
from 1998 and 2003, compared with that
observed on West African and European isolates since the 1970s. In Madagascar, it has
been suspected that the introduced virus
was adapted to local bush pigs and
Ornithodoros ticks, leading to its accelerated
diversification. Such diversification phenomena as reported in several African
36
Frank Koenen et al.
countries has been previously analysed by

Dixon and Wilkinson (1988) on Zambian
virus isolates from Ornithodoros ticks. A considerable genetic diversity was observed
between virus isolates from ticks collected
from the same regions and even from the
same warthog burrows. This diversity
resulted from peculiar point mutations all
along the length of the genome, instead of
insertions/deletions in the region close to
the left-hand terminus of the genome usually observed for host selection (Dixon and
Wilkinson, 1988). In this case, ticks would be
able to enhance the diversification of ASFV
and the emergence of new virulent isolates
to domestic pigs. However, no information
is yet available on the location and the
expression of this genetic diversification. In
addition, this process does not seem compatible with the persistence of ASFV in Iberian
Ornithodoros ticks and the observed genetic
homogeneity of ASFV in Europe.
4.2
Crimean-Congo Haemorrhagic
Fever
CCHF is a tick-borne zoonotic infection that

has public health concern in several regions
of the world including Africa, the Middle
East, the Balkans region, Greece, Turkey and
western Asia. The infection is caused by a
virus belonging to the genus Nairovirus (family Bunyaviridae) and is transmitted by several
species of hard (ixodid) ticks, particularly by
those belonging to the Hyalomma genus
(Horak et al., 2001).
Phylogenetic analyses performed on S-,
M- and L-RNA segments of the CrimeanCongo haemorrhagic fever virus (CCHFV)
showed virus strains grouped in seven different clades (Deyde et al., 2006). Three clades
are distributed in Africa, two in Europe and
the other two in Asia. Despite the potential
for dispersal of the virus in Africa and Eurasia,
it appears that circulation of the virus is
largely confined to within two specific regions
in these continents, and corresponds to the
distribution and dispersal of tick vectors of
the virus (Burt and Swanepoel, 2005; Paweska,
2007). It has also been concluded that viral
strains have a latitudinal relationship in

which there is not much interchange of the
strains between different latitudes (i.e. Africa
and Europe).
The virus may be maintained in tick
populations during inter-epizootic periods
through several mechanisms, such as transstadial and transovarial transmission, and
non-viraemic transmission of ticks aggregated on the same host (co-feeding).
Outbreaks usually take place during the
peak activity periods of Hyalomma ticks,
coinciding with the hot and dry season
(Swanepoel, 2006).
There are a large number of potential
vertebrate host reservoirs for CCHFV, reflecting the diverse feeding preferences of the
immature and adult tick vectors. Antibodies
against the virus have been detected in
domestic and wild animals, including hares,
hedgehogs, rodents, bats, and large mammals such as giraffes and rhinoceroses. The
most important source of virus transmission
is immature ticks of the Hyalomma genus
which have fed on the blood of viraemic
small vertebrates. Once infected, the tick
remains infected throughout its life (transstadial transmission), and the mature tick
may transmit the virus to large vertebrates
such as livestock. Domestic ruminants such
as cattle, sheep and goats will have viraemia
for approximately a week after becoming
infected. They may be the source of infection
for humans during slaughtering or veterinary procedures such as castration. In
Appendix 1, Table 1.24 contains data on the
serological surveillance of CCHF in domestic
animals. The level of viraemia in birds is usually low and unnoticeable, even though
migratory birds may play role in the epidemiology of the virus by disseminating
infected Hyalomma ticks. Tick-infested birds
migrating from Russia across the Black Sea
were suggested as a link to introduction of
the virus in Turkey (Karti et al., 2004).
Nevertheless, many social and environmental factors affect CCHF occurrence in Turkey.
The epidemiological features of the disease
are still under discussion.
The most common cause of infection in
humans is a bite from an infected tick.
Infection can also occur through direct contact
with blood or tissues from infected humans

or livestock. The highly pathogenic nature of
the virus occasionally results in serious nosocomial outbreaks (Swanepoel, 1995; Aradaib
et al., 2010; Elata et al., 2011; Naderi et al., 2011).
Clinical infection in humans is initially manifested as an acute febrile illness that can be
followed by a fatal haemorrhagic syndrome
with case-fatality rates of up to 50%
(Swanepoel et al., 1987). Misdiagnosis is frequently due to the nonspecific clinical signs
(Fisgin et al., 2010). The diagnosis is based on
serology or viral RNA detection by molecular
techniques. During human infection, viral
genomes are present in saliva and urine with
viral loads similar to those in blood (Bodur
et al., 2010). Two animal models have been
recently established by using genetically
defective mice in which the interferon
response had been altered (Bente et al., 2010;
Bereczky et al., 2010). These animal models
will be beneficial for the development of
treatments and vaccines (Keshtkar-Jahromi
et al., 2011).
Outbreaks in South Africa arose among
slaughterhouse operators during the slaughter of ostriches which were heavily infested
with ticks. The infection occurred when
the infected ticks on the carcasses were
squashed during skinning (Swanepoel, 1998).
Nevertheless, meat from butchered animals
does not pose a risk; in this substrate the
CCHFV is quickly inactivated by a drop in
pH, as occurs during the maturation process
that the meat undergoes after slaughter. A scientific report from the EFSA Panel and Animal
Health and Welfare (2010c) addresses the role
of tick vectors in the epidemiology of CCHF
and ASF in Eurasia.
4.3 Tick-borne Encephalitis Group

4.3.1 Tick-borne encephalitis
Tick-borne encephalitis (TBE) is one of the
most important and serious human infections occurring in Europe and many parts of
Asia. The aetiological agent, tick-borne
encephalitis virus (TBEV), is a member of
the genus Flavivirus of the family Flaviviridae.
37
TBEV is believed to cause at least 11,000

human cases of encephalitis in Russia and
more than 3000 cases in the rest of Europe
annually (Donoso Mantke et al., 2008).
Related viruses are louping ill virus (LIV),
Langat virus (LGTV) and Powassan virus
(POWV), which may also cause human
encephalitis, and Omsk haemorrhagic fever
virus (OHFV), Kyasanur Forest disease virus
(KFDV) and Alkhurma virus (ALKV), which
cause serious haemorrhagic fevers rather
than encephalitis (Gritsun et al., 2003).
Three subtypes of the TBEV are recognized: the Western or Central European
subtype, including the Kumlinge virus on
land in Finland; the Siberian subtype; and
the Far Eastern subtype. Recently, the
Siberian subtype was recognized as a
human pathogen in western Finland where
populations of the vector, Ixodes persulcatus,
were also recorded.
I. ricinus is the main tick vector involved
in TBEV infections in Europe. All its stages
can attack humans although the nymphs are
the most important ones as vectors of the virus
to humans. This is due to several factors: the
unfed larvae are usually not infected but may
become infected while taking their first blood
meal on a viraemic host or, which is more
important, by co-feeding with infective
nymphs; the nymphs are far more abundant
in nature than the adult ticks; and the colourful adult females are relatively large (3.5 mm)
compared with the dull-coloured smaller
(1.5 mm) nymphs and, therefore, are more
easily detected and removed when encountered on the human body. In TBE foci in central and northern Europe, the infection
prevalence of TBEV in nymphs ranges around
0.10.5%, with that in adults about 0.36.0%.
The infection prevalence in adult females of
I. persulcatus, which is the main vector stage
for the Eastern TBEV, tends to be much higher
(up to 40%) than in I. ricinus (Labuda and
Nuttall, 2008). Co-circulations of both Western
TBEV and Eastern TBEV occur in some foci in
the Baltic States, where the distributions of the
two tick species overlap. The Far Eastern subtype has been discovered not only in Siberia
but also in some European localities (Chausov
et al., 2010). In Estonia, all three humanpathogenic subtypes of TBEV have been
38
Frank Koenen et al.
found in the same areas (Golovljova et al.,

2004). Apart from the two main vector species, I. ricinus and I. persulcatus, several other
tick species, including I. hexagonus, I. arboricola, Haemaphysalis (Ha.) concinna, Ha. inermis
and Ha. punctata are competent but secondary
vectors (Labuda and Nuttall, 2008).
Until not long ago, it was believed that
viraemic small rodents, particularly the bank
vole, Clethrionomys glareolus, and the field
mouse, Apodemus flavicollis, and insectivores
were the principal reservoirs of TBEV that
infected the vectors. However, the viraemia
in these rodents is usually of short duration
(2 days). For the maintenance of TBEV in
I. ricinus populations, co-feeding transmission between infective nymphs and susceptible larvae feeding very close to and on
the same small rodent is now considered to
be much more important than transmission
via viraemic small mammals (Labuda and
Nuttall, 2008).
Many cases of TBE in humans are unrecognized and without clinical signs or symptoms. In some cases, however, the clinical
syndrome of TBE disease is severe, with a
life-threatening neurological syndrome and
high case fatality (5 to 35%) in its Eastern
form, mainly in Russia. In contrast, the case
fatality in Western Europe, mainly in Central
and Northern Europe, is usually comparatively low (approximately 1%) with nearly
all deaths confined to patients above
60 years of age. Patients infected with the
Siberian subtype may suffer from a milder
but often more chronic disease compared
with the disease caused by the Far Eastern
subtype.
The incidence of TBE usually fluctuates
from year to year, but an increased incidence
has been noted in some countries (Danielova
et al., 2006; Lindquist and Vapalahti, 2008),
and new TBE foci seem to have appeared,
especially in the last decade. This is presumably the result of a complex interaction of factors such as the changing climate affecting the
vector both directly and indirectly by affecting the plant and host communities, sociopolitical changes and technological factors,
e.g., better diagnostic methods and increased
awareness (Donoso Mantke et al., 2008;
Telford and Goethert, 2008).
4.3.2
Louping ill
Louping ill virus (LIV), also known as

ovine encephalitis/encephalomyelitis virus,
is closely related to TBEV and is the only
member of this virus complex present in the
British Isles, where the vector is I. ricinus
(Reid, 1988). Louping ill (LI) is endemic in
sheep-farming areas of northern England,
Scotland, Wales, Ireland and Norway. Several
tick hosts, such as the red grouse, willow
grouse, field vole and deer become viraemic
when infected with LIV. The viraemia, however, is usually too low to be infective to feeding tick larvae. In contrast to these wild hosts,
sheep and red grouse consistently develop
viraemia sufficient to infect tick larvae and
amplify the virus. Occasionally, horses, cattle
and goats develop viraemias sufficiently high
to be infective to tick larvae (Reid, 1988).
Mountain hares, Lepus timidus, may be maintenance hosts for LIV by non-viraemic
transmission between co-feeding ticks, as
experimentally demonstrated by Jones et al.
(1997). One of the main assumptions for the
transmission of a vector-borne pathogen is
usually that feeding by the vector is the sole
or main route of host infection. Gilbert et al.
(2004), however, demonstrated experimentally a transmission route whereby an important tick host, the red grouse (Lagopus lagopus
scoticus), became infected with LIV after eating infected I. ricinus ticks. These authors
estimated from field observations conducted
in Scotland that this mode of infection could
account for 7398% of all virus infections in
wild red grouse in their first season. Certainly,
this way of transmission has potential implications for the understanding of other vectorborne pathogens in which hosts may ingest
vectors through foraging or grooming.
LI is principally a disease of sheep and
red grouse, and less commonly of cattle,
other domesticated animals and birds.
Dogs, and particularly sheepdogs and hunting dogs in endemic areas, are occasionally
infected. Clinical signs include fever, ataxia,
trembling, salivation, coma and death. The
virus can cause severe encephalitis in
humans; about 35 cases have been recorded.
Most of these cases were due to accidents
while handling the virus in the laboratory
(Labuda and Nuttal, 2008). Definitive diagnosis is based upon the isolation and identification of the virus (Reid, 1988; Lobetti,
2007). LIV infection in sheep is exacerbated
by co-infection with Anaplasma phagocytophilum (Reid, 1988).
Experimental and trans-stadial transmission have been reported in Rhipicephalus
(Rh.) appendiculatus and Hyalomma (Hy.)
anatolicum, but there is no evidence that
they are natural vectors or that any other
tick species except I. ricinus play any significant role in the epidemiology of the disease
(Reid, 1988).
Although I. ricinus is the primary vector
and virus reservoir, because of the transstadial but presumably not transovarial
transmission, the vector efficiency of this
species is relatively restricted. Even when
virions are acquired by the feeding larvae,
only a few of the nymphs become infected.
Like the prevalence of TBEV in the I. ricinus
population, the prevalence of LIV in I. ricinus
is also low.
4.3.3
Other viruses related to TBE
Infections of domesticated animals similar to

TBE and LI also occur in other European
countries (Spain, Bulgaria, Greece and
Turkey). Greek goat encephalitis virus
(GGEV), which was isolated from the brain
of a newborn goat with neurological symptoms, is currently classified in the TBEV
group. The vector of GGEV has not yet been
specifically identified but it is considered as
likely to be I. ricinus. A study during 2003
2006 in goat and sheep farming rural areas of
northern Greece suggested the presence of
TBEV in two pools of I. ricinus ticks. Sequence
analysis showed that the virus was GGEV.
These virus-positive ticks were detected in
regions where a high prevalence of TBE
antibodies was present in humans. TBEV is
considered not to be endemic in Greece,
so most probably the seroprevalence of
TBE antibodies in humans is due to crossreactivity to GGEV (Grard et al., 2007). The
Turkish subtype (Turkish sheep encephalitis
virus) is more closely related to LIV and
should be reclassified.
39
4.4 Anaplasmoses
Alphaproteobacteria of the order Rickettsiales
are obligate intracellular organisms with a
wide range of eukaryotic hosts. There are two
well-characterized families: Anaplasmataceae
and
Rickettsiaceae.
In
the
family
Anaplasmataceae, four genera have been identified: Anaplasma, Ehrlichia, Wolbachia and
Neorickettsia. Several species of the genus
Anaplasma pose severe threats to livestock
and human health. The main Anaplasma species responsible for animal infections or zoonosis in Europe and the Mediterranean basin
are listed in Table 4.1. Anaplasmosis was formerly known as gall sickness, a disease of
ruminants caused by intraerythrocytic bacteria. As a result of a taxonomic reorganization
of the order Rickettsiales (Dumler et al., 2001),
some species of the genus Ehrlichia (E. equi,
E. phagocytophila and Ehrlichia spp. causing
human granulocytic ehrlichiosis HGE) were
renamed as A. phagocytophilum in the genus
Anaplasma. This species is the aetiological
agent of human and animal granulocytic anaplasmosis. Similarly, E. bovis and E. platys are
now known as A. bovis and A. platys. These
latter, recently added, species all invade blood
cells other than erythrocytes in their respective mammalian hosts.
Infection of domestic and wild animals
and humans with these organisms may lead
to a clinical disease collectively called anaplasmosis that manifests as a febrile systemic
illness with haematological abnormalities
and lymphadenopathy (Rikihisa, 2006).
4.4.1
Anaplasma phagocytophilum
A. phagocytophilum, formerly known as

E. phagocytophila and E. equi, and the agent of
HGE, is a commonly found bacterium causing tick-borne fever (TBF) in sheep; pasture
fever in cattle and wild ruminants (deer,
bison and wild goats); human granulocytic
anaplasmosis in humans; equine granulocytic anaplasmosis in horses; and canine
granulocytic anaplasmosis in dogs (Strle,
2004; Rymaszewska and Grenda, 2008). It
was first recognized in Scotland in 1932 and
is now identified in most other European
40
Frank Koenen et al.
Table 4.1. Anaplasmoses in Europe and in the Mediterranean basin transmitted by hard ticks.
Anaplasma spp.
Disease
Host range
A. phagocytophilum
Tick-borne fever (TBF)

Pasture fever
Human granulocytic
anaplasmosis
Equine granulocytic
anaplasmosis
Canine granulocytic
anaplasmosis
Sheep
Cattle, wild ruminants
Humans
A. marginale
Bovine anaplasmosis
A. centrale
Bovine anaplasmosis
A. bovis
Bovine mononuclear
or agranulocytic
anaplasmosis
Ovine anaplasmosis
A. ovis
A. platys
Canine infectious cyclic

thrombocytopenia
Tick involved
(in Europe and the
Mediterranean basin)
I. ricinus, Ha. punctata,
I. persulcatus,
I. trianguliceps,
Rh. sanguineus
Horses, llamas,
rodents
Dogs
I. ricinus, Ha. punctata,

I. persulcatus,
I. trianguliceps,
Rh. sanguineus
Ruminants
I. ricinus, I. persulcatus,
Rh. sanguineus,
Rh. bursa, Rh. annulatus
Cattle
I. ricinus, I. persulcatus,
Rh. sanguineus,
Rh. bursa, Rh. annulatus
Cattle, small mammals Hy. excavatum,
Rh. sanguineus,
Rh. turanicus
Goats, sheep, cattle
Rh. bursa,
Rh. sanguineus
Dogs
Rh. sanguineus,
Rh. turanicus
A., Anaplasma; Ha., Haemaphysalis; Hy., Hyalomma; I., Ixodes; Rh., Rhipicephalus.
countries. Apart from domestic ruminants,

free-living ruminants such as feral goats, and
red, fallow and roe deer tested positive for
A. phagocytophilum. Not only a tick-ruminant
cycle, but also a rodent-tick cycle, is believed
to maintain the TBF variants. The wood
mouse (Apodemus sylvaticus), yellow-necked
mouse (A. flavicollis), field vole (Microtus
agrestis) and bank vole (Myodes glareolus) are
found to be competent reservoirs of infection
(Barandika et al., 2007). The bacterium infects
granulocytic leucocytes (neutrophils, eosinophils and basophils), monocytes and tissue
macrophages.
TBF in sheep and pasture fever in cattle
are characterized by fever, neutropenia, lymphopenia, thrombocytopenia and general
immunosuppression (Woldehiwet, 2006).
Human granulocytic anaplasmosis is a multisystemic disease that occurs more in adults
than in children, especially in persons above
the age of 60 years. The disease is characterized by acute fever, headache, myalgia,
nausea and lethargy, similar to symptoms of

the common flu. In particular, immunocompromised patients are at high risk.
Meningoencephalitis, respiratory distress,
shock and opportunistic infections are occasional complications. In Europe, no fatal
cases have been reported, but the mortality
rate in the USA has been shown to be between
7 and 10% (reviewed by Bakken and Dumler,
2008 and Rymaszewska and Grenda, 2008).
Equine granulocytic anaplasmosis occurs in
horses as their natural host but also in llamas
and rodents. It is generally a benign disease
in these animals, yet fulminating cases have
been described. Mortality is low, and the disease is always acute, never chronic.
The main vector of A. phagocytophilum is
I. ricinus (Strle, 2004), and the prevalence of
infection varies among regions and with the
development stage of the tick (Stuen, 2007).
For example, in unfed nymphs the infection
rate varied between 0.25 and 25% (Walker
et al., 2001). The survival of the parasite is
believed to be over a year while ticks are

awaiting a new host. Only trans-stadial transmission occurs.
The transmission of A. phagocytophilum
has also been associated with other tick
species, such as Ha. punctata, in areas of the
UK where I. ricinus was not present
(MacLeod, 1936). It has also been linked
with I. persulcatus, I. trianguliceps and Rh.
sanguineus (Alekseev et al., 1998; Ogden
et al., 1998; Alberti et al., 2005). The role of
the latter species as a vector is not yet determined, as only one Rh. sanguineus was
found to be positive; this tick was removed
from a dog showing clinical signs of tickborne disease (TBD).
4.4.2 Anaplasma marginale
Bovine anaplasmosis, caused by A. marginale,
was formerly known as gall sickness, and is a
disease that affects domestic and wild ruminants (water buffalo, bison, African antelopes
and mule deer). The disease is characterized
by fever, anaemia, weight loss, reduction of
milk production and in pregnant females,
abortion; it may lead to death (Rymaszewska
and Grenda, 2008; Kocan et al., 2010). A. marginale is present in tropical and subtropical
regions although this bacterial species is frequently detected in Europe (Sicily, Hungary
and Spain) (de la Fuente et al., 2005; Naranjo
et al., 2006; Hornok et al., 2007b; Torina et al.,
2007, 2008). It is an obligate intracellular
species invading erythrocytes mostly in ruminants, both domestic and wild, i.e. calves,
water buffalo, bison, African antelopes and
mule deer.
In cattle, the disease in cattle causes considerable losses to dairy and beef industries
worldwide. Trans-stadial transmission of the
bacteria is effected by ticks of approximately
20 species; in Europe this mainly involves
I. ricinus, I. persulcatus, Rh. sanguineus,
Rh. bursa and Rh. (Boophilus) annulatus
(Kocan et al., 2004). Calves under the age of
6 months have innate resistance and will not
develop clinical anaplasmosis, no matter the
immune status of the mothers. Thereafter,
the risk for serious diseases increases with
age, unless sufficient contact in the first
41
months of life allowed for the development

of immunity. Hence, cattle reared in endemic
regions develop a naturally acquired immunity, quite often without passing through a
stage of clinical disease, as endemic stability
means that all calves need to come into contact with the disease, reservoirs and stable
vector populations.
Wild ruminants (antelopes, buffalo, deer,
eland) can function as reservoirs of A. marginale and the infection can be maintained in
game reserves (deer-to-deer transmission)
without bovine intervention being necessary
(Potgieter and Stoltsz, 2004).
It appears that A. marginale is often
introduced into a herd by ticks, but
subsequently mechanical transmission
(transmission by insects or by veterinary
interventions) may become more important. Mechanical transmission occurs via
the contaminated mouthparts of biting flies,
but can only be achieved within a few minutes after the initial bite, although the pathogen can remain viable and infective in
arthropods for several days after ingestion
(Ewing, 1981; Hornok et al., 2008). Horseflies
(Tabanus spp.) and stable flies (Stomoxys
spp.), and to a lesser extent mosquitoes
(Psorophora spp.), transmit A. marginale and
also A. centrale.
4.4.3
Anaplasma centrale
A. centrale is considered as a separate species

or subspecies of A. marginale, and is also an
intraerythrocytic tick-borne pathogen that
causes mild infections in cattle. A crossimmunity between the two bacteria exists,
and because of its mild virulence this naturally attenuated strain has been used for more
than 100 years in live-blood vaccines to protect cattle from the more virulent A. marginale
(Potgieter and Stoltsz, 2004). These vaccines
are mainly used in Africa, Australia, Latin
America and Israel (Rymaszewska and
Grenda, 2008).
Not much is known about the epidemiology of A. centrale; only few strains have been
characterized. In Europe, the pathogen has
been detected mainly in Italy (George et al.,
2001; Carelli et al., 2008).
42
Frank Koenen et al.
The first case of bovine anaplasmosis

caused by A. centrale in Europe was reported
in 2008 in Italy (Carelli et al., 2008). The case
involved a naturally occurring infection in a
dairy cow, and the clinical signs were typical
of acute anaplasmosis caused by A. marginale.
Molecular analysis of the Italian strains linked
these to the African A. centrale strains (Carelli
et al., 2008).
Recently, a complete genome sequence of
A. centrale was compared with that of virulent
A. marginale (Herndon et al., 2010) with the
aim of identifying possible outer membrane
protein candidates for the development of a
safer inactivated vaccine.
4.4.4
Anaplasma bovis
A. bovis is the aetiological agent of bovine

mononuclear or agranulocytic anaplasmosis,
a disease occurring mainly in cattle and small
mammals (Goethert and Telford, 2003). Goats
appear to be resistant. A. bovis infects the
monocytes of the peripheral blood and the
macrophages of the reticuloendothelial system. Infection may occur with limited or no
clinical signs. The disease is characterized by
weakness, weight loss, fever, enlargement of
the prescapular lymph nodes, paleness of the
mucous membranes and mucous nasal secretion (Uilenberg, 1997). This disease has been
reported in Italy (Georges et al., 2001) and
Israel (Harrus et al., 2011), but is most commonly present in South America, West, Central
and southern Africa, and the Indian subcontinent. The transmission of the disease is transstadial by known vectors: Amblyomma (Am.)
variegatum, Rh. appendiculatus and Hy. excavatum (Coetzer and Tustin, 2004). The pathogen
has recently been detected in Israel in unfed
Rh. sanguineus and Rh. turanicus adults
collected from vegetation (Harrus et al., 2011).
4.4.5 Anaplasma ovis

A. ovis mainly infects wild ruminants and
small ruminants like sheep and goats, and is
prevalent worldwide (Rymaszewska and
Grenda, 2008). In Europe, it has been detected
in Italy, Hungary and Turkey (de la Fuente

et al., 2002, 2005; Christova et al., 2003; Hornok
et al., 2007b; Aktas et al., 2009). This bacterium also infects erythrocytes, but in general
anaplasmosis due to A. ovis in small ruminants is a benign infection with low morbidity and mortality. Goats are normally more
susceptible than sheep or cattle. The biological vector of A. ovis in the Mediterranean
basin is Rh. bursa (Friedhoff, 1997), but possibly also Rh. sanguineus. A. ovis was identified molecularly in the salivary glands of two
Rh. sanguineus ticks collected from sheep in
Turkey (Aktas et al., 2009), although the
authors also suggest that D. marginatus could
play a role in the transmission because of
the high prevalence of ticks found in the
affected flock. In contrast to A. marginale,
vaccination with A. centrale does not protect
against A. ovis.
4.4.6
Anaplasma platys
Canine anaplasmosis or canine infectious

cyclic thrombocytopenia (CICT) is caused by
A. platys. The bacterium multiplies in platelets, but infected dogs may remain asymptomatic (Harvey et al., 1978). Clinical signs are
usually not observed, apart from occasional
haemorrhages after trauma or surgery. The
infection has been detected worldwide, and
in Europe cases have been reported in Spain,
France, Greece and Italy (Sainz et al., 1999;
Sparagano et al., 2003; Mylonakis et al., 2004;
Alberti and Sparagano, 2006; Torina et al.,
2008; Yabsley et al., 2008). Serological tests
may cross-react with other Anaplasma and in
vivo tests may be inaccurate because of low
bacteraemias (de la Fuente et al., 2006).
Cases of importation of both the infectious agent and the vector have been reported
in dogs visiting the Mediterranean region
(Heyman et al., 2007; Nijhof et al., 2007). The
implicated vector seems to be Rh. sanguineus
(Inokuma et al., 2000; Sanogo et al., 2003;
Sparagano et al., 2003), a cosmopolitan tick
species that also transmits E. canis, although
ticks of Rh. sanguineus fed on experimentally
infected dogs were not able to infect naive
dogs in the adult stage (Simpson et al., 1991).
Similarly, as mentioned for A. bovis, unfed Rh.

turanicus adults collected from the vegetation
also have been shown to harbour A. platys
(Harrus et al., 2011).
4.5
Ehrlichioses
4.5.1 Ehrlichia canis

E. canis is a tick-transmitted obligate intracellular Gram-negative bacterium; in dogs it
infects monocytes and causes classical canine
monocytic ehrlichiosis. It resides as a colony
within a membrane-lined intracellular vacuole or morula. Canine monocytic ehrlichiosis
is also known by other names, such as tracker
dog disease, tropical canine pancytopenia,
canine haemorrhagic fever and canine
typhus. The disease has been known since
1935 as a disease of dogs and other canids
(jackal, wolf, fox, coyote) (Donatein and
Lestoquard, 1935), but its importance was
not seriously considered until 1968 when an
epizootic occurred in Vietnam among military working dogs.
The bacterium is transmitted transstadially by the kennel tick (or brown dog
tick), Rh. sanguineus, and is widespread in
tropical and temperate areas of the world
(from 50N to 35S). Its distribution has
expanded with the distribution of its vector
and is maintained in nature by persistent
infections of wild and domestic canids
(Groves et al., 1975). In Europe, E. canis is
restricted to the Mediterranean and Balkan
countries (Hornok et al., 2010), although several cases of canine ehrlichiosis have been
detected in more northern regions in dogs
with a history of travelling (Hirsch and
Pantchev, 2008).
The disease has three clinicopathological
phases: acute, subclinical and chronic
(Skotarczak, 2003). German Shepherd dogs
and their crosses are particularly prone to
more severe signs of disease, and infections in
this breed are associated with a poorer prognosis (Raoult and Parola, 2007).
Several cases of human infections with
E. canis have been reported (Maeda et al., 1987;
Sambri et al., 2004; Perez et al., 2006). Clinical
43
signs of canine monocytic ehrlichiosis in

humans are very similar to those of human
monocytic ehrlichiosis, a zoonosis present in
the USA and caused by the closely related
E. chaffeensis (Perez et al., 2006).
4.5.2
Ehrlichia ruminantium
E. ruminantium, previously known as

Cowdria ruminantium, causes heartwater or
cowdriosis (other names are black gall
sickness, mad gall sickness, infectious exudative pericarditis or malignant rickettsiosis of ruminants). Heartwater is an
infectious tick-transmitted disease of ruminants, i.e. bovines, sheep, goats and various
wild species. Transmission is related to the
Amblyomma vector. In Africa, at least ten
species are capable of transmission; the most
important are Am. variegatum and Am.
hebraeum, and the adults of both species
parasitize cattle. The most widely distributed Amblyomma species in Africa is Am.
variegatum, and this even spreads outside
the continent. Am. hebraeum is the most
important vector in southern Africa.
Heartwater occurs in sub-Saharan Africa
and in several African islands, the islands in
the Indian Ocean and several Caribbean
islands, i.e. Guadeloupe and Antigua. In continental America, it has not been observed, in
spite of the presence of potential vectors
(Coetzer and Tustin, 2004). The control of Am.
variegatum in the Caribbean has been a daunting task. Although the ticks had been eradicated for a number of years, the islands
became infested again after the import of
immature stages on migrating cattle egrets
(Corn et al., 1993).
Am. variegatum sporadically occurs in the
Mediterranean basin, most probably imported
on migratory birds (Papadopoulos et al., 1996).
According to a predictive GIS (geographical
information system) model using temperature and land use, the survival of Am. variegatum and Am. hebraeum in this region would
be best suited to Sardinia, Sicily and the
south-western part of the Italian peninsula
(Pascucci et al., 2007). This poses a possible
risk for the introduction of E. ruminantium
into the Mediterranean region.
44
Frank Koenen et al.
4.6
Rickettsioses
Rickettsial diseases are among the oldest

known arthropod-borne diseases affecting
human health throughout the ages. These
diseases exist primarily in endemic and
enzootic foci and occasionally give rise to
sporadic or seasonal outbreaks, causing
illness and death worldwide (Maxey, 1899).
Evidence suggests that rickettsiae and
Rickettsia-like organisms have evolved and
survived as obligate intracellular bacteria,
cultivating long and well-established relationships with arthropods (lice, mites, fleas
and ticks) and vertebrate hosts (Azad and
Beard, 1998). The vectors of rickettsiae do
include lice, mites and fleas, but most
Rickettsia species are associated with ticks,
which are both their vectors and reservoirs.
Ticks become infected with rickettsiae
mainly by feeding on rickettsaemic host animals, particularly young rodents, and by the
passage of rickettsiae via eggs to the progeny of infected female ticks. Trans-stadial
persistence from larvae to nymphs and to
adults is also important for maintaining the
infection so that transovarial transmission is
successful (Burgdorfer and Brinton, 1975).
Geographic distribution and activity of
infected ticks are important determinants in
the epidemiology of tick-borne rickettsioses.
Humans are accidental hosts that become
infected when ticks containing virulent rickettsiae in their salivary glands take a blood
meal and inject the rickettsiae into the
feeding site. Currently, there are 12 ticktransmitted species of rickettsiae that are
known to cause disease in man. Eight are
present in Europe and the Mediterranean
basin, and the majority have been described
in the last 25 years. This boom in new or
rediscovery of Rickettsia species in recent
years is undoubtedly related to the development of cell culture systems and the advent
of molecular genetics technology. Likewise,
the detection and identification of longknown pathogens in patients from new and
distinct geographic regions has refuelled
interest in these agents. These emerging
infections often have diverse presentations
that underscore the need for increased
awareness among clinicians.
4.6.1
Rickettsiae as bacteria
The genus Rickettsia is classified in the

Alphaproteobacteria
subdivision,
order
Rickettsiales (which also includes the families
Anaplasmataceae and Holosporaceae), family
Rickettsiaceae (Dumler et al., 2001). This family
includes the genus Rickettsia along with
Orientia (Tamura et al., 1995).
Rickettsiae are coccobacillary Gramnegative, obligate intracellular bacteria that
reside free in the cytosol and occasionally in
the nucleus of host cells (Walker, 2006). This
lifestyle within a highly specialized niche, the
eukaryotic cell, has given rise to unique adaptations such as the reduction of bacterial
metabolism and the exploitation of host
metabolites. The ability to transport substrates
that are present in the host cell cytosol but
rarely available in the extracellular milieu is
likely to have contributed to the evolution of
the small-sized rickettsial genome (1.11
1.12 Mb). Many of the de novo biosynthetic
pathways characteristic of free-living bacteria
are no longer present in rickettsiae (Andersson
and Kurland, 1998).
The rickettsiae are traditionally divided
into two antigenically distinct groups based
on their lipopolysaccharide (LPS): the typhus
group (TG) that includes two species,
R. prowazekii, the agent of epidemic typhus
transmitted by the human body louse
(Pediculus humanus corporis), and R. typhi, the
agent of murine typhus transmitted by the
rat flea; and the spotted fever group (SFG)
that includes the majority of rickettsial species, which are transmitted by different
arthropod vectors.
Different phylogenetic studies using the
sequencing of 16S rDNA (Roux and Raoult,
1995) and diverse rickettsial genes gltA
(Roux et al., 1997), ompA (Fournier et al., 1998),
ompB (Roux and Raoult, 2000), sca2 (Sekeyova
et al., 2001), sca4 (Ngwamidiba et al., 2005) and
sca1 (Ngwamidiba et al., 2006) have shown
distinct groups of Rickettsia species, but some
authors still believe that it is difficult to define
the SFG species. This is because some species
of Rickettsia are so closely related that they
could be considered to be strains of a single
species (Walker, 2007). For example, the
sequence analysis of 16S rDNA for R. conorii,
R. sibirica, R. africae, R. parkeri and many others

has shown less than 0.5% of divergence
between them (Roux and Raoult, 1995).
The diagnosis of rickettsial disease is
most often confirmed by serological tests
because culture, rickettsial DNA detection by
PCR and other methods require specialized
laboratories, and also because serum is usually the most commonly available sample.
Indirect immunofluorescent assay remains
the gold standard serological test and is the
mostly widely used technique for diagnosis
of rickettsial diseases (Brouqui et al., 2004).
However, owing to the presence of shared
protein and LPS antigens among spotted
fever group rickettsiae, the use of serological
methods to distinguish between infections
due to closely related rickettsiae is extremely
difficult. Western blot (WB) and cross adsorption (CA) immunoassays can sometimes
assist in the differentiation of SFG or TG rickettsiae; however, interpretation should be
undertaken cautiously because some strains
are so closely related that they cannot be
distinguished (Teysseire and Raoult, 1992).
45
The advent of molecular methods based on

PCR has though enabled the development of
specific and rapid tools for the detection and
identification of Rickettsia species. It should
be noted that the sensitivity of PCR in skin
biopsy (eschar) is higher than in blood.
Culture of the agent is the ultimate criterion
to confirm the diagnosis and to identify the
species of Rickettsia from the patients blood
(Gouriet et al., 2005; de Sousa et al., 2008b).
This technique can reach a good level of success if the sample is collected and preserved
under appropriate conditions until isolation
procedures begin.
The main Rickettsia species responsible
for animal infections or zoonoses in Europe
and the Mediterranean basin are listed in
Table 4.2.
4.6.2
Rickettsia conorii
R. conorii, the causative agent of

Mediterranean spotted fever (MSF), is the
most frequently isolated rickettsia and has
Table 4.2. Rickettsioses in Europe and in the Mediterranean basin transmitted by hard ticks.
Rickettsia spp.
Disease
R. conorii Malish
Mediterranean spotted fever

(MSF)
Israeli spotted fever (ISF)
R. conorii Israeli spotted fever

strain
R. conorii Astrakhan strain
Astrakhan fever (AF)
R. sibirica mongolitimonae strain Lymphangitis-associated
rickettsiosis (LAR)
R. slovaca
Tick-borne lymphadenopathy
(TIBOLA)/Dermacentor-borne
necrosis erythema
lymphadenopathy (DEBONEL)
R. massiliae
Unnamed
R. aeschlimannii
Unnamed
R. helvetica
Unnamed
R. monacensis
Unnamed
Tick involved (based on

publications review)
Rh. sanguineus, Rh. turanicus
Rh. pumilio, Rh. sanguineus

Hy. anatolicum, Hy. excavatum,
Rh. pusillus
D. marginatus, D. reticulatus,
Ha. inermis, Ha. punctata
Rh. sanguineus, Rh. turanicus

D. reticulatus, Ha. punctata,
Ha. inermis, Hy. detritum (syn.
Hy. scupense), Hy. marginatum,
Rh. bursa, Rh. sanguineus,
Rh. turanicus
D. reticulatus, I. hexagonus,
I. ricinus, I. ventalloi
Ha. punctata, I. ricinus
D., Dermacentor; Ha., Haemaphysalis; Hy., Hyalomma; I., Ixodes; Rh., Rhipicephalus.
46
Frank Koenen et al.
the widest geographical distribution of the

SFG rickettsial species. Rh. sanguineus, commonly known as the brown dog tick, is the
main vector and reservoir for R. conorii
strains in the Mediterranean area, former
USSR, northern Africa and India (Rehacek
and Tarasevich, 1988). In some situations,
mainly in anthropozoonotic and domestic
cycles, dogs are the main host for feeding of
all the stages of Rh. sanguineus (Gilot, 1984).
Dogs are transient reservoirs because they
have a short-lived rickettsaemia after infection, but do not seem to be a reservoir for
R. conorii. One study performed in Portugal
also showed that dogs can present with
febrile illness related to infection with
R. conorii Malish and R. conorii Israeli spotted
fever strains (Alexandre et al., 2011).
MSF was described for the first time by
Conor and Brusch in Tunisia in 1910, and is
widely distributed in the Old World, being
endemic in southern Europe, Africa, the
Middle East, India and Pakistan. Case reports
and isolations of R. conorii from patients in
the Mediterranean area continue to be
described, including recent cases in Croatia
(Sardelic et al., 2003), Turkey (Kuloglu et al.,
2004) and Greece (Psaroulaki et al., 2005a).
Sporadic cases have also occurred in nonendemic countries in northern and central
Europe, such as Belgium (Lambert et al., 1984),
Switzerland (Chamot et al., 1987), Sweden
(Vene, 1989) and Germany (McDonald et al.,
1988), as well as in Japan (Yoshikawa, et al.,
2005), the USA (Anderson et al., 1981) and the
UK (Chai et al., 2008). Most of these cases
were attributed to the introduction of
imported vectors or travellers who acquired
the infection in endemic areas (Walker, 2003;
Menn et al., 2010).
In some countries, the incidence of MSF
is unknown, and most of the knowledge is
based on the human seroprevalence of antibodies to SFG rickettsiae. Inasmuch as the
reporting of cases of MSF is not obligatory in
all endemic countries, it is difficult to compare the incidence of the disease in different
regions. However, it seems that in the last
few decades there has been an increase in
reported MSF cases in Portugal, Italy, Spain,
France and Israel (Walker and Fishbein, 1991);
this is likely to be due to climate changes
influencing tick activity. In Portugal, the

national incidence rate during the period
19892003 was 8.9/105 inhabitants per year.
During that same period the incidence of
MSF in the Bragana and Beja districts of
Portugal reached rates of 56.8/105 inhabitants and 47.4/105 inhabitants, respectively
(de Sousa et al., 2003; de Sousa and Bacellar,
2004). A study by de Sousa et al. (2006a),
showed that the average winter temperatures
in Portugal over the last 10 years have been
the warmest on record and this, coupled with
low rainfall, possibly played a role in doubling the number of confirmed MSF cases
during 20002005.
In Italy, during a 5 year period from 1998
to 2002, a national incidence rate of 1.6/105
inhabitants was reported. However, the same
period of time in Sicily, accounting for 51.4%
of all clinical cases, had an incidence rate of
9.3/105 inhabitants (Ciceroni et al., 2006). In
France, a prospective study in the south of
Corsica showed a higher incidence of 48/105
inhabitants compared with other regions
(Raoult et al., 1985). Otero et al. (1982), estimated that in Israel the annual incidence
of MSF is 6.2/105 inhabitants and that the
highest incidences of the disease occur in
the western coastal area and the southern
Negev Desert.
MSF is seasonal, and in most of European
countries the cases are encountered in late
spring and summer (8188%), peaking in July
and August when immature stages of the tick
predominate. As the larvae and nymphs of
the associated ticks are small and their attachment to the body during feeding is painless,
they are more difficult to detect and so more
likely to transmit MSF organisms (Raoult and
Roux, 1997).
The disease is characterized by a generalized endothelial infection of the microvasculature and the main clinical features are due
to the injury of blood vessels. The histopathological phenomenon of vasculitis can involve
all the organs, not only the skin, and it has
particularly serious manifestations when the
lungs and brain are affected (Valbuena and
Walker, 2009). The incubation period ranges
from 3 to 7 days after the tick bite, but it can
be longer. The onset of MSF is generally
abrupt and the disease is characterized by
fever, a maculopapular rash involving the

entire body, including the palms and soles,
and the presence of an eschar at the site of the
tick bite. Occasionally, the eschar is not found
and is seen rarely in multiples. Recently, clinical signs and symptoms were reported in
a group of 71 Portuguese patients with
confirmed diagnosis either by PCR or isolation of R. conorii Malish. The main clinical
signs were fever (94%), maculopapular rash
(94%), eschar (60%), myalgias (84%), headache (78%), asthenia (95%) and anorexia
(73%). Purpuric or petechial rashes were
present in 6% of patients with severe forms;
this was indicative of a bad prognosis
(de Sousa et al., 2008b).
R. conorii has always been considered to
produce a less severe disease than R. rickettsii. However, severe forms of MSF have
been reported in 6% of patients, and a case
fatality rate of 1.413% was reported for hospitalized patients in France, Israel, Spain,
Algeria (Oran) and Portugal (Ruiz-Beltran
et al., 1985; Walker et al., 1987; Amaro et al., 2003;
de Sousa et al., 2008b). Moreover, in southern
Portugal the fatality rate in 1997 reached
32.3% in hospitalized patients (de Sousa et al.,
2003). The co-morbidity condition of diabetes mellitus was identified as the risk factor
for a fatal outcome. Statistical analysis of a
representative sample of Portuguese patients
with MSF also showed that alcoholism
increases the risk for a fatal outcome. Other
underlying conditions, such as heart failure,
increasing age and glucose-6-phosphate
dehydrogenase (G6PD) deficiency can be
also be implicated in severe illness (Walker,
1990; Regev-Yochay et al., 2000).
It is still unclear whether the more
severe and fatal cases are simply related to
host factors or whether there are differences
in the virulence of the bacteria causing the
disease.
4.6.3
Rickettsia conorii Israeli spotted

fever strain
Israeli spotted fever (ISF) was described for

the first time in 1946 in Israel, and some years
later the aetiological agent was isolated from
Rh. sanguineus ticks and from a patient
47
(Goldwasser et al., 1974). The disease that initially seemed restricted to that area is actually
widespread in the Mediterranean basin, and
has been detected in Rh. sanguineus collected
in Portugal and Sicily (Giammanco et al., 2003;
de Sousa et al., 2007). Several clinical cases
were described in those same countries, and
more recently patients have also been reported
in Tunisia and Libya (Bacellar et al., 1999;
Giammanco et al., 2005; de Sousa et al., 2008b;
Znazen et al., 2011). In general, clinical and
laboratory data are similar to those found in
patients infected with R. conorii strains.
However, studies in Portuguese and Sicilian
patients infected with the R. conorii ISF strain
showed that only 39% and 40% of patients
had eschars; this is less frequent than what is
reported by patients infected with the
R. conorii Malish strain (de Sousa et al., 2005;
Giammanco et al., 2005). In Israel, eschars
have been described in only 4% of cases, and
some clinical series reported the total absence
of an eschar (Gross and Yagupsky, 1987;
Wolach et al., 1989). A prospective study conducted in Portugal during 19942006 identified and compared two groups of patients
infected by Malish and ISF strains, confirmed
either by isolation or DNA detection by PCR
(de Sousa et al., 2008b). Although the comparison of the clinical manifestations of MSF
caused by different strains revealed a tremendous overlap, some differences were found.
Patients infected with ISF had a recognized
tick bite, but significantly fewer reported
presence of an eschar compared with patients
infected by the Malish strain. Also, significantly higher percentages of ISF patients had
nausea, vomiting and increased levels of total
bilirubin, g-glutamyl transferase and alkaline
phosphatase. The most important and statistically significant finding documented in the
study was that the ISF strain was associated
with a higher number of fatal cases. Severe
forms and fatal cases were also described in
patients in Sicily and Tunisia, and also in travellers that visited endemic areas (de Sousa
et al., 2003, 2008b; Giammanco et al., 2005;
Boillat et al., 2008; Chai et al., 2008). The differences in virulence between strains are not yet
understood and comparative analysis of
genomes will probably lead to better
comprehension.
48
Frank Koenen et al.
4.6.4 Rickettsia conorii Astrakhan strain

Astrakhan fever (AF) was first reported in the
1970s in patients living in rural areas of
Astrakhan, a region of Russia located on the
Caspian Sea (Tarasevich and Mediannikov,
2006). The appearance of the clinical cases
coincided with the construction of a petrochemical complex that was releasing enormous quantities of CO2 into the atmosphere.
It seems that the CO2 attracted ticks to this
area and somehow increased the probability
of exposure of the population to the vector
(Tarasevich and Mediannikov, 2006). Later,
the disease was serologically diagnosed, and
the R. conorii Astrakhan strain was isolated
from humans and Rh. pumilio ticks (Tarasevich
et al., 1991; Eremeeva et al., 1994). Since 1983,
more than 2000 cases of the disease have been
registered in a small area of the Astrakhan
region (Tarasevich et al., 1991). Infection
caused by the R. conorii Astrakhan strain
showed similar clinical manifestations to
those of the other strains of R. conorii;
however, eschars have been reported in only
23% of patients (Tarasevich et al., 1991).
R. conorii Astrakhan strain has recently been
isolated from a patient in Chad and from
Rh. sanguineus ticks in Kosovo (Fournier et al.,
2003a,b).
(Raoult et al., 1996). The patient had no travel

history, and disease manifestations similar
to MSF, but the unusual aspect of the case
was its occurrence in March when MSF is
rarely reported. Subsequently, other human
cases were described in France, and the diagnosis was confirmed by rickettsial isolation
and/or PCR detection of the agent in eschars
(Fournier et al., 2005). Clinical cases with isolation or DNA detection of the agent were
also reported in Greece, Portugal and Spain,
and in French travellers who visited
Algeria and Egypt (Fournier et al., 2005;
Psaroulaki et al., 2005b; de Sousa et al., 2006b;
Aguirrebengoa et al., 2008; Scolovschi et al.,
2010). In France, most of the reported cases
caused by the R. sibirica mongolitimonae strain
have occurred in the spring; this is in contrast with the majority of Portuguese cases
that occurred in summer during the MSF
season. The occurrence of these cases in different months could be related to the differences in seasonal activity and population
dynamics of the different vectors. The clinical presentation of R. sibirica mongolitimonae
infection has included fever, a rare or diffuse
maculopapular rash and eschars; 50% of
patients also have a lymphangitis expanding
from the inoculation eschar to the draining
lymph node (de Sousa et al., 2008a). The latter feature has led to the name given to this
disease: lymphangitis-associated rickettsiosis (LAR) (Fournier et al., 2005).
4.6.5 Rickettsia sibirica mongolitimonae

strain
R. sibirica mongolitimonae strain initially
named strain HA-91 was originally isolated from a Hy. asiaticum tick collected in
the Alashian region of Inner Mongolia in
1991 (Yu et al., 1993). In Europe and the
Mediterranean basin, the R. sibirica mongolitimonae strain was detected in Hy. anatolicum
from Greece, Hyalomma sp. from Israel and
Rh. pusillus from Portugal and Spain
(Psaroulaki et al., 2005b; de Sousa et al.,
2006a; Toledo et al., 2009b). In 1996, the first
human case of infection caused by this
Rickettsia strain was described in southern
France. The new strain was isolated from the
blood and skin of a patient admitted in
March to the Hospital La Timone in Marseille
4.6.6
Rickettsia slovaca
R. slovaca was first isolated in 1968 from a

Dermacentor marginatus tick in Slovakia
(Rehacek, 1984). In subsequent years, suspected cases of infection caused by R. slovaca
were reported in patients from Hungary and
Slovakia (Mittermayer et al., 1980; Rehacek,
1984; Raoult et al., 2002). However, the first
proven case of R. slovaca infection was
reported only in 1997 in a French patient who
presented with an eschar on the scalp and
enlarged cervical lymph nodes after receiving
a bite from a Dermacentor tick (Raoult et al.,
1997). The lymphadenopathy present in most
of the patients that have been seen infected
with R. slovaca has led to this clinical syndrome being named tick-borne lymphadenopathy (TIBOLA). Spanish researchers
have also coined the name Dermacentorborne necrosis erythema lymphadenopathy
(DEBONEL) (Lakos, 1997; Raoult et al., 1997).
R. slovaca has been identified in D. marginatus
and D. reticulatus in most European countries.
The prevalence rates of R. slovaca infection
that have been found in Dermacentor ticks in
Europe range from 21% in Hungary to higher
rates in Spain (40.6%), Portugal (41.5%) and
Switzerland (45.4%) (Beati et al., 1994; Bacellar
et al., 1995; Lakos and Raoult, 1999; Oteo et al.,
2006; Parola et al., 2009; Milhano et al., 2010).
The number of R. slovaca infections in
Europe is still under-evaluated; the majority
of patients have been reported from France,
Hungary, Spain, Slovakia and Italy (Lakos,
2002; Raoult et al., 2002; Ibarra et al., 2006;
Selmi et al., 2008; Parola et al., 2009). The
epidemiological and clinical findings on
R. slovaca infections among patients in France,
Hungary, Spain and Italy showed that the
infection occurred mainly during the colder
months of the year, mostly from October to
April, in accordance with the density and
activity of Dermacentor ticks. It seems that
children and women had a higher risk of
infection, and patients were more frequently
bitten on the scalp (68100%). Fever is present
in 1267% of the patients; rash is rare
(1423%) compared with other rickettsioses.
The enlargement of lymph nodes has been
reported in almost all of the patients
(74100%) (Lakos, 2002; Raoult et al., 2002;
Ibarra et al., 2006; Selmi et al., 2008, Parola
et al., 2009). No complications were observed,
but 2152% of patients developed a localized
alopecia at the site of the tick bite and around
37% suffered from persistent asthenia. Only
50% of the patients develop detectable antibodies, which may reflect the fact that this
disease is a localized infection.
R. raoultii (formerly genotypes RpA4,
DnS14, DnS28), the other aetiological agent of
TIBOLA/DEBONEL, was described for the
first time in Rh. pumilio collected in the
Astrakhan region and in D. nutallii in Siberia.
Later, this Rickettsia was detected in
Dermacentor ticks in other European countries
(Rydkina et al., 1999). The pathogenicity of
49
the species has been suggested by the amplification of its DNA from the blood and skin
biopsy samples of patients with a clinical
picture of R. slovaca-like infection. This
Rickettsia species was also found in
Dermacentor removed from Spanish and
French patients with cases of TIBOLA/
DEBONEL (Ibarra et al., 2006; Parola et al.,
2009). Nevertheless, R. slovaca remains the
main aetiological agent responsible for the
majority of TIBOLA/DEBONEL cases.
4.6.7
Rickettsia massiliae
R. massiliae was isolated in 1992 from ticks in

France near Marseille (Beati and Raoult,
1993). Subsequently, this rickettsia has been
detected by molecular methods and isolated
in several countries in Europe (Scolovschi
et al., 2010). R. massiliae has been found mainly
in ticks from the Rhipicephalus genus:
Rh. sanguineus and Rh. turanicus. The first human
case was described in a Sicilian patient who
was admitted at Palermo hospital in 1985.
However, it was not until 2005 that the isolate
discovered 20 years before was characterized
and identified as R. massiliae (Vitale et al.,
2006). The patient presented with an eschar
on his right ankle, and a maculopapular rash
involving the palms and soles, revealing similar manifestations to those found with other
rickettsioses. He recovered completely after
receiving tetracycline. Recently, another case
of R. massiliae infection was diagnosed in a
patient from Argentina. The diagnosis was
confirmed by PCR in a Spanish laboratory
(Garca-Garca et al., 2010).
4.6.8
Rickettsia aeschlimannii
R. aeschlimannii was first characterized in 1997

from Hy. marginatum ticks from Morocco
(Beati et al., 1997). Later, this rickettsia was
also described from Hy. marginatum ticks from
Portugal, France, Spain, Croatia, Algeria, Italy
and Egypt (Beati et al., 1997; Bacellar, 1999;
Punda-Polic et al., 2002; Fernndez-Soto et al.,
2003). However, until now the two described
human cases caused by R. aeschlimannii were
50
Frank Koenen et al.
only diagnosed in Africa. The first case was in

a French traveller who had visited Morocco,
and the second case was detected in a South
African patient who was bitten by a Rh. appendiculatus tick (Pretorius and Birtles, 2002).
Symptoms exhibited by the patients were
similar to those of MSF, and infection by
R. aeschlimannii was confirmed by PCR amplification of rickettsial DNA from serum and
skin biopsy (Pretorius and Birtles, 2002;
Raoult et al., 2002). Recent studies in Algeria,
Corsica, Greece, Spain, Germany and the
European part of Russia also revealed that
other tick species can harbour R. aeschlimannii, such as: Hy. anatolicum, Hy. detritum (syn.
Hy. scupense), Hy. rufipes, D. reticulatus, I. ricinus, Ha. punctata, Ha. inermis, Rh. bursa, Rh.
sanguineus and Rh. turanicus (Fernndez-Soto
et al., 2003, Matsumoto et al., 2004; Psaroulaki
et al., 2006; Mokrani et al., 2008; Bitam et al.,
2009; Shypnov et al., 2009).
4.6.9 Rickettsia helvetica
R. helvetica was detected for the first time in
Swiss I. ricinus in 1979, and confirmed as a
new member of the SFG Rickettsia in 1993
(Beati et al., 1993). R. helvetica has been
detected and isolated from I. ricinus in many
European countries, including Austria,
Bulgaria, Denmark, Eastern Ukraine, France,
Germany, Hungary, Italy, Moldova, the
Netherlands, Poland, Portugal, Slovenia,
Spain, Sweden and the UK (Bacellar et al.
1995; Parola et al., 1998; Nilsson et al., 1999;
Beninati et al., 2002; Christova et al., 2003;
Prosenc et al., 2003; Fernndez-Soto et al.,
2004; Srter-Lancz et al., 2005; Nijhof
et al., 2007; Skarphedinsson et al., 2007;
Blaschitz et al., 2008; Chmielewski et al., 2009;
Movila et al., 2009; Pluta et al., 2010; TijsseKlassen et al., 2011). In Portugal and Spain,
the organism has been also detected in
I. ventalloi parasitizing birds (Santos-Silva
et al. 2006; Movila et al., 2011). The prevalence
of R. helvetica in ticks from different countries
has been found to range from 2.8% in Poland
to 91.4% in the south of Germany (Chmielewski
et al., 2009; Silaghi et al., 2011). Detection of
co-infections with R. helvetica and Borrelia
burgdorferi s.l. in the same vector (I. ricinus)
has gained attention for possible exacerbation

of the illness arising from R. helvetica infection
or Lyme borreliosis (Fernndez- Soto et al.,
2004; Milhano et al., 2010).
R. helvetica infection has been progressively becoming a clinical entity of its own.
It was reported for the first time in 1999 in
Sweden in two patients with fatal perimyocarditis, and it was recently reported in two
Swedish patients with meningitis and septicaemia (Nilsson et al., 1999). Serological
associations with R. helvetica infections have
been reported in patients in Europe (France,
Italy, Switzerland) and Asia (Thailand), but
additional evaluation and isolation of the
bacterium from clinical samples are needed
to confirm the pathogenicity of R. helvetica
(Baumann et al., 2003; Fournier et al. 2004;
Ciceroni et al., 2006; Nilsson et al. 2010).
In 2002, an association of R. helvetica with sarcoidosis was proposed; however, the validity
of this association has been questioned and
later serological studies did not reveal the
presence of anti-rickettsial antibodies in a
group of Scandinavian sarcoidosis patients
(Nilsson et al., 2002; Planck et al., 2004).
4.6.10
Rickettsia monacensis
R. monacensis was isolated and characterized

from I. ricinus for the first time in Germany.
Since then, most European countries have
reported the presence of this agent, essentially based on molecular detection (Simser
et al., 2002). More recently, this Rickettsia species has also been isolated from I. ricinus in
Portugal, and it was shown that R. monacensis
was easily propagated and isolated in Vero
cell lines at 28C in conditions different from
that for the isolate from Germany (Milhano
et al., 2010). R. monacensis was recently associated with febrile disease in humans in northern Spain (Jado et al., 2007).
Table 1.25 in Appendix 1 elaborates on
the geographical distribution of Rickettsia
spp. and their reported diseases in Europe
and in the Mediterranean basin. This table
was generated independently of the main
systematic literature review that is described
in the introduction.
4.7
Lyme Borrelioses
Lyme borreliosis (LB) or Lyme disease is the

most common tick-borne disease of humans
in the northern hemisphere. It is a complex of
several different zoonotic infections of which
the aetiological agents are transmitted by
hard ticks. At least 18 species or genospecies
of spirochaetes in the B. burgdorferi s.l. complex have so far been described. Several are
pathogenic to humans and domestic animals.
They include B. burgdorferi, which is predominant in North America but also present in
Eurasia, and B. afzelii and B. garinii, which are
predominant in Eurasia. They are transmitted by tick species of the genus Ixodes, mainly
I. ricinus in Europe, I. persulcatus in Eurasia,
I. pacificus in the western USA and I. scapularis in the eastern USA. In Europe, at least
another four Borrelia species, i.e. B. bavariensis, B. valaisiana, B. spielmanii and B. lusitaniae
sometimes infect humans and may cause
human LB (Richter et al., 2004; Piesman and
Gern, 2008; Rudenko et al., 2011). In Europe,
three tick species are considered to be vectors
of LB spirochaetes, i.e. Ixodes ricinus, I. hexagonus and I. uriae (Piesman and Gern, 2008).
Although the level of infection in the adult
ticks in European populations of I. ricinus is
higher (mean 17.4%; range 358%) than in
the nymphs (mean 10.8%; range 243%), the
nymphs are usually more important than
the female adult ticks for transmission of
the pathogens to humans (Hublek and
Halouzka, 1998). Larvae are rarely infected
(mean 1.9%; range 011%; Hublek and
Halouzka, 1998). In I. persulcatus, however,
the nymphs rarely feed on humans, so in this
case it is the adult female ticks that are responsible for nearly all human infections with LB
spirochaetes.
The enzootic cycle in general involves
Ixodes spp. larvae and nymphs which become
infected when feeding on infective wild
bacteraemic mammals, particularly insectivores (shrews, hedgehogs), rodents (mice,
voles, rats and squirrels) or hares. Certain
bird species also serve as vertebrate
reservoirs to the spirochaetes. Co-feeding
transmission has been demonstrated to occur
when sheep serve as a Borrelia reservoir
(Ogden et al., 1997).
51
It is important to distinguish between

vertebrate hosts for the ticks and vertebrate
reservoirs of the spirochaetes. Cervids appear
to be refractory to the infection and usually
do not serve as Borrelia reservoirs, but are
extremely important hosts to I. ricinus females
(Jaenson and Tlleklint, 1992). Many species
of Borrelia may circulate in the same ecosystem, with the result that a single tick can be
infected with two or more species of Borrelia
and with the TBE virus and other species of
human-pathogenic bacteria. Throughout
Europe, 13% of Borrelia infections in I. ricinus
are mixed infections (Rauter and Hartung,
2005; Piesman and Gern, 2008). Multiple
infection of a tick may occur because the host
on which the tick was feeding had a multiple
infection or because the tick had fed two or
more times on hosts infected with different
Borrelia spp. In Europe, B. garinii and B. valaisiana are predominant in the mixed infections,
followed by mixed B. garinii/B. afzelii infection (Piesman and Gern, 2008). B. afzelii is
mainly associated with rodents, while some
serotypes of B. garinii and all serotypes of
B. valaisiana are associated with birds. B. lusitaniae is associated with lizards in the
Mediterranean countries, and often infects
vector ticks more frequently than do the other
genospecies in the complex (Richter and
Matuschka, 2006).
LB is prevalent in most parts of Europe,
but its prevalence is lower in southern Europe,
such as in Portugal and Italy than in the
former USSR to Japan, Mongolia and northwestern China (Rauter and Hartung, 2005).
The infection also occurs in some specific
locations in North Africa. In North America,
nearly all human LB cases are confined to the
north-eastern USA, but the infection also
occurs in other parts, including California,
but with a lower prevalence than the northeastern part of the country. LB is becoming
increasingly prevalent in southern Canada
and has also been reported from South
America, including Mexico and Brazil.
Climate change (not synonymous with global warming in the authors opinion, as
climate change is a broad concept that
includes all potential intentional and nonintentional changes in weather conditions)
and an increasing abundance of deer could
52
Frank Koenen et al.
be associated with the spread of LB in

Northern Europe (Gray et al., 2009).
Human LB infections sometimes cause
clinical disease, which can range from a relatively short influenza-like illness often accompanied by excruciating (nocturnal) pain, to a
severe syndrome with neurological involvement, including meningitis, chronic severe
arthritis and/or myocarditis. For instance, it
has been estimated that in Sweden alone
about 10,000 people annually contract the
infection, resulting in clinical disease
(Berglund, 2004). Among domesticated animals, clinical symptoms associated with LB
have been reported in dogs, cattle and horses.
Antibody titres against B. burgdorferi s.l. in
dogs, cats and livestock can be high, but it is
often difficult to establish a cause-and-effect
relationship between exposure to the spirochaetes and clinical signs.
4.8
Recurrent (Relapsing) Fever
Relapsing fever is an infection caused by several spirochaetes of the genus Borrelia (Cutler,
2006). Relapsing fever borrelioses are characterized by recurrent febrile episodes and spirochaetaemia. Other than the louse-borne
relapsing fever caused by B. recurrentis and
transmitted by the body louse Pediculus
humanus, endemic tick-borne relapsing fever
is a zoonotic disease transmitted worldwide
by soft tick species of the genus Ornithodoros.
Within each region, specific relationships usually exist between the Ornithodoros vector species, Borrelia species and their distribution
areas. Reservoir hosts are usually wild
rodents. Ornithodoros ticks are included in the
family Argasidae. They live close to their host,
although the time spent on the host is relatively short. After each blood meal they are
found in their habitats, typically in cracks and
crevices of rodent burrows, but also in human
shelters or just below the soil surface. Ticks
become infected during a blood meal on a
vertebrate with spirochaetaemia. Spirochaetes
then invade all tissues of the tick, including
ovaries (responsible for transmission between
generations), salivary glands and excretory
organs. Vertebrates and humans become
infected during a blood meal through

contamination of the feeding site by salivary
and/or coxal secretions of the tick (Parola and
Raoult, 2001).
B. hispanica is found in Spain, Portugal,
Cyprus, Greece and North Africa. It has been
isolated in O. erraticus, a tick commonly found
in south-western Europe. This tick species
usually lives in the burrows of wild rodents,
its natural host. In Spain and Portugal; however, it has adapted to bite domestic pigs that
are kept in continuous grazing and sometimes overnight in large burrows or inside old
buildings, and this tick species has adapted to
live in these habitats (Estrada-Pea and
Jongejan, 1999). Humans may be bitten, and
hence relapsing fever was sporadically
reported in countries such as Spain during
the 20th century, probably with an underestimated incidence (Snchez-Yebra et al., 1997).
The disease caused by B. hispanica is one of
the less severe in the relapsing fever group,
and presents with neurological signs in less
than 5% of cases (Cadavid and Barbour, 1998).
In 1996, a new Borrelia species was isolated in
southern Spain from three patients with
relapsing fever and from O. erraticus ticks
found in nearby areas (Anda et al., 1996). The
reservoir of this bacterium is still unknown
and further records of the pathogen are unavailable. Although this new Borrelia species
has not yet been cultured, molecular analyses
have shown that it is closely related to B. hispanica, B. duttoni (an African species not
present in Europe) and B. crocidurae.
On the borders of Europe, several other
relapsing borrelioses are present. B. persica,
the agent of Persian relapsing fever, is found
in Israel, Syria, Egypt, Iran and Central Asia.
It is transmitted by O. tholozani (Rodhain,
1998). This tick commonly lives in localities
where livestock are housed, for example manmade shelters, caves and rocky overhangs
(Estrada-Pea and Jongejan, 1999). The disease is sometimes severe (Cadavid and
Barbour, 1998). B. caucasica, present in the
Caucasus and Iraq, is transmitted by O. verrucosus, another argasid parasite of rodents.
B. latyschevii is transmitted by O. tartakovskyi
in Central Asia, the former USSR and Iran
(Estrada-Pea and Jongejan, 1999; Rebaudet
and Parola, 2006).
4.9
Piroplasmoses
4.9.1
Babesioses
Babesioses are caused by naturally ticktransmitted and generally host-specific intraerythrocytic protozoan parasites of the genus
Babesia
(phylum Apicomplexa,
order
Piroplasmida). Babesiae, which are the second most common blood-borne parasites of
mammals after the trypanosomes, are capable of infecting a wide variety of vertebrate
mammalian species, including humans. More
than 100 species have been identified, and
these are traditionally divided on the basis of
their morphology into the small and large
babesiae (Telford et al., 1993; Homer et al.,
2000). Molecular analysis suggests that the
host range of many Babesia species is less
restricted than previously believed and that
still-unrecognized species may cause zoonotic
infections in a variety of animals and humans
(Gray and Weiss, 2008).
To date, only ixodid ticks have been identified as vectors for Babesia species. Some
Babesia species can infect more than one genus
of ticks; others can infect only ticks from the
genus Ixodes. Several tick vectors can carry
more than one Babesia species. The vectors
become infected when ingesting the infected
blood cells from a vertebrate reservoir that is
competent in maintaining the Babesia organisms in an infectious state. Certain species of
Babesia, such as B. divergens and B. canis, are
transmitted transovarially to the next generation because they invade the female ticks
ovaries. These species may persist in several
generations of ticks, even without new infections. Some other Babesia species (e.g.
B. microti) are only transmitted trans-stadially.
All species of Babesia are naturally transmitted
by the bite of infected female ticks. It has been
demonstrated that male ticks may transmit
Babesia species, although the epidemiological
importance of male ticks in transmission has
yet to be established.
Infection is initiated by inoculation of the
sporozoites with the saliva of the vector tick
into the bloodstream of the host. Transmission
only occurs a few days after the tick has
attached, because maturation of the sporozoites in the salivary glands of the vector is
53
stimulated by feeding. Babesia species directly

invade red blood cells, where their asexual
multiplication most often results in two,
sometimes four, daughter cells; these then
leave the host cell and each enters another red
cell (Homer et al., 2000; Uilenberg, 2006).
Animals and humans can also acquire the
infection through the transfusion of contaminated blood products.
The two major factors involved in the
pathogenesis of babesiosis are the release of
pharmacologically active agents and intravascular haemolysis. The relative importance
of each varies with the species of Babesia. The
clinical features of babesiosis vary substantially from asymptomatic to life threatening,
depending on the condition of the host and
the parasite involved. During the acute babesial infection, the host may become severely
ill as a result of host-mediated immunopathological mechanisms and erythrocyte lysis.
Typically, the infected host can suffer high
fever, anaemia and hyperbilirubinuria, possibly followed by alterations in the kidneys and
other organs. All mammalian hosts examined
have been able to develop immunity to Babesia
species in which both humoral and cellular
factors are involved. In endemic areas, all or
almost all individuals of the host population
are infected when they are young, with no or
minimal clinical disease. The introduction of
susceptible animals into endemic regions
could lead to the recrudescence of babesiosis
(Telford et al., 1993; Homer et al., 2000;
Uilenberg, 2001; Hunfeld et al., 2008).
Babesioses are well-recognized diseases
of veterinary importance in Europe and North
Africa. One of the most important and widespread Babesia species affecting cattle in temperate Europe is B. divergens, and this species
probably occurs wherever the vector I. ricinus
is present, which includes North Africa
(Bouattour and Darghouth, 1996). B. major,
another species in Europe, gives a much
milder infection that is transmitted by Ha.
punctata in western Europe. A low or nonpathogenic bovine Babesia species, B. occultans, was detected for the first time in unfed
Hy. marginatum ticks collected in three bioclimatic regions of Tunisia. It was supposed that
this species may have a wide distribution in
the Mediterranean region, and not only in
54
Frank Koenen et al.
sub-Saharan Africa as previously described

(Ros-Garca et al., 2011). The taxonomy and
even the geographical distribution of babesial
parasites of small ruminants are not quite settled. The main species are B. motasi and B. ovis,
which are transmitted by Ha. punctata and
Rh. bursa, respectively. B. motasi in northwestern Europe produces a clinically milder
disease than it does in the Mediterranean basin,
and there are serological differences as well
(Uilenberg, 2006). Babesiosis in the horse is
mentioned with theileriosis (see Section 4.9.2).
On the basis of differences in vector
specificity, geographical distribution, pathogenicity, antigenic properties and molecular
investigations, the species B. canis of dogs has
been subdivided into three subspecies,
namely B. c. canis, B. c. vogeli and B. c. rossi
(Kjemtrup et al., 2000). These subspecies are
currently considered to be separate species.
Babesia c. canis, transmitted by D. reticulatus
ticks, is the most common canine Babesia
subspecies in temperate regions of Europe
(Cacci et al., 2002; Irwin, 2009). Babesia
c. vogeli, transmitted by Rh. sanguineus, has
also been reported in continental Europe
(Cacci et al., 2002; Duh et al., 2004; Cardoso
et al., 2008). This subspecies is considered to
be a mildly virulent subspecies, and commonly induces moderate clinical signs in
dogs. Besides these large parasites, small
babesiae such as B. gibsoni, transmitted by the
brown dog tick, Rh. sanguineus, and
the B. microti-like or Spanish isolate, with
the proposed name of Theileria annae, have
been reported from some European countries
(Zahler et al., 2000; Camacho, 2006; Beck et al.,
2009). Several studies have proved that B. gibsoni infection can be transmitted from dog to
dog via bite wounds, saliva or ingested blood
independently of the limitations of vector tick
infestation (Jefferies et al., 2007).
Babesioses are an emerging zoonotic
problem caused by several species of protozoans in the genus Babesia (Homer et al., 2000;
Kjemtrup and Conrad, 2000; Herwaldt et al.,
2003; Hunfeld et al., 2008).The first confirmed
case of human babesiosis was diagnosed in a
splenectomized Yugoslavian cattle farmer
who died of a fatal B. divergens infection in
1956 (Skrabalo and Deanovic, 1957). To date,
more than 60 cases of human babesiosis have
been reported from Europe. At least 70% of

the cases in Europe are associated with the
cattle piroplasm, B. divergens (Genchi, 2007).
In recent years, molecular studies have confirmed the responsibility of B. divergens in two
cases of human babesiosis in the Canary
Islands (Olmeda et al., 1997) and in Portugal
(Centeno-Lima et al., 2003). A new European
B. divergens-like organism (EU1), named
B. venatorum, has been described from deer
and from I. ricinus (Duh et al., 2005; Gray,
2006; Bonnet et al., 2007). This parasite species
was involved in the first documented cases of
human babesiosis in asplenic men in Italy,
Austria and Germany (Herwaldt et al., 2003;
Hselbarth et al., 2007). A human infection in
a splenectomized patient, caused by a strain
named Babesia EU3 that has high homology
with Babesia EU1, was reported from Germany
(Hselbarth et al., 2007). B. microti infection
has also been reported in Europe (Hildebrandt
et al., 2007). Sero-surveys suggest that a low
percentage of Europeans (< 3.4%) from several countries may be infected with B. microti.
Most infected patients share splenectomy as a
risk factor for acquiring the disease, but the
rising number of HIV-positive individuals
and the increasing population of immunocompromised patients may also serve to boost
the number of human cases. The clinical
course of human babesiosis varies according
to the aetiological agent and ranges from
subclinical infection to a severe disease with
sudden onset. In many individuals, however,
babesiosis is a mild, self-limited disease that
requires only supportive therapy. Splenectomized or elderly patients infected with
B. microti or B. divergens tend to develop
severe and sometimes fatal illnesses (Homer
et al. 2000; Gray, 2006).
4.9.2 Theilerioses
Theileria spp. (phylum Apicomplexa, order
Piroplasmida) are tick-borne intracellular protozoan haemoparasites causing infection and
often disease of veterinary and economic
importance in livestock and wild animals in
different regions of the world (Preston, 2001;
Uilenberg, 2001). The genus Theileria differs
from Babesia in that Theileria first penetrates

lymphocytes or macrophages and develops
there, and then enters red blood cells where
the parasites multiply, forming tetrads often in
the shape of a Maltese cross (Uilenberg, 2006).
The clinical signs of theileriosis in animals differ from babesioses in the absence of
haemoglobinuria and the occurrence of a less
severe anaemia. Ticks can only transmit these
haemoparasites trans-stadially. There is no
transovarial transmission because theileriae
do not passage the ovaries and the eggs of
the vectors. The newly hatched larvae of the
ticks are never infected. Nymphs and adults
become infective only if they were infected in
the previous developmental stage. The transmission of parasites takes place by the injection of infected saliva of ticks, but it only
occurs a few days after the tick has attached;
the parasites have first to mature before they
become infective (Mehlhorn and Schein, 1984;
Preston, 2001).
A mild disease of domestic cattle is called
tropical or Mediterranean theileriosis; this is
caused by T. annulata, which is distributed in
many areas of the world, extending from
southern Europe to southern Asia (Brown,
1990). T. annulata is transmitted by a number
of Hyalomma species, which are found in large
numbers in the Mediterranean region, especially in semi-arid areas (Viseras and GarcaFernndez, 1999). Hy. anatolicum anatolicum,
Hy. detritum detritum and Hy. dromedarii are
considered as the main vectors in the field.
Other species, such as Hy. anatolicum excavatum and Hy. marginatum marginatum may also
play a role in the epidemiology of the disease
in the field (Estrada-Pea et al., 2004). T. sergenti/buffeli/orientalis causes a mild or
asymptomatic disease in cattle known as
bovine benign theileriosis (Uilenberg, 1981).
T. ovis and T. lestoquardi (formerly T. hirci) are
recognized as the species that can cause
serious theileriosis in small ruminants,
particularly sheep, where the disease occurs
in the Old World (Schnittger et al., 2000;
Preston, 2001).
Equine piroplasmosis (EP) is a tick-borne
intraerythrocytic protozoal disease of equids
(horses, donkeys, mules and zebras) caused
by B. caballi and Theileria (syn. Babesia) equi.
B. c. canis of dogs has been reported in horses,
55
but no clinical signs attributable to this parasitic species were described (Criado-Fornelio
et al., 2003; Hornok et al., 2007a). This economically important protozoan disease of
horses has been reported in many countries,
thus making this disease a cause of great concern in the global horse industry. For this reason, testing of horses for EP is mandatory for
the international movement of horses, either
for participation in international events or for
export. Only horses seronegative for both
T. equi and B. caballi are qualified for importation to some countries, such as the USA,
Canada, Australia and Japan (Bruning, 1996;
Knowles, 1996). Within Europe, equine theileriosis is more prevalent in Portugal
(Bashiruddin et al., 1999), Spain (Camacho
et al., 2005) and Italy (Moretti et al., 2009). The
disease agent is mainly spread by competent
ticks. To date, up to 12 species of ixodid ticks
belonging to the genera Hyalomma,
Dermacentor and Rhipicephalus have been
identified as vectors of both B. caballi and
T. equi (Bruning, 1996; Massaro et al., 2003).
Transmission of T. equi appears only to occur
trans-stadially (de Waal, 1992). The parasites
are also spread by the transfer of blood from
infected to naive equids through shared needles, improperly shared equipment, and
blood or serum transfusions (de Waal and
Van Heerden, 2004). Transplacental transmission of T. equi from carrier mares to asymptomatic foals was recently confirmed (Allsopp
et al., 2007).
Clinical signs of infection with EP are
not pathognomonic, especially in endemic
areas, and vary from mild to severe. Acute
and subacute cases are the most commonly
observed. The mild form of the disease can
cause equids to appear weak or show lack
of appetite, while more severe cases may
have fever, anaemia, jaundice, swollen
abdomen, haemoglobinuria and bilirubinuria, and sometimes result in death (de Waal
and Van Heerden, 2004). In the chronic
phase, the horse can appear normal. In
some cases of acute or chronic disease, mortality can reach up to 50% (de Waal, 1992).
Infected animals that recover from acute or
primary infection of T. equi remain lifelong
carriers because anti-theilerial drugs suppress but do not eliminate the parasite.
56
Frank Koenen et al.
The detection of apparently healthy carrier

horses therefore remains a worldwide challenge for controlling the spread of the disease. Carrier mares may transmit the
organism to their offspring and this may
result in abortion or neonatal piroplasmosis. Some researchers suggest that foals may
be born as carriers yet remain apparently
healthy as colostral T. equi antibody may act
to suppress parasitaemia in the newborn,
reducing the incidence of clinical neonatal
piroplasmosis (Allsopp et al., 2007).
4.10
Hepatozoonosis
Hepatozoonosis is an arthropod-borne infection of both wild and domestic animals,

including mammals, birds, reptiles and
amphibians. It is caused by about 300 apicomplexan protozoal species from the genus
Hepatozoon (Smith, 1996). The vertebrates are
intermediate hosts, and several blood-sucking
invertebrates, including hard ticks, are definitive hosts in which the sexual reproduction
and sporogony of the protozoans occurs.
Transmission of H. canis to the dog takes
place by ingestion of a tick or parts of ticks
containing Hepatozoon oocysts; this is because
the protozoan is not disseminated within the
tick but remains in the haemocoel. No salivary transfer of these parasites has been
documented. In this respect, Hepatozoon differs from many other tick-borne protozoal
and bacterial pathogens. The ticks become
infected when they ingest infected neutrophils and gametocytes while feeding on
the blood of a parasitaemic dog. Trans-stadial
transmission of H. canis from the nymph to
the adult stage in the tick vector has been
recorded, whereas transovarial transmission
could not be demonstrated (Baneth, 2006;
Baneth et al., 2007). Japanese scientists
(Murata et al., 1993) have reported that vertical transmission of H. canis occurred in puppies born from an infected dam and raised
in a tick-free environment.
In the Old World, canine hepatozoonosis caused by H. canis is of veterinary importance. H. canis has been reported from the
Mediterranean region of Europe (Spain,
Portugal, Italy, Greece and France), Africa,

the Middle East, the Far East and South
America, where its main vector, the brown
dog tick, Rh. sanguineus, is enzootic
(Vincent-Johnson et al., 1997; Baneth et al.,
2003). Recently, Italian scientists reported
that I. ricinus might also be implicated in
parasite transmission, thus explaining the
occurrence of hepatozoonosis in areas considered Rh. sanguineus free (Gabrielli et al.,
2010). However, further studies are needed
to confirm this hypothesis. Hepatozoonosis
is of increasing importance in dogs in
regions that have previously been considered free of the infection. H. canis is regularly introduced by dogs into north-west
Europe after ingestion of infected Rh. sanguineus during visits to endemic regions
(Holland, 2001). The first detection of H.
canis in Slovakia (Majlthov et al., 2007)
and Italy (Gabrielli et al., 2010) has been
reported in naturally infected red fox
(Vulpes vulpes). Prevalence data for dogs are
still restricted to small areas and range from
0.9% in France (Criado-Fornelio et al., 2009)
to 71% in Croatia (Vojta et al., 2009).
According to a sero-survey of H. canis in
Israel, this protozoal infection appears to be
endemic in red fox populations and these
may serve as reservoirs of the parasite for
domestic dogs (Fishman et al., 2004).
When the infected vector is ingested by
the dog, H. canis mainly infects the haemolymphatic tissues and blood-forming organs,
including the bone marrow, lymph nodes
and spleen. The pathogenicity and clinical
manifestations of hepatozoonosis vary
according to the age of the host and the
degree of infection. Canine hepatozoonosis is
typically a mild clinical disease. However,
clinical signs vary from an apparently subclinical infection with a low parasitaemia, to
a life-threatening disease with lethargy, fever,
cachexia and anaemia with a large number of
circulating parasites. The grave and potentially fatal disease occurs mostly in young
animals or in dogs suffering from a concurrent infection or immunosuppressive conditions. H. canis is commonly associated with
co-infection with other diseases, in particular
ehrlichiosis and leishmaniosis in endemic
areas (Baneth, 2006).
4.11
Other Potentially Tick-borne

Infections
The following infections are associated or

suspected to be associated with ticks as one of
the potential but not the only modes of
transmission.
4.11.1 African horse sickness
African horse sickness (AHS) is caused by
African horse sickness virus (AHSV), a member of the genus Orbivirus, subfamily
Sedoreovirinae and the family Reoviridae. The
virion is approximately 70 nm in diameter
(Polson and Deeks, 1963), is non-enveloped
and consists of a double-layered icosahedral
capsid with 32 capsomeres (Pringle and
Wickner, 2000). The double-capsid particles
contain seven structural proteins (VP1VP7),
with the outer capsid made up of VP2 and
VP5, and the inner capsid (the core) made up
of VP3 and VP7 (as major proteins) and VP1,
VP4 and VP6 (minor proteins) (Roy et al.,
1994). Among the capsid proteins, VP2 has
been shown to be the most variable, and
responsible for most of the antigenic variation of the virus (Iwata et al., 1992; Mellor and
Hamblin, 2004); hence VP2 is the major target
of the hosts neutralizing antibodies response
(Burrage et al., 1993, Roy et al., 1994).
AHSV infects all equids, with horses
being the most susceptible to clinical disease,
which has a severe morbidity and high case
fatality rate of 5095%. Mules have a similar
morbidity to horses, but with a lower case
fatality rate of 5070% (Coetzer and Guthrie,
2004). Donkeys and zebras are very resistant
to the disease, and usually just develop subclinical infection (Theiler, 1921; Barnard, 1993;
Coetzer and Guthrie, 2004). Donkeys in the
Middle East can have a case fatality rate of up
to 10%, and may be more susceptible to clinical disease than the South African donkey
(Alexander, 1948; Hamblin et al., 1998; Coetzer
and Guthrie, 2004). Zebras only show mild
fever when experimentally infected with the
virus (Erasmus et al., 1978; Coetzer and
Guthrie, 2004); they maintain year-round
infections and are considered a reservoir host
for the virus in endemic regions (Davies and
57
Otieno, 1977; Barnard, 1993). AHSV has also

been isolated in blood samples 40 days post
infection in zebras. In comparison, in experimentally challenged horses, viraemia usually
lasts 48 days, but does not exceed 21 days
(Barnard et al., 1994; Coetzer and Guthrie,
2004). Donkeys were considered to be potential reservoirs, but the absence of viral antigens after 1419 days post infection makes
them unlikely long-term hosts, although they
may play a small role in spread of the virus.
Theiler demonstrated that AHSV can be
transmitted to dogs and cause a similar
pathology to that found in horses (Theiler,
1906). Dogs, however, are likely to be deadend hosts for the virus (Braverman and
Chizov-Ginzburg, 1996). The most common
way for a dog to become infected with the
virus is through the consumption of uncooked
meat from an infected horse carcass, as was
the case in Pretoria in 1980 (Van Rensburg
et al., 1981).
AHSV is considered to be endemic in
north-eastern parts of South Africa, primarily
Mpumalanga Province, but outbreaks regularly occur in other parts of South Africa
(Lord et al., 2002; Coetzer and Guthrie, 2004).
In addition, major outbreaks of the virus have
recently occurred in regions with close proximity to the Mediterranean Sea, such as in the
Middle East (19591961), North Africa (1965,
1989, 1991), Spain (1966, 19871990) and
Portugal (1989) (Rodriguez et al., 1992;
Coetzer and Guthrie, 2004; Mellor and
Hamblin, 2004).
Clinical disease develops in susceptible
animals after an incubation of about 57 days,
depending on virulence and dose of virus
(Theiler, 1921; Coetzer and Guthrie, 2004).
While AHSV causes severe morbidity and
mortality in the majority of infected horses,
the pathogenesis and clinical disease that
develops differs among cases. This difference
is not completely understood, but is a function of both host factors (genetics and immune
status) and virus factors (dose, route of infection in experimental inoculations and virulence phenotype) (Burrage and Laegreid,
1994).
The insect Culicoides imicola is the principal vector responsible for AHSV transmission, and its importance in AHSV transmission
58
Frank Koenen et al.
has been recognized for over 50 years. C. imicola has historically been found in Africa and
South-east Asia (Meiswinkel et al., 2004).
Nevertheless tick species in general are also
capable of transmitting the virus (Meiswinkel
et al., 2000; Mellor and Hamblin, 2004).
4.11.2
Bartonelloses
Bartonella infections are widespread in wild

and domesticated mammals, and several new
species have been described during the last
few decades. These alphaproteobacteria infect
erythrocytes and endothelial cells, leading to
persistent infections of their mammalian
hosts. As Bartonella species tend to infect the
blood of their vertebrate hosts chronically,
these microparasites can be ingested and
potentially transmitted by blood-feeding
arthropods. Confirmed vectors of B. henselae
(the aetiological agent of cat-scratch disease),
B. bacilliformis, B. quintana, B. grahamii and
B. taylorii are Ctenocephalides felis, Lutzomyia
verrucarum, Pediculus humanus humanus and
Ctenophthalmus nobilis, respectively (Billeter
et al., 2008). Bartonella bacteria have been
detected based mainly on PCR in several
tick species, including I. ricinus, I. scapularis,
I. persulcatus, D. reticulatus, Rh. sanguineus and
Carios kelleyi (Billeter et al., 2008). Some of the
reasons why Bartonella species might be transmitted by ticks have been listed by Telford
and Wormser (2010): other arthropods can
transmit Bartonella species; the DNA of
Bartonella species is often detected in ticks;
human cases of bartonellosis preceded by tick
bites are on record; and Bartonella species are
commonly present in important hosts of
Ixodes ticks, i.e. deer and rodents.
The mere detection of Bartonella DNA by
PCR in blood-feeding arthropods is no
evidence that these bacteria are viable and
infective, or that these arthropods are competent vectors of Bartonella species. In fact, there
is no conclusive evidence that any Bartonella
species can, under natural circumstances,
infect a vertebrate via tick bite, although a
recent laboratory investigation using a membrane feeding technique suggested that I. ricinus is a competent vector for B. henselae (Cott
et al., 2008). Trans-stadial transmission of the
bacteria, their multiplication within the ticks

salivary glands after a second blood meal,
and transmission of viable and infective
B. henselae from ticks to blood were recorded.
However, this study may not be relevant to
establishing the vector competence of ticks
for bartonellae as certain of its parameters
were unnatural: the ticks were fed continuously on blood containing exceedingly
high numbers of bacteria; and the strain of
B. henselae used is highly adapted to laboratory conditions and grows easily in vitro.
As suggested by Telford and Wormser (2010),
a more reliable proof of vector competence
would be to feed an uninfected Ixodes sp. on a
B. henselae-infected cat and then, after the tick
has hatched, determine whether the nymph
can transmit B. henselae by bite to an uninfected cat. However, additional (epidemiological) data would be necessary to
conclusively prove that ticks are of importance as natural vectors of bartonellae.
4.11.3
Q fever
The aetiological agent of Q fever is the rickettsial parasite, Coxiella burnetii. This is an obligate intracellular bacterium, although it can
survive for months or years outside host cells
in such media as water, dried or frozen tissues
and soil. The organism has been described as
a possible biological weapon (Madariaga
et al., 2003). Ticks are one of a broad range of
reservoirs for the organism. More than 40 species of ticks including D. marginatus, are naturally infected with C. burnetii. After feeding
on bacteraemic hosts, nymphs or adults can
transmit the pathogen trans-stadially, and
females can pass it transovarially (Lang, 1990;
Toledo et al., 2009a).
Q fever is a worldwide zoonosis affecting mammals (including domestic animals),
birds and arthropods in most areas in the
world, including Europe (Aitken et al., 1987;
Lang, 1990). The most common reservoirs for
human infections are cattle, sheep and goats.
Dogs, cats, birds and reptiles are also susceptible to infection and may play a role in maintaining the infection in natural habitats.
Although C. burnetii does not usually cause
clinical disease in animals, it occasionally
induces reproductive disorders such as stillbirths, abortion and metritis in pregnant

goats and sheep, but is rarely documented in
dairy cows (Aitken et al., 1987; Lang, 1990).
Infected animals develop high rickettsaemias
and excrete large numbers of C. burnetii in
their faeces for weeks, as well as shedding
organisms via tissues and fluids.
The disease is epidemic, especially on
farms or in farming communities when
infected domestic animals are being handled,
such as during wool shearing, lambing, calving and slaughtering. Therefore, farmers,
abattoir workers, meat-packing workers, veterinarians and laboratory workers in contact
with livestock are at high risk of infection.
Humans can also be infected by inhalation of
the organism or by the ingestion of infected
milk and/or fresh dairy products. The organism may survive in contaminated milk and
butter for up to 3 months. Close contact with
infected animals is not required; infection
may be from contaminated dust, straw or
manure, as was reported for an outbreak of Q
fever in an urban area in southern Wales, UK
(Ayres et al., 1996). After an incubation period
of about 20 days, Q fever is characterized by
mainly limited flu-like illness, pneumonia or
hepatitis. It may become chronic and manifests with fatigue syndrome, chronic hepatitis, endocarditis and other endovascular
infections. The fatality rate is less than 1% in
acute cases but may rise to 30% in chronic
cases. A scientific opinion on Q fever has been
published by the EFSA Panel on Animal
Health and Animal Welfare (2010b).
4.11.4 Tularaemia
Tularaemia is a zoonotic disease caused by
one of the most infectious bacteria, Francisella
tularensis, a Gram-negative obligate intracellular agent (Ellis et al., 2002). This bacterial
species causes great concern as a potential
bioterrorism agent, and is listed among Class
A biothreat agents (Dennis et al., 2001; Oyston
et al., 2004). F. tularensis can be recovered from
contaminated water, soil and vegetation. Four
closely related subspecies of F. tularensis have
been identified: F. t. tularensis, F. t. holarctica,
F. t. mediasiatica and F. t. novicida (Forsman
59
et al., 1994). Subspecies F. t. tularensis and F. t.

holarctica cause most human illness. F. t. holarctica is found throughout much of Europe
except for the UK, Ireland and Iceland.
Tularaemia is typically a disease of northern
and central Europe and the countries of the
former Soviet Union (Ellis et al., 2002; Tarnvik
et al., 2004). Natural infections with F. tularensis have been reported in over 250 animal species and in a range of vertebrates, including
mammals, birds, amphibians and fish, as well
as in certain invertebrates (Morner and
Addison, 2001; Oyston et al., 2004; Santic et al.,
2006). However, tularaemia is primarily a disease of the orders Lagomorpha (rabbits and
hares) and Rodentia (Friend, 2006). The
European brown hare (Lepus europaeus) is a
common host of F. tularensis in central Europe,
where it causes a public health problem
(Morner and Addison, 2001; Strauss and
Pohlmeyer, 2001; Pikula et al., 2004). In
Scandinavia and Russia, tularaemia occurs
frequently in mountain hares (L. timidus)
(Morner and Addison, 2001).
Haematophagous arthropods such as
deer and horse flies, ticks and mosquitoes are
common vectors of F. tularensis (Petersen et al.,
2009). Ticks are believed to be the most important arthropods for F. tularensis as both
mechanical and biological vectors (Hopla and
Hopla, 1994). D. reticulatus plays an important role in the maintenance and transmission
of F. tularensis among small and mediumsized mammals in central Europe (Gurycov
et al., 1995, 2001; Hubalek et al., 1996). Other
ticks, such as I. ricinus, I. persulcatus, D. marginatus, Rh. rossicus and Ha. concinna, have
also been found to be naturally infected with
F. tularensis in Europe (Hopla and Hopla,
1994; Keim et al., 2007).
Tularaemia, in a range from subclinical
infection to death, rarely occurs among
domestic animals, and is most frequently
observed in sheep during the lambing season, and in cats and dogs (Friend, 2006;
OToole et al., 2008). Humans are highly susceptible to F. tularensis. Infections in humans
are typically sporadic, but outbreaks do
occur (Matyas et al., 2007). The disease is not
contagious and is most often transmitted
to humans by the bite of an infected arthropod vector, by direct contact with infected
60
Frank Koenen et al.
animals (e.g. voles, mice, water rats, squirrels, rabbits and hares), by contact with
infected animal tissues or fluids, by ingestion
of contaminated water or food, or by inhaling infected materials (Dennis et al., 2001).
Several emergences or re-emergences of
tularaemia have recently been seen all over
the world (Petersen and Schriefer, 2005): the

highest incidences occurred in confined geographical areas of Finland and Sweden
(Eliasson et al., 2002), and the disease also
appeared in Spain (Perez-Castrillon et al.,
2001), Kosovo (Reintjes et al., 2002) and
Germany (Kaysser et al., 2008).
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Maps of Reported Occurrence

of Tick-borne Pathogens*
Agustn Estrada-Pea,1 Frank Koenen,2 Ilaria Pascucci,3

Thomas G.T. Jaenson,4 Maxime Madder,5
Rita de Sousa6 and Rbert Farkas7
1
University of Zaragoza, Spain; 2Veterinary and Agrochemical Research Centre,
CODA-CERVA, Brussels, Belgium; 3Istituto Zooprofilattico Sperimentale dellAbruzzo
e del Molise G. Caporale, Teramo, Italy; 4Uppsala University, Sweden; 5Institute
of Tropical Medicine, Antwerp, Belgium and University of Pretoria, South Africa;
6
National Institute of Health Doutor Ricardo Jorge, IP (INSA), guas de Moura,
Portugal; and 7Szent Istvn University, Budapest, Hungary
The maps that are presented in this chapter

on the geographical occurrence of tickborne pathogens have been produced using
three sources of data: (i) a systematic literature review based on scientific papers
retrieved from the databases integrated in
the ISI Web of Knowledge and PubMed,
published from January 2000 to March 2010
(see Introduction); (ii) a pool of scientific
papers considered relevant by the experts
of the Working Group on Ticks and Tickborne Pathogens of the Animal Health and
Animal Welfare (AHAW) Unit of the
European Food Safety Authority (EFSA),
and derived from their private collections
regardless of the time frame concerned;
and (iii) published historical data (for the
period of approximately 19702000) of the
Integrated Consortium on Ticks and Tickborne Diseases (the Sixth Framework
Programme funded ICTTD-3 European
project). Consequently, the data presented
in the maps were classified according to

their chronology into those published in the
years 2000 to 2010, or as historical data
(published before the year 2000).
The data included in the maps were classified according to the reported geographic
location using either coordinates (latitude/
longitude) or, in the cases where coordinates
were not indicated or could not be found
because the name of the location was not specified in the corresponding paper, according to
the smallest administrative region in which
the findings were made. In this last case, the
symbols used in the maps are placed either in
the middle of the smallest administrative
region described in the scientific paper (for
the countries that do not have official nomenclature of territorial units for statistics
NUTS), or in the middle of the NUTS
containing the specified location. Countries
or areas not showing cases of the tickborne pathogen under investigation are not
* Frank Koenen wrote Section 5.1; Ilaria Pascucci, Section 5.2; Thomas G.T. Jaenson, Sections 5.3, 5.6; Maxime
Madder, Section 5.4; Rita de Sousa, Section 5.5; Rbert Farkas, Sections 5.7, 5.8, 5.9, 5.11, 5.12; Agustn
Estrada-Pea produced the maps with the advice of the authors and Thomas G.T. Jaenson, Section 5.10.
76
necessarily free of that pathogen. Rather, data

on that species may not be presented in the
literature available for this report. The presence of the pathogens represented in the maps
is based on confirmed detection of the antigen
for that pathogen. Serological evidence was
not included, so these maps may underestimate the real distribution of the pathogens.

The few existing records for African swine
fever virus (ASFV) have been obtained from
active surveillance of the infection in domestic pigs, wild boars and soft ticks associated
with foci of outbreaks. The exception to this
is the island of Sardinia, which is considered
to be endemic for ASFV. The Ornithodoros tick
is not present on this island, and no records
exist of wild boar infested with ticks.
Therefore, ticks do not play a role in the
maintenance of ASFV on that island. The
endemicity is mainly related to the traditional
extensive production system. The distribution recorded on the map (Fig. 5.1) displays
77
the records for ASFV that were found in the

available literature of the last 10 years (solid
black circles) and data from before 2000 (solid
black diamonds). Both symbols indicate
records in which the coordinates were not
provided, just the name of the location (see
the introduction to this chapter). The distribution is expected to be biased because the
culinary habits in wide regions of northern
Africa and parts of the eastern Mediterranean
erase the domestic pig as the main host of the
virus. The most recent outbreaks are in the
Caucasus. ASFV in Portugal and Spain was
eradicated in 1999 and 1995, respectively
(Snchez-Vizcano, 2006).
5.2 Crimean-Congo Haemorrhagic

Fever
Clinical cases of Crimean-Congo haemorrhagic fever (CCHF) in humans, as well as
ticks infected with the CCHF virus (CCHFV)
under natural (field) conditions have been
recorded only in an area covering southern
Russia, Turkey, Albania, Kosovo, Greece,
Fig. 5.1. Reported occurrence of African swine fever virus (ASFV). Smallest administrative region or
territorial unit for statistics (NUTS), data from 20002010; Smallest administrative region or territorial unit
for statistics (NUTS), historical data (before 2000).
78
Agustn Estrada-Pea et al.
Romania and Bulgaria (Fig. 5.2). The distribution map displays the records for CCHFV that
were found in the available literature of the
last 10 years, and are indicated by solid black
circles. These symbols represent records in
which the coordinates were not provided, just
the name of the location (see the introduction
to this chapter). The disease is widespread in
Turkey, following the areas where the main
tick vector, Hyalomma marginatum, exists. No
published reports exist for the distribution
and incidence of the disease in the Balkans
and Russian areas other than reports collated
from ProMED Mail (http://www.promedmail.org, from the International Society for
Infectious Diseases). It is likely that the virus
circulates in the Balkans (as suggested by the
recent first report of CCHF in humans in
Greece; Papa et al., 2008). This is probably the
most exposed area for spread of the virus to
other parts of Europe. Both the virus and
signs of clinical infection in humans have
been partly surveyed in Europe, but with
negative results. This is a cause of concern,
because we then ignore the reasons why the
disease (or the virus) is absent in places
where the tick vectors are widespread (see the

comments on Hy. marginatum and other hard
ticks in Chapter 2, Section 2.2, and Chapter 6,
Section 6.2).
5.3 Tick-borne Encephalitis

Group Viruses
The map shown in Fig. 5.3 displays the
records of the tick-borne encephalitis (TBE)
group viruses that were found in the available
literature of the last 10 years. The solid black
circles indicate the coordinates (latitude/
longitude). The stars indicate records in which
the coordinates were not provided, just the
name of the location (see the introduction
to this chapter).
Tick-borne encephalitis virus (TBEV)
includes three viral subtypes, i.e. Western
TBEV, Siberian TBEV and Far Eastern TBEV.
Ixodes ricinus is the main vector of the Western
subtype and the Siberian and Far Eastern
subtypes are vectored mainly by I. persulcatus. Although the main vector of the Western
Fig. 5.2. Reported occurrence of Crimean-Congo haemorrhagic fever virus (CCHFV) (20002010).
Smallest administrative region or territorial unit for statistics (NUTS).
79
Fig. 5.3. Reported occurrence of tick-borne encephalitis group viruses (20002010). Smallest
administrative region or territorial unit for statistics (NUTS); Coordinate (latitude/longitude).
subtype is I. ricinus, several other species, including I. hexagonus, I. arboricola, Haemaphysalis

(Ha.) concinna, Ha. inermis and Ha. punctata are
competent but secondary vectors (Labuda
and Nuttall, 2008). Records of TBE group
viruses published during the last 10 years and
shown on the map originate from the following countries: Czech Republic, Denmark,
Estonia, Finland (including the land Islands),
Germany, Greece, Hungary, Italy, Latvia,
Liechtenstein, Lithuania, Norway, Poland,
Russia, Sweden, Switzerland and Turkey. The
map (Fig. 5.3) does not include records published before 2000. Countries with increased
risk of TBE include Austria, Slovakia,
Hungary, the Czech Republic, the Baltic
States, southern Germany and southern and
eastern Sweden (ECDC, 2010). TBE is prevalent in Russia all the way from the western
border with Europe to its eastern border.
The TBE virus subtypes in Russia are in general vectored by I. persulcatus and usually
cause more severe disease than the central
European strain.
The map does not show the presence of
the louping ill virus (LIV). This virus is
endemic in sheep farming areas of Northern
Ireland, the UK (Scotland, Cornwall and

Wales) and Norway.
The geographical distribution of TBEinfected ticks within regions and countries is
such that the risk of infection can change dramatically from one area to another within
short distances. TBE incidence fluctuates from
year to year, but the number of reported
human TBE cases has increased in Europe in
the last two decades. The mean number of
TBE infections in Europe, excluding Russia,
between 1990 and 2007 was 2805 cases per
year, up 193% from a mean of 1452 cases per
year (19761989) (ECDC 2010). An increase
in TBE incidence has been noted in some
northern countries and new TBE foci have
appeared, especially in the last decade, in
many areas, e.g. Denmark, Norway, Sweden
and Finland (Jskelinen et al., 2006;
Formsgaard et al., 2009). The main determinants for such changes in TBE incidence may
vary from one time to another and from one
region to another region. Also, the determining factors are often many and they
interact often in complex manners. Among
the main determinants should be mentioned:
the changing climate, which may affect the
80
vector directly as well as indirectly via the

plant and host communities; sociopolitical
changes, which may increase or decrease
human activities in TBEV foci; and technological factors, e.g. better diagnostic
methods and increased awareness (Donoso
Mantke et al., 2008; Telford and Goethert,
2008; Randolph, 2010).
5.4 Anaplasmoses and Ehrlichioses

records of Anaplasma and Ehrlichia species
that were found in the available literature of
the last 10 years. The solid black circles indicate the coordinates (latitude/longitude).
The stars indicate records in which the coordinates were not provided, just the name of
the location (see the introduction to this
chapter).
The distribution of E. canis has expanded
with the distribution of its vector, Rhipicephalus (Rh.) sanguineus. For E. ruminantium,
no records have yet been found in Europe
and its vector ticks seem to be absent from
this continent, except for some imports of

immature stages on birds.
For anaplasmoses, A. phagocytophilum
has been recorded in most European countries. A. marginale, normally recorded from
tropical and subtropical regions, has a more
limited distribution in Europe with records
from Sicily, Hungary and Spain. A. ovis has so
far been found in the same countries as
A. marginale. In Italy, A. centrale, a similar parasite to A. marginale, was first recorded in
2008 (Carelli et al., 2008). Similarly, A. bovis
has also been recorded in Italy although it is
most commonly present in other continents,
including South America, Africa and the
Indian subcontinent.
A. platys has been detected worldwide,
and in Europe cases of infection have been
reported from Spain, France, Greece and Italy
(de la Fuente et al., 2006).
5.5
Rickettsioses

records of Rickettsia species that were found
Fig. 5.4. Reported occurrence of Anaplasma spp. and Ehrlichia spp. (20002010). Smallest administrative
region or territorial unit for statistics (NUTS); Coordinate (latitude/longitude).
81
Fig. 5.5. Reported occurrence of Rickettsia spp. (20002010). Smallest administrative region
or territorial unit for statistics (NUTS).
in the available literature of the last 10 years.

chapter).
The map represents records of all the
tick-transmitted rickettsiae recorded in
Europe, irrespective of the species or strains.
Each of the species has a specific area of distribution, R. conorii Malish, which causes
Mediterranean spotted fever (MSF) is mainly
present in the Mediterranean, where its vector, Rh. sanguineus, is present. The R. conorii
Israeli spotted fever strain was first reported
from Israel but was later also detected in
Portugal, Sicily, Tunisia and Libya. As most of
the Rickettsia species are transmitted by different tick species, of which the distribution is
not always overlapping, these rickettsial
pathogens are found in areas determined by
their vector.
Although no detailed information is
available about the presence and absence of
the different rickettsiae, it must be said that
more and more recent studies, especially
molecular studies, reveal the presence of the
Rickettsia species in new member states of the

European Union (EU), and not necessarily in
adjacent countries from which the parasite
was originally described.
5.6
Borrelioses

records of Borrelia species that were found in
the available literature of the last 10 years.
The solid black circles indicate the coordinates (latitude/longitude). The stars indicate
provided, and only the name of the location
was included in the original report (see the
introduction to this chapter).
As described above, this map shows the
more recent records of B. afzelii, B. burgdorferi s.s., B. garinii, B. lusitaniae, B. spielmanii,
B. valaisiana and B. burgdorferi s.l. B. burgdorferi s.l. is considered to consist of at least 15
genospecies, eight of which occur in Europe,
namely: B. burgdorferi s.s., present in Europe
and the USA, but rare in Russia and apparently absent from Asia; and B. afzelii,
82
Fig. 5.6. Reported occurrence of Borrelia spp. (20002010). Smallest administrative region or territorial
unit for statistics (NUTS); Coordinate (latitude/longitude).
B. bavariensis (formerly B. garinii OspA serotype 4), B. garinii, B. valaisiana, B. spielmanii,

B. lusitaniae and B. bissettii in Eurasia. B. garinii and B. afzelii are the most commonly
encountered and most widely distributed of
the European Lyme disease genospecies. In
Portugal, Tunisia and Morocco, B. lusitaniae is
common and predominates over the other
genospecies in I. ricinus populations (Piesman
and Gern, 2008).
Although I. ricinus and I. persulcatus are
apparently the only tick species responsible
for infecting humans with the Lyme disease
spirochaetes in Europe and Asia, B. burgdorferi s.l. has been found in several other tick
species (EUCALB, 2010). These carrier species are not considered to be competent natural vectors of B. burgdorferi s.l.
There are indications from northern
Europe about an increasing incidence of Lyme
borrelioses and TBE as a result of increased
abundance of the vector, I. ricinus, and the
expansion of its range, probably due to an
extended vegetation period caused by climate change (Tlleklint and Jaenson, 1998;
Lindgren et al., 2000; Gray et al., 2009). Thus,
the ranges of at least some of the Lyme
borreliosis genospecies are likely to change

significantly in the near future.
Note that, as mentioned above, the
present map is based solely on recent records
(published after 1 January 2000). Therefore,
many earlier records of B. burgdorferi s.l. and
some genospecies, especially B. afzelii, B. garinii and B. burgdorferi s.s., from large areas of
Europe are not shown on the map. Thus,
while not revealed by this map, it should be
noted that B. burgdorferi s.l. is common in
most or all I. ricinus populations in southern
and central Sweden, southern and central
Finland, Estonia, Latvia, Russia, Byelorussia,
Kirghizia, Moldavia, Ukraine and Ireland.
5.7
Babesioses

records of Babesia species that were found in
the literature of the last 10 years. The empty
circles indicate the coordinates (latitude/
longitude). The solid black circles indicate
the introduction to this chapter).
83
Fig. 5.7. Reported occurrence of Babesia spp. (20002010). Smallest administrative region or territorial
The occurrence of Babesia species has

been reported from many European countries. Several tick vectors can carry more than
one species of Babesia. Some Babesia species
can infect more than one genus of ticks; others can only infect ticks from the genus Ixodes.
The most common species is B. canis canis
which exists in several foci of wide areas in
Europe, from France to Poland, where its
known vector species, Dermacentor reticulatus,
is present. Recently a new zoonotic Babesia
species (Babesia EU-1; proposed name B. venatorum) has been detected in roe deer and in
humans, with I. ricinus identified as its vector.
In endemic areas, all or almost all individuals
of the host population are infected when they
are young, with no or minimal clinical disease. The introduction of susceptible animals
into endemic regions could lead to a recrudescence of babesiosis.
5.8 Theilerioses
records of Theileria species that were found in

The empty circles indicate the coordinates (latitude/longitude). The solid black
circles indicate records in which the coordinates were not provided, just the name of
chapter).
There have been very few reports on
theileriosis of domesticated ruminants from
the European countries. Mediterranean theileriosis, caused by T. annulata is known to occur
in southern Europe, North Africa and Turkey.
However, no published reports exist for its
incidence and distribution in the Balkans and
Iberian Peninsula where ixodid ticks such as
Hy. anatolicum, Hy. marginatum and Hy. excavatum, which are known to transmit T. annulata,
are found in large numbers, especially in semiarid areas. Most papers published in the last
10 years reported the occurrence of horse
theileriosis caused by T. (Babesia) equi in
Morocco, Turkey, Italy, Croatia and Slovenia
(see Fig. 5.9, equine piroplasmoses). Based on
these reports, the exact geographical distribution and the incidence of theileriosis in
domesticated and wild animals cannot be
mapped properly in Europe.
84
Fig. 5.8. Reported occurrence of Theileria spp. (20002010). Smallest administrative region or territorial
Fig. 5.9. Reported occurrence of equine piroplasmoses (20002010). Smallest administrative region or
territorial unit for statistics (NUTS).
5.9
Equine Piroplasmoses

records of equine piroplasmoses that were
found in the available literature of the last
10 years. The solid black squares indicate

The distribution and incidence of equine
piroplasmoses caused by two species of apicomplexan protozoa, Theileria (syn. Babesia)
equi and Babesia caballi, have been reported
from a few European countries. Piroplasmosis
seems to be more prevalent in France, Italy

and Turkey than in other countries. However,
the disease may occur in other countries
because several species of ixodid ticks belonging to the genera Hyalomma, Dermacentor and
Rhipicephalus have been identified as vectors
of both B. caballi and T. equi in a wide area in
Europe. Besides infected tick vectors, both
parasite species can be spread by infected
horses without clinical signs, by the transfer
of blood from infected to naive equids through
shared needles, improperly shared equipment, and blood or serum transfusions.
5.10
Bartonelloses

records of Bartonella species that were
found in the available literature of the last
10 years. The empty circles indicate the
coordinates (latitude/longitude). The solid
black circles indicate records in which the
coordinates were not provided, just the
name of the location (see the introduction
to this chapter).
85
This map shows the geographical locations only of recent (20002010) records of
B. henselae, B. quintana, B. vinsonii and unidentified Bartonella species. The genus at present
comprises 28 species and subspecies (DSMZ,
2010). Thus, the map does not give a complete
picture of the geographical distribution in
Europe of the genus Bartonella.
Because Bartonella species usually infect
their vertebrate hosts chronically, these
microparasites can be ingested and potentially transmitted by blood-feeding arthropods. Confirmed vectors of B. henselae (the
aetiological agent of cat-scratch disease),
B. bacilliformis, B. quintana, B. grahamii and
B. taylorii are Ctenocephalides felis, Lutzomyia
verrucarum, Pediculus humanus humanus and
Ctenophthalmus nobilis, respectively (Billeter
et al., 2008). Bartonella bacteria have been
detected based mainly on PCR in several
tick species, including I. ricinus, I. scapularis,
I. persulcatus, D. reticulatus, Rh. sanguineus and
Carios kelleyi (Billeter et al., 2008). Some of the
reasons that Bartonella species might be transmitted by ticks have been listed by Telford
and Wormser (2010): other arthropods can
transmit Bartonella species; the DNA of
Fig. 5.10. Reported occurrence of Bartonella spp. (20002010). Smallest administrative region or
territorial unit for statistics (NUTS); Coordinate (latitude/longitude).
86
Bartonella species is often detected in ticks;

human cases of bartonellosis preceded by tick
bites are on record; and Bartonella species are
commonly present in important hosts of
Ixodes ticks, i.e. deer and rodents.
Although many researchers have indicated that ticks might be vectors of bartonelloses, it should be clear that there is, so far, no
evidence that any tick species is a natural,
competent vector of any Bartonella species
(Telford and Wormser, 2010).
5.11 Tularaemia
records of Francisella tularensis infection in
humans or animals that were found in the
available literature of the last 10 years. The
solid black squares indicate records in which
name of the location (see the introduction to
this chapter).
Not all of these cases are necessarily related
to tick transmission. Although vector-borne
transmission of tularaemia to mammalian

hosts has an important role in the pathogenesis of the disease (Petersen et al., 2009), there
are several vectors in addition to ticks that are
related to the transmission and maintenance
of the pathogen. The map indicates the sporadic reported cases by locations; the sparsity
of the data is mainly due to the low percentage of compliance with reporting this disease
and the similarity of clinical signs to those of
several other diseases.
5.12
Q Fever

records of Coxiella burnetii infection in animals
and Q fever disease in humans or animals
that were reported in the available literature
of the last 10 years. The solid black squares
indicate records in which the coordinates
were not provided, just the name of the location (see the introduction to this chapter).
In domestic animals, C. burnetii is
endemic in most European member states,
Fig. 5.11. Reported occurrence of Francisella tularensis (20002010). Smallest administrative region or
87
Fig. 5.12. Reported occurrence of Coxiella burnetii (20002010). Smallest administrative region or
but has a limited impact on human and

animal health. Most countries have a long
history of infection in domestic animals and
sporadic human cases. In the Netherlands, a
Q fever outbreak in humans first emerged

in 2007 and is considered to have been the
largest community outbreak ever recorded
(EFSA, 2010).
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de la Fuente, J., Torina, A., Naranjo, V., Nicosia, S., Alongi, A., La Mantia, F. and Kocan, K.M. (2006) Molecular
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Maps of Reported Occurrence of Ticks*
Agustn Estrada-Pea,1 Thomas G.T. Jaenson,2

Rbert Farkas3 and Ilaria Pascucci4
1
University of Zaragoza, Spain; 2Uppsala University, Sweden;
3
Szent Istvn University, Budapest, Hungary; and 4Istituto Zooprofilattico
Sperimentale dellAbruzzo e del Molise G. Caporale, Teramo, Italy
The maps that are presented in this chapter on

the occurrence of ticks have been drawn up
using three sources of data: (i) a systematic literature review previously published by EFSA
(2010) based on scientific papers published
from January 2000 to March 2010 and retrieved
from the databases integrated in the ISI Web of
Knowledge and PubMed; (ii) a pool of scientific papers considered relevant by the experts
of the Working Group on Ticks and Tick-borne
Pathogens of the Animal Health and Animal
Welfare (AHAW) Unit of the European Food
Safety Authority (EFSA), and derived from
their private collections regardless of the time
frame concerned; and (iii) published historical
data (for the period of approximately years
19702000) of the Integrated Consortium on
Ticks and Tick-Borne Diseases (the Sixth
Framework Programme funded ICTTD-3
European project).
The data included in the maps were classified according to their geographic accuracy
using either coordinates (latitude/longitude)
or, in the cases where coordinates were not
indicated or could not be found because
the name of the location was not specified in
the corresponding paper, according to the
smallest administrative region in which the
findings were made. In the latter case, the

symbol representing the data was placed
either in the middle of that region (for the
countries that do not have official nomenclature of territorial units for statistics NUTS),
or in the middle of the NUTS containing the
specified location. Countries or areas not
showing the presence of a tick genus or species are not necessarily free of it. Rather, that
genus or species may not be represented in
the literature that was available for this
monograph.
Owing to the lack of reported data in the
area of concern, no maps were produced for
the genera Argas and Amblyomma.
6.1
Soft Ticks (Argasidae), Genus

Ornithodoros
The map shown in Fig. 6.1 covers the known

distribution of the species in the genus
Ornithodoros in the Palaearctic region, and
shows historical data (before 2000). The solid
black circles indicate the coordinates (latitude/longitude). The stars indicate records in
which the coordinates were not provided, just
* Agustn Estrada-Pea produced the maps with the advice of the authors and wrote Sections 6.1, 6.2.4;
Thomas G.T. Jaenson, Section 6.2.1; Rbert Farkas, Sections 6.2.2, 6.2.5; Ilaria Pascucci, Section 6.2.3.
89
90
Fig. 6.1. Reported occurrence of Ornithodoros spp. Smallest administrative region or territorial unit for
statistics (NUTS), historical data (before 2000); Coordinate (latitude/longitude), historical data (before 2000).
the name of the location (see the introduction

to this chapter). As mentioned previously
(Section 2.1.2), the lack of adequate morphological features renders the separation of possible species a difficult task. Hence, the map
includes only the accurate records of the species in the group of O. erraticus. The reported
distribution of these ticks is poor, as they
seem to be restricted to parts of the western
Mediterranean and portions of Georgia,
Armenia and Turkey. Species in the genus are
also known to be present in parts of Morocco,
Tunisia and Egypt. Currently, there is fragmentary knowledge about the distribution of
this species, although it has been hypothesized that O. erraticus would be present in the
Iberian Peninsula and parts of Morocco, while
O. sonrai would be the species found in Egypt
and southern Morocco. The mechanisms
operating in the separation of these species in
Morocco are unknown. It is interesting to
note, however, that there are no recent records
(years 20002010) of the ticks in Europe, so, as
noted above, the mapping has been carried
out with historical records (years 19502000).
This is because the tick has received some
attention owing to the occurrence of the epidemics of African swine fever (ASF) in
Europe. All of the species in the O. erraticus
group are xerophilous ticks, and therefore
live in open and dry habitats commonly associated with rodent burrows.
The apparent lack of records of the tick in
the centre of the target area seems to be the
result both of a lack of interest in the soft ticks
and the lack of knowledge on the morphology of these species. Further sampling efforts
should be done in wide areas of the European
Mediterranean region, with special emphasis
on nests of rodents, which are the preferred
hosts of this group of species.
6.2
6.2.1
Hard Ticks (Ixodidae)

Genus Ixodes: Ixodes ricinus
The map shown in Plate 20 displays the

records of I. ricinus ticks that were found in
the available literature of the last 10 years
(solid red circles and black stars) and those
that are from historical data older than 2000

(solid green circles). The circles indicate the
coordinates (latitude/longitude). The stars
I. ricinus is the common tick of Europe
and is widely distributed in north-western
Europe through much of the Palaearctic, from
Iceland and Ireland through central and
southern Europe eastwards to central Asia
(Iran) and southwards to North Africa. It is
present in relatively dry Mediterranean
habitats in northern Africa and the Iberian
Peninsula, in the damp sheep pastures of
Ireland, Scotland, Wales and England, and
in relatively humid, mixed coniferous/
deciduous woodland biotopes throughout
most of Europe, including Scandinavia,
Finland and western Russia. In Europe,
changes in climate during the last few decades appear to have caused an expansion in
the ticks range to higher altitudes (Materna
et al., 2005) and latitudes (Tlleklint and
Jaenson, 1998; Lindgren et al., 2000; Lindgren
and Jaenson, 2006; Jaenson and Lindgren,
91
2011; Jaenson et al., 2012), as well as affecting

its population density. Moreover, with the
changing range of I. ricinus, its role as an
important vector of infections of humans
and/or domesticated mammals, e.g. tickborne encephalitis (TBE), including louping
ill of sheep, Lyme borrelioses due to Borrelia
burgdorferi s.l., anaplasmosis (Anaplasma
phagocytophilum), babesiosis (Babesia divergens), rickettsiosis (Rickettsia helvetica), and
tularaemia (Francisella tularensis), needs to be
monitored in the future.
6.2.2
Genus Haemaphysalis
Haemaphysalis punctata
records of Haemaphysalis (Ha.) punctata ticks
the last 10 years (stars), and in historical data
published before 2000 (solid black circles).
The circles indicate the coordinates (latitude/
longitude). The stars show records in which
Fig. 6.2. Reported occurrence of Haemaphysalis punctata. Smallest administrative region or territorial unit
for statistics (NUTS), data for 20002010; Coordinate (latitude/longitude), historical data (before 2000).
92
name of the location (see the introduction to

this chapter).
Ha. punctata is widely present in Europe
and in the whole Mediterranean area. It
occurs mainly in a dense forest environment.
Adults feed on domestic and wild ungulates;
immature stages, besides feeding on small
mammals, can also feed on birds, including
migratory birds. Migratory birds are carriers
of immature ticks and could potentially introduce them into free areas. Nevertheless, most
reports show this tick species to be concentrated in southern-central Europe while
northern Africa appears to be the southern
limit of its distribution.
Haemaphysalis concinna
The map shown in Fig. 6.3 displays the records
of Ha. concinna ticks that were found in the
available literature of the last 10 years. The
stars indicate records in which the coordinates
Ha. concinna is a palaearctic species scattered in the Eurasian region from the Atlantic
coast to Japan. It is mainly present in temperate climates and is well adapted to different
biotopes. Adults feed on domestic and wild
ungulates; immature stages, besides feeding
on small mammals, can also feed on birds,
including migratory birds.
Haemaphysalis inermis
records of Ha. inermis ticks that were found in
chapter).
Ha. inermis is present in the southerncentral Eurasian region. It is an ancient species and its geographic origin has been
identified in the Caucasus. Ha. inermis is well
adapted to different biotopes. Adults feed on
domestic and wild ungulates, but also on
canids and lagomorphs. Immature stages,
besides feeding on small mammals, can also
feed on lizards, on ground-feeding birds, and
migratory birds.
Fig. 6.3. Reported occurrence of Haemaphysalis concinna. Smallest administrative region or territorial
unit for statistics (NUTS), data for 20002010.
93
Fig. 6.4. Reported occurrence of Haemaphysalis inermis. Smallest administrative region or territorial unit
for statistics (NUTS), data for 20002010.
6.2.3
Genus Hyalomma
Hyalomma marginatum
Hyalomma marginatum is well known from
the Mediterranean region, where it can be
found populating areas in northern Africa
and southern Europe. The map shown in
Plate 21 displays the records of Hy. marginatum ticks that were found in the available
literature of the last 10 years (solid red circles
and black stars) and those that are from
historical data older than 2000 (solid green
circles). The circles indicate the coordinates
(latitude/longitude). The stars indicate
The tick is restricted to the oroMediterranean (temperate) zone; it appears
that a too-cold or too-dry climate may restrict
the colonization of potentially available areas.
Accurate records exist for the tick in wide
areas from Turkey to Portugal. The presence
of the tick has been recorded from southern
Russia and neighbouring Republics (but has
not been documented with any geographical
information). The tick is also known from

other sites in northern Africa, but those
records lack adequate referencing. Scattered
records of the tick exist in northern Europe,
but these records came from migratory birds,
and they do not represent established populations. Migratory birds may bring immature
ticks into free areas in spring, but the current
climate in the Scandinavian countries jeopardizes their establishment. Concerns exist
about the probable spread of the tick into
northern latitudes where the tick is currently
absent. Permanent populations of the tick are
currently established at around 46N, and
these seem to be restricted in movement
because of the Alps and the Balkan mountain
ranges. In any case, considerable interest
exists about the possible spread of the tick,
and more field studies are necessary to understand the basic phenological processes behind
the observed distribution.
6.2.4
Genus Rhipicephalus
There are serious concerns about the identity

of the ticks of the genus Rhipicephalus in the
94
Palearctic region. Currently, the presence has

been reported of Rhipicephalus (Rh.) annulatus
(formerly in the genus Boophilus) and of three
more species, namely Rh. bursa, Rh. sanguineus
and Rh. turanicus. There is no problem in differentiating specimens of Rh. bursa, but there
are serious troubles in the differentiation of
both Rh. sanguineus and Rh. turanicus.
Therefore, we prefer to report on specimens
of both Rh. sanguineus and Rh. turanicus as the
Rh. sanguineus complex (group).
Rhipicephalus sanguineus complex
(Rh. sanguineus and Rh. turanicus)
records of ticks of the Rh. sanguineus complex
the last 10 years (stars and solid black circles)
and those that are from historical data older
than 2000 (open circles). The open circles indicate the coordinates (latitude/longitude). The
stars and solid black circles indicate records
in which the coordinates were not provided,
just the name of the location (see the introduction to this chapter).
Species in the Rh. sanguineus complex
have a wide Mediterranean distribution (see
Fig. 6.5) linked to an endophilous (nidicolous)
habitat associated mainly with domestic carnivores. Other than this anthropophilous habitat, the ticks of this complex are common in
open areas with different degrees of moisture
content, but are always associated with warm
areas. It is difficult to ascertain the degree to
which Rh. sanguineus ticks can colonize northern latitudes. While the climate is much too
cold for the tick at such latitudes, the association of the tick with endophilous sites may
protect its populations from the extremely cold
temperatures. Therefore, it could be expected
that the tick may be found at more northern
latitudes than the currently reported range.
Rhipicephalus bursa
records of Rh. bursa ticks that were found in
Fig. 6.5. Reported occurrence of Rhipicephalus sanguineus group (Rh. sanguineus and Rh. turanicus).
Smallest administrative region or territorial unit for statistics (NUTS), data for 20002010; Smallest
administrative region or territorial unit for statistics (NUTS), data for 20002010; Coordinate (latitude/
longitude), historical data (before 2000).
the available literature of the last 10 years (red

stars and solid red circles) and those that are
from historical data older than 2000 (green
stars and solid green circles). The circles
indicate the coordinates (latitude/longitude).
The stars indicate records in which the
coordinates were not provided, just the name
of the location (see the introduction to this
chapter).
The distribution of Rh. bursa overlaps
with that of Hy. marginatum and Dermacentor
marginatus in parts of its Mediterranean range.
Such a distribution extends to approximately
45N latitude. However, Rh. bursa prefers drier
areas while Hy. marginatum and D. marginatus
colonize colder sites. It is interesting to note
the different distribution areas of D. marginatus and Rh. bursa in France, where similar tick
field collection efforts on both species have
been carried out in the last 3040 years.
Rh. bursa is more abundant in the oroMediterranean zone than D. marginatus, while
D. marginatus can colonize colder and more
humid sites. The tick seems to be absent from
many parts of southern Russia and is probably
95
replaced by other tick species in this region,

but this extreme has not been reported.
Rhipicephalus annulatus
(formerly Boophilus)
records of Rh. annulatus ticks that were found
in the available literature of the last 10 years
(stars), and in historical data older than 2000
(solid black circles). The circles indicate the
coordinates (latitude/longitude). The stars
The data gathered about the distribution
of Rh. annulatus in the Mediterranean region
seems to be discontinuous in nature. The tick
is well represented in many areas of the
Mediterranean region (see Fig. 6.6). It seems
that the apparent lack of Rh. annulatus at
sites that should support populations of the
species may be a result of the control of ticks
on cattle, which has resulted in an apparent
absence of the species. In any case, Rh. annulatus
Fig. 6.6. Reported occurrence of Rhipicephalus (Boophilus) annulatus. Smallest administrative region or
territorial unit for statistics (NUTS), data for 20002010; Coordinate (latitude/longitude), historical data
(before 2000).
96
has a patchy area of distribution because it is

restricted to sites with adequate moisture
content. In many parts of northern Africa, the
tick is found only in areas where a permanent
or temporal flow of water is available.
6.2.5
Genus Dermacentor
Dermacentor marginatus
records of D. marginatus ticks that were found
(solid red circles and black stars) and those
that are from historical data published before
2000 (solid green circles). The circles indicate
the coordinates (latitude/longitude). The
stars show records in which the coordinates
There are no conclusive studies about the
involvement of D. marginatus in the transmission of Crimean-Congo Haemorrhagic Fever
(CCHF) virus among animals and humans.
However, the area of distribution of this tick
species overlaps in many parts with that of
Hy. marginatum (the proven vector) and

Rh. bursa (a suspected vector of some CCHF
viral strains). D. marginatus is well known
from the Mediterranean region (see Plate 23),
and it seems that this species can spread further than its previously known distribution
area. It is present in areas north of the Balkans,
being recorded from as far north as Hungary
and southern Germany. The tick D. reticulatus, which has a northern distribution in
Europe, has been wrongly reported as D. marginatus in parts of Germany. Therefore, the
actual northern limits of D. marginatus are not
well known, although the species does seem
to be widespread in France in areas of a climate similar to that found in southern
Germany. D. marginatus has not been recorded
from Belgium or the Netherlands. Established
populations are well known in southern
Russia, but there are no geographically accurate reports about this.
Dermacentor reticulatus
records of D. reticulatus ticks that were found
Fig. 6.7. Reported occurrence of Dermacentor reticulatus. Smallest administrative region or territorial
unit for statistics (NUTS), data for 20002010; Coordinate (latitude/longitude), data for 20002010;
Coordinate (latitude/longitude), historical data (before 2000).
(stars and open circles) and those that are

from historical data older than 2000 (solid
black circles). The circles indicate the coordinates (latitude/longitude). The stars show
Accurate records of D. reticulatus (syn.
D. pictus) also known as the ornate dog tick
exist in wide areas in Europe, from France to
Moldavia. The area of distribution of this tick
species (Fig. 6.7) overlaps in many parts of
Europe with that of D. marginatus and it has
been wrongly reported as the latter species.
97
D. reticulatus seems to be widespread in France,

Germany, Hungary and the eastern part of
Poland. It prefers cold regions with an adequate
amount of air relative humidity, therefore it is
absent in the Mediterranean region where the
relative humidity is not conducive to survival.
It is absent from mountain regions, but very
abundant in low-altitude hills. The actual geographical distribution and northern limits of
this species are not well known. Concerns exist
about the probable spread of D. reticulatus into
other areas of countries that are already infested
(e.g. Germany) and into new countries where
the tick is currently absent.
References
EFSA Panel on Animal and Welfare (AHAW) (2010) Scientific opinion on the role of tick vectors in the
epidemiology of Crimean Congo hemorrhagic fever and African swine fever in Eurasia. EFSA Journal
8(8), 1703.
Jaenson, T.G.T. and Lindgren, E. (2011) The range of Ixodes ricinus and the risk of contracting Lyme borreliosis
will increase northwards when the vegetation period becomes longer. Ticks and Tick-Borne Diseases
2, 4449.
distribution and abundance of the tick Ixodes ricinus during the past 30 years in Sweden. Parasites and
Vectors 5, 8, doi:10.1186/1756-3305-5-8.
Lindgren, E. and Jaenson, T.G.T. (2006) Lyme borreliosis in Europe: influences of climate and climate change,
epidemiology, ecology and adaptation measures. Report No. EUR/04/5046250, Regional Office for
Europe, World Health Organization, Copenhagen.
Lindgren, E., Tlleklint, L. and Polfeldt, T. (2000) Impact of climatic change on the northern latitude limit and
population density of the disease-transmitting European tick Ixodes ricinus. Environmental Health
Perspectives 108, 119123.
Materna, J., Daniel, M. and Danielov, V. (2005) Altitudinal distribution limit of the tick Ixodes ricinus shifted
considerably towards higher altitudes in central Europe: results of three years monitoring in the Krkonose
Mts. (Czech Republic). Central European Journal of Public Health 13, 2428.
Tlleklint, L. and Jaenson, T.G. (1998) Increasing geographical distribution and density of Ixodes ricinus
(Acari: Ixodidae) in central and northern Sweden. Journal of Medical Entomology 35, 521526.
7
1
Surveillance and Control Measures*
Rbert Farkas,1 Mo Salman2 and Agustn Estrada-Pea3

Szent Istvn University, Budapest, Hungary; 2Colorado State University,
USA; and 3University of Zaragoza, Spain
Tick surveys often are conducted to determine

whether tick control is needed or not. The
species that are collected can provide an indication of the potential risks of tick-borne diseases in a given area. The most common
method for sampling ticks is the use of a flag
or drag cloth pulled or dragged through the
vegetation. Ticks collected on the cloth are
counted as the number of a given species per
unit of distance dragged (e.g. 100 m). Absolute
measures of tick population densities cannot
be obtained with this method, although the
relative abundance of ticks in the sampling
area can be determined. An alternative to
dragging/flagging is the use of carbon dioxide traps. The gas is an attractant for crawling ticks, which then adhere to the trap.
When more reliable estimates of tick abundance are required, a mark-and-recapture
method can be used. Ticks can also be sampled from wild or domestic hosts, or from
people (Lindenmayer et al., 1991; Nicholson
and Mather, 1996). For tick-infested cattle,
horses, mules and other livestock, a timetested method is the scratching technique,
whereby livestock inspectors pass their
hands over different regions of the animals
body to detect attached ticks. A similar technique is used in combination with visual
inspection to examine wild animals. For

example, investigators can be assigned to
deer-check stations during the hunting season to count all ticks on hunter-killed animals.
Another technique for sampling ticks is to
trap small and medium-sized wild animals
and hold them over trays filled with water or
alcohol to catch fed ticks as they detach.
These methods provide more accurate information, but are expensive and labour intensive. Passive surveillance utilizes information
collected from tick laboratories or hospitals
to assess tick or disease distribution (White,
1993); this passive approach is less expensive
and requires less effort, and it can provide
useful information of appropriate types,
although the value of the results is sometimes limited by unidentifiable biases in data
collection (Johnson et al., 2004).
7.1
Introduction to Surveillance
and its Aim
7.1.1 What are surveillance and

monitoring?
A surveillance system is defined as the systematic, ongoing or repeated, measurement,
* Mo Salman wrote Section 7.1; Rbert Farkas, Sections 7.2, 7.3.1; Agustn Estrada-Pea, Section 7.3.2.
98
collection, collation, analysis, interpretation

and timely dissemination of health-related
data that is essential for describing health
hazard occurrence and for the planning,
implementation and evaluation of risk mitigation measures (OIE, 2010). The term monitoring is defined as the systematic, ongoing
or repeated, measurement, collection, collation, analysis, interpretation and timely dissemination information of health-related data
without an associated predefined plan of
action. Thus, the difference between these
two systems is related to the anticipated
action plan for the expected outcome from
the corresponding system. It should be
noticed that monitoring the presence, species, density and geographical distribution of
ticks is an essential component for an early
warning system for specific tick-borne
diseases (TBDs) in selected regions of the
world. This type of monitoring is considered
as part of a surveillance system for TBDs in
order to increase the likelihood that there
will be timely detection of undefined (emerging) or unexpected (exotic or re-emerging)
threats of TBDs.
A surveillance system for TBDs may
require a combination of measurements
related to the host as well as the ticks that are
involved in the transmission of the pathogens. The collection of tick specimens can
support at least the essential components of
the surveillance: (i) tick species identification;
and (ii) presence or absence of the pathogens
in the collected specimens. Tick collection,
however, should follow specific field protocol
in order to maximize the available specimens
to be collected.
The key feature of surveillance is that it
has an associated corrective action when the
presence of disease exceeds a predetermined
threshold. Because of the requirement of action
at certain thresholds, surveillance is a critical
component of control programmes (Salman,
2003). The surveillance process can be broken
down into three distinct components based on
the fact that it is an action-oriented programme.
There is a data gathering and analysis component via a defined monitoring system, a dissemination of the collected data or analysis
findings to stakeholders and a predefined
response or action component based on when
99
the findings of the data analysis exceed a predefined disease level threshold (Salman, 2003;
Thrusfield, 2007a). Further objectives of surveillance include identifying hypotheses for
research, evaluating the effectiveness of intervention programmes, disease forecasting, and
developing policy and public education campaigns (Akhtar and White, 2003).
This requirement that surveillance has
an action associated with a predefined threshold is its key difference from disease monitoring. Disease monitoring is the continuous
adaptable efforts used to assess the disease or
health status of a population and the risk factors associated with disease (Salman, 2003).
This can be a continuous effort to gather data
about general health or production metrics,
or it can be the monitoring of a specific health
or production metric. The data gathered
under general health monitoring are commonly known as vital statistics (Thrusfield,
2007a). The use of monitoring data is in its
ability to provide information on the prevalence of a particular disease or changes in the
rate or direction of the spread of that disease.
Typically, the same approaches used in surveillance are used in monitoring, and the
terms have been used interchangeably, but
the key difference is in the objectives of the
system (Salman, 2003). For example, the US
Department of Agriculture conducts monitoring programmes through its National
Animal Health Monitoring System in which
it investigates and collects data over time on
disease states, animal characteristics and production characteristics. This body of data can
be used to recognize a need for further
research or the need for a surveillance programme, but in and of itself is only informative. In monitoring, the objectives are purely
an observational undertaking.
7.1.2 Necessary components and
considerations of monitoring
and surveillance systems
A monitoring and surveillance system (MoSS) is
built on three basic principles. First is the identification of the disease(s) to be assessed based on
a risk assessment, the epidemiology of the
selected diseases, the disease pathophysiology
100
Rbert Farkas et al.
and the inherent agent characteristics (Zepeda

and Salman, 2003). This surveillance can be
general, focusing on a broad range of diseases
or syndromes, or specific to a particular disease of interest (OIE, 2010). Secondly, there is
the identification of the goals and objectives
of the surveillance system. These can be monitoring for trends, disease control or disease
eradication; the latter two tasks use surveillance systems specifically because an action is
implied. Therefore, thresholds need to be
identified that will result in action should a
disease-positive animal be found or should
population levels of disease exceed a predefined prevalence or incidence threshold
(Christensen, 2003; Salman, 2003). The identification of the sampling scheme must also be
decided upon, which also has an impact on
the type of data collection done: either passive or active. Finally, the limitations of diagnostic tests and other disease measurement
methods need to be incorporated into the system in order to understand its overall validity
(Stark, 2003; Jebara, 2004).
A risk assessment can be used to identify
the diseases of concern for an area. The protocol consists of the identification of disease
hazards, the characterization of those hazards and the likelihood of exposure to the
ranges of hazards or outcomes that can be
caused by a disease. Ideally, the assessment
quantifies the probability of experiencing the
full range of hazards associated with a disease threat. The hazards that are identified
can then be prioritized based on their public
health, production and international trade
implications. The risk management of the
prioritized hazards uses a monitoring and
surveillance system (Dennis et al., 2002). Risk
communication is the role of the OIE (World
Organisation for Animal Health), public
health systems and appropriate national
government organizations. The role of the
OIE in risk communications is through its
requirement that OIE-listed diseases and
significant animal health concerns be reported
to the organization on a weekly, monthly and
annual basis via the required reports for all
member states (OIE, 2010).
Monitoring and surveillance is demanding of resources. It does not just include
finding cases, but also requires the following
components: public education campaigns,

indemnities, producer awareness, response
resources such as vaccines, and the appropriate training of personnel involved in sample
collection
and
disease
determination
(Christensen, 2003; Salman, 2003; Jebara,
2004; Thrusfield, 2007a).
Proper training of personnel is even more
important when attempting to exclude or
prevent diseases with a public health and
socio-economic impact. This is because few
people have experience with the disease in
their day-to-day clinical life and they are
unfamiliar with its clinical manifestations.
The design of a surveillance system has
many parts that are necessary to define before
initiating a surveillance programme. These
are: an established case definition; standardized methods of collecting data; a defined
sampling method and sample size; an understanding of how the data will be analysed
and used by decision makers; methods of
communication of the results from the surveillance system; and an evaluation system to
allow for regular critiquing and improvement
of the programme.
Case definitions increase the specificity
of a surveillance programme, especially of a
disease-specific programme. Typically, a case
is defined by the clinical manifestation of disease and laboratory results. Definitions that
include the serological testing authorized to
determine suspect and confirmed cases are
most specific when the tests used are defined
and the order of laboratory tests is explicitly
stated (Akhtar and White, 2003; OIE, 2010).
Screening tests are used to identify suspect
cases. By nature, screening tests are typically
of higher sensitivity so as to reduce false negatives, but are also known for having lower
specificity than other tests and thus have a
higher rate of false positives. Therefore, it is
necessary to confirm suspect cases with tests
of higher specificity or diagnostic tests (Dohoo
et al., 2003b). Those developing the surveillance system should also consider the reproducibility of test sensitivity and specificity
between different laboratories, as well as the
laboratory burden and capacity to test for the
disease(s) of interest. Another issue that needs
to be considered for TBDs is the species identification and the detection of the pathogens
among the collected ticks associated with

specific TBDs.
Data collection methods and, more
importantly, storage and tracking methods
must also be standardized. Using the same
collection forms across all areas of the surveillance system will prevent differences in
reporting from different sources and will
increase consistency. This will also ensure
that all necessary information is captured and
reported by the reporting agencies. Lastly, it
will reduce errors when data are recorded
into a database by narrowing the need to
interpret different questions and answers as
to where they fit into the database. When a
standardized form is used, the data entry can
be standardized too (Thrusfield, 2007b). The
need for this standardization comes from the
need of the surveillance system and its results
to be reliable. Decision makers will be more
likely to consider the MoSS information when
they are confident that what it is reporting
has internal validity and, when necessary,
external validity as well.
External validity, or the extrapolation of
the surveillance results to the greater population of interest, is determined by the sampling
method (Dohoo et al., 2003c). There are different populations that can be considered for
surveillance based on whether probability or
non-probability sampling is used. Validity
requires probability sampling and, more specifically, random selection, sampling from the
population of interest, such as the general
population, and sample size.
The external validity of a programme is
greatest when a census of the population is
performed. External validity is important in
monitoring and surveillance when it is critical that the output from the sample population mirrors the true situation in the study
population. For example, estimating disease
prevalence would make this important. In the
situation of a census, all animals or people in
the population of interest are assessed for the
presence of disease (Cameron et al., 2003).
Populations are groups of interest with a
common characteristic. If populations of
interest are quite small, this may be manageable, but if they are quite large, such
censuses are not feasible. Yet, when the population of interest is too large to census, then
101
representative sampling is needed to estimate

the level of disease in that larger population.
If the sample is not representative of the population of interest, then the results of the
surveillance within that sample will be also
invalid. The most reliable method to ensure a
representative sample is to use random sampling techniques (Cameron et al., 2003).
Random sampling is sampling which
ensures that the probability of being selected
is the same for every entity in the study
population (Cameron et al., 2003). Random
sampling often results in representative samples and allows for statistical evaluation of
the results as well (Dohoo et al., 2003a). This is
also known as probability sampling. Nonrandom sampling or non-probability sampling, such as purposive or convenience
sampling, can also be used for specific events,
but not for measures of disease occurrence.
Targeted samples are a type of purposive
sampling sampling of populations at high
risk. Convenience sampling can save resources
and time. Yet neither of these methods yields
representative samples of the greater population (Cameron et al., 2003).
Sample populations are more advantageous than censuses for multiple reasons.
They are smaller, less costly and more manageable, while being representative of the
study population. The results from an appropriately selected sample population can
then be extrapolated to the study population
without conducting surveillance on the entire
population (Cameron et al., 2003; Dohoo et al.,
2003a). For measures of disease such as prevalence, a census gives you a specific and reliable measure of disease, but a sample gives
you a value and its variability. The variability
is the measure of uncertainty that comes with
using a sample of the population to estimate
the level of disease rather than using a census
to assess the entire population. This variability can be reduced as the sample size increases
(Dohoo et al., 2003a). If one is using the surveillance system to assess risk factors, a certain sample size is needed to assure that there
is truly no association when the results find
no association; this is the concept of the power
of a study and is also related to sample size.
Sample size is determined through calculations that consider the allowable error,
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Rbert Farkas et al.
the confidence of the estimate, the variability

of the measure of interest in the population
and the power wanted in the study (Cameron
et al., 2003; Dohoo et al., 2003a). Other factors
such as the study population size and the
accuracy of tests must also be considered.
However, there are also other considerations when evaluating disease presence.
Diseases often cluster based on political,
social and environmental factors. This clustering needs to be considered not only in
relation to the sample size determination,
but also the sampling method. If it is not considered, there are three possibilities. First,
the surveillance may accurately estimate the
level of disease by chance. Secondly, the
sampling method oversamples from clusters
without disease, thus underestimating disease. Thirdly, the sampling method may
oversample from clusters with disease, and
thus overestimate disease (Buehler et al.,
2003). In terms of sample size, the greater the
similarity of the animals within the clusters
in terms of the measure of interest, the greater
the sample size (Dohoo et al., 2003a). Clearly,
clustering has many impacts on a surveillance system that must be considered.
The results of a surveillance system are
used for multiple purposes. The collection of
the data should be fitted to the desired analysis instead of fitting the analysis to the data
collections. This requires that the purpose of
the data is predetermined. It also allows the
data to appropriately feed into pertinent
statistical analysis, spatio-temporal analysis
or simulation models that are often used for
risk assessments. The impact of a surveillance
system depends upon the usefulness of its
output. If the output does not measure
variables of interest, the surveillance system
is limited in its overall utility.
All systems should be continuously evaluated and improved to meet current needs and
threats. Assessment and validation of a system
must consider the needs of the stakeholders,
the ability to detect disease and the usefulness
of the data to the stakeholders. In order to adequately assess a system, the objectives and
aims of the system, the disease under surveillance, the legal basis of the programme, the
various components of the programme and the
responsible organizations need to be defined.
By using these definitions and the needs of the
stakeholders, even as they change, the baseline

purpose of the surveillance assessment can be
defined. A needs assessment of the stakeholders can be used to identify areas of weakness in the surveillance system or areas that
need to be addressed that currently may not be
in the system (Martell-Moran et al., 2011).
There are numerous methods for conducting assessments. These include questionnaires
of stakeholders, qualitative methods such as
graphical methods, descriptive assessments or
classification analysis and quantitative methods such as scoring systems. Graphical methods include fault tree analysis and scenario
analysis. Classification analysis is a descriptive
assessment of the surveillance system through
its component parts. If standardized, classification analysis can compare various surveillance
systems, such as those in different regions of
countries, or a surveillance system over time.
The next step in a classification analysis is to
assign a scoring system to the effectiveness of
each component, thereby making it a quantitative analysis (Stark, 2003). Lastly, another
potential quantitative assessment is via benchmarks or indicators. These can supply quality
assurance to the system too by setting benchmarks for the number of tests run or disease
reports generated in a period of time (Stark,
2003). Of course, benchmarks are meant to be a
tool, and the ability to reach them must adhere
to sufficient quality standards as well.
Finally, the results of the analysis from a
surveillance system must be communicated to
all stakeholders; otherwise the entire programme is for naught. An important component of dissemination of the results is in
identifying all of the critical stakeholders in
federal, state and local governments, industry,
private citizens where appropriate (pet owners, domestic flock owners), veterinarians,
physicians, laboratories and the general public (Martell-Moran et al., 2011). Furthermore,
the form in which the data are distributed to
the various stakeholders may need to differ
based on how such stakeholders will use the
information. The general public may need
information on how to protect themselves
from disease; producers may need information on how to enhance their biosecurity in
the face of an epidemic; and veterinary public
health personnel may need information on
the rate of spread and scope of the disease to
implement appropriate disease control strategies on a regional or national level.

As different populations with different
backgrounds will be using the surveillance
results, there need to be various reports that
present different types and levels of information: public information requires the statement
of easily understandable facts about the disease situation; for those familiar with surveillance systems and epidemiology, reports using
tabular summaries that require some level of
interpretation can be used; and for stakeholders and policy makers, interpretative reports
are needed that include a description of the
programme and the results (Wineland and
Dargatz, 2003). The latter reports are useful to
policy makers and industry key players who
have a larger stake in the results of the surveillance system, but do not have the appropriate
training to interpret the data alone.
There are numerous components and considerations that go into a surveillance system
and its methods. Without consideration of all
these parts, from development to assessment
and dissemination of the data, the surveillance
system will fail to reach its objectives or purpose.
A comprehensive surveillance system should be
reasonably sensitive and specific for the disease
or diseases of interest, have an adequate positive predictive value especially in the face of
endemic diseases, be timely and useful to the
stakeholders for developing policy, and be able
to dictate when interventions for disease control should be implemented in biosecurity and
bio-containment programmes. In order to fully
advise stakeholders the programme must be
timely, acceptable to those who provide the data
and those who use the data, accurate in its depictions of the disease situation/s, and even flexible
in its ability to adapt as situations change
(Akhtar and White, 2003). There is no right
answer or recipe for a surveillance system,
but these guidelines can assist in the development of an effective programme.
7.1.3 Types of data collection methods
and surveillance techniques
Passive and active data collection systems are
both useful, but their usefulness depends
upon the purpose of the surveillance programme. Passive surveillance is the collection
103
of laboratory results, tick specimens, slaughterhouse information or reporting by clinicians (veterinarians and physicians) through
health departments or similar organizations.
The data are second-hand and generated for
purposes of diagnosis and not for the sole
reason of surveillance. Making critical diseases notifiable can assist in ensuring that
they are better reported and recognized by
clinicians and veterinarians. Passive data
collection is useful for recognizing trends
and emerging diseases, but it should not be
relied upon as a rapid early warning system.
Active collection requires data collection from
the apparently normal population for the sole
purpose of surveillance and monitoring
(Akhtar and White, 2003; Salman, 2003;
Thrusfield, 2007a).
In active surveillance, the sampling
frame can be derived from different types of
populations. These are the general population, sentinel populations and targeted populations. General populations are best used
when the disease is very common and can
easily be found with a reasonable sample size
that is economically feasible within manpower and laboratory constraints. General
population surveillance makes use of random
sampling and systematic sampling to assess
the level of disease in a population (Thrusfield,
2007a; OIE, 2010). General population-based
surveillance programmes allow for the calculation of incidence and prevalence if the
sampling is representative. This is based on
the concept that a representative sample of
the population at risk is followed over a time
period and all new cases in the representative
sample are identified. The results can, therefore, be extrapolated to the general population because there is external validity. The
more common a disease is in a population
the smaller the sample size that is needed.
Sample sizes increase as the disease becomes
rare (Salman, 2003). Thus for rare diseases
alternative sampling populations are used.
For very rare diseases that are being eradicated, a targeted sample population will most
likely need to be used. The targeted population should have a higher prevalence of
disease than the general population (Cameron
et al., 2003). This reduces the sample size and
hence the burden on resources needed to successfully eradicate disease.
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Surveillance programmes can also use

sentinel populations. Sentinel populations
are useful in programmes of disease exclusion or prevention. Sentinel populations are
populations that are monitored for the presence of a seasonal or foreign animal disease.
These populations should have a higher rate
of disease than the general population
(McCluskey, 2003). If the sentinel population
is sufficiently sensitive to disease, it can act as
an early warning system to the presence of
disease (Rabinowitz et al., 2006). Further, sentinel populations can be used as a public
health tool, acting as an early warning system
when the sentinel population incubation
period is shorter than the human incubation
period for the disease. When sentinel populations indicate the presence of a zoonotic
disease, this finding can be used by the public
health professionals in the area as a way to
alert the human population to the disease
threat. West Nile fever and eastern equine
encephalitis in horses act as an indicator for
the human population in this way.
Participatory epidemiology is a specialty
that uses local knowledge to understand the
local disease situation. Often, when surveillance is done using participatory epidemiology, syndromic surveillance is utilized. The
reason for such an approach is that in underresourced countries there is often poor laboratory support, poor access to remote regions
and a lack of appropriate levels of manpower
(Thrusfield, 2007a). Syndromic surveillance
requires a detailed clinical case definition.
The clinical case definition is the only method
of capturing all potential suspect cases.
Ideally, it is tailored to a specific disease. The
purpose of syndromic surveillance is to capture all of these potential cases so that laboratory confirmation (in a field or diagnostic
laboratory) can be carried out. Such participatory disease surveillance programmes were
used in the Rinderpest eradication campaign,
which determined the absence of disease
in remote locations (Jost et al., 2007).
7.2
Control Options for Hard Ticks
Tick control has stimulated intense interest

for more than a century. There is a variety of
strategies for the control of ticks and TBDs

(Sonenshine et al., 2006; Willadsen, 2006). Tick
prophylaxis should cover the entire period
during which ticks are active. Treatment with
synthetic chemicals known as acaricides is
still the most widely used means to control or
prevent hard tick attacks, and has the aim of
preventing pathogen transmission (Polar
et al., 2005). Control of ticks with acaricides
can either be directed against the ticks on the
host or against the free-living stages of those
ticks in the environment.
7.2.1
Chemical control of ticks on animals
Acaricides include several groups of pesticides:

organophosphates (e.g. coumaphos, diazinon),
carbamates (e.g. propoxur), pyrethroids (e.g.
permethrin, deltamethrin, flumethrin), formamidines (e.g. amitraz), two classes (avermectins
and milbemycins) of macrocyclic lactones (e.g.
ivermectin, doramectin, moxidectin, eprinomectin), phenylpyrazoles (e.g. fipronil) and
natural acaricides such as botanical materials
(e.g. extracts of the neem tree containing
azadirachtin).
Several methods are used for the application of acaricides to animals, such as
dips, sprays using manual or motorized highpressure sprayers, dusts, pour-ons, spot-ons,
the intraruminal bolus and injections. Dipping
vats have been used extensively for tick control on livestock including cattle and sheep.
When these are used for several animals over
extended periods of time, dipping vats need
to be maintained with the proper solution
levels and percentage of active ingredient.
Small numbers of livestock can be treated
with hand-held sprayers but manual spraying depends on the skill of the person applying it. Retreatment may be needed several
times during a season when sprays and dips
are used. Pour-on formulations of acaricides
contain high-quality oil that spreads through
the greasy hair coat of livestock, and these
products can also be used with applicators to
treat wild ungulates in game reserves (Latif
and Walker, 2004). Pour-on formulations are
relatively expensive, but there is little wastage of acaricide and they may be cheaper in
the long term. The other topical application
methods are often employed owing to their

advantage in limiting the amount of acaricide
used; the cattle pass through a heavy low
pressure spray and become soaked with
acaricidal fluid, but body parts such as the
ears and groin may not be effectively treated
by this method (Latif and Walker, 2004). To
achieve long-lasting efficacy, acaricides can
be incorporated into plastic or other suitable
matrices that provide a slow release of the
toxicant over a period of weeks or months.
Systemic acaricides offer another means of
providing long-lasting and effective tick control. These preparations can be divided into
injectable, oral and topically applied products, all of which are delivered to the tick
during its feeding activity on the skin. Each
application method has its advantages and
disadvantages.
Oil, creams or alcohol should not be used
for killing the ticks. The application of acaricides is advisable because not all of the ticks,
especially the larval and nymphal stages and
unengorged adults, may be detected on the
animal. When treating dogs and cats against
ticks, only approved veterinary medicinal
ectoparasiticides should be used. There are
several different formulations of acaricides
for pets, and both dogs and cats can be
sprayed with approved products. Topical
spot-on treatments containing fipronil or
selamectin can be used on both dogs and cats;
however, permethrin can be used only on
dogs. Insecticide-impregnated pet collars are
also available for dogs and cats, although
those containing amitraz cannot be used on
cats. These collars offer at least partial protection but may not provide total protection
against ticks (Elfassy et al., 2001; Spencer et al.,
2003). Dogs and cats travelling to regions
with ticks and endemic TBDs should also
receive a regular application of acaricidal
products, particularly if those TBDs are not
endemic in the home country. To advise pet
owners and improve owner compliance, the
duration of efficacy for an individual product
should be established from the relevant product data sheet so that the owners are given the
correct retreatment intervals. It is advisable
that animals are checked regularly, in particular towards the end of the protection period,
to ensure that any visible ticks are removed
105
and that early repeat treatment is considered

if appropriate. It should also be remembered
that the duration of efficacy of an acaricide
may differ between tick species, again highlighting the importance of visual checking to
verify that the treatment remains effective
(ESCCAP, 2009). If permanent infestations
with Rhipicephalus (Rh.) sanguineus occur
within a kennel or household environment,
regular acaricidal treatment of pet animals
coupled with environmental treatment using
a compound from a different chemical group
can be used. There are special formulations of
acaricides for use in kennel or household
environments. The active compounds mostly
belong to the same chemical classes as the
acaricides used for pets.
For wildlife, an important concept called
host-targeted tick control has been introduced
in recent years as a way to destroy ticks without harming their hosts. The hosts can be
attracted (using feed, nesting materials or
other attractants) to devices that apply acaricides to them. Several such devices have been
developed to attract deer or small mammals
to feeding stations where they are treated.
Examples include bait boxes, permethrintreated cotton balls, and so-called four-poster
devices (Stafford and Kitron, 2002). One of
the most promising of these devices is the
four-poster device. It includes a central bin,
containing bait to attract deer, with a bait
dispenser and applicator station on either
side. As the deer inserts its head to feed on
the bait, it contaminates its fur with acaricide
from the applicators. Four-poster devices
have the advantage of providing well-targeted
applications using far lower amounts of pesticide than are used in broadcast applications
(Pound et al., 2000). The effectiveness of any
tick-control approach tends to depend on the
ecological conditions at the application site,
but these methods can be important tools in
integrated pest management programmes,
especially when combined with other management techniques appropriate for the local
conditions of tick distribution and transmission dynamics. Another novel hosttargeted method is the use of rodent-targeted
bait boxes containing fipronil. This has
proven effective in eliminating immature ticks
on mice, thereby reducing the subsequent
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Rbert Farkas et al.
population of questing nymphs and adults,

as well as reducing the proportion of these
ticks infected with the Lyme disease spirochaete Borrelia burgdorferi (Dolan et al., 2004).
The chemical control of ticks on livestock poses several problems. Besides leaving residues in meat and milk, the use of
many acaricides (e.g. organophosphates, carbamates) is associated with risks of side
effects or poisoning resulting from overdoses on specific species or from breed
sensitivity (de Oliveira Monteiroa et al.,
2010). Environmental contamination and
effects on non-target animals have also been
well documented for organochlorines, organophosphates, carbamates and pyrethroids
(Kunz and Kemp, 1994). The development of
acaricide resistance by some tick species is an
increasing and continuing concern in tropical and subtropical areas, where these ticks
have been found to be resistant to chlorinated hydrocarbons, organophosphorus
insecticides, pyrethroids and formamidines
(Kunz and Kemp, 1994). Acaricide resistance
in tick species occurring in Europe has not
yet been reported, but to avoid the onset of
this problem there are certain rules that
should be followed:
When acaricides are used they should be

stored as recommended and used as soon
as possible so that they are of full strength
as specified by the manufacturer.
All ticks on treated animals should be
destroyed or eliminated.
The acaricide type recommended by veterinary authorities should be used until
official advice to change to another type
is given.
Novel acaricides should only be used when
the older types of acaricides have become
ineffective (Latif and Walker, 2004).
For the above-mentioned reasons, major

alternatives to conventional acaricide treatments have been developed in recent years
which can be used in integrated management programmes for hard ticks in order
to minimize the environmental impacts
and acaricide resistance, and to reduce
production costs (Samish et al., 2000). The promising alternative methods described in
Sections 7.2.27.2.8 are being investigated.
7.2.2
Pheromone or hormone-assisted
control
Pheromone-assisted tick control is one of the

novel strategies being used for the control of
hard ticks. Research with tick pheromones
suggests that combinations of pheromones
and acaricides can be significantly more effective in controlling ticks than the acaricide alone
because ticks are unlikely to develop resistance to their own pheromones. A pheromone
acaricide combination applied to a single spot
on cattle can be effective in killing ticks.
Another promising device is the tick decoy,
in which the sex pheromone 2,6-dichlorophenol and an acaricide are impregnated into
plastic beads on the surface of which mounting sex pheromone is smeared. Male ticks are
attracted to these decoys on the animals hair
coat and killed. This approach also disrupts
mating activity so that any surviving females
cannot lay viable eggs (Norval et al., 1996).
Another novel technology for killing Ixodes
scapularis ticks in their natural habitats was
developed by incorporating the components
of the tick arrestment pheromone (guanine,
xanthine and haematin) along with permethrin into an oily matrix for dispersal on vegetation. These paste-like droplets attracted
and killed ticks before they could infest
humans or animals (Sonenshine et al., 2003,
2006; Sonenshine, 2008). Hormones and
insect growth regulators (IGRs) such as
methoprene have also been used to disrupt
tick development in laboratory experiments.
Analogues or mimics of ecdysteroids and
juvenile hormone are effective in killing ticks
by delaying their development, disrupting
oviposition or killing the larvae when they
hatch from eggs deposited by treated
females. However, these compounds do not
appear to be uniformly effective against all
types of ticks.
7.2.3
Biological control
Biological control of ticks seems an appropriate alternative that may reduce the frequency
of use of chemical acaricides and the need for
treatment of TBDs. Biological control agents
are in principle highly desirable, but their
narrow host specificity, often relatively low

efficacy, costs of manufacture, certain application problems and sometimes low stability
present serious challenges. Numerous pathogens attack ticks, including bacteria, fungi
and nematodes (Samish et al., 2004). Several
papers have reported the testing of entomopathogens for the control of ticks in laboratories, and the use of these novel biocontrol
techniques on animals (Alonso-Daz et al.,
2007) or vegetation (Kaaya, 2000). Among the
potential biocontrol agents, entomopathogenic fungi, nematodes and parasitic wasps
are the most promising candidates (Samish
and Rehacek, 1999).
Entomopathogenic fungi have been studied as control agents of ticks mainly in laboratory assays (Kaaya, 2000; Samish et al., 2001,
2004; Polar et al., 2005). These fungi invade
their host by direct penetration of the cuticle.
After germination of the attached spore,
appressoria are formed which then secrete
histolytic enzymes and produce hyphae.
After successful penetration by the hyphae,
yeast-like blastospores are formed and propagate. After the death of the host, the fungus
grows out of the cadaver and sporulates
(Kleespies, 1993). The comparative ease with
which the spores of these fungi can be produced and artificially disseminated makes
them promising potential agents for the control of ticks (Norval and Horak, 2004). It has
been reported that temperature and host
secretions (e.g. sweat) may affect the virulence of entomopathogenic fungi on animals
treated with biopesticides to control ticks
(Polar et al., 2005). Alternative targeted methods of applying fungal pathogens to the
environment for the control of ticks are
needed. Studies of the effects of entomopathogenic fungi on ticks under field conditions
are scarce (Benjamin et al., 2002; AlonsoDaz et al., 2007), but at present one of the best
candidates for tick biocontrol is Metarhizium
anisopliae (Samish et al., 2001). Preliminary
field trials have had modest results. The use
of an oil-based carrier enhanced tick mortality when compared with a water-based solution (Kaaya and Hassan, 2000). The pathogens
that affect ticks typically also affect other
arthropods (Brownbridge and Glare, 2007),
so effects on non-target arthropods must be
107
considered in strategies for the application of

biocontrol materials. There are opportunities
to use auto-dissemination devices to deliver
pathogens to ticks (Maniania et al., 2007).
Such devices use visual cues, pheromones
and kairomones to attract host pests to a
pathogen source (Vega et al., 2000).
A promising alternative to the use of
entomopathogenic fungi is biological control
through the use of entomopathogenic nematodes (EPNs) (Samish et al., 2001). The thirdstage infective larvae of EPNs of the families
Heterorhabditidae and Steinernematidae live
freely in the soil. They enter their hosts through
natural body openings such as the genital pore
and release symbiotic bacteria carried in their
intestines (Kocan et al., 1998). These bacteria
proliferate in the haemolymph and produce
toxins and other metabolites. The tick then
dies from septicaemia and the bacteria produce suitable conditions for the nematodes to
complete their life cycle (Ehlers, 2001).
Arthropods can also be used for biological control of ticks: chalcid wasps of the genus
Ixodiphagus are obligatory parasitoids of ixodid ticks and most species will oviposit and
develop only in the nymphal stage of the tick.
Several wasp larvae can successfully develop
in a single engorged nymph, which is killed
during this process (Mwangi et al., 1997; Hu
et al., 1998; Norval and Horak, 2004).
Domestic chickens are opportunistic predators of ticks and can be used for tick control in
rural areas. In particular, the indigenous breeds
of Galliformes, if allowed to scavenge among
cattle, can consume considerable numbers of
ticks (Latif and Walker, 2004).
7.2.4
Genetic resistance
In general, tick tolerance or tick resistance

exhibited by certain livestock breeds might
vary with the species of infesting tick; heterospecific resistance appears to be low or even
absent among different genera of ticks, while
a certain degree of cross resistance is expressed
to tick species belonging to the same genus
(de Castro and Newson, 1993). Genetic resistance has been described in West African
NDama cattle. This breed has a higher degree
of natural resistance to ticks with a long
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Rbert Farkas et al.
hypostome, such as some Amblyomma and

Hyalomma species, than to tick genera with a
short hypostome. In Bos indicus breeds, the
evidence for a genetic resistance trait to multihost ticks, such as Rh. appendiculatus, is not as
strong as for the one-host ticks, i.e. Rh. microplus (Mattioli et al., 2000).
(de la Fuente and Kocan, 2006). Other promising vaccines target tick-cement protein, disrupting the attachment success as well as
causing midgut injury and affecting the ticks
ability to transmit pathogens (Labuda et al.,
2006); affect the ability to disrupt the male
engorgement factor; or apply a combination
of anti-tick and anti-pathogen vaccines
(Sonenshine et al., 2006, Willadsen, 2008).
7.2.5 Vaccine
In Australia, a commercial recombinant antigen vaccine has been developed for the control of the cattle tick Rh. (Boophilus) microplus
based on a so-called concealed antigen (Bm86)
in cells of the tick gut. A similar recombinant
vaccine has been developed in Cuba. Recent
reports suggest that the recombinant Bm86
can reduce tick fecundity by as much as 90%
(Willadsen, 2006, 2008). Although it is possible that antigen-resistant strains of cattle ticks
may appear, large-scale vaccination of cattle
herds with these recombinant vaccines offers
a promising alternative or supplement to acaricides. Although the vaccine uses an antigen
from Rh. (Bo.) microplus, it is even more efficacious against Rh. (Bo.) annulatus than against
the homologous species (Fragoso et al., 1998).
Such effects appear not to correlate with the
degree of sequence conservation of the antigen across tick species (Willadsen, 2006).
Because vaccines are expensive and
involve considerable risk, a high level of efficacy is required to offset these negative
aspects of their use. Research on other antigens and other tick species is in progress.
Many more potential antigens have been proposed than have been tested. Tick antigen
targets studied to date are from a restricted
range of functional classes. They include
structural proteins, particularly from salivary
glands, hydrolytic enzymes and their inhibitors, especially those involved in haemostatic
processes, and a range of membraneassociated proteins of unknown function
(Willadsen, 2006, 2008). Of special interest is
the development of novel combinations using
RNA interference (RNAi) to silence subolesin
and the tick-protective antigen Rs86 (similar
to Bm86) against Rh. sanguineus; the synergistic effect of silencing both genes causes a
much greater reduction of tick feeding and
oviposition than targeting either one alone
7.2.6
Personal protection
Preventive measures are the most effective

means of protection from ticks. Ticks can
be avoided by avoiding exposure to fields,
forests and other tick-infested habitats, especially in areas of known disease foci (Ginsberg
and Stafford, 2005). Maintaining a shortclipped lawn in gardens and parks, and establishing barriers to prevent access to woods
can minimize human exposure to ticks in
these environments. The use of clearly defined
paths can help people avoid contact with tickinfested vegetation. People in tick-prone areas
should wear boots, socks, long trousers and
light-coloured clothing. Trousers should be
tucked into the boots, socks drawn over trousers, and the socks taped to form a tight seal.
A recent study showed that wearing protective clothing was 40% effective in preventing
Lyme disease (Vsquez et al., 2008). Clothing
should also be treated with an effective repellent or acaricide such as permethrin before
entering tick-infested habitats. Exposed skin
also should be treated with repellents or acaricides suitable for use on humans. The most
widely used personal protectant is the repellent DEET (diethyl toluamide or N,N-diethyl3-methyl benzamide), which is available as a
lotion or a spray. Repellents should be applied
repeatedly as needed to maintain maximum
protection, but because of absorption or abrasion the interval should be longer than a few
hours. Each person should conduct selfexaminations for ticks during and after exposure to tick-infested areas. Early removal of
attached ticks is important in minimizing the
risk of contracting TBD. There is a large variety
of purpose-designed tick removal tools available, and these may be used for the removal of
ticks attached to the skin. Oil, creams or alcohol should not be used for killing the ticks.
7.2.7
Habitat modification
Tick control primarily involves the treatment

of animals. However, habitat modifications of
the infested land can aid in reducing tick
abundance. Ticks have species-specific habitat
requirements that are often associated with
habitats of hosts and the need to avoid desiccation. Therefore, habitats can be manipulated to make them unsuitable for ticks or to
minimize encounters between ticks and
animals and people (Stafford, 2007). Several
acaricides, including organophosphorus and
synthetic pyrethroid preparations can be
applied directly to vegetation in gardens,
parks and other areas that are suitable sites
for ticks. Using acaricides off the host is of
limited value because ticks commonly occur
in microhabitats covered by vegetation, leaf
litter and other natural materials, or in the
nests, burrows and other cavities used by
their hosts. For these reasons, the ticks often
do not come into direct contact with acaricides. Therefore, the chemicals must reach the
ticks when they are seeking hosts. However,
public opposition to the treatment of natural
habitats with pesticides has made it unpopular to use this form of tick control. In the
European Union (EU) it is prohibited to use
any acaricides in the environment.
Selective grazing and pasture rotations
often reduce cattle exposure to tick populations. By keeping animals out of certain
pastures (e.g. by the removal of deer by hunting and deer-exclusion fences), the number of
fed ticks that would serve to build up a population is reduced (Daniels et al., 1993). Although
lowering deer populations by hunting can
also lower tick numbers, this approach is not
generally practical, because deer populations
must be reduced to extremely low levels to
have a reliable effect on the transmission of
tick-borne pathogens (Ginsberg and Stafford,
2005). An extreme case of pasture management is zero grazing by dairy cattle, but there
is a risk of the unexpected reintroduction of
ticks on cut fodder, on wild mammals or birds,
or on newly introduced animals. This practice
of zero (or non-continuous) grazing has
often been referred to as pasture spelling
(Ginsberg and Stafford, 2005). Mowing and
burning vegetation in natural areas also lowers tick numbers temporarily, but ticks reinfest
109
treated areas as the vegetation grows back

(Wilson, 1986).
For minimizing human and pet exposure
to ticks in outdoor areas, efforts should first
be made to keep overgrown and heavy vegetation cleared and cut in potentially tickinfested areas. A study on Lyme disease New
York state showed that lawns that were cut
short and were open to the sun had minimal
numbers of ticks, while tick densities
increased incrementally in gardens, wood
edges and forests (Maupin et al., 1991). Indoor
tick control, as well as tick control in and
around kennels, with cleaning or changing of
pet bedding to prevent or remove ticks, is
directed primarily at the brown dog tick,
Rh. sanguineus.
7.2.8
Integrated tick management
There is no single, ideal solution to the control

of ticks, but integrated control scenarios
involving increased scientific and practical
complexity can be developed and recommended. The integrated approach is probably
the most effective way to control ticks
(Jongejan and Uilenberg, 1994; Willadsen,
2006). Integrated pest management (IPM) is
an approach to the management of arthropod
pests that fosters the integration of various
pest control methods so as to minimize reliance on individual environmentally damaging approaches and to provide sustained
management of pest populations. Given the
many tick control techniques currently available and the numerous novel techniques
being developed, it is important to refine the
theory and practice of integrating these methods so they can be efficiently and effectively
applied to prevent human and animal
disease.
In conclusion, controlling hard ticks can
be an effective way to break the cycle of the
tick-borne pathogens and subsequently to
control the spread of disease. There are
numerous conventional and relatively new
approaches to control hard ticks and their
spread, and several factors should be considered in determining the most effective options.
These factors can be summarized as: the tick
species involved, the animal species involved,
the environmental and ecological conditions,
110
Rbert Farkas et al.
animal management and the feasibility of

the measures. Integration of these factors is
required in order to most efficiently control ticks and the potential pathogens that
they carry.
7.3 Control Possibilities for Soft Ticks

7.3.1 Argasidae
The control of argasid ticks is a difficult
task because they possess a range of biological features (e.g. morphological, physiological and ethological) that make them
difficult to eradicate. These soft ticks can
form long-term aggregations deep inside
the fissures, cracks and crevices of buildings where it is difficult to spray with
chemicals to destroy them. Thus, efforts to
eradicate these ticks from old buildings can
be extremely costly and are generally
unsuccessful.
In the case of Argas reflexus (the European
pigeon tick), a low level of metabolism that
allows the resting tick to close its spiracles for
several hours, or perhaps even for days,
might help this particular species to withstand the application of toxic gases (Dautel
et al., 1994). Even when using long-term residual pesticides, A. reflexus is difficult to exterminate owing to its long life cycle. A successful
control measure of this tick species usually
involves three stages (Dautel and Kahl, 1999):
(i) the complete uncovering of all potential
tick hiding places (in some instances, plaster
must be removed from the walls); (ii) the
application of a pyrethrum spray to expel
ticks; followed by (iii) the use of a propoxurbased contact pesticide. A second control
measure is often indicated 34 months later
because of the resistance of the egg stage to
control. Indeed, repeated treatments by a pest
control professional are often necessary
(Herrmann, 2000).
Several pesticides have been reported
to be effective against argasid ticks, including permethrin (Dusbabek et al., 1997),
ivermectin (Montasser et al., 2005), and
Bacillus thuringiensis (Hassanain et al., 1997).
It was reported that entomopathogenic
fungi have also good potential for the control
of A. persicus (Sewify and Habib, 2001).

Repellents (e.g. permethrin and DEET)
have been shown to be effective against
A. persicus, the fowl or poultry tick (Kumar
et al., 1992; Haag-Wackernagel, 2005).
No vaccine against argasid ticks exists yet.
The various alternatives need further
investigation because no effective method
for the long-term control of these ticks is
known. At present, the only practical measure is to avoid housing domestic fowls in
old, infested, buildings and to prevent
access to this type of building by freeranging herds. In order to control argasid
ticks in poultry houses, all cracks and
crevices which provide shelter to these
ticks must be eliminated.
7.3.2
Ornithodoros
Eradication of O. erraticus from old pig sties

is invariably unsuccessful. This is because
of the ticks long life and long survival
without feeding, the existence of accidental
hosts other than pigs, and the possibility of
penetrating into the cracks and surfaces of
the buildings where the ticks are not accessible to acaricides or to the use of a fire
torch on these surfaces or on the ground.
These factors have been the major reason
for abandoning O. erraticus-infested pig
farms after an outbreak of African swine
fever (ASF), and they have led to avoiding
the use of this type of building to shelter
pigs on the Iberian peninsula (Boinas,
1994). The use of endectocides in pigs can
reduce the level of infestation in the
premises, but does not prevent infection by
the ASF virus in the pigs if they are bitten
by an infective tick.
Practical measures include not housing
pigs in infested buildings. When such buildings are located near a pig herd, the premises
can be isolated with fences to prevent access
by the pigs, or the buildings can even be
destroyed and new premises built in another
location (Arias and Snchez-Vizcano, 2002).
No effective vaccine against the ticks exists
but there are promising studies on salivary
glands extracts and concealed gut antigen
extracts (Astigarraga et al., 1995, 1997;
Manzano-Romn et al., 2006, 2007).
111
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Glossary
Argasid ticks: soft ticks.

Co-feeding: a phenomenon in which ticks become infected with a pathogen during feeding
adjacent to infective ticks on the same vertebrate host, even when the vertebrate host has not
developed a systemic infection.
Competence: the ability of a vector to transmit a pathogen to a susceptible host in such a way
that the host becomes infected.
Conscutum: the hard, sclerotized protective shield (plate) which covers most of the dorsal
surface of Ixodidae males.
Diapause: a neurohormonally mediated dynamic state of low metabolic activity. Associated
with this is reduced morphogenesis, increased resistance to environmental extremes,
and altered or reduced behavioural activity. Diapause occurs during a genetically determined stage(s) of metamorphosis, and its full expression develops in a species-specific
manner, usually in response to a number of environmental stimuli that precede unfavourable conditions. Once diapause has begun, metabolic activity is suppressed even if conditions
favourable for development prevail.
Enamelling: is often called ornamentation. It is most conspicuous on the conscutum of males.
The colour is mainly pink, orange or red; the enamel looks like paint on the surface of the
integument.
Endophilic and Exophilic: when not feeding, endophilic (= nidiculous) ticks live in the nest,
burrow or den of the host. Exophilic ticks live in the open environment away from the hosts
nest or burrow.
Ixodid ticks: hard ticks.
Kairomone: a compound (chemical messenger) emanating from a potential host, e.g. carbon
dioxide in ox breath that may induce appetitive behaviour in blood-feeding arthropods (ticks,
mosquitoes, etc.). Kairomones enable an interspecific system of chemical communication.
Monotropic: ticks which as immature stages or adults feed on the same type of host.
Nidiculous: endophilic and nidiculous are synonymous.
Questing: many exophilic ticks cling to plant stems or similar substrates and await passing
animals. The ticks front legs, which have chemosensillae, are held out. Such questing ticks
may be collected by flagging or dragging a cloth to which they try to attach.
Reservoir: an animate or inanimate object on or in which an infectious agent usually lives, and
which therefore is often a source of such an infectious agent.
115
116
Glossary
Scutum: the hard plate on the anterior dorsal surface of the larva, nymph and adult female
ixodid tick. Soft ticks (Argasidae) do not have a scutum.
Telotropic: when the immature stages of a tick are able to feed on both different types of hosts
and the same types of host as the adult ticks. For instance, both rodents and ruminants can
support the feeding of immature stages of a certain telotropic tick.
Transovarial transmission: the transmission of microorganisms, including viruses, from
mother to offspring via the ovaries.
Trans-stadial transmission: the passage of microorganisms and viruses in arthropods from
one stage (stadium or instar) to the next.
Appendix 1 Tables of Geographic

Distribution Data for Ticks
and Tick-borne Diseases
Table A1.1. Geographic distribution data for African swine fever (ASFV). See Appendix 2 (hosted on
http://www.cabi.org/vetmedresource/) for complete reference indicated by the Reference ID.
Country
Administrative region 1
Armenia
Armenia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Italy
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Russia
Russia
Russia
Russia
Russia
Russia
Spain
Spain
Lori
Tavush
Ajaria
Guria
Imereti
Kakheti
Kvemo Kartli
Mtskheta-Mtianeti
Racha-Lechkhumi-Kvemo Svaneti
Samegrelo-Zemo Svaneti
Samtskhe-Javakheti
Shida Kartli
Tbilisi
Sardegna
Beja
Beja
vora
vora
Faro
Portalegre
Portalegre
Astrakhan
Kalmyk
Krasnodar
Rostov
Stavropol
Volgograd
Andaluca
Extremadura
Batumi
Chokhatauri
Bagdati
Akhmeta
Bolnisi
Akhalgori
Ambrolauri
Abasha
Adigeni
Gori
Tbilisi
Cagliari
Aljustrel
Alandroal
Albufeira
Alter do Cho
Akhtubinsk
Chernozemelskiy rayon
Abinskiy rayon
Aksayskiy rayon
Aleksandrovskiy rayon
Alekseevskiy rayon
Reference ID
4000
4000
4000
4000
4000
4000
4000
4000
4000
4000
4000
4000
4000
4000
122
122
122
122
122
122
122
4000
4000
4000
4000
4000
4000
0
0
117
118
Appendix 1
Table A1.2. Geographic distribution data for Crimean-Congo haemorrhagic fever virus (CCHFV).
See Appendix 2 (hosted on http://www.cabi.org/vetmedresource/) for complete reference indicated by the
Reference ID.
Country
Albania
Bulgaria
Bulgaria
Greece
Greece
Kosovo
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Russia
Kuks
Blagoevgrad
Blagoevgrad
Anatoliki Makedonia kai Thraki
Pristina
Astrakhan
Astrakhan
Astrakhan
Dagestan
Dagestan
Ingush
Kalmyk
Kalmyk
Kalmyk
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Rostov
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Stavropol
Russia
Russia
Russia
Russia
Russia
Russia
Stavropol
Stavropol
Stavropol
Stavropol
Volgograd
Volgograd
Gotse Delchev
Gotse Delchev
Rodopi
Dubovskiy rayon
Martynovskiy rayon
Orlovskiy rayon
Orlovskiy rayon
Peschanokopskiy rayon
Proletarskiy rayon
Remontnenskiy rayon
SalskSalskiy rayon
Tselinnyy rayon
Tsimlyanskiy rayon
Zimovnikovskiy rayon
Zimovnikovskiy rayon
Apanasenkovskiy rayon
Apanasenkovskiy rayon
Arzgirskiy rayon
Blagodarnenskiy rayon
Budennovsk
Budennovskiy rayon
Ipatovskiy rayon
Izobilnenskiy rayon
Krasnogorodskiy rayon
Krasnogorodskiy rayon
Novoaleksandrovskiy
rayon
Novoselitskiy rayon
Petropavlovskiy rayon
Rostov
Stepnovskiy rayon
Reference ID
No. entriesa
723
1109
1412
1413
721
1402
1068
1404
635
1403
1411
1411
1401
1404
1408
1068
634
1407
1414
1405
1414
1407
1407
1407
1405
1406
1406
1406
1414
1404
1404
635
1410
1414
1414
1410
1408
1414
1414
1410
1409
1410
1414
1410
1414
1400
1414
1068
1404
Continued
Distribution for Ticks and Tick-borne Diseases
119
Table A1.2. Continued.

Country
Russia
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Volgograd
Adana
Adiyaman
Afyon
Aksaray
Amasya
Ankara
Artvin
Artvin
Aydn
Balkesir
Bayburt
Bilecik
Bingl
Bolu
Bursa
anakkale
ankr
orum
Erzincan
Erzurum
Erzurum
Eskisehir
Giresun
Giresun
Gmshane
Gmshane
Hatay
I el
Isparta
Istanbul
Istanbul
I zmir
Karabuk
Kars
Kastamonu
Kayseri
Krkkale
Krklareli
Krsehir
Konya
Ktahya
Malatya
Manisa
Mugla
Mus
Nevsehir
Ordu
Osmaniye
Samsun
Sanlurfa
Sinop
Oktyabrskiy
Reference ID
1408
1399
1399
1399
1399
1342
1399
1399
708
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
708
1399
1102
1399
1399
991
1399
1399
1399
1399
649
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1399
1102
1399
1399
1399
1399
No. entriesa
3
4
44
20
31
3
14
2
3
3
14
38
117
21
73
2
6
8
25
70
2
2
4
4
18
42
5
4
2
3
4
8
4
2
3
2
2
15
11
3
3
Continued
120
Appendix 1

Country
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Sivas
Tokat
Tokat
Trabzon
Tunceli
Van
Yozgat
Zinguldak
Reference ID
1399
1102
708
708
1399
1399
1399
1399
No. entriesa
114
219
2
2
2
95
If more than one.
Table A1.3. Geographic distribution data for the tick-borne encephalitis (TBE) group. See Appendix 2
(hosted on http://www.cabi.org/vetmedresource/) for complete reference indicated by the Reference ID.
Country
land Islands
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Jihocesk
Jihocesk
Jihocesk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Kraj Vysocina
Krlovhradeck
Krlovhradeck
Plzensk
rhus
Bornholm
Bornholm
Bornholm
Fyn
Ringkbing
South Jutland
Storstrm
Storstrm
Vejle
Vejle
Harju
Harju
Hiiu
Ida-Viru
Ida-Viru
Ida-Viru
Jrva
Jgeva
Lne
Lne
Lne-Viru
Prnu
Plva
Rapla
Administrative region 2 Reference ID No. entriesa
Hasle
Otterup
Ringkbing
Fakse
Vejle
1329
1045
226
525
1338
227
525
227
225
230
230
888
1290
345
1325
582
582
888
888
582
888
582
1317
377
1317
1317
376
377
1317
1317
1317
377
1317
1317
1317
1317
Continued
121

Country
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Finland
Finland
Finland
Finland
Finland
Finland
France
France
France
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Greece
Saare
Tartu
Tartu
Tartu
Valga
Valga
Valga
Viljandi
Vru
Lapland
Southern Finland
Southern Finland
Western Finland
Western Finland
Alsace
Bourgogne
Champagne-Ardenne
Franche-Comt
Lorraine
Baden-Wrttemberg
Baden-Wrttemberg
Baden-Wrttemberg
Baden-Wrttemberg
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Brandenburg
Brandenburg
Hessen
Hessen
Hessen
Hessen
Mecklenburg-Vorpommern
Nordrhein-Westfalen
Rheinland-Pfalz
Rheinland-Pfalz
Rheinland-Pfalz
Rheinland-Pfalz
Rheinland-Pfalz
Sachsen
Sachsen-Anhalt
Sachsen-Anhalt
Thringen
Thringen
Lapland
Uusimaa
Central Ostrobothnia
Freiburg
Mittelfranken
Niederbayern
Niederbayern
Oberbayern
Oberpfalz
Unterfranken
Brandenburg
Rheinhessen-Pfalz
1317
1317
1348
376
1317
376
377
1317
1317
409
1341
409
1329
34
465
1285
1285
1285
1285
1285
1058
1327
969
968
1058
1327
969
1287
1287
968
1287
1287
439
1058
424
1058
1327
968
969
1121
523
1064
1058
1327
968
969
1059
969
1121
523
1121
969
736
Continued
122
Appendix 1

Country
Greece
Greece
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Dytiki Makedonia
Kentriki Makedonia
Borsod-Abaj-Zempln
Fejr
Gyor-Moson-Sopron
Heves
Heves
Komrom-Esztergom
Ngrd
Ngrd
Somogy
Tolna
Vas
Veszprm
Zala
Friuli-Venezia Giulia
Piemonte
Trentino-Alto Adige
Trentino-Alto Adige
Trentino-Alto Adige
Trentino-Alto Adige
Veneto
Veneto
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Latgale
Latgale
Latgale
Latgale
Latgale
Latgale
Latgale
Latgale
Latgale
Riga
Riga
Riga
Riga
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Torino
Trento
Trento
Trento
Trento
Belluno
Belluno
Dobele
Kuldiga
Kuldiga
Liepaja
Liepaja
Saldus
Tukums
Ventspils
Balvi
Balvi
Daugavpils
Jekabpils
Kraslava
Ludza
Madona
Preili
Rezekne
Limbai
Riga
Riga
Riga
Aizkraukle
Aluksne
Aluksne
Gulbene
Limbai
Madona
Valka
Valmiera
736
736
1286
784
784
1286
784
784
1286
784
784
784
784
784
784
336
780
1304
160
804
806
1304
201
968
1317
968
1301
968
968
968
968
1317
968
968
968
968
968
968
968
968
1317
126
1301
968
1301
968
1317
968
968
968
968
968
968
3
3
4
Continued
123

Country
Latvia
Latvia
Latvia
Latvia
Liechtenstein
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Norway
Norway
Norway
Norway
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Russia
Sweden
Sweden
Sweden
Sweden
Zemgale
Zemgale
Zemgale
Zemgale
Aizkraukle
Bauska
Jelgava
Ogre
Alytaus
Alytaus
Kauno
Kauno
Klaipedos
Marijampoles
Paneveio
Paneveio
iauliai
iauliai
Taurages
Taurages
Teliai
Teliai
Utenos
Vilniaus
Vilniaus
Aust-Agder
Aust-Agder
Aust-Agder
Sr-Trndelag
Greater Poland
Kuyavian-Pomeranian
Lesser Poland
Ldz
Lower Silesian
Lublin
Lublin
Lublin
Lublin
Lubusz
Masovian
Opole
Podlachian
Podlachian
Podlachian
Pomeranian
Silesian
Subcarpathian
Swietokrzyskie
Warmian-Masurian
West Pomeranian
Blekinge
Blekinge
Blekinge
Blekinge
Alytaus
Birtono
Klaipedos
Marijampoles
Biru
Akmenes
Jurbarko
Maeikiu
Anykciu
alcininku
Arendal
Arendal
Lublin
Radzyn
Biaystok
Karlskrona
Olofstrm
1317
968
968
968
541
489
489
1317
489
1317
1317
1317
413
1317
1317
413
489
1317
1317
489
1317
489
1317
887
209
210
1290
942
942
942
942
942
193
942
403
192
942
942
942
532
942
1350
942
942
942
942
942
942
1293
1054
1289
1289
1289
2
2
Continued
124
Appendix 1

Country
Sweden
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Turkey
Turkey
Blekinge
Skne
Stockholm
Stockholm
Stockholm
Aargau
Bern
Bern
Neuchtel
Sant Gallen
Schaffhausen
Thurgau
Thurgau
Thurgau
Zrich
Tokat
Yozgat
Ronneby
If more than one.
Sderhamn
Sderhamn
Upplands-Bro
1289
483
636
636
636
1328
1328
165
1328
1328
1328
1328
541
855
556
308
1351
Table A1.4. Geographic distribution data for Anaplasma spp. and Ehrlichia spp. now included in Anaplasma spp. See Appendix 2 (hosted on http://www.cabi.org/
vetmedresource/) for complete reference indicated by the Reference ID.
Anaplasma bovis
Anaplasma centrale
Anaplasma centrale
Anaplasma centrale
Anaplasma centrale
Anaplasma centrale
Anaplasma centrale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Italy
Egypt
Italy
Italy
Italy
Italy
Italy
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Basilicata
Al Wadi al Jadid
Basilicata
Basilicata
Calabria
Calabria
Sicily
Borsod-Abaj-Zempln
Abruzzo
Apulia
Apulia
Basilicata
Basilicata
Basilicata
Basilicata
Calabria
Calabria
Calabria
Campania
Campania
Lazio
Lombardia
Marche
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Potenza
Matera
Potenza
Crotone
Vibo Valentia
Palermo
Bari
Foggia
Matera
Potenza
Crotone
Vibo Valentia
Agrigento
Caltanissetta
Catania
Enna
Messina
Palermo
Palermo
Reference ID
156
1343
156
156
156
156
366
443
156
207
156
207
996
156
156
996
156
156
207
996
996
996
996
994
996
992
992
992
992
992
242
244
No. entriesa
2
2
2
Continued
125
Country
Species
126

Country
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma marginale
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Anaplasma ovis
Italy
Italy
Italy
Italy
Italy
Spain
Switzerland
Switzerland
Switzerland
Cyprus
Cyprus
Cyprus
Cyprus
Cyprus
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Romania
Spain
Turkey
Albania
Belarus
Bulgaria
Bulgaria
Croatia
Cyprus
Czech Republic
Sicily
Sicily
Sicily
Toscana
Umbria
Castilla-La Mancha
Graubnden
Graubnden
Graubnden
Famagusta
Larnaca
Limassol
Nicosia
Paphos
Borsod-Abaj-Zempln
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Tulcea
Andaluca
Elzig
Tiran
Minsk
Soa
Soa
Koprivnicko-Krizevacka
Famagusta
Jihomoravsk
Palermo
Palermo
Trapani
Ciudad Real
Agrigento
Caltanissetta
Catania
Enna
Messina
Palermo
Palermo
Trapani
Cdiz
Stolichna
Reference ID
No. entriesa
366
992
992
996
996
246
1211
270
438
775
775
775
775
775
443
994
992
992
992
992
992
242
992
992
59
241
24
409
1336
448
185
656
1139
1338
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Appendix 1
Species
Moravskoslezsk
Stredocsk
Stredocsk
Copenhagen
Frederiksborg
Fyn
North Jutland
South Jutland
South Jutland
Al Jizah
Prnu
Saare
Alsace
Alsace
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Auvergne
Auvergne
Auvergne
Auvergne
Basse-Normandie
Basse-Normandie
Basse-Normandie
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bretagne
Bretagne
Bretagne
Bretagne
Centre
Bas-Rhin
Bas-Rhin
Dordogne
Gironde
Landes
Lot-et-Garonne
Pyrnes-Atlantiques
Allier
Cantal
Haute-Loire
Puy-de-Dme
Calvados
Manche
Orne
Cte-dOr
Nivre
Sane-et-Loire
Sane-et-Loire
Yonne
Ctes-dArmor
Finistre
Ille-et-Vilaine
Morbihan
Cher
1338
1091
1092
888
888
888
888
1120
888
1343
1348
1336
484
719
484
484
484
484
484
484
484
484
484
484
484
484
484
484
484
719
484
484
484
484
484
484
Continued
127
Czech Republic
Czech Republic
Czech Republic
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Egypt
Estonia
Estonia
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
128

Country
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Centre
Centre
Centre
Centre
Centre
Centre
Centre
Champagne-Ardenne
Champagne-Ardenne
Champagne-Ardenne
Corse
Franche-Comt
Franche-Comt
Franche-Comt
Franche-Comt
Franche-Comt
Haute-Normandie
Haute-Normandie
le-de-France
le-de-France
le-de-France
le-de-France
Languedoc-Roussillon
Limousin
Limousin
Limousin
Limousin
Limousin
Lorraine
Lorraine
Lorraine
Lorraine
Lorraine
Eure-et-Loir
Eure-et-Loir
Indre
Indre-et-Loire
Indre-et-Loire
Loir-et-Cher
Loiret
Ardennes
Haute-Marne
Marne
Corse-du-Sud
Doubs
Haute-Sane
Haute-Sane
Jura
Territoire de Belfort
Eure
Seine-Maritime
Essonne
Seine-et-Marne
Seine-et-Marne
Yvelines
Bouches-du-Rhne
Lozre
Corrze
Corrze
Creuse
Haute-Vienne
Haute-Vienne
Meurthe-et-Moselle
Meurthe-et-Moselle
Meuse
Meuse
Moselle
Reference ID
484
719
484
484
719
484
484
484
484
484
484
484
484
719
484
484
484
484
719
484
719
484
561
484
484
719
484
484
719
484
719
484
484
484
No. entriesa
Appendix 1
Species
Lorraine
Lorraine
Lorraine
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Nord-Pas-de-Calais
Nord-Pas-de-Calais
Nord-Pas-de-Calais
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Picardie
Picardie
Picardie
Picardie
Poitou-Charentes
Poitou-Charentes
Poitou-Charentes
Provence-Alpes-Cte-dAzur
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Moselle
Vod
Vosges
Aveyron
Haute-Garonne
Haute-Garonne
Hautes-Pyrnes
Lot
Tarn
Tarn
Tarn-et-Garonne
Nord
Pas-De-Calais
Pas-De-Calais
Loire-Atlantique
Maine-et-Loire
Maine-et-Loire
Mayenne
Sarthe
Vende
Aisne
Oise
Oise
Somme
Charente
Deux-Svres
Vienne
Alpes-Maritimes
Bouches-Du-Rhne
Bouches-Du-Rhne
Var
Var
Vaucluse
Ain
Haute-Savoie
Isre
719
484
719
484
484
719
484
484
484
719
484
484
484
719
484
484
719
484
484
484
719
484
719
484
484
484
484
484
484
719
484
719
484
484
484
484
Continued
129
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
130

Country
France
France
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Greece
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Lithuania
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Baden-Wrttemberg
Bayern
Berlin
Berlin
Brandenburg
Thringen
Kriti
Basilicata
Lazio
Lazio
Sardegna
Sardegna
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Trentino-Alto Adige
Trentino-Alto Adige
Klaipedos
Loire
Rhne
Rhne
Savoie
Berlin
Potenza
Agrigento
Caltanissetta
Catania
Enna
Messina
Palermo
Palermo
Palermo
Ragusa
Syracuse
Trapani
Trento
Trento
Reference ID
484
1032
484
484
718
1352
540
1294
540
1352
1108
156
191
639
93
832
833
28
619
999
992
992
992
992
992
242
243
992
992
992
992
158
806
1336
No. entriesa
8
2
4
4
4
4
4
4
Appendix 1
Species
Chisinau
Chisinau
Taza-Al Hoceima-Taounate
Aust-Agder
Sr-Trndelag
Telemark
Greater Poland
Lublin
Lublin
Lublin
Lublin
Lublin
Masovian
Masovian
Podlachian
Podlachian
Podlachian
Podlachian
Pomeranian
Warmian-Masurian
West Pomeranian
Faro
Madeira
Madeira
Madeira
Vologda
Branicevski
Grad Beograd
Macvanski
Niavski
Pomoravski
Raki
Sremski
Zajecarski
Banskobystrick
Hitra
Krager
Lublin
Warsaw
Augustw
Hajnwka
Hajnwka
Biaystok
1134
528
666
643
1323
1323
1323
1131
1062
11
179
990
194
1119
1098
1129
0
385
386
931
0
822
836
1143
238
838
35
652
652
652
652
652
652
652
652
529
6
2
6
Continued
131
Moldova
Moldova
Moldova
Morocco
Norway
Norway
Norway
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Portugal
Portugal
Portugal
Portugal
Russia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Slovakia
132

Country
Anaplasma platys
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovenia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Switzerland
Switzerland
Switzerland
Switzerland
Tunisia
Turkey
Turkey
Turkey
Ukraine
UK
UK
UK
Italy
Banskobystrick
Bratislavsk
Koick
Koick
Koick
Nitriansk
Presov
Zilinsky
Andaluca
Castilla y Len
Comunidad de Madrid
Galicia
Galicia
Galicia
Galicia
Galicia
La Rioja
La Rioja
Pas Vasco
Pas Vasco
Bern
Graubnden
Graubnden
Neuchtel
Jendouba
Giresun
Rize
Trabzon
Kharkiv
England
England
England
Abruzzo
Cdiz
Madrid
Ourense
Ourense
Pontevedra
Pontevedra
La Rioja
La Rioja
Cheshire
Hampshire
Northumberland
Reference ID
No. entriesa
941
1349
256
499
936
1349
936
1347
191
241
327
18
360
52
53
52
53
701
773
78
79
586
1211
438
586
643
26
26
26
1134
134
808
131
912
12
2
2
3
4
2
2
4
3
Appendix 1
Species
If more than one.
Italy
Italy
Italy
Tunisia
Turkey
Albania
Albania
Cyprus
Germany
Slovakia
Slovakia
Turkey
Turkey
Turkey
Sicily
Sicily
Sicily
Jendouba
Mugla
Durrs
Kuks
Berlin
Bratislavsky
Nitriansky
Antalya
Kayseri
Malatya
Palermo
Durrsit
Kuksit
Berlin
994
999
245
643
1012
185
185
1118
1330
124
124
185
185
185
2
4
2
3
Anaplasma platys
Anaplasma platys
Anaplasma platys
Anaplasma platys
Anaplasma platys
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
Anaplasma spp.
133
Country
Rickettsia aeschlimanni
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Algeria
Croatia
Egypt
France
Russia
Spain
Spain
Spain
Spain
Spain
Albania
Albania
Algeria
Algeria
France
France
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Splitsko-Dalmatinska
Shamal Sina
Corse
Stavropol
Castilla y Len
Castilla y Len
Castilla y Len
La Rioja
La Rioja
Durrs
Kuks
Oran
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kriti
Stere Ellda
Thessalia
Corse-du-Sud
La Rioja
La Rioja
Durrsit
Kuksit
Hrault
Var
Drama
Kavala
Florina
Grevena
Kastoria
Kozani
Imathia
Khalkidiki
Kilkis
Pella
Pieria
Serrai
Thessaloniki
Fokis
Trikala
Reference ID
110
781
1343
624
1144
327
327
328
700
772
185
185
110
664
1292
1299
720
222
222
222
222
222
222
222
222
222
222
222
222
222
367
778
222
No. entriesa
4
2
3
Appendix 1
Species
134
Table A1.5. Geographic distribution data for Rickettsia spp. See Appendix 2 (hosted on http://www.cabi.org/vetmedresource/) for complete reference indicated
by the Reference ID.
Piemonte
Piemonte
Sicily
Kosovska Mitrovica
Kosovska Mitrovica
Pecki
Grand Casablanca
Grad Beograd
Pcinjski
Raski
Srednje-Banatski
Zlatiborski
Andaluca
Castilla y Len
Catalua
Catalua
Galicia
Galicia
Islas Baleares
La Rioja
Antalya
Edirne
Kayseri
Krklareli
Malatya
Tekirdag
Sicily
Albania
Albania
Austria
Bulgaria
Croatia
Denmark
Durrs
Kuks
Burgenland
Soa
Medimurska
North Jutland
Cuneo
Cuneo
Kosovska Mitrovica
Kosovska Mitrovica
Pec
Stari Grad
Vranje
Tutin
Zrenjanin
Uzice
Sevilla
Soria
Barcelona
Tarragona
Ourense
Pontevedra
La Rioja
Durrsit
Kuksit
Stolichna
416
607
608
999
346
827
827
1104
827
827
827
827
827
95
681
907
907
53
53
907
1295
185
1122
185
1122
185
1122
369
4
4
4
2
2
185
185
265
185
264
0
2
3
Continued
135
Israel
Italy
Italy
Italy
Kosovo
Kosovo
Kosovo
Morocco
Serbia
Serbia
Serbia
Serbia
Serbia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Italy
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii
Rickettsia conorii israelensis
(Rickettsia conorii Israeli
spotted fever strain)
Rickettsia helvetica
136

Country
Denmark
Denmark
France
Germany
Germany
Italy
Italy
Italy
Moldova
Netherlands
Netherlands
Netherlands
Netherlands
Poland
Poland
Poland
Poland
Slovakia
Spain
Spain
Spain
Switzerland
Tunisia
Turkey
Turkey
Turkey
Ukraine
Algeria
Greece
Morocco
Spain
Spain
Spain
Spain
Vestsjlland
Vestsjlland
Auvergne
Bayern
Berlin
Toscana
Trentino-Alto Adige
Veneto
Flevoland
Friesland
Noord-Holland
Overijssel
Greater Poland
Greater Poland
Lesser Poland
Pomeranian
Banskobystrick
Andaluca
Andaluca
Andaluca
Zrich
Jendouba
Antalya
Kayseri
Malatya
Kharkiv
Stere Ellda
Gharb-Chrarda-Bni Hssen
Andaluca
Andaluca
Andaluca
La Rioja
Korsr
Puy-de-Dme
Berlin
Trento
Granada
Huelva
Jan
Fokis
Granada
Jan
Sevilla
La Rioja
Reference ID
1146
973
727
1063
1294
1281
1281
1281
1134
917
917
917
917
1145
932
932
932
941
1127
1127
611
125
0
185
185
185
1134
110
778
1104
1127
611
614
700
No. entriesa
3
7
3
2
12
2
2
2
Appendix 1
Species
Ticino
El Tarf
Bayern
Bayern
Madeira
Andaluca
Andaluca
Andaluca
La Rioja
Pas Vasco
Jendouba
Kharkiv
Kriti
Baden-Wrttemberg
Berlin
Toscana
Toscana
Podlachian
Andaluca
Andaluca
Andaluca
Stere Ellda
Medimurska
Bretagne
Bayern
Kentriki Makedonia
Toscana
Toscana
Andaluca
Andaluca
Castilla y Len
Castilla y Len
Granada
Huelva
Jan
Bouches-du-Rhne
Freiburg
Berlin
Lucca
Lucca
Granada
Huelva
Jan
Fokis
Khalkidiki
Lucca
Lucca
Granada
Jan
Burgos
96
1111
1112
878
1134
1104
238
1127
1127
611
468
468
0
1134
348
776
1137
1330
1332
862
1129
1127
1127
611
778
264
781
1130
1137
1303
1332
862
1127
611
330
587
2
2
2
2
2
2
Continued
137
Switzerland
Algeria
Germany
Germany
Moldova
Morocco
Portugal
Spain
Spain
Spain
Spain
Spain
Tunisia
Ukraine
France
Greece
Germany
Germany
Italy
Italy
Poland
Spain
Spain
Spain
Greece
Croatia
Croatia
France
Germany
Greece
Italy
Italy
Spain
Spain
Spain
Spain
Rickettsia mongolotimonaeb
Rickettsia mongolotimonae
Rickettsia raoultii
Rickettsia raoultii
Rickettsia raoultii
Rickettsia raoultii
Rickettsia raoultii
Rickettsia raoultii
Rickettsia raoultii
Rickettsia raoultii
Rickettsia rhipicephali
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
138

Country
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia slovaca
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Rickettsia spp.
Spain
Spain
Spain
Switzerland
Albania
Cyprus
Germany
Germany
Germany
Germany
Germany
Germany
Italy
Italy
Poland
Poland
Portugal
Russia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Spain
Castilla y Len
La Rioja
La Rioja
Ticino
Tiran
Palencia
La Rioja
La Rioja
Bayern
Bayern
Berlin
Brandenburg
Sachsen
Sachsen-Anhalt
Veneto
Podlachian
Warmian-Masurian
Madeira
Vologda
Banskobystrick
Banskobystrick
Bratislavsk
Nitriansk
Andaluca
Pas Vasco
Berlin
Belluno
Granada
Reference ID
587
1295
700
96
409
1118
236
236
1330
236
236
236
337
752
928
928
238
35
124
529
124
124
1127
79
No. entriesa
5
2
2
15
If more than one.

Rickettsia mongolotimonae is the name rst proposed for this species. It was later identied as a member of the R. sibirica species complex, and is now called both R. sibirica
mongolotimonae and R. sibirica mongolitimonae.
3
2
Appendix 1
Species
Table A1.6. Geographic distribution data for Borrelia spp. See Appendix 2 (hosted on http://www.cabi.org/vetmedresource/) for complete reference indicated by
the Reference ID.
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Bulgaria
Croatia
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Denmark
Finland
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Krnten
Niedersterreich
Obersterreich
Steiermark
Steiermark
Tirol
Vorarlberg
Wien
Soa
Primorsko-Goranska
Jihocesk
Jihocesk
Jihomoravsk
Jihomoravsk
Krlovhradeck
Krlovhradeck
Libereck
Moravskoslezsk
Plzensk
steck
Zlnsk
North Jutland
Western Finland
Alsace
le-de-France
Baden-Wrttemberg
Bayern
Bayern
Berlin
Niedersachsen
Nordrhein-Westfalen
Administrative
region 2
Stolichna
Haut-Rhin
Yvelines
Tbingen
Berlin
Hannover
Reference ID No. entriesa

114
511
114
114
963
114
114
511
185
1011
224
255
550
742
225
230
271
220
230
550
550
1026
34
331
753
794
332
333
1294
431
600
8
2
2
2
4
2
6
5
3
2
5
Continued
139
Country
Species
140
Country
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Hungary
Ireland
Ireland
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Latvia
Moldova
Moldova
Netherlands
Netherlands
Netherlands
Netherlands
Norway
Norway
Norway
Norway
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Russia
Serbia
Serbia
Serbia
Serbia
Pest
Galway
Kerry
Lazio
Lazio
Toscana
Toscana
Trentino-Alto Adige
Trentino-Alto Adige
Riga
Chisinau
Chisinau
Flevoland
Gelderland
Noord-Holland
Overijssel
Aust-Agder
Sr-Trndelag
Telemark
Telemark
Greater Poland
Lublin
Lublin
Silesian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Vologda
Branicevski
Grad Beograd
Macvanski
Niavski
Administrative
region 2
Pisa
Trento
Trento
Arnhem
Hitra
Krager
Lublin City
Lublin City
Tarnowskie

344
380
754
639
831
833
1331
98
609
741
319
528
666
1291
365
1050
1050
1323
1323
475
1323
603
194
1337
1288
384
738
739
35
652
652
652
652
3
2
3
3
Appendix 1
Species
2
2
2
2
Pomoravski
Raki
Sremski
Zajecarski
Bratislavsk
Koick
Nitriansk
Trenciansk
Trenciansk
Zilinsk
La Rioja
Pas Vasco
Gvleborg
Kalmar
Skne
Skne
Stockholm
Vsterbotten
Graubnden
Neuchtel
Neuchtel
Neuchtel
Ticino
Ticino
Valais
Valais
ankiri
Istanbul
Ordu
Burgenland
Burgenland
Burgenland
Burgenland
Borrelia burgdorferi s.l.
Austria
Burgenland
La Rioja
Nynshamn
Ume
Eisenstadt
Eisenstadt
Umgebung
Gssing
652
652
652
652
1320
1349
602
1349
1349
603
1347
315
78
350
350
350
696
470
470
163
143
163
368
1284
163
1284
163
396
395
396
114
258
172
172
16
2
172
Continued
141
Serbia
Serbia
Serbia
Serbia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Spain
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Turkey
Turkey
Turkey
Austria
Austria
Austria
Austria
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
Borrelia afzelii
142
Country

Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Belgium
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Burgenland
Burgenland
Burgenland
Burgenland
Burgenland
Krnten
Niedersterreich
Obersterreich
Salzburg
Steiermark
Tirol
Vorarlberg
Wien
Antwerp
Brussels
East Flanders
East Flanders
Flemish Brabant
Flemish Brabant
Hainaut
Liege
Limburg
Luxembourg
Namur
Namur
Walloon Brabant
Walloon Brabant
West Flanders
West Flanders
Soa
Soa
Stara Zagora
Administrative
region 2
Jennersdorf
Mattersburg
Neusiedl am See
Oberpullendorf
Oberwart
Namur
Stolichna

172
172
172
172
172
114
114
114
114
258
114
114
114
580
580
580
668
580
615
580
580
580
580
580
1298
1333
580
580
668
184
448
1319
1007
2
2
2
2
2
2
2
2
2
Appendix 1
Species
Jihocesk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Kraj Vysocina
Krlovhradeck
Libereck
Libereck
Moravskoslezsk
Moravskoslezsk
Moravskoslezsk
Olomouck
Pardubick
Pardubick
Plzensk
Prague
steck
Zlnsk
Zlnsk
Copenhagen
Frederiksborg
Fyn
North Jutland
North Jutland
South Jutland
Western Finland
Alsace
Alsace
Alsace
Alsace
Aquitaine
Basse-Normandie
Brno
Liberec
st nad Orlic
Bas-Rhin
656
491
1077
1338
490
491
669
472
491
491
491
472
1035
491
1338
491
491
472
491
669
550
491
550
888
888
888
888
946
888
34
1285
572
851
719
572
572
4
2
4
4
Continued
143
Croatia
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Finland
France
France
France
France
France
France

144
Country

France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Bourgogne
Bourgogne
Bretagne
Bretagne
Centre
Centre
Centre
Centre
Centre
Champagne-Ardenne
Champagne-Ardenne
Champagne-Ardenne
Franche-Comt
Franche-Comt
Haute-Normandie
Haute-Normandie
le-de-France
le-de-France
le-de-France
Limousin
Limousin
Lorraine
Lorraine
Lorraine

France
France
France
France
France
France
France
France
Lorraine
Lorraine
Midi-Pyrnes
Nord-Pas-de-Calais
Pays de la Loire
Pays de la Loire
Picardie
Administrative
region 2
Morbihan
Eure-et-Loir
Indre-et-Loire
Indre-et-Loire
Loiret
Ardennes
Eure
Seine-et-Marne
Meurthe-etMoselle
Meuse
Moselle
Nord
Maine-et-Loire
Alpes-Maritimes

1285
572
572
626
572
719
626
719
626
1285
572
626
1285
572
572
719
262
572
626
572
851
1285
572
719
626
626
572
408
572
626
572
719
Appendix 1
Species

France
France
France
France
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Greece
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Baden-Wrttemberg
Baden-Wrttemberg
Bayern
Bayern
Berlin
Berlin
Berlin
Brandenburg
Brandenburg
Hessen
Hessen
Sachsen-Anhalt
Thringen
Thringen
Attiki
Calabria
Lazio
Marche
Molise
Toscana
Toscana
Toscana
Trentino-Alto Adige
Veneto
Bouches-duRhne
Ain
Haute-Savoie
Loir-et-Cher
Rhne
Freiburg
Stuttgart
Unterfranken
Berlin
Darmstadt
Attica
Arezzo
Florence
Siena
Trento
Belluno
626
572
851
782
719
782
782
691
691
564
439
540
633
1330
540
633
930
496
633
633
268
633
476
381
191
49
93
832
325
325
1283
1283
1283
806
201
10
3
4
3
Continued
145
France
146
Country

Italy
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Netherlands
Netherlands
Veneto
Alytaus
Alytaus
Kauno
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Marijampoles
Marijampoles
Panevezio
Panevezio
Panevezio
iauliai
iauliai
iauliu
iauliu
iauliu
Utenos
Utenos
Utenos
Vilniaus
Vilniaus
Friesland
Noord-Holland
Administrative
region 2
Belluno
Birzu
Joniskio
Kupiskio
Pakruojo
Pasvalio
Rokiskio
Varenos
ilutes
Klaipedos
Klaipedos
Neringos
Skuodo
Marijampoles
Paneveio
Rokikio
iauliu
Jonikio
Utenos
Utenos
Ukmerges
Ameland

752
1100
1100
1100
1100
1100
1100
1009
732
1009
1009
732
1100
732
732
1100
1009
732
1009
732
732
732
732
1009
1009
1009
1009
732
732
1009
732
467
1050
2
3
Appendix 1
Species
Aust-Agder
Hordaland
Rogaland
Telemark
Vest-Agder
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Ldz
Ldz
Lower Silesian
Lower Silesian
Lower Silesian
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lubusz
Masovian
Birkenes
Kvinnherad
Krager
Kristiansand
Czarnkw-Trzcianka
Jarocin
Krotoszyn
Leszno
Miedzychd
Poznan
Poznan
Naklo
Seplno
Wocawek
Lowicz
Lubartw
Lublin City
Lublin City
Lublin City
Radzyn
Wodawa
732
732
707
732
732
645
107
107
107
107
107
107
644
1324
283
283
284
1324
284
267
272
513
105
1062
178
193
925
990
192
192
194
196
192
284
1324
1324
6
2
4
Continued
147
Norway
Norway
Norway
Norway
Norway
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland

148
Country

Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Portugal
Russia
Russia
Russia
Russia
Serbia
Serbia
Serbia
Serbia
Slovakia
Slovakia
Masovian
Masovian
Masovian
Masovian
Masovian
Opole
Podlachian
Podlachian
Podlachian
Pomeranian
Pomeranian
Silesian
Swietokrzyskie
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
Lisboa
Leningrad
Moskva
Novgorod
Vologda
Grad Beograd
Grad Beograd
Toplicki
Zapadno-Backi
Banskobystrick
Banskobystrick
Administrative
region 2
Kozienice
Kozienice
Przasnysz
Warsaw
Hajnwka
Lidzbark
Stargard
Szczecin
Szczecin
Mafra

531
284
285
284
1098
1324
1324
715
334
1324
931
956
1324
1324
738
739
283
1324
687
1296
138
891
76
533
522
533
35
654
655
655
655
529
87
2
2
5
2
2
3
Appendix 1
Species
Banskobystrick
Bratislavsk
Bratislavsk
Koick
Koick
Koick
Koick
Koick
Koick
Koick
Nitriansk
Preovsk
Preovsk
Preovsk
Preovsk
Trenciansk
Trenciansk
Trnavsk
Zilinsk
Zilinsk
Galicia
Galicia
La Rioja
Pas Vasco
Pas Vasco
Dalarna
Gvleborg
Gotland
Halland
Halland
Jmtland
Jnkping
Kalmar
Kronoberg
Orebro
Ourense
Pontevedra
La Rioja
Varberg
939
87
939
256
382
87
934
936
940
945
87
87
939
936
940
87
939
87
1347
87
191
53
53
705
78
79
1312
1312
1312
1312
470
1312
1312
1312
1312
1312
22
18
3
4
6
3
Continued
149
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovenia
Spain
Spain
Spain
Spain
Spain
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden

150
Country

Borrelia burgdorferi s.s.
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Switzerland
Turkey
Turkey
Turkey
Turkey
Austria
Austria
Austria
Austria
Austria
Austria
Bulgaria
Croatia
Czech Republic
Czech Republic
Czech Republic
Czech Republic
stergtland
Skne
Sdermanland
Stockholm
Stockholm
Uppsala
Uppsala
Vrmland
Vsterbotten
Vsterbotten
Vsterbotten
Vsternorrland
Vstmanland
Vstra Gtaland
Genve
Neuchtel
Valais
Bursa
Dzce
Isparta
Izmir
Niedersterreich
Niedersterreich
Obersterreich
Salzburg
Tirol
Wien
Soa
Primorsko-Goranska
Jihocesk
Jihocesk
Jihomoravsk
Jihomoravsk
Administrative
region 2
Norrtlje
Uppsala
Ume
Ume
Stolichna

1312
1312
1312
1312
470
1312
470
1312
1318
1322
1322
1312
1312
1312
798
1326
1321
101
507
252
101
114
511
114
114
114
511
185
1011
224
255
550
742
Appendix 1
Species
2
3
Krlovhradeck
Krlovhradeck
Libereck
Moravskoslezsk
Plzensk
North Jutland
Alsace
le-de-France
Baden-Wrttemberg
Bayern
Bayern
Berlin
Niedersachsen
Nordrhein-Westfalen
Pest
Kerry
Lazio
Toscana
Toscana
Trentino-Alto Adige
Riga
Riga
Chisinau
Chisinau
Telemark
Greater Poland
Greater Poland
Lublin
Lublin
Silesian
Warmian-Masurian
Branicevski
Grad Beograd
Macvanski
Haut-Rhin
Yvelines
Tbingen
Berlin
Hannover
Pisa
Trento
Taza
Poznan
Lublin City
Lublin City
Tarnowskie
225
230
271
220
230
1026
331
753
794
332
333
1294
431
600
344
754
639
831
1331
98
741
319
319
528
666
642
475
603
645
194
1337
1288
384
652
652
652
2
4
Continued
151
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Denmark
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Hungary
Ireland
Italy
Italy
Italy
Italy
Italy
Latvia
Latvia
Moldova
Moldova
Morocco
Norway
Poland
Poland
Poland
Poland
Poland
Poland
Serbia
Serbia
Serbia

152
Country

Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Serbia
Serbia
Serbia
Serbia
Serbia
Slovakia
Spain
Sweden
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Switzerland
Turkey
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Bulgaria
Croatia
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Niavski
Pomoravski
Raki
Sremski
Zajecarski
Kosicky
Pas Vasco
Blekinge
Gvleborg
Kalmar
Skne
Stockholm
Neuchtel
Neuchtel
Valais
Istanbul
Burgenland
Krnten
Niedersterreich
Niedersterreich
Obersterreich
Salzburg
Steiermark
Tirol
Vorarlberg
Wien
Wien
Soa
Primorsko-Goranska
Jihocesk
Jihocesk
Jihomoravsk
Krlovhradeck
Administrative
region 2
Stockholm
Stolichna

652
652
652
652
652
602
80
350
350
350
350
350
143
163
163
395
114
114
114
511
114
114
963
114
114
114
511
185
1011
224
255
742
225
2
2
2
6
2
2
2
2
3
Appendix 1
Species
Krlovhradeck
Libereck
Moravskoslezsk
steck
North Jutland
Western Finland
Alsace
le-de-France
Baden-Wrttemberg
Bayern
Bayern
Berlin
Niedersachsen
Nordrhein-Westfalen
Baranya
Somogy
Veszprm
Galway
Kerry
Lazio
Lazio
Toscana
Toscana
Trentino-Alto Adige
Trentino-Alto Adige
Trentino-Alto Adige
Veneto
Riga
Chisinau
Chisinau
Gelderland
Noord-Holland
Overijssel
Aust-Agder
Haut-Rhin
Yvelines
Tbingen
Berlin
Hannover
Pisa
Trento
Trento
Trento
Belluno
Taza
Arnhem
230
271
220
550
1026
34
331
753
794
332
333
1294
431
600
341
341
341
380
754
639
831
833
1331
98
1344
609
741
1344
319
528
666
642
365
1050
1050
1323
2
5
2
Continued
153
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Denmark
Finland
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Hungary
Hungary
Hungary
Ireland
Ireland
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Latvia
Moldova
Moldova
Morocco
Netherlands
Netherlands
Netherlands
Norway
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
154
Country
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Norway
Norway
Norway
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Russia
Serbia
Serbia
Serbia
Finnmark
Sr-Trndelag
Telemark
Greater Poland
Greater Poland
Lublin
Silesian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Aveiro
Azores
Beja
Braga
Castelo Branco
Coimbra
vora
Faro
Guarda
Leiria
Lisboa
Lisboa
Madeira
Portalegre
Porto
Santarm
Setbal
Viana do Castelo
Viseu
Vologda
Branicevski
Grad Beograd
Macvanski
Administrative
region 2
Hitra
Poznan
Lublin City
Tarnowskie
Mafra

558
1323
475
603
645
1337
1288
384
738
739
597
597
597
597
597
597
597
597
597
597
597
76
597
597
597
597
597
597
597
35
652
652
652
2
2
Appendix 1
Species
Niavski
Pomoravski
Raki
Sremski
Zajecarski
Bratislavsky
Kosicky
Kosicky
Nitriansky
Trenciansky
Zilinsky
La Rioja
Pas Vasco
Gvleborg
Gotland
Halland
Jnkping
Kalmar
Skne
Skne
Skne
Uppsala
Uppsala
Neuchtel
Neuchtel
Ticino
Ticino
Ticino
Valais
Valais
Istanbul
Soa
North Jutland
Pest
La Rioja
Gotland
Kungsbacka
Bstad
lvkarleby
Tierp
Stolichna
652
652
652
652
652
1349
1349
602
1349
1349
1349
315
80
695
350
470
470
426
350
350
696
470
470
470
143
163
1284
163
485
1284
163
395
185
1026
1320
344
3
3
Continued
155
Serbia
Serbia
Serbia
Serbia
Serbia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Spain
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Turkey
Bulgaria
Denmark
Hungary
Hungary
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia garinii
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
156
Country
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Italy
Italy
Moldova
Moldova
Morocco
Morocco
Poland
Poland
Portugal
Portugal
Portugal
Romania
Romania
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Switzerland
Toscana
Trentino-Alto Adige
Chisinau
Chisinau
Greater Poland
Lublin
Lisboa
Madeira
Setbal
Tulcea
Branicevski
Macvanski
Niavski
Pomoravski
Raki
Sremski
Zajecarski
Zajecarski
Banskobystricky
Bratislavsky
Kosicky
Kosicky
Nitriansky
Trenciansky
Trenciansky
Zilinsky
Zilinsky
Pas Vasco
Ticino
Administrative
region 2
Pisa
Trento
Taza
Lublin City
Mafra
Grndola

98
741
528
666
425
642
603
1337
76
238
76
1320
603
652
652
652
652
652
652
652
652
1320
602
1349
1349
602
1349
1349
603
1347
1349
80
1284
Appendix 1
Species
Ticino
Valais
Jendouba
Istanbul
Gvleborg
Kalmar
Skne
Bayern
Bayern
Niedersachsen
Burgenland
Niedersterreich
Obersterreich
Salzburg
Vorarlberg
Wien
Soa
Jihocesk
Jihomoravsk
Krlovhradeck
Krlovhradeck
Libereck
Moravskoslezsk
Plzensk
Alsace
Bayern
Berlin
Niedersachsen
Nordrhein-Westfalen
Galway
Kerry
Lazio
Lazio
Toscana
Trentino-Alto Adige
Trentino-Alto Adige
Veneto
Hannover
Stolichna
Haut-Rhin
Berlin
Hannover
Trento
Trento
Belluno
163
1284
425
395
350
350
350
333
333
431
114
114
114
114
114
114
185
255
742
225
230
271
220
230
331
333
1294
431
600
380
754
831
833
1331
609
741
1344
7
2
2
2
2
3
157
Switzerland
Switzerland
Tunisia
Turkey
Sweden
Sweden
Sweden
Germany
Germany
Germany
Austria
Austria
Austria
Austria
Austria
Austria
Bulgaria
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
France
Germany
Germany
Germany
Germany
Ireland
Ireland
Italy
Italy
Italy
Italy
Italy
Italy
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia lusitaniae
Borrelia miyamotoi
Borrelia miyamotoi
Borrelia miyamotoi
Borrelia spielmanii
Borrelia spielmanii
Borrelia spielmanii
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Continued
158
Country
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Borrelia valaisiana
Italy
Latvia
Moldova
Moldova
Netherlands
Netherlands
Netherlands
Netherlands
Poland
Poland
Portugal
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Slovakia
Slovakia
Spain
Spain
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Turkey
Veneto
Riga
Chisinau
Chisinau
Flevoland
Gelderland
Noord-Holland
Overijssel
Greater Poland
Lublin
Lisboa
Branicevski
Grad Beograd
Macvanski
Niavski
Pomoravski
Raki
Sremski
Zajecarski
Kosicky
Zilinsky
La Rioja
Pas Vasco
Neuchtel
Neuchtel
Neuchtel
Ticino
Ticino
Ticino
Valais
Valais
Istanbul
If more than one.
Administrative
region 2
Belluno
Arnhem
Lublin City
Mafra
La Rioja

324
319
528
666
1291
365
1050
1050
603
1337
76
652
652
652
652
652
652
652
652
602
1347
315
80
143
163
368
1284
163
485
1284
163
395
2
2
Appendix 1
Species
Table A1.7. Geographic distribution data for Babesia spp. See Appendix 2 (hosted on http://www.cabi.org/vetmedresource/) for complete reference indicated by
the Reference ID.
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bigemina
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Egypt
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Morocco
Morocco
Morocco
Morocco
Spain
Spain
Switzerland
Turkey
Turkey
Egypt
Italy
Italy
Italy
Italy
Italy
Al Jizah
Apulia
Basilicata
Calabria
Calabria
Campania
Emilia-Romagna
Lazio
Lazio
Marche
Sicily
Sicily
Sicily
Umbria
Umbria
Veneto
Veneto
Doukkala-Abda
Marrakech-Tensift-Al Haouz
Tadla-Azilal
Islas Baleares
Pas Vasco
Graubnden
Antalya
Kayseri
Al Jizah
Calabria
Calabria
Campania
Emilia-Romagna
Lazio
Crotone
Bologna
Roma
Palermo
Ragusa
Perugia
Padua
Crotone
Bologna
Reference ID
No. entriesa
1101
207
207
997
156
207
755
997
755
997
997
366
366
997
755
997
755
291
291
291
291
41
363
434
508
457
1101
997
156
997
755
997
2
3
2
3
3
Continued
159
Country
Species
160

Country
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia bovis
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia caballi
Babesia canis
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Spain
Spain
Spain
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Spain
Spain
Turkey
Turkey
Albania
Lazio
Marche
Sicily
Sicily
Umbria
Umbria
Veneto
Veneto
Islas Baleares
Pas Vasco
Pas Vasco
Jendouba
Nabeul
Nabeul
Kayseri
Tekirdag
Hajd-Bihar
Calabria
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Marche
Sardegna
Sicily
Toscana
Umbria
Umbria
Veneto
Galicia
Pas Vasco
Ankara
Malatya
Tiran
Roma
Palermo
Perugia
Padua
Bologna
Roma
Perugia
Padua
Reference ID
755
997
997
366
997
755
997
755
41
363
364
641
641
641
457
43
441
997
997
755
997
755
997
997
997
606
997
755
755
146
363
390
25
409
No. entriesa
3
3
3
3
3
4
2
2
2
2
2
2
2
3
2
Appendix 1
Species
Asyut
Baden-Wrttemberg
Baden-Wrttemberg
Bayern
Bayern
Berlin
Brandenburg
Bremen
Hamburg
Hessen
Niedersachsen
Nordrhein-Westfalen
Rheinland-Pfalz
Saarland
Sachsen
Sachsen-Anhalt
Schleswig-Holstein
Thringen
Attiki
Hajd-Bihar
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Lazio
Lombardia
Marche
Sicily
Sicily
Toscana
Umbria
Umbria
Umbria
Freiburg
Attica
Bologna
Bologna
Roma
Perugia
510
83
576
576
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
476
441
1107
755
755
1107
1107
997
755
997
997
997
999
1107
1107
997
755
2
2
3
Continued
161
Egypt
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Greece
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis
162

Country
Babesia canis
Babesia canis
Babesia canis
Babesia canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis canis
Babesia canis vogeli
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Italy
Italy
Poland
UK
Croatia
Croatia
Croatia
Croatia
Hungary
Hungary
Hungary
Hungary
Italy
Italy
Italy
Italy
Poland
Spain
Croatia
Croatia
Croatia
Croatia
Italy
Spain
Belgium
Estonia
France
Germany
Hungary
Hungary
Italy
Italy
Poland
Slovakia
Veneto
Veneto
Masovian
England
Bjelovarska-Bilogorska
Grad Zagreb
Medimurska
Varadinska
Budapest
Gyor-Moson-Sopron
Pest
Veszprm
Lazio
Umbria
Veneto
Pomeranian
Catalua
Grad Zagreb
Medimurska
Varadinska
Emilia-Romagna
Catalua
Flemish Brabant
Ida-Viru
Rhne-Alpes
Bayern
Borsod-Abaj-Zempln
Heves
Sicily
Veneto
West Pomeranian
Bratislavsk
Padua
Warsaw
Kent
Barcelona
Barcelona
Rhne
Palermo
Szczecin
Reference ID
1107
755
1098
440
88
88
88
88
340
341
341
341
1107
1107
1107
1107
190
978
88
88
88
88
1107
978
321
1335
261
564
442
442
366
997
890
1349
No. entriesa
Appendix 1
Species
Nitriansk
Pas Vasco
Graubnden
Graubnden
Ticino
Kayseri
Emilia-Romagna
Lazio
Umbria
Veneto
Malatya
Grad Zagreb
Medimurska
Varadinska
Baden-Wrttemberg
Budapest
Catalua
Apulia
Basilicata
Calabria
Pas Vasco
Amasya
Giresun
Gmshane
Tokat
Trabzon
Jihomoravsk
Harju
Ida-Viru
Prnu
Tartu
Tartu
Western Finland
Emilia-Romagna
Bologna
Roma
Perugia
Padua
Tbingen
Barcelona
1349
364
435
622
435
457
755
755
755
755
25
88
88
88
88
417
323
978
157
157
157
364
46
46
46
46
46
820
1335
1335
1335
1335
1348
34
1107
549
2
2
2
2
Continued
163
Slovakia
Spain
Switzerland
Switzerland
Switzerland
Turkey
Italy
Italy
Italy
Italy
Turkey
Croatia
Croatia
Croatia
Croatia
Germany
Hungary
Spain
Italy
Italy
Italy
Spain
Turkey
Turkey
Turkey
Turkey
Turkey
Czech Republic
Estonia
Estonia
Estonia
Estonia
Estonia
Finland
Italy
Poland
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia divergens
Babesia equi
Babesia equi
Babesia equi
Babesia equi
Babesia equi
Babesia gibsoni
Babesia gibsoni
Babesia gibsoni
Babesia gibsoni
Babesia gibsoni
Babesia gibsoni
Babesia gibsoni
Babesia major
Babesia major
Babesia major
Babesia major
Babesia major
Babesia major
Babesia major
Babesia major
Babesia major
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
164

Country
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia microti
Babesia motasi
Babesia motasi
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Babesia ovis
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Slovakia
Slovakia
Spain
Switzerland
Turkey
UK
Romania
Spain
Romania
Spain
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Lower Silesian
Lublin
Lublin
Lublin
Lublin
Masovian
Podlachian
Podlachian
Pomeranian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
West Pomeranian
West Pomeranian
West Pomeranian
Bratislavsk
Nitriansk
La Rioja
Graubnden
Nigde
England
Tulcea
Pas Vasco
Tulcea
Pas Vasco
Afyon
Diyarbakr
Elzig
Elzig
Erzincan
Erzurum
Igdr
Kayseri
Malatya
Katowice City
Parczew
Puawy
Wodawa
Biaystok
Sopot
Szczecin
Szczecin
La Rioja
Northumberland
Reference ID
504
1062
1061
1061
1061
504
504
72
504
1046
737
879
504
890
891
1349
1349
315
1305
188
131
59
677
59
677
186
23
23
45
23
23
23
458
23
No. entriesa
8
4
2
2
4
2
2
2
2
2
Appendix 1
Species
Turkey
Turkey
Turkey
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Estonia
Estonia
Estonia
Estonia
France
Greece
Greece
Italy
Italy
Netherlands
Netherlands
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Turkey
Turkey
Mardin
Mus
Sanlurfa
Burgenland
Krnten
Niedersterreich
Obersterreich
Salzburg
Salzburg
Steiermark
Steiermark
Tirol
Vorarlberg
Wien
Harju
Ida-Viru
Prnu
Tartu
Nord-Pas-de-Calais
Ipeiros
Thessalia
Calabria
Veneto
Gelderland
Zuid-Holland
Graubnden
Graubnden
Neuchtel
Ticino
Valais
Zrich
Diyarbakr
Istanbul
Oberwart
Wolfsberg
Voitsberg
Hallein
Sankt Johann im Pongau
Graz Umgebung
Mrzzuschlag
Imst
Bludenz
Wien
Nord
Ioannina
Trikala
Belluno
Arnhem
23
23
303
115
115
115
115
115
115
115
115
115
115
115
1335
1335
1335
1335
408
985
985
997
752
447
447
1211
438
166
166
166
166
1066
394
2
2
2
2
2
2
Babesia ovis
Babesia ovis
Babesia ovis
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia spp.
Babesia vogeli
a
If more than one.
165
166
Appendix 1
Table A1.8. Geographic distribution data for Theileria spp. See Appendix 2 (hosted on http://www.cabi.
org/vetmedresource/) for complete reference indicated by the Reference ID.
Species
Country
Administrative
region 1
Theileria annae
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria annulata
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Spain
Egypt
Egypt
Egypt
Italy
Italy
Morocco
Spain
Spain
Spain
Spain
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Italy
Italy
Italy
Italy
Spain
Spain
Tunisia
Tunisia
Tunisia
Turkey
Pas Vasco
Ad Daqahliyah
Ad Daqahliyah
Al Gharbiyah
Sicily
Sicily
Doukkala-Abda
Islas Baleares
Islas Baleares
Pas Vasco
Pas Vasco
Bja
Bizerte
Bizerte
Jendouba
Nabeul
Zaghouan
Adyaman
Amasya
Ankara
Ankara
Antalya
Bingl
Bingl
Elzig
Elzig
Erzincan
Erzurum
Erzurum
Giresun
Gmshane
Kars
Kayseri
Malatya
Mus
Mus
Sanlurfa
Tekirdag
Tokat
Trabzon
Van
Apulia
Basilicata
Basilicata
Sicily
Pas Vasco
Pas Vasco
Bizerte
Nabeul
Zaghouan
Ankara
Administrative
region 2
Palermo
Ragusa
Foggia
Matera
Potenza
Palermo

363
295
301
295
366
366
701
40
41
363
364
641
641
641
641
641
641
282
46
844
845
508
22
282
22
282
282
22
282
46
46
282
456
282
22
282
282
43
46
46
282
156
156
156
366
363
364
641
641
641
254
3
2
2
3
3
3
3
3
3
3
3
4
2
2
2
Continued
167
Species
Country
Administrative
region 1
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria buffeli
Theileria caballi
Theileria caballi
Theileria caballi
Theileria caballi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria equi
Theileria orientalis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria ovis
Theileria sergenti
Theileria sergenti
Turkey
Turkey
Turkey
Turkey
Turkey
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Morocco
Morocco
Morocco
Morocco
Spain
Spain
Turkey
Turkey
Turkey
Turkey
Turkey
Croatia
Spain
Spain
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Bingl
Elzig
Erzurum
Kayseri
Mus
Grad Zagreb
Medimurska
Varadinska
Grad Zagreb
Medimurska
Varadinska
Calabria
Lazio
Marche
Sardegna
Sicily
Toscana
Umbria
Veneto
Fs-Boulemane
Grand Casablanca
Galicia
Pas Vasco
Ankara
Bingl
Elzig
Erzurum
Mus
Pas Vasco
Pas Vasco
Aksaray
Bingl
ankr
ankr
Diyarbakr
Elzig
Elzig
Erzincan
Erzurum
Kars
Malatya
Mardin
Mersin
Mus
Van
Bingl
Elzig
Administrative
region 2
Fs
Rabat
Marrakech
Taza

22
22
22
456
22
88
88
88
88
88
88
88
88
997
997
997
997
997
606
997
997
800
800
800
800
146
363
390
22
22
22
22
277
363
677
1353
1353
1353
1353
44
1353
21
44
44
44
44
44
1353
44
1353
22
22
2
2
2
2
2
2
2
2
2
2
6
2
2
2
2
2
2
2
Continued
168
Appendix 1
Species
Country
Administrative
region 1
Theileria sergenti
Theileria sergenti
Theileria spp.
Theileria spp.
Theileria spp.
Theileria spp.
Theileria spp.
Theileria spp.
Theileria spp.
Turkey
Turkey
Switzerland
Switzerland
Turkey
Turkey
Turkey
Turkey
Turkey
Erzurum
Mus
Graubnden
Graubnden
Amasya
Giresun
Gmshane
Tokat
Trabzon
Administrative
region 2

22
22
1211
438
46
46
46
46
46
If more than one.
Table A1.9. Geographic distribution data for equine piroplasmoses. See Appendix 2 (hosted on http://
www.cabi.org/vetmedresource/) for complete reference indicated by the Reference ID.
Species
Country
Administrative
region 1
Theileria caballi
Theileria equi
Theileria caballi
Theileria equi
Theileria caballi
Theileria equi
Theileria caballi
Theileria equi
Ehrlichia equi
Babesia caballi
Babesia caballi
Theileria equi
Babesia caballi
Babesia caballi
Babesia equi
Babesia caballi
Theileria equi
Babesia caballi
Babesia equi
Babesia caballi
Theileria equi
Babesia caballi
Theileria equi
Babesia caballi
Theileria equi
Babesia caballi
Theileria equi
Babesia caballi
Theileria equi
Babesia caballi
Babesia equi
Theileria equi
Babesia caballi
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Denmark
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Grad Zagreb
Grad Zagreb
Medimurska
Medimurska
Varazdinska
Varazdinska
North Jutland
Hajd-Bihar
Calabria
Calabria
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Lazio
Lazio
Marche
Marche
Sardegna
Sardegna
Sicily
Sicily
Toscana
Toscana
Umbria
Umbria
Umbria
Umbria
Veneto
Veneto
Administrative
region 2
Bologna
Bologna
Roma
Roma
Perugia
Perugia
Padua

88
88
88
88
88
88
88
88
946
441
997
997
997
755
755
997
997
755
755
997
997
997
997
997
997
606
606
997
997
755
755
997
755
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Continued
169
Species
Country
Babesia equi
Theileria equi
Theileria equi
Theileria equi
Italy
Morocco
Morocco
Morocco
Theileria equi
Morocco
Babesia caballi
Theileria equi
Babesia caballi
Theileria equi
Babesia caballi
Theileria equi
Babesia caballi
Babesia equi
Spain
Spain
Spain
Spain
Turkey
Turkey
Turkey
Turkey
Administrative
region 1
Administrative
region 2
Veneto
Fs-Boulemane
Grand Casablanca
Marrakech-Tensift-Al
Haouz
Taza-Al HoceimaTaounate
Galicia
Galicia
Pas Vasco
Pas Vasco
Ankara
Ankara
Malatya
Malatya
Padua
Fs
Rabat
Marrakech
755
800
800
800
Taza
800
146
146
363
363
390
390
25
25
3
2
2
2
If more than one.
Table A1.10. Geographic distribution data for Bartonella spp. See Appendix 2 (hosted on http://www.
cabi.org/vetmedresource/) for complete reference indicated by the Reference ID.
Species
Country
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Croatia
Denmark
Denmark
France
France
France
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
France
Greece
Italy
Italy
Italy
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Bartonella henselae
Italy
Italy
Italy
Italy
Italy
Norway
Poland
Poland
Poland
Poland
Poland
Spain
Spain
Turkey
Administrative
region 1
Zadarska
Frederiksborg
Fyn
le-de-France
Lorraine
Provence-AlpesCte-dAzur
Rhne-Alpes
Kriti
Emilia-Romagna
Emilia-Romagna
Friuli-Venezia
Giulia
Lombardia
Sardegna
Toscana
Toscana
Toscana
Sr-Trndelag
Ldz
Lublin
Masovian
Masovian
Andaluca
Catalua
Ankara
Administrative
region 2
Helsingr
Snders
Meurthe-et-Moselle
Bouches-du-Rhne
Rhne
Modena
Warsaw
Barcelona
No.
Reference ID entriesa
1194
304
304
1179
1179
1179
1179
1246
1182
1181
830
1182
1182
1182
1195
1237
1161
1177
1138
1138
1138
763
1128
1261
1175
2
4
Continued
170
Appendix 1
Species
Country
Bartonella quintana
Bartonella quintana
France
France
Bartonella quintana
Bartonella quintana
Bartonella quintana
Bartonella spp.
Bartonella spp.
Bartonella spp.
Bartonella spp.
Bartonella spp.
Bartonella spp.
Bartonella spp.
Italy
Poland
Poland
Albania
Algeria
Algeria
Algeria
Algeria
Czech
Republic
Czech
Republic
France
Bartonella spp.
Bartonella spp.
Bartonella spp.
Bartonella spp.
Bartonella vinsonii
Greece
Italy
Italy
Spain
Denmark
Bartonella spp.
Administrative
region 1
le-de-France
Toscana
Administrative
region 2
No.
Reference ID entriesa
1206
1265
Bouches-du-Rhne
Masovian
Tiran
Alger
Biskra
Mascara
Tiaret
Jihomoravsk
Prague
Nord-Pas-deCalais
Thessalia
Apulia
Basilicata
Andaluca
Fyn
Nord
1195
1177
1138
409
1167
1167
1167
1167
427
4
4
4
4
4
427
408
Thessaloniki
Taranto
1191
1191
1191
1128
304
Snders
2
2
If more than one.
Table A1.11. Geographic distribution data for Francisella tularensis. See Appendix 2 (hosted on http://
Country
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Croatia
Czech Republic
Czech Republic
France
Germany
Germany
Slovakia
Slovakia
Spain
Spain
Spain
Spain
Sweden
Sweden
Burgenland
Niedersterreich
Niedersterreich
Niedersterreich
Niedersterreich
Niedersterreich
Steiermark
Jihomoravsk
Jihomoravsk
Pays de la Loire
Hessen
Niedersachsen
Banskobystrick
Trnavsk
Castilla y Len
Castilla y Len
Castilla y Len
Castilla y Len
Dalarna
Gvleborg
Gnserndorf
Gnserndorf
Korneuburg
Mistelbach
Vende
Soria
Valladolid
Valladolid
Reference ID
259
259
1053
401
401
1053
259
656
1053
171
882
509
509
401
401
39
31
617
91
300
300
No. entriesa
2
7
2
4
3
Continued
171

Country
Sweden
Sweden
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Jmtland
Vsternorrland
Amasya
Ankara
Antalya
Bursa
Dzce
Edirne
Edirne
Kars
Kars
Kastamonu
Krklareli
Krklareli
Kocaeli
Samsun
Tekirdag
Tekirdag
Van
Zinguldak
Reference ID
No. entriesa
300
300
560
19
19
19
710
19
250
19
825
19
19
250
19
19
19
250
19
19
If more than one.
Table A1.12. Geographic distribution data for Coxiella burnetii. See Appendix 2 (hosted on http://www.
Country
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Algeria
Belgium
Belgium
Belgium
Belgium
Belgium
Bosnia and Herzegovina
Administrative
region 1
Berat
Dibr
Dibr
Elbasan
Elbasan
Elbasan
Fier
Fier
Gjirokastr
Gjirokastr
Kor
Lezh
Tiran
Tiran
Stif
Antwerp
East Flanders
Flemish Brabant
Hainaut
West Flanders
Federacija Bosna i
Hercegovina
Administrative
region 2
Dibrs
Matit
Gramshit
Librazhdit
Lushnjs
Prmetit
Kolonjs
Kavajs
Tirans

1174
1174
1174
1174
1174
1174
1174
1174
1174
1174
1174
1174
1174
1174
1221
1198
1198
1198
1198
1198
1209
2
3
3
3
2
2
3
2
3
9
Continued
172
Appendix 1
Country
Bulgaria
Cyprus
Cyprus
Cyprus
Cyprus
Cyprus
Cyprus
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Egypt
Egypt
Egypt
France
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Greece
Greece
Greece
Greece
Greece
Administrative
region 1
Federacija Bosna i
Hercegovina
Republika Srpska
Soa
Famagusta
Larnaca
Limassol
Nicosia
Paphos
rhus
Frederiksborg
Fyn
North Jutland
Ribe
Ringkbing
South Jutland
Storstrm
Vejle
Vestsjlland
Viborg
Al Jizah
Qina
Shamal Sina
Centre
Baden-Wrttemberg
Bayern
Berlin
Brandenburg
Bremen
Hamburg
Hessen
MecklenburgVorpommern
Niedersachsen
Nordrhein-Westfalen
Rheinland-Pfalz
Saarland
Sachsen
Sachsen-Anhalt
Schleswig-Holstein
Thringen
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Administrative
region 2

1276
Botevgrad
1276
1254
1118
777
777
777
777
777
1149
1149
1149
1149
1149
1149
1149
1149
1149
1149
1149
1343
1343
1343
1164
599
599
599
599
599
599
599
599
Drama
599
599
599
599
599
599
599
599
1254
Evros
1254
Kavala
1254
Rodopi
1254
Xanthi
1254
4
4
3
4
5
Continued
173
Country
Administrative
region 1
Administrative
region 2
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Israel
Israel
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Montenegro
Netherlands
Netherlands
Poland
Slovakia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Tunisia
Turkey
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Haifa
Jerusalem
Apulia
Apulia
Basilicata
Campania
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Lombardia
Lombardia
Lombardia
Sardegna
Sicily
Veneto
abljak
Andrijevica
Berane
Cetinje
Danilovgrad
Mojkovac
Nikoic
Pljevlja
Pluine
Podgorica
Ulcinj
Gelderland
Noord-Brabant
Lublin
Koick
Castilla y Len
Castilla-La Mancha
Castilla-La Mancha
Catalua
Pas Vasco
Pas Vasco
Pas Vasco
Pas Vasco
Sousse
Ankara
Florina
Grevena
Grevena
Kastoria
Kozani
Imathia
Khalkidiki
Kilkis
Pella
Pieria
Serrai
Thessaloniki
Bari
Modena
Parma
Reggio NellEmilia
Cremona
Mantua
Padua
Soria
Albacete
Toledo
Barcelona

1254
1254
1254
1254
1254
1254
1254
1254
1254
1254
1254
1254
1252
1275
1257
1247
1257
1259
1354
1354
1354
1197
1300
1300
1236
999
1300
1222
1222
1222
1222
1222
1222
1222
1222
1222
1222
1222
1271
1271
1184
1193
1250
1158
989
1153
1201
1266
78
79
1159
1214
3
2
12
Continued
174
Appendix 1
Country
Administrative
region 1
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
Ankara
Antalya
Antalya
Aydn
Bolu
Diyarbakr
Kayseri
Nigde
Samsun
Tokat
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Administrative
region 2
Antrim
Ballymena
Banbridge
Craigavon
Derry
Down
Dungannon
Fermanagh
Limavady
Magherafelt
Moyle
Omagh
Strabane

1216
1160
1204
1215
1213
1160
1214
1214
1160
1203
1239
1239
1239
1239
1239
1239
1239
1239
1239
1239
1239
1239
1239
If more than one.
Table A1.13. Geographic distribution data for Ornithodoros spp. See Appendix 2 (hosted on http://www.
Country
Algeria
Armenia
Armenia
Armenia
Armenia
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
El Tarf
Lake Sevan
Lori
Shirak
Tavush
Ad Daqahliyah
Al Buhayrah
Al Gharbiyah
Al Iskandariyah
Al Ismailiyah
Al Jizah
Al Minuyah
Al Qahirah
Al Qalyubiyah
Ash Sharqiyah
Bur Said
Dumyat
Kafr ash Shaykh
Matruh
Shamal Sina
Ain El Assel
Reference ID
0
0
0
0
0
0
0
0
0
0
1101
0
0
0
0
0
0
0
0
0
Continued
175

Country
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Georgia
Israel
Israel
Israel
Israel
Israel
Israel
Israel
Jordan
Jordan
Jordan
Jordan
Jordan
Jordan
Jordan
Jordan
Jordan
Lebanon
Lebanon
Lebanon
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Palestine
Palestine
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Abkhazia
Ajaria
Guria
Imereti
Kakheti
Kvemo Kartli
Mtskheta-Mtianeti
Racha-Lechkhumi-Kvemo Svaneti
Samegrelo-Zemo Svaneti
Samtskhe-Javakheti
Shida Kartli
Tbilisi
Golan
HaDarom
Haifa
HaMerkaz
HaZafon
Jerusalem
Tel Aviv
Ajlun
Amman
Aqaba
Balqa
Irbid
Jarash
Karak
Madaba
Talah
An Nabatiyah
Beirut
South Lebanon
Chaouia-Ouardigha
Doukkala-Abda
Grand Casablanca
Guelmim-Es-Semara
Oriental
Rabat-Sal-Zemmour-Zaer
Souss-Massa-Dra
Tadla-Azilal
Tanger-Ttouan
Gaza
West Bank
Beja
Castelo Branco
Coimbra
Faro
Leiria
Lisboa
Portalegre
Santarm
Gagra
Batumi
Chokhatauri
Bagdati
Akhmeta
Bolnisi
Akhalgori
Ambrolauri
Abasha
Adigeni
Gori
Tbilisi
Ajloun
Amman
Aqaba
Al-Balqa
Aghwar Shamaliyyeh
Jarash
Ayy
Dhiban
Bsaira
Hasbaya
Beirut
Jezzine
Ben Slimane
El Jadida
Knitra
Casablanca
Assa-Zag
Al Haouz
Berkane Taourirt
Khmisset
Agadir-Ida ou Tanane
Azilal
Chefchaouen
Deir Al-Balah
Bethlehem
Aljustrel
Belmonte
Arganil
Albufeira
Alcobaa
Alenquer
Alter do Cho
Mao
Reference ID
1398
1398
1398
1398
1398
1398
1398
1398
1398
1398
1398
1398
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Continued
176
Appendix 1

Country
Portugal
Portugal
Russia
Russia
Russia
Russia
Spain
Spain
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Setbal
vora
Chechnya
Ingush
Kabardin-Balkar
North Ossetia
Andaluca
Extremadura
Ariana
Ben Arous (Tunis Sud)
Bizerte
Bja
Gabs
Jendouba
Manubah
Mdenine
Nabeul
Tunis
Zaghouan
Artvin
Rize
Trabzon
Alccer do Sal
Alandroal
Achkhoy-Martanovskiy rayon
Malgobekskiy rayon
Baksanskiy rayon
Alagirskiy rayon
Almera
Badajoz
Ariana Mdina
Ben Arous
Bizerte Nord
Amdoun
Gabs Mdina
An Draham
Borj El Amri
Ben Guerdane
Beni Khalled
Bab Bhar
Bir Mchergua
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table A1.14. Geographic distribution data for Ixodes ricinus. See Appendix 2 (hosted on http://www.cabi.
org/vetmedresource/) for complete reference indicated by the Reference ID.
Country
Algeria
Algeria
Algeria
Algeria
Algeria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Austria
Alger
Constantine
Guelma
Guelma
Tlemcen
Burgenland
Burgenland
Krnten
Krnten
Niedersterreich
Obersterreich
Obersterreich
Salzburg
Salzburg
Steiermark
Steiermark
Steiermark
Steiermark
Steiermark
Tirol
Tirol
Vorarlberg
Ain Benian
Didouche Mourad
Hammam Debagh
Hammam Debagh
Tlemcen
Oberpullendorf
Oberwart
Villach
Wolfsberg
Mistelbach
Braunau am Inn
Urfahr Umgebung
Hallein
Sankt Johann im Pongau
Deutschlandsberg
Graz Umgebung
Mrzzuschlag
Voitsberg
Weiz
Imst
Schwaz
Bludenz
Reference ID
No. entriesa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
Continued
177

Country
Austria
Belgium
Belgium
Belgium
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Croatia
Croatia
Croatia
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Vorarlberg
Antwerp
Lige
West Flanders
Blagoevgrad
Blagoevgrad
Burgas
Burgas
Burgas
Burgas
Dobrich
Grad Soya
Kardzhali
Kardzhali
Kyustendil
Lovech
Montana
Pazardzhik
Pazardzhik
Plovdiv
Plovdiv
Sliven
Smolyan
Soa
Soa
Stara Zagora
Stara Zagora
Stara Zagora
Varna
Vidin
Vratsa
Yambol
Medimurska
Primorsko-Goranska
Varadinska
Jihocesk
Jihocesk
Jihocesk
Jihocesk
Jihocesk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsk
Jihomoravsky
Kraj Vysocina
Feldkirch
Turnhout
Lige
Brugge
Gotse Delchev
Sandanski
Burgas
Kameno
Malko Tarnovo
Tsarevo
Shabla
Stolichna
Kirkovo
Momchilgrad
Rila
Letnitsa
Montana
Pazardzhik
Strelcha
Krichim
Plovdiv
Sliven
Zlatograd
Svoge
Zlatitsa
Chirpan
Haskovo
Pavel Banya
Valchi Dol
Belogradchik
Vratsa
Bolyarovo
Cesk Budejovice
Jindrichuv Hradec
Psek
Tbor
Blansko
Breclav
Brno
Brno
Brno-Venkov
Vykov
Znojmo
dr nad Szavou
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
69
0
0
0
0
0
0
0
0
216
0
0
0
0
1082
427
490
742
0
0
0
472
0
0
0
0
No. entriesa
4
2
12
2
2
2
10
2
2
2
2
2
2
2
4
2
8
2
2
6
2
2
2
2
2
4
4
10
4
4
4
4
2
6
12
4
6
3
13
Continued
178
Appendix 1

Country
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Czech Republic
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Denmark
Kraj Vysocina
Krlovhradeck
Krlovhradeck
Krlovhradeck
Krlovhradeck
Libereck
Libereck
Libereck
Moravskoslezsk
Moravskoslezsk
Moravskoslezsk
Olomouck
Olomouck
Olomouck
Pardubick
Pardubick
Plzensk
Plzensk
Plzensk
Plzensk
Plzensk
Plzensk
Plzensk
Prague
Prague
Stredocesk
Stredocesk
Stredocesk
Stredocesk
Stredocesk
Stredocesk
Zlnsk
rhus
rhus
rhus
rhus
Bornholm
Copenhagen
Copenhagen
Copenhagen
Frederiksborg
Frederiksborg
Fyn
North Jutland
Ribe
Ringkbing
Roskilde
South Jutland
Storstrm
Vejle
Vestsjlland
Vestsjlland
Trebc
Nchod
Trutnov
Trutnov
Trutnov
Liberec
Liberec
Bruntl
Opava
Jesenk
Olomouc
Prostejov
st nad Orlic
st nad Orlic
Domalice
Klatovy
Plzen
Plzen-Jih
Plzen-Sever
Rokycany
Tachov
Beneov
Beroun
Koln
Praha-v chod
Praha-zpad
Prbram
Kromer
Ebeltoft
Hammel
Skanderborg
Helsinge
Korsr
Reference ID
0
0
0
620
621
271
0
472
220
0
0
0
0
0
0
472
0
0
0
0
0
0
0
216
427
0
0
0
0
0
0
0
888
0
0
0
888
478
479
888
888
0
888
888
888
888
888
888
888
888
888
973
No. entriesa
3
2
24
5
4
4
2
6
7
2
2
5
3
2
2
2
4
2
Continued
Plate 6. Ixodes canisuga: adult female dorsal view (a) and ventral view (b).
Plate 7. Ixodes uriae: adult male dorsal view (a) and ventral view (b); adult female dorsal view (c)
and ventral view (d).
7c
Plate 20. Reported occurrence of Ixodes ricinus. Smallest administrative region or territorial unit for statistics (NUTS), data for 20002010;
Coordinate (latitude/longitude), data for 20002010; Coordinate (latitude/longitude), historical data (before 2000).
7d
7a
7b
6b
18b
18c
18d
19a
19c
6a
18a
8a
8b
8c
8d
9a
9b
19b
19d
Plate 18. Dermacentor marginatus: adult male dorsal view (a) and ventral view (b); adult female dorsal view
(c) and ventral view (d).
Plate 19. Dermacentor reticulatus: adult male dorsal view (a) and ventral view (b); adult female dorsal view
Plate 8. Haemaphysalis punctata: adult male dorsal view (a) and ventral view (b); adult female dorsal view (c)
Plate 9. Haemaphysalis concinna: adult male dorsal view (a) and ventral view (b).
Plate 23. Reported occurrence of Dermacentor marginatus. Smallest administrative region or territorial unit for statistics (NUTS), data for 20002010;
Coordinate (latitude/longitude), data for 20002010. Coordinate (latitude/longitude), historical data (before 2000).
Plate 1. Argas persicus: adult male dorsal view (a) and ventral view (b).
Plate 2. Argas reflexus: adult male dorsal view (a) and ventral view (b).
Plate 3. Ornithodoros erraticus: adult (sex undermined) dorsal view (a) and ventral view (b).
3a
3b
2a
2b
1a
1b
13a
13b
14a
14b
13c
13d
14c
14d
Plate 13. Hyalomma scupense: adult male dorsal view (a) and ventral view (b); adult female dorsal view
Plate 14. Rhipicephalus sanguineus: adult male dorsal view (a) and ventral view (b); adult female dorsal view
Plate 4. Ixodes ricinus: adult male dorsal view (a) and ventral view (b); adult female dorsal view (c) and
ventral view (d).
Plate 12. Hyalomma excavatum: adult male dorsal view (a) and ventral view (b); adult female dorsal view
4a
Plate 15. Rhipicephalus turanicus, adult male dorsal view (a) and ventral view (b); adult female dorsal view
4c
4d
Plate 22. Reported occurrence of Rhipicephalus bursa. Smallest administrative region or territorial unit for statistics (NUTS), historical data (before 2000);
Smallest administrative region or territorial unit for statistics (NUTS), data for 20002010; Coordinate (latitude/longitude), historical data (before 2000);
Coordinate (latitude/longitude), data for 20002010.
15a
15b
12a
12b
15c
15d
12c
12d
4b
Plate 21. Reported occurrence of Hyalomma marginatum. Smallest administrative region or territorial unit for statistics (NUTS), data for 20002010;
Plate 5. Ixodes hexagonus: adult male dorsal view (a) and ventral view (b); adult female dorsal view (c) and
ventral view (d).
5d
5c
5b
5a
11a
Coordinate (latitude/longitude), data for 20002010; Coordinate (latitude/longitude), historical data (before 2000).
10
16a
16b
16c
16d
11b
17a
11c
17b
11d
17c
Plate 10. Haemaphysalis inermis: adult female ventral view.

Plate 11. Hyalomma marginatum: adult male dorsal view (a) and ventral view (b); adult female dorsal view
17d
Plate 16. Rhipicephalus bursa: adult male dorsal view (a) and ventral view (b); adult female dorsal view (c)
Plate 17. Rhipicephalus annulatus: adult male dorsal view (a) and ventral view (b); adult female dorsal view
179

Country
Denmark
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Estonia
Viborg
Harju
Hiiu
Jrva
Jgeva
Lne
Lne-Viru
Prnu
Plva
Rapla
Saare
Tartu
Valga
888
0
0
0
0
0
0
0
0
0
0
0
0
Estonia
Estonia
Faroe Islands
Finland
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Viljandi
Vru
0
0
471
0
0
0
331
0
0
0
0
0
0
0
0
0
132
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Southern Finland
Alsace
Alsace
Alsace
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Auvergne
Auvergne
Auvergne
Auvergne
Auvergne
Basse-Normandie
Basse-Normandie
Basse-Normandie
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bretagne
Bretagne
Bretagne
Bretagne
Centre
Centre
Centre
Centre
Centre
Centre
Champagne-Ardenne
Champagne-Ardenne
Champagne-Ardenne
Champagne-Ardenne
Corse
Uusimaa
Bas-Rhin
Haut-Rhin
Haut-Rhin
Dordogne
Gironde
Landes
Lot-et-Garonne
Pyrnes-Atlantiques
Allier
Cantal
Haute-Loire
Puy-de-Dme
Puy-de-Dme
Calvados
Manche
Orne
Cte-dOr
Nivre
Sane-et-Loire
Yonne
Ctes-dArmor
Finistre
Ille-et-Vilaine
Morbihan
Cher
Eure-et-Loir
Indre
Indre-et-Loire
Loir-et-Cher
Loiret
Ardennes
Aube
Haute-Marne
Marne
Corse-du-Sud
Reference ID
No. entriesa
27
13
18
13
17
26
30
8
23
24
11
12
17
11
5
2
60
28
39
21
24
7
18
33
31
7
7
22
24
14
11
9
45
30
88
37
43
50
30
10
21
29
12
10
3
38
24
16
2
Continued
180
Appendix 1

Country
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Franche-Comt
Franche-Comt
Franche-Comt
Haute-Normandie
Haute-Normandie
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
Limousin
Limousin
Limousin
Lorraine
Lorraine
Lorraine
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Nord-Pas-de-Calais
Nord-Pas-de-Calais
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Picardie
Picardie
Picardie
Poitou-Charentes
Poitou-Charentes
Poitou-Charentes
Poitou-Charentes
France
France
France
France
France
France
Rhne-Alpes
Doubs
Haute-Sane
Jura
Eure
Seine-Maritime
Essonne
Seine-et-Marne
Val-dOise
Val-de-Marne
Ville de Paris
Yvelines
Yvelines
Aude
Gard
Hrault
Pyrnes-Orientales
Corrze
Creuse
Haute-Vienne
Meurthe-et-Moselle
Meuse
Vosges
Arige
Aveyron
Gers
Haute-Garonne
Hautes-Pyrnes
Lot
Tarn
Tarn-et-Garonne
Nord
Pas-de-Calais
Loire-Atlantique
Maine-et-Loire
Mayenne
Sarthe
Vende
Aisne
Oise
Somme
Charente
Charente-Maritime
Deux-Svres
Vienne
Alpes-De-HauteProvence
Alpes-Maritimes
Bouches-du-Rhne
Hautes-Alpes
Var
Vaucluse
Ain
Reference ID
No. entriesa
0
0
0
0
0
0
0
0
0
0
0
1024
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
8
15
20
4
18
31
10
8
8
40
0
0
0
0
0
0
8
2
10
14
21
4
24
50
19
17
20
11
13
14
2
8
4
6
2
23
16
14
24
7
18
26
2
11
9
8
10
6
4
14
8
10
2
208
Continued
181

Country
France
France
France
France
France
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Baden-Wrttemberg
Baden-Wrttemberg
Baden-Wrttemberg
Baden-Wrttemberg
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Bayern
Berlin
Brandenburg
Brandenburg
Hessen
Hessen
Hessen
Ardche
Drme
Haute-Savoie
Isre
Loire
Rhne
Savoie
Freiburg
Karlsruhe
Stuttgart
Tbingen
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Niedersachsen
Nordrhein-Westfalen
Nordrhein-Westfalen
Nordrhein-Westfalen
Nordrhein-Westfalen
Rheinland-Pfalz
Rheinland-Pfalz
Saarland
Sachsen
Sachsen
Sachsen-Anhalt
Sachsen-Anhalt
Sachsen-Anhalt
Thringen
Thringen
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Mittelfranken
Niederbayern
Oberbayern
Oberfranken
Oberpfalz
Schwaben
Unterfranken
Brandenburg
Darmstadt
Gieen
Kassel
MecklenburgVorpommern
Braunschweig
Kln
Munster
Koblenz
Rheinhessen-Pfalz
Saarland
Chemnitz
Leipzig
Dessau
Halle
Magdeburg
Thringen
Drama
Evros
Kavala
Rodopi
Florina
Grevena
Reference ID
0
0
0
0
0
0
0
0
0
0
0
332
333
564
0
0
0
0
0
0
0
236
236
0
496
0
0
0
0
600
857
0
0
0
0
0
0
0
0
0
0
268
0
735
0
0
0
0
735
0
0
No. entriesa
6
27
87
159
4
4
83
36
18
16
17
2
5
6
52
29
2
14
4
4
294
5
5
114
2
15
12
121
2
15
9
12
2
3
12
Continued
182
Appendix 1

Country
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Dytiki Makedonia
Ipeiros
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Bcs-Kiskun
Bcs-Kiskun
Bcs-Kiskun
Baranya
Baranya
Baranya
Bks
Borsod-Abaj-Zempln
Borsod-Abaj-Zempln
Borsod-Abaj-Zempln
Borsod-Abaj-Zempln
Budapest
Csongrd
Fejr
Fejr
Gyor-Moson-Sopron
Gyor-Moson-Sopron
Gyor-Moson-Sopron
Hajd-Bihar
Hajd-Bihar
Heves
Jsz-Nagykun-Szolnok
Komrom-Esztergom
Pest
Pest
Somogy
Szabolcs-Szatmr-Bereg
Vas
Veszprm
Veszprm
Zala
Cavan
Clare
Cork
Donegal
Dublin
Galway
Galway
Kerry
Kerry
Kerry
Kerry
Kilkenny
Kozani
Preveza
Khalkidiki
Khalkidiki
Kilkis
Pieria
Serrai
Thessaloniki
Reference ID
No. entriesa
0
0
735
0
1303
0
0
0
0
0
322
341
0
322
341
341
0
322
341
443
322
341
0
322
0
322
341
0
322
0
0
341
0
344
341
341
341
0
341
0
0
0
0
0
0
0
380
0
419
420
754
0
8
2
3
6
3
16
8
24
7
2
2
8
2
7
4
3
3
2
5
2
3
10
8
Continued
183

Country
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Ireland
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Limerick
Longford
Mayo
Meath
Monaghan
Offaly
Roscommon
Sligo
Waterford
Wexford
Wicklow
Abruzzo
Abruzzo
Apulia
Apulia
Basilicata
Basilicata
Calabria
Calabria
Campania
Campania
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Lazio
Lazio
Lazio
Liguria
Liguria
Liguria
Lombardia
Lombardia
Lombardia
Lombardia
Lombardia
Italy
Italy
Italy
Italy
Italy
Italy
Lombardia
Lombardia
Lombardia
Piemonte
Piemonte
Piemonte
LAquila
Teramo
Bari
Lecce
Matera
Potenza
Catanzaro
Reggio di Calabria
Avellino
Bologna
Ferrara
Forl-Cesena
Parma
Piacenza
Ravenna
Reggio nellEmilia
Rimini
Pordenone
Trieste
Udine
Frosinone
Latina
Rieti
Roma
Viterbo
Genova
Imperia
Savona
Brescia
Como
Lecco
Monza and Brianza
Pavia
Sondrio
Varese
Alessandria
Biella
Cuneo
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
802
0
0
0
0
0
0
0
0
0
215
0
0
0
0
0
0
0
0
0
0
0
847
0
0
0
0
0
0
0
0
0
0
No. entriesa
2
2
6
2
2
2
32
4
2
2
6
8
2
2
8
12
3
12
9
6
2
2
10
7
9
5
18
10
23
13
8
2
12
4
2
2
4
4
2
2
2
6
Continued
184
Appendix 1

Country
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Kosovo
Kosovo
Kosovo
Kosovo
Kosovo
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Piemonte
Piemonte
Piemonte
Sardegna
Sicily
Sicily
Toscana
Toscana
Toscana
Toscana
Toscana
Toscana
Toscana
Toscana
Trentino-Alto Adige
Trentino-Alto Adige
Trentino-Alto Adige
Trentino-Alto Adige
Trentino-Alto Adige
Umbria
Umbria
Veneto
Veneto
Veneto
Veneto
Veneto
Veneto
Gnjilane
Kosovska Mitrovica
Kosovska Mitrovica
Prizren
Prizren
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Kurzeme
Latgale
Latgale
Riga
Riga
Riga
Riga
Riga
Riga
Riga
Riga
Novara
Torino
Vercelli
Oristano
Palermo
Palermo
Florence
Grosseto
Livorno
Lucca
Pisa
Pistoia
Siena
Bolzano
Trento
Trento
Trento
Trento
Perugia
Terni
Belluno
Padua
Treviso
Verona
Verona
Vicenza
Kosovska Kamenica
Kosovska Mitrovica
Kosovska Mitrovica
Draga
Prizren
Kuldiga
Kuldiga
Liepaja
Liepaja
Saldus
Saldus
Talsi
Talsi
Ventspils
Ventspils
Daugavpils
Kraslava
Limbai
Limbai
Ogre
Ogre
Riga
Riga
Tukums
Tukums
Reference ID
No. entriesa
0
0
0
0
0
242
1331
0
0
0
0
0
0
0
0
0
160
609
741
0
0
0
0
0
0
0
0
0
0
346
0
0
0
126
0
126
0
126
0
126
0
126
126
126
0
126
0
126
0
126
0
126
12
8
2
2
6
3
6
2
6
2
2
28
150
11
6
2
9
2
2
9
14
4
8
7
4
3
5
5
Continued
185

Country
Latvia
Vidzeme
Cesis
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Latvia
Libya
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Lithuania
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Vidzeme
Zemgale
Zemgale
Zemgale
Zemgale
Zemgale
Zemgale
Zemgale
Zemgale
Al Hizam Al Akhdar
Alytaus
Alytaus
Alytaus
Kauno
Kauno
Kauno
Kauno
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Klaipedos
Marijampoles
Marijampoles
Panevezio
Panevezio
Panevezio
Panevezio
Panevezio
iauliai
iauliai
iauliai
iauliai
iauliai
iauliai
iauliai
Utenos
Utenos
Utenos
Utenos
Vilniaus
Vilniaus
Vilniaus
Vilniaus
Cesis
Cesis
Valka
Valka
Valmiera
Valmiera
Aizkraukle
Aizkraukle
Bauska
Bauska
Dobele
Dobele
Jelgava
Jelgava
Alytaus
Varenos
Varenos
Kauno
Kauno
Prienu
Prienu
Klaipedos
Klaipedos
Neringos
Neringos
ilutes
ilutes
Marijampoles
Marijampoles
Paneveio
Paneveio
Rokikio
Rokikio
Jonikio
Jonikio
Kelmes
Kelmes
Pakruojo
iauliu
iauliu
Ignalinos
Utenos
Utenos
Zarasu
Ukmerges
Ukmerges
Vilniaus
Reference ID
No. entriesa
0
0
126
0
126
0
126
0
126
0
126
0
126
0
126
0
0
0
732
0
732
0
732
733
0
732
0
732
0
732
0
732
733
0
732
0
732
0
732
0
732
0
0
732
0
0
732
0
733
0
732
0
2
4
4
3
2
4
2
2
2
2
Continued
186
Appendix 1

Country
Lithuania
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Vilniaus
Anenii Noi
Balti
Basarabeasca
Bender
Briceni
Cahul
Calarasi
Cantemir
Causeni
Chisinau
Cimislia
Criuleni
Donduseni
Drochia
Dubasari
Edinet
Falesti
Floresti
Gagauzia
Glodeni
Hncesti
Ialoveni
Leova
Nisporeni
Ocnita
Orhei
Rezina
Rscani
Sngerei
Soldanesti
Soroca
Stefan Voda
Straseni
Taraclia
Telenesti
Transnistria
Ungheni
Chaouia-Ouardigha
Mekns-Talalet
Tadla-Azilal
Tanger-Ttouan
Drenthe
Friesland
Friesland
Friesland
Gelderland
Gelderland
Gelderland
Gelderland
Vilniaus
Ben Slimane
Knitra
Ifrane
Khmisset
Azilal
Chefchaouen
Westerveld
Ameland
Opsterland
Terschelling
Brummen
Buren
Epe
Nijkerk
Reference ID
732
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
13
2
7
13
3
3
2
2
Continued
187

Country
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Netherlands
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Gelderland
Noord-Brabant
Noord-Brabant
Noord-Brabant
Noord-Holland
Noord-Holland
Noord-Holland
Overijssel
Overijssel
Overijssel
Utrecht
Utrecht
Utrecht
Utrecht
Utrecht
Utrecht
Utrecht
Utrecht
Zeeland
Zeeland
Zeeland
Zuid-Holland
Zuid-Holland
Zuid-Holland
Akershus
Akershus
stfold
stfold
stfold
stfold
stfold
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Aust-Agder
Buskerud
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Nunspeet
Halderberge
Moerdijk
Oisterwijk
Amsterdam
Heemskerk
Hilversum
Deventer
Raalte
Zwolle
Bunnik
De Bilt
Doorn
Leersum
Maarn
Soest
Utrecht
Zeist
Middelburg
Schouwen-Duiveland
Sluis
Pijnacker-Nootdorp
Waddinxveen
Warmond
Aurskog-Hland
Srum
Aremark
Halden
Marker
Sarpsborg
Spydeberg
Arendal
Birkenes
Birkenes
Froland
Grimstad
Lillesand
Lillesand
Tvedestrand
Vegrshei
Flesberg
Bergen
Eidfjord
Etne
Kvinnherad
Kvinnherad
Linds
Masfjorden
Masfjorden
Modalen
Odda
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
733
0
0
732
0
0
0
732
0
0
0
0
0
0
0
732
0
0
0
0
0
No. entriesa
3
4
2
4
2
Continued
188
Appendix 1

Country
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Norway
Hordaland
Hordaland
Hordaland
Hordaland
Hordaland
Mre og Romsdal
Mre og Romsdal
Mre og Romsdal
Mre og Romsdal
Mre og Romsdal
Mre og Romsdal
Mre og Romsdal
Mre og Romsdal
Rogaland
Rogaland
Rogaland
Rogaland
Rogaland
Rogaland
Rogaland
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sogn og Fjordane
Sr-Trndelag
Telemark
Telemark
Telemark
Telemark
Telemark
Telemark
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vest-Agder
Vestfold
Vestfold
Vestfold
Vestfold
Samnanger
Ullensvang
Vaksdal
Vaksdal
Voss
Norddal
rsta
Rauma
Sunndal
Surnadal
Ulstein
Volda
Bjerkreim
Gjesdal
Hjelmeland
Lund
Sandnes
Sauda
Suldal
Askvoll
Aurland
Frde
Gaular
Gloppen
Gulen
Gulen
Hyllestad
Hyllestad
Stryn
Hitra
Drangedal
Drangedal
Krager
Krager
Skien
Audnedal
Flekkefjord
Hgebostad
Kristiansand
Kristiansand
Kvinesdal
Marnardal
Songdalen
Vennesla
Larvik
Vaale
Reference ID
0
0
0
0
0
1116
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
732
0
732
0
1323
1116
0
732
0
732
0
1116
733
0
0
0
0
732
0
0
0
0
1116
810
0
0
No. entriesa
4
8
2
2
3
3
2
8
3
2
8
2
2
2
7
2
Continued
189

Country
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Greater Poland
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Kuyavian-Pomeranian
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Chodziez
Czarnkw-Trzcianka
Gniezno
Gostyn
Gostyn
Grodzisk Wielkopolski
Kalisz
Kalisz City
Kepno
Koscian
Krotoszyn
Leszno
Miedzychd
Nowy Tomysl
Oborniki
Ostrw Wielkopolski
Ostrzeszw
Pleszew
Poznan
Poznan
Poznan
Poznan City
Rawicz
Supca
Srem
Sroda Wielkopolska
Turek
Wagrowiec
Wolsztyn
Zlotw
Bydgoszcz City
Chelmno
Grudziadz
Grudziadz City
Inowrocaw
Lipno
Nako
Rypin
Swiecie
Torun City
Tuchola
Bochnia
Brzesko Brzeg
Chrzanw
Gorlice
Krakw
Krakw City
Limanowa
Miechw
Myslenice
Reference ID
603
645
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
644
645
0
0
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0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
2
2
4
3
2
2
2
2
2
3
13
3
3
4
3
3
2
2
3
2
4
4
12
3
2
5
2
Continued
190
Appendix 1

Country
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Lesser Poland
Ldz
Ldz
Ldz
Ldz
Ldz
Ldz
Ldz
Ldz
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lower Silesian
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Lublin
Nowy Sacz
Nowy Targ
Olkusz
Oswiecim
Proszowice
Sucha
Tarnw
Tarnw City
Tatra
Wieliczka
Kutno
Lowicz
Ldz
Pabianice
Sieradz
Tomaszw Mazowiecki
Wieruszw
Zgierz
Bolesawiec
Dzierzoniw
Jelenia Gra
Kodzko
Legnica
Lubin
Lwwek Slaski
Milicz
Olesnica
Sroda Slaska
Strzelin
Swidnica
Trzebnica
Wow
Wrocaw
Wrocaw
Wrocaw
Zabkowice
Zgorzelec
Bilgoraj
Chelm
Hrubieszw
Janw
Krasnik
Lubartw
Lublin
Lublin City
Puawy
Radzyn
Tomaszw
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
513
0
0
102
104
105
953
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
3
6
5
4
3
2
4
2
3
3
4
4
8
2
4
2
6
4
3
3
2
4
2
2
4
4
2
2
3
3
Continued
191

Country
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Lublin
Lublin
Lubusz
Lubusz
Lubusz
Lubusz
Lubusz
Lubusz
Lubusz
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Masovian
Opole
Opole
Opole
Opole
Opole
Opole
Opole
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Podlachian
Pomeranian
Pomeranian
Wodawa
Zamosc
Gorzw
Miedzyrzecz
Slubice
Strzelce-Drezdenko
Sulecin
wiebodzin
S
Zielona Gra
Ciechanw
Garwolin
Legionowo
Losice
Minsk
Mawa
Nowy Dwr Mazowiecki
Ostroeka
Otwock
Piaseczno
Pock
Przysucha
Radom
Warsaw
Warsaw
Warsaw West
Wegrw
Wyszkw
Zuromin
Glubczyce
Kedzierzyn-Kozle
Namysw
Nysa
Opole
Prudnik
Strzelce
Augustw
Biaystok
Bielsk
Grajewo
Hajnwka
Hajnwka
Lomza
Monki
Siemiatycze
Sokka
Suwaki
Zambrw
Bytw
Reference ID
No. entriesa
0
0
0
0
0
0
0
0
0
964
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1099
0
0
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0
0
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0
0
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0
0
1340
464
0
0
0
0
0
385
0
0
0
0
0
0
843
0
5
6
3
3
2
2
3
2
3
4
3
4
2
3
9
4
3
2
8
2
5
2
2
4
Continued
192
Appendix 1

Country
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Pomeranian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Silesian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Subcarpathian
Swietokrzyskie
Swietokrzyskie
Swietokrzyskie
Swietokrzyskie
Swietokrzyskie
Chojnice
Czluchw
Gdansk
Gdansk City
Gdynia
Kartuzy
Koscierzyna
Kwidzyn
Lebork
Malbork
Nowy Dwr Gdanski
Puck
Supsk
Supsk City
Sopot
Starogard
Sztum
Tczew
Wejherowo
Bytom
Chorzw
Cieszyn
Czestochowa
Gliwice
Katowice City
Kobuck
Lubliniec
Mikolw
Myslowice
Myszkw
Rybnik
Siemianowice Slaskie
Zabrze
Zawiercie
Zywiec
Bieszczady
Jaso
Kolbuszowa
Krosno
Lesko
Lezajsk
Lubaczw
Nisko
Przemysl
Przeworsk
Rzeszw City
Sanok
Busko
Kielce
Kielce City
Sandomierz
Skarzysko
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
5
3
6
3
5
9
5
4
4
2
6
7
5
3
2
2
9
4
3
2
2
3
7
2
3
3
2
2
3
2
2
Continued
193

Country
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Swietokrzyskie
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
Warmian-Masurian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
West Pomeranian
Braga
vora
vora
Faro
Faro
Lisboa
Lisboa
Staszw
Bartoszyce
Braniewo
Dziadowo
Elblag
Ek
Gizycko
Iawa
Ketrzyn
Lidzbark
Mragowo
Nidzica
Olecko
Olsztyn
Olsztyn
Olsztyn City
Ostrda
Pisz
Szczytno
Wegorzewo
Biaogard
Choszczno
Drawsko
Goleniw
Gryce
Gryno
Kamien
Koobrzeg
Koszalin
Koszalin City
Lobez
Myslibrz
Police
Pyrzyce
Sawno
Stargard
Swidwin
Szczecin
Szczecin
Szczecin
Szczecin
Szczecinek
Wacz
Montemor-o-Novo
Viana do Alentejo
Monchique
Vila do Bispo
Mafra
Reference ID
0
738
739
0
0
0
0
0
0
0
0
0
0
0
0
0
546
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
138
893
894
0
0
1143
0
0
0
0
1143
0
No. entriesa
2
3
2
3
3
3
18
4
5
2
2
2
2
3
3
3
8
3
2
2
2
8
5
2
8
3
3
2
2
2
2
2
Continued
194
Appendix 1

Country
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Russia
Russia
Russia
Serbia
Madeira
Portalegre
Portalegre
Santarm
Setbal
Setbal
Setbal
Setbal
Setbal
Setbal
Viana do Castelo
Alba
Arad
Arges
Bihor
Botosani
Braila
Buzau
Calarasi
Caras-Severin
Cluj
Constanta
Covasna
Dmbovita
Dolj
Galati
Giurgiu
Gorj
Hunedoara
Iasi
Mehedinti
Mures
Satu Mare
Suceava
Suceava
Suceava
Suceava
Teleorman
Teleorman
Timis
Timis
Tulcea
Tulcea
Tulcea
Tulcea
Tulcea
Vlcea
Vrancea
City of St Petersburg
Kaliningrad
Moskva
Grad Beograd
Avis
Alpiara
Alccer do Sal
Barreiro
Grndola
Montijo
Palmela
Arcos de Valdevez
Reference ID
1143
1143
0
0
1143
0
0
0
0
0
0
769
769
769
769
769
769
769
769
769
769
769
0
460
769
769
769
769
0
0
769
769
769
0
460
462
769
460
462
0
177
0
198
460
603
769
0
769
35
35
522
654
No. entriesa
2
2
8
2
6
2
10
2
4
4
2
3
Continued
195

Country
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Grad Beograd
Grad Beograd
Grad Beograd
Grad Beograd
Grad Beograd
Moravicki
Moravicki
Niavski
Niavski
Pirotski
Raki
Raki
umadijski
umadijski
umadijski
Zajecarski
Zajecarski
Zajecarski
Zapadno-Backi
Zlatiborski
Zlatiborski
Zlatiborski
Banskobystrick
Banskobystrick
Banskobystrick
Banskobystrick
Banskobystrick
Bratislavsk
Bratislavsk
Bratislavsk
Bratislavsk
Koick
Koick
Koick
Koick
Koick
Koick
Koick
Nitriansk
Nitriansk
Nitriansk
Nitriansk
Nitriansk
Nitriansk
Preovsk
Preovsk
Trenciansk
Trenciansk
Trenciansk
Trenciansk
Trenciansk
Trenciansk
Grocka
Mladenovac
Rakovica
Stari Grad
Vodovac
Cacak
Gornji Milanovac
Aleksinac
Svrljig
Bela Palanka
Kraljevo
Novi Pazar
Arandelovac
Kragujevac
Topola
Boljevac
Knjaevac
Zajecar
Sombor
Bajina Bata
Priboj
Prijepolje
arnovica
iar nad Hronom
Bratislava II
Malacky
Senec
Koice I
Koice III
Koice IV
Koice-okolie
Levice
Nitra
Nov Zmky
Topolcany
Zlat Moravce
Kemarok
Star Lubovna
Ilava
Myjava
Nov Mesto nad Vhom
Partiznske
Reference ID
0
0
785
785
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
140
449
0
0
0
0
0
1349
140
602
914
0
0
0
0
1349
0
0
0
0
0
0
0
140
603
0
0
0
0
No. entriesa
2
2
2
6
3
2
2
10
14
16
5
5
9
2
3
5
2
2
4
4
4
Continued
196
Appendix 1

Country
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Slovenia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Trenciansk
Trenciansk
Trenciansk
Trenciansk
Trnavsk
Trnavsk
Trnavsk
Trnavsk
Trnavsk
Trnavsk
ilinsk
ilinsk
ilinsk
ilinsk
ilinsk
ilinsk
Gorenjska
Gorenjska
Gorenjska
Gorenjska
Gorenjska
Gorenjska
Gorika
Gorika
Jugovzhodna Slovenija
Jugovzhodna Slovenija
Notranjsko-kraka
Notranjsko-kraka
Obalno-kraka
Obalno-kraka
Obalno-kraka
Obalno-kraka
Osrednjeslovenska
Osrednjeslovenska
Osrednjeslovenska
Osrednjeslovenska
Osrednjeslovenska
Osrednjeslovenska
Savinjska
Savinjska
Zasavska
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Aragn
Aragn
Cantabria
Castilla y Len
Povask Bystrica
Prievidza
Pchov
Trencn
Dunajsk Streda
Pietany
Senica
Skalica
Trnava
Bytca
Cadca
Martin
Nmestovo
Tvrdon
Bled
Cerklje na Gorenjskem
Cerklje na Gorenjskem
elezniki
Gorenja Vas-Poljane
Preddvor
Brda
Cerkno
Kocevje
Sodrazica
Ilirska Bistrica
Pivka
Divaca
Hrpelje-Kozina
Koper
Grosuplje
Kamnik
Ljubljana
Medvode
Medvode
Velike Lac
Mozirje
Tabor
Zagorje ob Savi
Cdiz
Crdoba
Granada
Huelva
Jan
Jan
Mlaga
Huesca
Zaragoza
Cantabria
Burgos
Reference ID
0
0
0
0
140
0
0
0
0
0
140
0
0
0
0
0
0
0
527
0
0
0
0
0
0
527
0
0
527
0
0
0
0
527
0
0
527
0
527
0
0
0
0
0
0
0
611
0
0
0
0
0
No. entriesa
4
3
3
3
3
3
8
2
2
2
2
2
3
2
2
8
4
12
12
8
3
2
6
2
9
9
Continued
197

Country
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Sweden
Castilla y Len
Castilla y Len
Castilla-La Mancha
Castilla-La Mancha
Catalua
Catalua
Comunidad Foral de Navarra
Extremadura
Extremadura
Galicia
Islas Baleares
La Rioja
La Rioja
Pas Vasco
Pas Vasco
Pas Vasco
Pas Vasco
Principado de Asturias
Regin de Murcia
Blekinge
Blekinge
Dalarna
Gvleborg
Gvleborg
Gotland
Gotland
Halland
Halland
Halland
Jmtland
Jnkping
Kalmar
Kalmar
Kronoberg
Norrbotten
Orebro
stergtland
Skne
Skne
Skne
Skne
Sdermanland
Stockholm
Stockholm
Stockholm
Stockholm
Uppsala
Uppsala
Uppsala
Uppsala
Vrmland
Vsterbotten
Len
Toledo
Ciudad Real
Cuenca
Barcelona
Girona
Navarra
Badajoz
Cceres
Lugo
Baleares
La Rioja
La Rioja
lava
Guipzcoa
Vizcaya
Murcia
Gotland
Kungsbacka
Varberg
Kalmar
Bstad
Simrishamn
Norrtlje
Nynshamn
Stockholm
lvkarleby
Tierp
Uppsala
Reference ID
No. entriesa
0
821
0
821
0
0
0
0
0
0
1345
0
315
77
0
0
0
821
616
1339
350
1339
1339
350
1339
470
1339
470
470
1339
1339
1339
350
1339
1339
1339
1339
1339
350
470
584
1339
1339
470
470
350
1339
470
470
470
1339
1339
3
2
2
2
4
4
14
51
2
21
17
38
9
41
2
2
3
2
2
2
2
2
2
2
2
2
2
2
2
Continued
198
Appendix 1

Country
Sweden
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Switzerland
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
UK
UK
Vsterbotten
Vsterbotten
Vsternorrland
Vstmanland
Vstra Gtaland
Aargau
Basel-Landschaft
Bern
Fribourg
Graubnden
Jura
Lucerne
Neuchtel
Neuchtel
Neuchtel
Sankt Gallen
Schaffhausen
Ticino
Ticino
Valais
Valais
Valais
Vaud
Zrich
Bja
Bja
Bja
Bja
Bizerte
Jendouba
Jendouba
Jendouba
Jendouba
Kairouan
Le Kef
Siliana
Siliana
Sousse
Tataouine
Zaghouan
Artvin
Burdur
Burdur
Giresun
Istanbul
Istanbul
Rize
Samsun
Sinop
Trabzon
England
England
Ume
Ume
Bja Nord
Mejez El Bab
Nefza
Testour
Sejnane
An Draham
Balta Bou Aouane
Ghardimaou
Tabarka
Chebika
Nebeur
Bouarada
Kesra
Kondar
Ghomrassen
Zriba
Bedfordshire
Berkshire
Reference ID
1322
470
1339
1339
1339
0
524
0
0
435
0
0
0
450
487
0
0
0
435
0
1284
1321
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1008
0
0
145
0
0
0
0
0
0
No. entriesa
2
2
7
7
5
2
2
2
3
6
8
2
4
19
2
6
2
7
10
3
3
3
2
3
2
5
3
4
7
12
5
3
4
Continued
199

Country
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
England
Buckinghamshire
Cambridgeshire
Cheshire
Cornwall
Croydon
Cumbria
Derbyshire
Devon
Dorset
Durham
East Sussex
Essex
Gloucestershire
Hampshire
Herefordshire
Hounslow
Isle of Wight
Kensington and Chelsea
Kent
Kingston upon Thames
Lambeth
Lancashire
Lancashire
Leicester
Leicestershire
Lincolnshire
Luton
Merseyside
Merton
Milton Keynes
Norfolk
North Somerset
North Yorkshire
North Yorkshire
Northamptonshire
Northumberland
Northumberland
Nottinghamshire
Oxfordshire
Peterborough
Plymouth
Poole
Portsmouth
Richmond upon Thames
Somerset
Staffordshire
Suffolk
Surrey
Sutton
Thurrock
Torbay
Tyne and Wear
Reference ID
No. entriesa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
559
0
0
0
0
0
0
0
0
0
0
1297
0
0
131
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
15
17
17
13
41
2
12
3
28
7
2
7
4
2
2
53
5
3
20
2
3
2
2
4
4
17
8
Continued
200
Appendix 1

Country
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
England
England
England
England
England
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Northern Ireland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Scotland
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Wales
Waltham Forest
Warwickshire
West Sussex
Wiltshire
Worcestershire
Down
Dungannon
Fermanagh
Newry and Mourne
Omagh
(*) If more than one.
Aberdeen
Aberdeenshire
Angus
Argyll and Bute
Dumfries and Galloway
East Ayrshire
Eilean Siar
Fife
Highland
Moray
North Ayshire
Perthshire and Kinross
Renfrewshire
Scottish Borders
Shetland Islands
South Ayrshire
South Lanarkshire
Stirling
West Dunbartonshire
Anglesey
Bridgend
Cardiff
Carmarthenshire
Ceredigion
Conwy
Denbighshire
Gwynedd
Monmouthshire
Neath Port Talbot
Pembrokeshire
Powys
Swansea
Torfaen
Vale of Glamorgan
Reference ID
0
0
0
0
0
0
0
0
0
0
1114
302
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
631
0
0
No. entriesa
2
12
8
19
8
40
17
2
5
104
5
6
17
11
3
4
2
8
2
6
14
19
5
30
2
2
9
16
201
Table A1.15. Geographic distribution data for Haemaphysalis punctata. See Appendix 2 (hosted on
Country
Albania
Albania
Albania
Albania
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Bulgaria
Bulgaria
Croatia
Croatia
Croatia
Croatia
Croatia
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Elbasan
Gjirokastr
Kor
Vlor
Alger
Bouira
Bouira
Bouira
Mda
Relizane
Stif
Tiaret
Tizi Ouzou
Tizi Ouzou
Tlemcen
Montana
Vratsa
Karlovacka
Zadarska
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Auvergne
Auvergne
Basse-Normandie
Bourgogne
Bourgogne
Centre
Centre
Centre
Centre
Corse
Corse
Corse
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
Limousin
Elbasanit
Gjirokastrs
Pogradecit
Vlors
Bouzareah
Ain el Hadjar
Ain Turk
Lakhdaria
Medea
Oued Rhiou
Ain Arnat
Sougueur
Boghni
Tizi-Ghenif
Tlemcen
Montana
Vratsa
Dordogne
Dordogne
Gironde
Gironde
Landes
Landes
Pyrnes-Atlantiques
Pyrnes-Atlantiques
Puy-de-Dme
Puy-de-Dme
Manche
Cte-dOr
Cte-dOr
Eure-et-Loir
Indre
Indre
Indre-et-Loire
Corse-du-Sud
Haute-Corse
Seine-et-Marne
Seine-et-Marne
Seine-Maritime
Val-de-Marne
Ville de Paris
Aude
Hrault
Pyrnes-Orientales
Pyrnes-Orientales
Haute-Vienne

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
277
781
0
0
1280
0
1280
0
1280
0
1280
0
0
1280
0
1280
0
0
1280
1280
1280
0
0
0
1280
1280
0
1280
0
0
0
1280
0
2
2
2
2
2
2
2
7
5
2
10
3
Continued
202
Appendix 1

Country
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Pays de la Loire
Poitou-Charentes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Attiki
Dytiki Makedonia
Dytiki Makedonia
Ipeiros
Ipeiros
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Stere Ellda
Stere Ellda
Abruzzo
Abruzzo
Abruzzo
Apulia
Apulia
Campania
Campania
Campania
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Lazio
Lazio
Liguria
Aveyron
Aveyron
Lot
Lot
Tarn
Vende
Vienne
Alpes-de-Haute-Provence
Alpes-de-Haute-Provence
Alpes-Maritimes
Bouches-du-Rhne
Hautes-Alpes
Var
Ain
Ain
Ardche
Drme
Isre
Savoie
Drama
Evros
Kavala
Attica
Grevena
Kozani
Ioannina
Preveza
Khalkidiki
Kilkis
Pella
Pieria
Serrai
Thessaloniki
Boeotia
Evritania
Chieti
LAquila
Teramo
Foggia
Lecce
Avellino
Caserta
Ferrara
Forl-Cesena
Modena
Parma
Piacenza
Latina
Rieti
Roma
Viterbo
Genova

0
1280
0
1280
0
0
1280
0
1280
0
0
0
0
0
1280
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
802
0
0
0
0
0
0
0
0
0
0
0
0
5
4
7
9
7
3
3
2
8
14
3
6
6
6
2
10
5
2
2
2
3
3
8
3
2
26
5
5
3
4
2
2
2
3
3
10
3
2
Continued
203

Country
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Kosovo
Kosovo
Kosovo
Libya
Libya
Libya
Libya
Macedonia
Macedonia
Macedonia
Macedonia
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Montenegro
Morocco
Morocco
Morocco
Morocco
Morocco
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Lombardia
Marche
Molise
Piemonte
Sardegna
Sardegna
Sardegna
Sardegna
Sardegna
Sardegna
Sicily
Sicily
Sicily
Sicily
Sicily
Toscana
Toscana
Toscana
Toscana
Trentino-Alto Adige
Trentino-Alto Adige
Umbria
Umbria
Veneto
Kosovska Mitrovica
Pristina
Uroevac
Ajdabiya
Al Jabal al Akhdar
An Nuqat al Khams
Tarhunah-Masallatah
Pelagonia
Polog
Skopje
Southwestern
Cahul
Cantemir
Causeni
Cimislia
Leova
Taraclia
Kolacin
Grand Casablanca
Mekns-Talalet
Mekns-Talalet
Mekns-Talalet
vora
Lisboa
Santarm
Setbal
Setbal
Setbal
Varese
Ancona
Isernia
Alessandria
Cagliari
Carbonia-Iglesias
Nuoro
Olbia-Tempio
Oristano
Sassari
Palermo
Palermo
Palermo
Palermo
Trapani
Florence
Pisa
Pistoia
Bolzano
Trento
Perugia
Terni
Verona
Kosovska Mitrovica
Pritina
Uroevac
Bitola
Tetovo
Centar
Ohrid
Knitra
Mohammedia
Ifrane
Khnifra
Mekns
Montemor-o-Novo
Coruche
Alccer do Sal
Grndola

0
0
0
0
0
0
0
0
0
0
0
242
366
998
0
1331
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
667
667
667
667
667
667
0
0
0
0
0
0
0
1143
0
1143
0
0
3
4
11
3
2
3
2
2
2
2
2
Continued
204
Appendix 1

Country
Romania
Serbia
Serbia
Serbia
Slovakia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Sweden
Sweden
Sweden
Sweden
Switzerland
Switzerland
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Turkey
Timis
Macvanski
Pcinjski
Toplicki
Koick
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Aragn
Aragn
Aragn
Cantabria
Castilla y Len
Castilla y Len
Castilla y Len
Catalua
Catalua
Catalua
Extremadura
Extremadura
Islas Baleares
La Rioja
Pas Vasco
Pas Vasco
Pas Vasco
Pas Vasco
Blekinge
Gotland
Kalmar
Vstra Gtaland
Ticino
Ticino
Ariana
Bja
Bja
Bja
Bizerte
Bizerte
Bizerte
Jendouba
Jendouba
Le Kef
Nabeul
Nabeul
Zaghouan
Aksaray
Ankara
Ankara
Elzig
If more than one.
Ljubovija
Surdulica
Prokuplje
Cdiz
Crdoba
Granada
Huelva
Jan
Jan
Huesca
Teruel
Zaragoza
Cantabria
Burgos
Burgos
Soria
Girona
Lleida
Tarragona
Navarra
Badajoz
Cceres
Baleares
La Rioja
lava
Guipzcoa
Vizcaya
Kalaat El Andalous
Bja Nord
Nefza
Tboursouk
Bizerte Sud
Mateur
Sejnane
An Draham
Ghardimaou
Nebeur
Haouaria
Menzel Temime
Zriba

177
0
0
0
140
0
0
0
0
0
611
0
0
0
0
0
821
0
0
0
0
0
0
0
1345
0
77
0
0
0
821
1339
1339
1339
1339
435
435
0
0
0
0
0
0
0
0
0
0
0
0
0
1353
0
1353
0
3
2
7
3
2
2
2
3
3
6
3
2
4
25
14
15
2
2
2
3
2
8
205
Table A1.16. Geographic distribution data for Haemaphysalis concinna. See Appendix 2 (hosted on
Country
Czech Republic
France
France
France
France
France
France
France
France
France
France
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Romania
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Spain
Jihomoravsk
Aquitaine
Aquitaine
Centre
Centre
le-de-France
le-de-France
Limousin
Midi-Pyrnes
Picardie
Poitou-Charentes
Bacs-Kiskun
Baranya
Csongrad
Fejer
Nograd
Somogy
Zala
Timis
Banskobystrick
Bratislavsk
Bratislavsk
Nitriansk
Trenciansk
Trnavsk
Pas Vasco
Landes
Pyrnes-Atlantiques
Indre
Indre-et-Loire
Seine-et-Marne
Seine-Maritime
Haute-Vienne
Arige
Oise
Vienne
Reference ID
490
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1417
1417
1417
1417
1417
1417
1417
177
140
1349
1349
1349
1349
900
77
821
Table A1.17. Geographic distribution data for Haemaphysalis inermis. See Appendix 2 (hosted on http://
Country
Bulgaria
Bulgaria
France
France
France
France
France
France
Hungary
Hungary
Hungary
Italy
Portugal
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Smolyan
Yuzhen tsentralen
Centre
Centre
le-de-France
Limousin
Poitou-Charentes
Borsod-Abauj-Zemplen
Fejer
Heves
Campania
Lisboa
Banskobystrick
Bratislavsk
Koick
Trenciansk
Zilinsk
Pas Vasco
Reference ID
884
Indre
Indre-et-Loire
Seine-et-Marne
Pyrnes-Orientales
Haute-Vienne
Vienne
1280
1280
1280
1280
1280
1280
1417
1417
1417
802
1143
140
1349
1349
1349
1349
77
206
Appendix 1
Table A1.18. Geographic distribution data for Hyalomma marginatum. See Appendix 2 (hosted on http://
Country
Administrative
region 1
Administrative
region 2
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Elbasan
Elbasan
Kor
Kor
Lezh
Vlor
Vlor
An Dea
An Dea
An Dea
An Tmouchent
Alger
Alger
Annaba
Batna
Batna
Batna
Batna
Batna
Batna
Beja
Biskra
Biskra
Biskra
Biskra
Blida
Blida
Blida
Bouira
Bouira
Bouira
Bouira
Bouira
Bouira
Boumerds
Boumerds
Boumerds
Chlef
Chlef
Chlef
Chlef
Chlef
Chlef
Constantine
Constantine
Djelfa
El Tarf
Guelma
Guelma
Jijel
Jijel
Elbasanit
Librazhdit
Kors
Pogradecit
Lezhs
Sarands
Vlors
Ain Benian
Ain Lechiakh
Arib
Ain Kihel
Ain Benian
Ouled Chebel
Annaba
Ain Touta
Batna
Chemora
Ksar Bellezma
Ras El Aioun
Seriana
Bejaia
Besbes
El Ouitaya
Ouled Djellal
Tolga
Bouaarfa
Bouinan
Soumaa
Ain El Hadjar
Ain Turk
Djebahia
El Asnam
Lakhdaria
Sour El Ghouzlane
Boudouaou
Corso
Isser
Bouzeghaia
Chlef
Sendjas
Sidi Akkacha
Taougrit
Tenes
Constantine
El Khroub
MLiliha
El Aioun
Ain Rekada
Hammam Debagh
Chekfa
Jijel
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
2
3
Continued
207
Country
Administrative
region 1
Administrative
region 2
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Jijel
Jijel
Laghouat
MSila
Mascara
Mda
Mda
Mda
Mila
Oran
Oran
Oum el Bouaghi
Relizane
Relizane
Jijel
Jijel
Laghouat
Ain Errich
Ain Ferah
Berrouaghia
Boughezoul
Medea
Chelghoum Laid
Oran
Oued Tlelat
Meskiana
Relizane
Sidi MHamed
Benaouda
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Bosnia and
Herzegovina
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Stif
Stif
Stif
Stif
Stif
Stif
Stif
Stif
Stif
Sidi Bel Abbs
Skikda
Tbessa
Tbessa
Tbessa
Tiaret
Tipaza
Tissemsilt
Tissemsilt
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Federacija Bosna
i Hercegovina
Blagoevgrad
Burgas
Burgas
Burgas
Haskovo
Kardzhali
Kardzhali
Kardzhali
Kyustendil
Lovech
Montana
Ain Arnat
Ain Lahdjar
El Eulma
Guellal
Mezloug
Ouled Saber
Setif
Dhaya
Ouled Attia
Bir El Ater
Boulhaf Dyr
Tebessa
Hadjout
Lardjem
Theniet El Had
Boghni
Draa-Ben-Khedda
Tadmait
Tizi Ouzou
Tizi-Ghenif
Zekri
Tuzla
Bansko
Karnobat
Malko Tarnovo
Tsarevo
Svilengrad
Ardino
Kardzhali
Momchilgrad
Rila
Troyan
Montana
Reference ID
No. entriesa
110
110
0
0
0
0
0
0
0
0
0
0
0
0
110
110
0
0
0
0
0
0
0
0
0
0
0
0
129
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
2
Continued
208
Appendix 1
Country
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Cyprus
Cyprus
Cyprus
France
France
France
France
France
France
France
France
France
France
France
France
Germany
Greece
Greece
Administrative
region 1
Administrative
region 2
Pazardzhik
Pernik
Pleven
Plovdiv
Plovdiv
Shumen
Sliven
Soa
Soa
Stara Zagora
Stara Zagora
Stara Zagora
Varna
Varna
Varna
Vidin
Vratsa
Yambol
Yambol
Grad Zagreb
Istarska
Karlovacka
Primorsko-Goranska
ibensko-Kninska
Zadarska
Famagusta
Nicosia
Paphos
Aquitaine
Aquitaine
Centre
Corse
Corse
Corse
Provence-Alpes-CtedAzur
Baden-Wrttemberg
Anatoliki Makedonia kai
Thraki
Thraki
Pazardzhik
Breznik
Pleven
Plovdiv
Saedinenie
Preslav
Sliven
Samokov
Zlatitsa
Chirpan
Haskovo
Radnevo
Dolni Chiik
Provadiya
Varna
Dimovo
Vratsa
Tundzha
Yambol
Pyrnes-Atlantiques
Pyrnes-Atlantiques
Indre-et-Loire
Corse-du-Sud
Haute-Corse
Lozre
Pyrnes-Orientales
Alpes-Maritimes
Reference ID
0
0
0
0
0
0
0
0
0
0
69
0
0
0
0
0
0
0
0
0
0
0
0
0
0
781
0
0
0
0
0
1280
0
1280
0
0
0
1280
0
Bouches-du-Rhne
Var
Var
1280
Freiburg
1302
735
No. entriesa
3
3
2
2
3
3
6
2
2
2
14
7
735
Continued
209
Country
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Administrative
region 1
Administrative
region 2

Thraki
Thraki
Thraki
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Ipeiros
Ipeiros
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kriti
Peloponnisos
Peloponnisos
Stere Ellda
Stere Ellda
Stere Ellda
Thessalia
Abruzzo
Abruzzo
Apulia
Apulia
Apulia
Apulia
Apulia
Apulia
Basilicata
Calabria
Calabria
Campania
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Lazio
Lazio
Liguria
Liguria
Lombardia
Drama
Evros
Kavala
Grevena
Kozani
Ioannina
Preveza
Khalkidiki
Khalkidiki
Khalkidiki
Khalkidiki
Kilkis
Pella
Serrai
Thessaloniki
Heraklion
Arcadia
Messinia
Boeotia
Euboea
Fthiotis
Larisa
LAquila
Teramo
Bari
Barletta-Andria-Trani
Brindisi
Foggia
Lecce
Taranto
Matera
Cosenza
Reggio di Calabria
Napoli
Forl-Cesena
Parma
Ravenna
Rimini
Udine
Latina
Rieti
Roma
Viterbo
Genova
Savona
Brescia
Reference ID
735
735
0
0
0
0
735
0
1303
1303
1303
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
4
2
3
2
11
2
2
19
2
2
7
3
3
4
3
8
14
5
2
Continued
210
Appendix 1
Country
Administrative
region 1
Administrative
region 2
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Kosovo
Kosovo
Kosovo
Kosovo
Kosovo
Kosovo
Kosovo
Kosovo
Libya
Macedonia
Macedonia
Macedonia
Macedonia
Macedonia
Macedonia
Macedonia
Morocco
Morocco
Morocco
Morocco
Morocco
Marche
Molise
Piemonte
Piemonte
Piemonte
Sardegna
Sardegna
Sardegna
Sardegna
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Toscana
Toscana
Toscana
Toscana
Trentino-Alto Adige
Umbria
Umbria
Veneto
Veneto
D
akovica
Gnjilane
Gnjilane
Kosovska Mitrovica
Kosovska Mitrovica
Kosovska Mitrovica
Pristina
Pristina
Al Marqab
Eastern
Pelagonia
Polog
Skopje
Southeastern
Southwestern
Vardar
Chaouia-Ouardigha
Chaouia-Ouardigha
Doukkala-Abda
Doukkala-Abda
Fs-Boulemane
Macerata
Campobasso
Alessandria
Torino
Verbano-Cusio-Ossola
Cagliari
Carbonia-Iglesias
Ogliastra
Oristano
Agrigento
Enna
Messina
Palermo
Palermo
Palermo
Palermo
Palermo
Palermo
Palermo
Ragusa
Trapani
Florence
Grosseto
Pisa
Trento
Perugia
Terni
Venezia
Verona
Decani
Gnjilane
Vitina
Kosovska Mitrovica
Kosovska Mitrovica
Kosovska Mitrovica
Lipljan
Pritina
Pehcevo
Bitola
Tetovo
Centar
Lake Dojran
Ohrid
Kavadartsi
Ben Slimane
Settat
El Jadida
Sa
Zouagha-Moulay
Yacoub
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
242
366
366
366
366
998
366
0
1331
0
0
0
0
0
0
0
0
0
0
0
0
346
346
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
6
2
2
2
2
2
2
2
2
2
5
2
2
3
3
3
Continued
211
Country
Administrative
region 1
Administrative
region 2
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Romania
Russia
Russia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Spain
Spain
Grand Casablanca
Mekns-Talalet
Souss-Massa-Dra
Tanger-Ttouan
Aveiro
Coimbra
vora
vora
Guarda
Leiria
Lisboa
Portalegre
Porto
Santarm
Santarm
Setbal
Setbal
Setbal
Setbal
Viana do Castelo
Alba
Bucharest
Constanta
Dolj
Gorj
Mehedinti
Mures
Timis
Tulcea
Tulcea
Tulcea
Stavropol
Stavropol
Borski
Macvanski
Pcinjski
Pirotski
Raki
Raki
Toplicki
Andaluca
Andaluca
Knitra
Mohammedia
Al Haouz
El Kela des Sraghna
Marrakech
Marrakech
Mekns
Khmisset
Skhirate-Tmara
Taroudannt
Chefchaouen
Al Hocema
Taza
Oliveira de Azemis
Coimbra
Montemor-o-Novo
Portel
Guarda
Peniche
Alenquer
Marvo
Marco de Canaveses
Benavente
Coruche
Alccer do Sal
Grndola
Palmela
Setbal
Paredes de Coura
Kladovo
Ljubovija
Surdulica
Pirot
Novi Pazar
Vrnjacka Banja
Kurumlija
Cdiz
Cdiz
Reference ID
No. entriesa
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
769
769
769
0
0
460
769
1144
1144
0
0
0
0
0
0
0
0
241
3
3
6
5
2
7
4
3
2
2
4
15
8
3
2
2
6
Continued
212
Appendix 1
Country
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Sweden
Sweden
Sweden
Switzerland
Switzerland
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Administrative
region 1
Administrative
region 2
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Aragn
Castilla y Len
Castilla y Len
Castilla y Len
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Comunidad Foral de
Navarra
Extremadura
Extremadura
Gotland
Kalmar
Skne
Ticino
Ticino
Ariana
Ariana
Ariana
Bja
Bja
Bja
Bja
Ben Arous (Tunis Sud)
Bizerte
Bizerte
Bizerte
Bizerte
Gabs
Gabs
Gafsa
Jendouba
Jendouba
Jendouba
Kairouan
Kairouan
Kebili
Le Kef
Mahdia
Manubah
Manubah
Manubah
Mdenine
Monastir
Cdiz
Cdiz
Crdoba
Granada
Jan
Sevilla
Zaragoza
Salamanca
Toledo
Toledo
Albacete
Ciudad Real
Cuenca
Toledo
Toledo
Navarra
Badajoz
Cceres
Ettadhamen
Kalaat El Andalous
Sidi Thabet
Amdoun
Bja Nord
Bja Sud
Nefza
Ezzahra
Bizerte Nord
Bizerte Sud
Ghar El Melh
Ghazala
Hamma
Matmata
Ksar
An Draham
Fernana
Tabarka
Kairouan Sud
Sbikha
Kebili Nord
Sakiet Sidi Youssef
Chebba
El Battan
Oued Ellil
Tebourba
Houmt Souk
Jammel
Reference ID
241
821
0
0
0
0
0
0
821
821
821
246
821
0
989
0
0
0
1339
1339
1339
1284
1284
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
3
3
5
8
8
14
50
3
2
Continued
213
Country
Administrative
region 1
Administrative
region 2
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Nabeul
Nabeul
Nabeul
Nabeul
Sidi Bou Zid
Tunis
Tunis
Tunis
Zaghouan
Zaghouan
Aksaray
Ankara
Ankara
Ankara
Ankara
Artvin
Balkesir
Burdur
orum
Elzig
Erzurum
Giresun
Gmshane
I zmir
Kayseri
Krklareli
Ordu
Samsun
Sivas
Tokat
Yozgat
Haouaria
Korba
Soliman
Takelsa
Meknassi
Cit El Khadra
La Goulette
Sidi El Bchir
Fahs
Zriba
Reference ID
0
0
0
0
0
0
0
0
0
0
1353
0
1353
1353
844
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
2
3
2
2
25
6
3
2
3
2
12
45
6
If more than one.
Table A1.19. Geographic distribution data for Rhipicephalus sanguineus group (Rh. sanguineus and
Rh. turanicus). See Appendix 2 (hosted on http://www.cabi.org/vetmedresource/) for complete reference
indicated by the Reference ID.
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Algeria
Alger
Algeria
El Tarf
El Tarf
Bulgaria
Stara Zagora
Haskovo
Egypt
Al Jizah
Egypt
Al Wadi al
Jadid
Reference ID
No. entriesa
110
110
69
1101
595
Continued
214
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
France
Aquitaine
Dordogne
1280
France
Aquitaine
Gironde
1280
France
Aquitaine
Landes
1280
France
Aquitaine
Lot-et-Garonne
1280
France
Auvergne
Allier
1280
France
Bourgogne
Cte-dOr
1280
France
Corse
France
Corse
Corse-du-Sud
1306
France
le-de-France
1280
France
le-de-France
Seine-SaintDenis
Ville de Paris
France
Alpes-Maritimes
1280
Bouches-duRhne
Var
1280
France
Rhne-Alpes
Isre
1280
Greece
Dytiki Makedonia
735
Greece
735
Greece
Kentriki
Makedonia
Kentriki
Makedonia
Kentriki
Makedonia
Stere Ellda
Hungary
Ngrd
445
Hungary
Ngrd
829
Italy
Sicily
Palermo
242
Italy
Sicily
Palermo
998
Italy
Toscana
Kosovo
Kosovska
Mitrovica
Kosovska
Mitrovica
France
France
Greece
Greece
Kosovo
Reference ID
No. entriesa
1280
4
1280
1280
735
Khalkidiki
Fokis
1303
778
1331
Kosovska
Mitrovica
Kosovska
Mitrovica
346
346
Continued
215
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Romania
Timis
Spain
Andaluca
Jan
611
Spain
Andaluca
Sevilla
614
Spain
Toledo
989
Spain
Castilla-La
Mancha
Islas Baleares
1345
Spain
Pas Vasco
Spain
Sweden
Regin de
Murcia
stergtland
Sweden
Sdermanland
1339
Sweden
Stockholm
1339
Sweden
Uppsala
1339
Sweden
Vsternorrland
1339
Sweden
Vstmanland
1339
Turkey
Afyon
186
Turkey
Elzig
22
Turkey
Erzurum
32
Turkey
Istanbul
145
Turkey
Sanlurfa
303
Albania
Durrs
Durrsit
Albania
Elbasan
Librazhdit
Albania
Gjirokastr
Prmetit
Albania
Kor
Kors
Albania
Lezh
Lezhs
Reference ID
No. entriesa
177
Baleares
77
Murcia
616
1339
Continued
216
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Albania
Shkodr
Shkodrs
Albania
Tiran
Tirans
Albania
Vlor
Sarands
Albania
Vlor
Vlors
Algeria
An Tmouchent Oued Sebbah
Algeria
Alger
Ain Benian
Algeria
Alger
Bouzareah
Algeria
Alger
Ouled Chebel
Algeria
Batna
Ain Touta
Algeria
Batna
Chemora
Algeria
Batna
Ras El Aioun
Algeria
Bchar
Beni Abbes
Algeria
Blida
Blida
Algeria
Blida
Boufarik
Algeria
Bouira
Ain Turk
Algeria
Bouira
Guerrouma
Algeria
Bouira
Lakhdaria
Reference ID
No. entriesa
Continued
217
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Algeria
Bouira
Sour El
Ghouzlane
Algeria
Chlef
Chlef
Algeria
El Bayadh
Boualem
Algeria
El Bayadh
Bougtoub
Algeria
El Bayadh
El Bayadh
Algeria
El Bayadh
Stitten
Algeria
Guelma
Hammam
Debagh
Algeria
Jijel
Chekfa
Algeria
Laghouat
Hassi RMel
Algeria
Laghouat
Laghouat
Algeria
MSila
Sidi MHamed
Algeria
Mascara
Mascara
Algeria
Mostaganem
Mostaganem
Algeria
Oran
Oran
Algeria
Relizane
Sidi MHamed
Benaouda
Algeria
Stif
Ain Arnat
Algeria
Stif
Setif
Reference ID
No. entriesa
Continued
218
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Algeria
Sidi Bel Abbs
Dhaya
Algeria
Souk Ahras
Ouled Moumen
Algeria
Tiaret
Meghila
Algeria
Tindouf
Tindouf
Algeria
Tissemsilt
Theniet El Had
Algeria
Tizi Ouzou
Ain El Hammam
Bulgaria
Burgas
Burgas
Bulgaria
Burgas
Malko Tarnovo
Bulgaria
Burgas
Primorsko
Bulgaria
Kardzhali
Ardino
Bulgaria
Kardzhali
Momchilgrad
Bulgaria
Shumen
Shumen
Bulgaria
Smolyan
Dospat
Bulgaria
Soa
Ihtiman
Bulgaria
Soa
Zlatitsa
Croatia
DubrovackoNeretvanska
Croatia
Istarska
Reference ID
No. entriesa
Continued
219

Administrative
region 1
Administrative
region 2
Reference ID
No. entriesa
SplitskoDalmatinska
Croatia
Zadarska
Cyprus
Famagusta
Cyprus
Larnaca
Cyprus
Limassol
Cyprus
Nicosia
Cyprus
Paphos
Egypt
Al Bahr al Ahmar
Egypt
Al Iskandariyah
Egypt
Al Jizah
Egypt
Al Minuyah
Egypt
Al Minya
Egypt
Aswan
Egypt
Bani Suwayf
Egypt
Janub Sina
France
Alsace
Bas-Rhin
France
Aquitaine
Dordogne
Species
Country
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Croatia
Continued
220
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
France
Aquitaine
France
Reference ID
No. entriesa
Gironde
Aquitaine
Lot-et-Garonne
France
Aquitaine
PyrnesAtlantiques
France
Auvergne
Allier
France
Auvergne
Cantal
France
Auvergne
Puy-de-Dme
France
BasseNormandie
Calvados
France
BasseNormandie
Orne
France
Centre
Indre
France
Centre
Indre-et-Loire
France
ChampagneArdenne
Ardennes
France
ChampagneArdenne
Marne
France
Corse
Corse-du-Sud
France
Corse
Haute-Corse
France
Franche-Comt Doubs
France
le-de-France
Hauts-de-Seine
France
le-de-France
Seine-et-Marne
Continued
221
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
France
le-de-France
Seine-SaintDenis
France
le-de-France
Val-de-Marne
France
le-de-France
Ville de Paris
France
le-de-France
Yvelines
France
LanguedocRoussillon
Aude
France
LanguedocRoussillon
Gard
France
LanguedocRoussillon
Hrault
France
LanguedocRoussillon
PyrnesOrientales
10
France
Lorraine
Meurthe-etMoselle
France
Midi-Pyrnes
Arige
France
Midi-Pyrnes
Aveyron
France
Midi-Pyrnes
Haute-Garonne
France
Midi-Pyrnes
Lot
France
Midi-Pyrnes
Tarn
France
Midi-Pyrnes
Tarn-et-Garonne
France
Pays de la Loire Vende
France
Picardie
Somme
Reference ID
No. entriesa
Continued
222
Appendix 1

Administrative
region 1
Administrative
region 2
Reference ID
No. entriesa
Provence-Alpes- Alpes-de-HauteCte-dAzur
Provence
France
Provence-Alpes- Alpes-Maritimes
Cte-dAzur
11
France
Provence-Alpes- Bouches-duCte-dAzur
Rhne
35
France
Provence-Alpes- Var
Cte-dAzur
26
France
Provence-Alpes- Vaucluse
Cte-dAzur
24
France
Rhne-Alpes
Ain
France
Rhne-Alpes
Ardche
11
France
Rhne-Alpes
Drme
France
Rhne-Alpes
Isre
France
Rhne-Alpes
Loire
France
Rhne-Alpes
Rhne
Greece
Anatoliki
Makedonia
kai Thraki
Anatoliki
Makedonia
kai Thraki
Anatoliki
Makedonia
kai Thraki
Attiki
Drama
Evros
Kavala
Attica
Greece
Dytiki
Makedonia
Grevena
Greece
Dytiki
Makedonia
Kastoria
Species
Country
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
France
Greece
Greece
Greece
Continued
223
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Greece
Ionioi Nisoi
Corfu
Greece
Ionioi Nisoi
Kefallinia
Greece
Ipeiros
Ioannina
Greece
Kentriki
Makedonia
Imathia
Greece
Kentriki
Makedonia
Khalkidiki
10
Greece
Kentriki
Makedonia
Kilkis
Greece
Kentriki
Makedonia
Pella
Greece
Kentriki
Makedonia
Pieria
Greece
Kentriki
Makedonia
Serrai
Greece
Kentriki
Makedonia
Thessaloniki
13
Greece
Notio Aigaio
Cyclades
Greece
Peloponnisos
Arcadia
Greece
Peloponnisos
Messinia
Greece
Stere Ellda
Boeotia
Greece
Thessalia
Larisa
Greece
Thessalia
Magnesia
Greece
Voreio Aigaio
Lesvos
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Reference ID
No. entriesa
Continued
224
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Italy
Abruzzo
LAquila
Italy
Abruzzo
Teramo
Italy
Apulia
Bari
29
Italy
Apulia
Barletta-AndriaTrani
Italy
Apulia
Brindisi
20
Italy
Apulia
Foggia
49
Italy
Apulia
Lecce
11
Italy
Apulia
Taranto
Italy
Basilicata
Matera
18
Italy
Basilicata
Potenza
Italy
Calabria
Cosenza
12
Italy
Calabria
Reggio di
Calabria
10
Italy
Campania
Italy
Campania
Avellino
Italy
Campania
Caserta
Italy
EmiliaRomagna
Forl-Cesena
Rhipicephalus
sanguineus
group
Italy
EmiliaRomagna
Modena
Reference ID
No. entriesa
802
Continued
225

Administrative
region 1
Administrative
region 2
Italy
EmiliaRomagna
Parma
Italy
EmiliaRomagna
Ravenna
Italy
EmiliaRomagna
Reggio
nellEmilia
Italy
Friuli-Venezia
Giulia
Udine
Italy
Lazio
Latina
Italy
Lazio
Rieti
Italy
Lazio
Roma
14
Italy
Lazio
Viterbo
Italy
Liguria
Genova
Italy
Liguria
Imperia
Italy
Marche
Ascoli Piceno
Italy
Molise
Campobasso
15
Italy
Piemonte
Cuneo
Italy
Sardegna
Cagliari
Italy
Sardegna
Carbonia-Iglesias
Italy
Sardegna
Nuoro
Italy
Sardegna
Ogliastra
Species
Country
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Reference ID
No. entriesa
Continued
226
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Reference ID
No. entriesa
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Italy
Sardegna
Olbia-Tempio
Italy
Sardegna
Oristano
Italy
Sardegna
Sassari
Italy
Sicily
Agrigento
Italy
Sicily
Catania
Italy
Sicily
Enna
Italy
Sicily
Palermo
17
Italy
Sicily
Trapani
Italy
Toscana
Florence
Italy
Toscana
Grosseto
Italy
Toscana
Livorno
Italy
Toscana
Pisa
Italy
Umbria
Perugia
Italy
Umbria
Terni
Italy
Veneto
Belluno
Italy
Veneto
Verona
Libya
Ajdabiya
Continued
227
Species
Country
Administrative
region 1
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Libya
Administrative
region 2
Reference ID
No. entriesa
Al Butnan
Libya
Al Hizam al
Akhdar
13
Libya
Al Jabal al
Akhdar
Libya
Al Jabal al
Akhdar
Libya
Al Kufrah
Libya
Al Marj
Libya
Al Marqab
Libya
Al Qubbah
Libya
Al Wahah
Libya
An Nuqat al
Khams
Libya
Ghadamis
Libya
Gharyan
Libya
Misratah
Libya
Nalut
Libya
Surt
Libya
Tajura wa an
Nawahi al Arba
Libya
Tarabulus
Continued
228
Appendix 1

Administrative
region 1
Administrative
region 2
Country
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Libya
TarhunahMasallatah
Libya
Yafran-Jadu
Monaco
Monaco
Morocco
ChaouiaOuardigha
Ben Slimane
Morocco
ChaouiaOuardigha
Settat
Morocco
Fs-Boulemane Zouagha-Moulay
Yacoub
Morocco
Gharb-Chrarda- Knitra
Bni Hssen
Morocco
Gharb-Chrarda- Sidi Kacem

Bni Hssen
Morocco
Grand
Casablanca
Mohammedia
Morocco
Guelmim-EsSemara
Assa-Zag
Morocco
Guelmim-EsSemara
Guelmim
Morocco
MarrakechTensift-Al
Haouz
MarrakechTensift-Al
Haouz
MarrakechTensift-Al
Haouz
MarrakechTensift-Al
Haouz
MeknsTalalet
Al Haouz
El Kela Des
Sraghna
Essaouira
Marrakech
Ifrane
MeknsTalalet
Khnifra
Morocco
Morocco
Morocco
Morocco
Morocco
Reference ID
No. entriesa
Species
Continued
229

Administrative
region 1
Administrative
region 2
Morocco
MeknsTalalet
Morocco
Oriental
Morocco
Rabat-SalKhmisset
Zemmour-Zaer
Morocco
Rabat-SalSkhirate-Tmara
Zemmour-Zaer
Morocco
Souss-MassaDra
Agadir-Ida Ou
Tanane
Morocco
Souss-MassaDra
Chtouka-At
Baha
Morocco
Souss-MassaDra
Ouarzazate
Morocco
Souss-MassaDra
Taroudannt
Morocco
Souss-MassaDra
Zagora
Morocco
Tadla-Azilal
Azilal
Morocco
Tadla-Azilal
Bni Mellal
Morocco
Tanger-Ttouan Chefchaouen
Morocco
Tanger-Ttouan Larache
Morocco
Tanger-Ttouan Tanger-Assilah
Morocco
Taza-Al Hocei- Taounate

ma-Taounate
Portugal
Beja
Beja
Portugal
Beja
Ferreira do
Alentejo
Species
Country
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Reference ID
No. entriesa
Mekns
Oujda Angad
Continued
230
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Portugal
Beja
Mrtola
Portugal
Beja
Odemira
Portugal
Braga
Cabeceiras de
Basto
Portugal
Bragana
Portugal
Castelo
Branco
Idanha-a-Nova
Portugal
vora
Montemor-oNovo
Portugal
vora
Portel
Portugal
Faro
Lagos
Portugal
Faro
Loul
Portugal
Faro
Monchique
Portugal
Faro
Vila do Bispo
Portugal
Faro
Vila Real de
Santo Antnio
Portugal
Guarda
Guarda
Portugal
Leiria
Portugal
Leiria
Ansio
Portugal
Leiria
Ansio
Portugal
Leiria
Pombal
Reference ID
No. entriesa
7
1143
1143
Continued
231
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Portugal
Lisboa
Sintra
Portugal
Lisboa
Sobral de Monte
Agrao
Portugal
Lisboa
Torres Vedras
Portugal
Portalegre
Portugal
Portalegre
Marvo
Portugal
Portalegre
Ponte de Sr
Portugal
Porto
Matosinhos
Portugal
Santarm
Portugal
Santarm
Benavente
Portugal
Santarm
Coruche
Portugal
Santarm
Santarm
Portugal
Setbal
Portugal
Setbal
Alccer do Sal
12
Portugal
Setbal
Almada
Portugal
Setbal
Barreiro
Portugal
Setbal
Grndola
Portugal
Setbal
Moita
Reference ID
No. entriesa
1143
1143
1143
Continued
232
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Portugal
Setbal
Portugal
Reference ID
No. entriesa
Palmela
18
Setbal
Seixal
Portugal
Setbal
Setbal
Portugal
Setbal
Sines
Portugal
Viana do
Castelo
Melgao
Portugal
Viana do
Castelo
Ponte da Barca
Romania
Constanta
Romania
Dolj
Romania
Timis
Slovenia
Obalno-kraka
Slovenia
Osrednjesloven- Ljubljana
ska
Spain
Andaluca
Almera
Spain
Andaluca
Cdiz
Spain
Andaluca
Granada
18
Spain
Andaluca
Huelva
Spain
Andaluca
Jan
Spain
Andaluca
Mlaga
Piran
Continued
233
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Spain
Andaluca
Spain
Reference ID
No. entriesa
Sevilla
Aragn
Zaragoza
Spain
Castilla y Len
Soria
Spain
Castilla y Len
Valladolid
Spain
Castilla y Len
Zamora
Spain
Castilla-La
Mancha
Guadalajara
Spain
Catalua
Barcelona
Spain
Catalua
Girona
12
Spain
Catalua
Lleida
Spain
Catalua
Lleida
Spain
Catalua
Tarragona
Spain
Extremadura
Badajoz
Spain
Extremadura
Cceres
46
Spain
Pas Vasco
lava
Spain
Pas Vasco
Vizcaya
Tunisia
Ariana
Sidi Thabet
Tunisia
Bja
Amdoun
Continued
234
Appendix 1
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Tunisia
Bja
Bja Nord
Tunisia
Bja
Mejez El Bab
Tunisia
Ben Arous
(Tunis Sud)
Fouchana
Tunisia
Bizerte
Bizerte Sud
Tunisia
Bizerte
Ghar El Melh
Tunisia
Bizerte
Sejnane
Tunisia
Gabs
Gabs Mdina
Tunisia
Gabs
Gabs Ouest
Tunisia
Gabs
Hamma
Tunisia
Gabs
Matmata
Tunisia
Gafsa
Ksar
Tunisia
Gafsa
Metlaoui
Tunisia
Jendouba
An Draham
Tunisia
Jendouba
Tabarka
Tunisia
Kairouan
Bouhajla
Tunisia
Kairouan
Kairouan Sud
Tunisia
Kairouan
Sbikha
Reference ID
No. entriesa
Continued
235
Species
Country
Administrative
region 1
Administrative
region 2
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Tunisia
Kebili
Faouar
Tunisia
Kebili
Kebili Nord
Tunisia
Le Kef
Kef Ouest
Tunisia
Mdenine
Houmt Souk
Tunisia
Nabeul
Menzel Temime
Tunisia
Nabeul
Soliman
Tunisia
Nabeul
Takelsa
Tunisia
Sfax
Sfax Sud
Tunisia
Sidi Bou Zid
Meknassi
Tunisia
Siliana
Siliana Nord
Tunisia
Tataouine
Tataouine Nord
Tunisia
Tozeur
Tamaghza
Tunisia
Tozeur
Tozeur
Tunisia
Tunis
Carthage
Tunisia
Tunis
La Goulette
Tunisia
Tunis
Sidi El Bchir
Turkey
Ankara
21
Reference ID
No. entriesa
Continued
236
Appendix 1
Species
Country
Administrative
region 1
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
sanguineus
group
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Turkey
Burdur
14
Turkey
orum
Turkey
Edirne
Turkey
Elzig
14
Turkey
Erzincan
Turkey
Kayseri
Turkey
Krklareli
Turkey
Samsun
Turkey
Sivas
Turkey
Tokat
Turkey
Van
Turkey
Yozgat
Algeria
Tizi Ouzou
Tizi Ouzou
Bulgaria
Stara Zagora
Haskovo
Croatia
SplitskoDalmatinska
Al Ismailiyah
Rhipicephalus
turanicus
Egypt
France
Greece
Greece
Administrative
region 2
Provence-Alpes- Bouches-duCte-dAzur
Rhne
Anatoliki
Makedonia
kai Thraki
Kentriki
Makedonia
Reference ID
No. entriesa
110
69
277
595
1280
735
735
Continued
237

Administrative
region 1
Administrative
region 2
Khalkidiki
Greece
Kentriki
Makedonia
Stere Ellda
Israel
Jerusalem
Italy
Sicily
Palermo
242
Spain
Andaluca
Jan
611
Spain
Andaluca
Jan
611
Spain
Islas Baleares
Baleares
Turkey
Afyon
Turkey
Aksaray
1353
Turkey
Ankara
1353
Turkey
Sanlurfa
Species
Country
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Rhipicephalus
turanicus
Greece
Reference ID
No. entriesa
1303
Fokis
778
1071
1345
186
303
If more than one.
Table A1.20. Geographic distribution data for Rhipicephalus bursa. See Appendix 2 (hosted on http://
Country
Administrative
region 1
Administrative
region 2
Reference ID
Albania
Albania
Albania
Albania
Albania
Albania
Albania
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Durrs
Elbasan
Kor
Lezh
Tiran
Vlor
Vlor
An Dea
An Dea
An Dea
An Dea
An Dea
An Tmouchent
An Tmouchent
Alger
Alger
Alger
Alger
Alger
Batna
Biskra
Durrsit
Librazhdit
Kors
Lezhs
Tirans
Sarands
Vlors
Ain Benian
Ain Lechiakh
Arib
Ben Allal
Rouina
Ain Kihel
Terga
Ain Benian
Ain Benian
Bordj El Kiffan
Bouzareah
Ouled Chebel
Ain Touta
El Ouitaya
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
2
3
2
4
Continued
238
Appendix 1
Country
Administrative
region 1
Administrative
region 2
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Blida
Bouira
Bouira
Bouira
Boumerds
Boumerds
Boumerds
Boumerds
Boumerds
Chlef
Chlef
Chlef
Chlef
Chlef
Chlef
Chlef
Constantine
Djelfa
El Tarf
Guelma
Jijel
Jijel
MSila
MSila
Mascara
Mda
Mila
Oran
Oran
Relizane
Relizane
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Algeria
Stif
Stif
Sidi Bel Abbs
Sidi Bel Abbs
Tbessa
Tbessa
Tiaret
Tiaret
Tiaret
Tipaza
Tissemsilt
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tizi Ouzou
Tlemcen
Boufarik
Ain Turk
Lakhdaria
Sour El Ghouzlane
Boudouaou
Corso
Dellys
Isser
Sidi Daoud
Abou El Hassen
Boukadir
Bouzeghaia
Chlef
Ouled Fares
Sidi Akkacha
Tenes
Constantine
Benhar
El Aioun
Hammam Debagh
Jijel
Taher
Ain Errich
Sidi MHamed
Ain Ferah
Medea
Mila
Misserghin
Oran
Relizane
Sidi MHamed
Benaouda
Ain Arnat
Setif
Dhaya
Sidi Bel Abbes
Bir El Ater
Tebessa
Meghila
Sougueur
Douaouda
Theniet El Had
Ain-El-Hammam
Boghni
Draa-Ben-Khedda
Larbaa-Nath-Irathen
Tadmait
Tizi Ouzou
Tizi-Rached
Zekri
Tlemcen
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
129
0
0
0
0
110
0
0
0
0
0
0
0
0
0
No. entriesa
2
Continued
239
Country
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Croatia
Cyprus
Egypt
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Administrative
region 1
Administrative
region 2
Blagoevgrad
Petrich
Blagoevgrad
Sandanski
Burgas
Burgas
Burgas
Kameno
Burgas
Malko Tarnovo
Kardzhali
Ardino
Montana
Montana
Pazardzhik
Septemvri
Plovdiv
Asenovgrad
Shumen
Preslav
Shumen
Shumen
Stara Zagora
Haskovo
Stara Zagora
Pavel Banya
Veliko Tarnovo
Veliko Tarnovo
Vratsa
Vratsa
Yambol
Tundzha
Yambol
Yambol
BjelovarskaBilogorska
DubrovackoNeretvanska
Grad Zagreb
Istarska
Primorsko-Goranska
Varadinska
Zadarska
Famagusta
Al Wadi al Jadid
Aquitaine
Dordogne
Aquitaine
Landes
Aquitaine
PyrnesAtlantiques
Aquitaine
PyrnesAtlantiques
Bourgogne
Sane-et-Loire
Corse
Corse
Corse-du-Sud
Corse
Haute-Corse
LanguedocAude
Roussillon
LanguedocPyrnes-Orientales
Roussillon
Midi-Pyrnes
Aveyron
Midi-Pyrnes
Lot
Provence-AlpesAlpes-de-HauteCte-dAzur
Provence
Provence-AlpesBouches-du-Rhne
Cte-dAzur
Reference ID
0
0
0
0
0
0
0
0
0
0
0
69
0
0
0
0
0
0
No. entriesa
2
2
2
2
2
2
2
5
0
0
0
0
0
277
781
0
0
0
0
0
0
0
2
2
5
6
2
2
2
5
1280
0
1280
0
0
0
12
9
3
0
0
0
2
3
4
Continued
240
Appendix 1
Country
France
France
France
France
France
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Israel
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Administrative
region 1
Administrative
region 2
Rhne-Alpes
Rhne-Alpes
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Attiki
Dytiki Ellada
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Dytiki Makedonia
Ipeiros
Ipeiros
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kriti
Stere Ellda
Stere Ellda
Stere Ellda
Thessalia
Thessalia
Jerusalem
Abruzzo
Abruzzo
Abruzzo
Apulia
Apulia
Apulia
Apulia
Apulia
Apulia
Bouches-du-Rhne
Reference ID
1280
Var
Vaucluse
Ardche
Drme
No. entriesa
0
0
735
2
4
2
Drama
10
Evros
Kavala
10
0
0
735
735
0
0
0
0
0
0
735
0
1303
1303
1303
0
0
0
0
0
0
0
0
778
0
0
1071
0
0
0
0
0
0
0
0
0
8
2
Attica
Achaea
Florina
Grevena
Kastoria
Kozani
Ioannina
Preveza
Khalkidiki
Khalkidiki
Khalkidiki
Khalkidiki
Kilkis
Pella
Pieria
Serrai
Thessaloniki
Rethymnon
Boeotia
Euboea
Fokis
Karditsa
Larisa
Chieti
LAquila
Teramo
Bari
Barletta-Andria-Trani
Brindisi
Foggia
Lecce
Taranto
10
2
8
3
13
8
4
5
14
12
7
7
7
7
2
8
2
17
3
26
2
7
Continued
241
Country
Administrative
region 1
Administrative
region 2
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Kosovo
Kosovo
Kosovo
Kosovo
Lebanon
Libya
Macedonia
Macedonia
Macedonia
Macedonia
Macedonia
Macedonia
Macedonia
Montenegro
Morocco
Morocco
Basilicata
Calabria
Calabria
Campania
Campania
Campania
Emilia-Romagna
Lazio
Lazio
Lazio
Liguria
Liguria
Marche
Molise
Molise
Sardegna
Sardegna
Sardegna
Sardegna
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Toscana
Toscana
Toscana
Umbria
Veneto
Veneto
Dakovica
Kosovska Mitrovica
Pristina
Uroevac
Mount Lebanon
Al Jabal al Akhdar
Pelagonia
Polog
Skopje
Skopje
Southeastern
Southwestern
Vardar
Roaje
Doukkala-Abda
Fs-Boulemane
Matera
Cosenza
Reggio di Calabria
Morocco
Gharb-ChrardaBni Hssen
Avellino
Caserta
Ravenna
Latina
Rieti
Roma
Genova
Savona
Ascoli Piceno
Campobasso
Isernia
Cagliari
Carbonia-Iglesias
Oristano
Sassari
Agrigento
Enna
Palermo
Palermo
Palermo
Palermo
Palermo
Palermo
Palermo
Trapani
Grosseto
Livorno
Pisa
Perugia
Treviso
Verona
Deani
Kosovska Mitrovica
Obilic
Uroevac
Matn
Bitola
Tetovo
Centar
Zelenikovo
Lake Dojran
Ohrid
Kavadartsi
Sa
Zouagha-Moulay
Yacoub
Knitra
Reference ID
No. entriesa
0
0
0
802
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
242
366
366
366
366
998
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
12
3
4
3
4
14
7
2
4
2
10
3
2
2
3
3
2
4
2
2
2
2
6
Continued
242
Appendix 1

Administrative
region 1
Administrative
region 2
Al Haouz
El Kela des
Sraghna
Essaouira
Marrakech
Mekns
Bni Mellal
Chefchaouen
Fahs Anjra
Tanger-Assilah
Ttouan
Taounate
0
0
0
0
0
0
0
Taza
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Marrakech-TensiftAl Haouz
Mekns-Talalet
Tadla-Azilal
Tanger-Ttouan
Tanger-Ttouan
Tanger-Ttouan
Tanger-Ttouan
Braga
Castelo Branco
vora
vora
Portalegre
Portalegre
Porto
Portugal
Portugal
Portugal
Portugal
Romania
Romania
Romania
Romania
Romania
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Serbia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Santarm
Setbal
Setbal
Viana do Castelo
Constanta
Dmbovita
Dolj
Timis
Tulcea
Borski
Juno-Backi
Moravicki
Niavski
Pcinjski
Pirotski
Raki
Raki
Raki
Rasinski
Zlatiborski
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Country
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Castelo Branco
Montemor-o-Novo
Portel
Marvo
Marco de
Canaveses
Coruche
Alccer do Sal
Palmela
Arcos de Valdevez
Kladovo
Bac
Cacak
Doljevac
Surdulica
Pirot
Kraljevo
Novi Pazar
Vrnjacka Banja
Brus
Uice
Cdiz
Cdiz
Crdoba
Granada
Huelva
Jan
Jan
Reference ID
1143
0
0
0
1143
0
0
0
0
0
0
0
460
0
0
769
0
0
0
0
0
0
0
0
0
0
0
0
821
0
0
0
0
611
No. entriesa
2
2
2
2
5
4
4
2
2
3
21
14
9
12
Continued
243
Country
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Turkey
Administrative
region 1
Administrative
region 2
Andaluca
Jan
Andaluca
Jan
Andaluca
Jan
Andaluca
Sevilla
Aragn
Huesca
Aragn
Teruel
Aragn
Zaragoza
Castilla y Len
Salamanca
Castilla y Len
Toledo
Castilla y Len
Toledo
Castilla y Len
vila
Castilla y Len
Zamora
Castilla-La Mancha Albacete
Castilla-La Mancha Albacete
Castilla-La Mancha Ciudad Real
Castilla-La Mancha Guadalajara
Castilla-La Mancha Toledo
Catalua
Girona
Catalua
Tarragona
Ceuta y Melilla
Ceuta
Navarra
Extremadura
Badajoz
Extremadura
Cceres
La Rioja
La Rioja
Pas Vasco
Pas Vasco
lava
Pas Vasco
Guipzcoa
Pas Vasco
Vizcaya
Principado de
Asturias
Ariana
Kalaat El Andalous
Bja
Bja Sud
Bja
Nefza
Bizerte
Bizerte Sud
Bizerte
Ghazala
Bizerte
Sejnane
Jendouba
An Draham
Jendouba
Ghardimaou
Jendouba
Jendouba Nord
Jendouba
Tabarka
Le Kef
Nebeur
Le Kef
Sakiet Sidi Youssef
Nabeul
Soliman
Nabeul
Takelsa
Sidi Bou Zid
Meknassi
Tunis
Carthage
Tunis
Sidi El Bchir
Zaghouan
Fahs
Afyon
Aksaray
Ankara
Ankara
Reference ID
611
611
611
0
0
0
0
0
821
821
0
0
0
821
0
0
0
0
0
0
0
0
0
0
77
0
0
0
821
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
186
1353
0
1353
No. entriesa
12
2
2
3
3
5
3
17
110
10
8
5
20
Continued
244
Appendix 1
Country
Administrative
region 1
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Ankara
Artvin
Burdur
Burdur
orum
Elazig
Giresun
Gmshane
Istanbul
Kayseri
Mus
Ordu
Samsun
Sivas
Tokat
Van
Yozgat
Administrative
region 2
Reference ID
1353
0
0
1008
0
0
0
0
145
0
22
0
0
0
0
0
0
No. entriesa
16
2
4
18
8
5
If more than one.
Table A1.21. Geographic distribution data for Rhipicephalus (Boophilus) annulatus. See Appendix 2
(hosted on http://www.cabi.org/vetmedresource/) for complete reference indicated by the Reference ID.
Country
Albania
Algeria
Algeria
Algeria
Algeria
Algeria
Bulgaria
Bulgaria
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
Egypt
France
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Administrative
region 1
Administrative
region 2
Tiran
Alger
Guelma
Mascara
Sidi Bel Abbs
Tiaret
Montana
Vratsa
Al Iskandariyah
Al Iskandariyah
Al Ismailiyah
Al Ismailiyah
Al Jizah
Al Jizah
Al Minuyah
Al Minya
Bur Said
Corse
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Tirans
Ain Benian
Hammam Debagh
Mascara
Sidi Bel Abbes
Reference ID
No. entriesa
Drama
0
0
0
0
0
129
0
0
0
595
1039
595
0
1101
0
0
595
1280
0
Kavala
735
0
1303
0
0
0
Montana
Vratsa
Khalkidiki
Khalkidiki
Kilkis
Serrai
Thessaloniki
2
2
2
Continued
245
Country
Greece
Italy
Libya
Libya
Libya
Libya
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Morocco
Romania
Spain
Spain
Spain
Spain
Spain
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
a
If more than one.
Administrative
region 1
Administrative
region 2
Thessalia
Sicily
Al Butnan
Al Hizam Al Akhdar
Al Jifarah
Al Wahah
Chaouia-Ouardigha
Gharb-Chrard-Bni
Hssen
Grand Casablanca
Mekns-Talalet
Tanger-Ttouan
Tanger-Ttouan
Tulcea
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Bizerte
Jendouba
Jendouba
Le Kef
Nabeul
Tozeur
Tunis
Zaghouan
Ankara
Artvin
Burdur
orum
Edirne
Elzig
Elzig
Erzurum
Erzurum
Kayseri
Krklareli
Mus
Samsun
Sivas
Tokat
Larisa
Palermo
Ben Slimane
Knitra
Reference ID
0
366
0
0
0
0
0
0
Mohammedia
El Kela des
Sraghna
Marrakech
0
0
Mekns
Khmisset
0
0
Larache
Tanger-Assilah
Taounate
0
0
0
Taza
Cdiz
Cdiz
Cdiz
Huelva
Sevilla
Sejnane
Fernana
Tabarka
Nebeur
Takelsa
Tozeur
Sidi El Bchir
Zriba
No. entriesa
2
2
2
3
769
0
241
821
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
22
0
22
0
0
22
0
0
0
2
2
3
3
2
4
95
7
246
Appendix 1
Table A1.22. Geographic distribution data for Dermacentor marginatus. See Appendix 2 (hosted on
Country
Administrative
region 1
Administrative
region 2
Albania
Algeria
Algeria
Algeria
Bulgaria
Bulgaria
Bulgaria
Bulgaria
Croatia
Croatia
Croatia
Croatia
France
France
France
France
France
France
France
France
France
France
France
Vlor
Mostaganem
Nama
Tlemcen
Grad Soya
Montana
Stara Zagora
Vratsa
Primorsko-Goranska
Zadarska
Alsace
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Vlors
Mostaganem
El Biod
Tlemcen
Stolichna
Montana
Haskovo
Vratsa
France
Aquitaine
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Auvergne
Auvergne
Auvergne
Auvergne
Auvergne
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bretagne
Bretagne
Bretagne
Centre
Centre
Centre
Centre
Centre
Centre
Centre
ChampagneArdenne
Bas-Rhin
Dordogne
Dordogne
Gironde
Gironde
Landes
Landes
Lot
Lot-et-Garonne
Lot-et-Garonne
PyrnesAtlantiques
PyrnesAtlantiques
Allier
Allier
Cantal
Puy-de-Dme
Puy-de-Dme
Cte-dOr
Cte-dOr
Nivre
Nivre
Sane-et-Loire
Yonne
Yonne
Ille-et-Vilaine
Ille-et-Vilaine
Morbihan
Cher
Cher
Eure-et-Loir
Indre
Indre-et-Loire
Indre-et-Loire
Loiret
Aube
Reference ID
0
0
0
0
0
0
69
0
0
0
781
0
0
0
1280
0
1280
0
1280
1280
0
1280
0
No. entriesa
2
3
4
11
17
1280
0
1280
0
0
1280
0
1280
0
1280
0
0
1280
0
1280
0
0
1280
0
0
0
1280
0
0
2
3
3
10
2
3
9
2
2
2
2
Continued
247
Country
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Administrative
region 1
Administrative
region 2
ChampagneArdenne
Corse
Corse
Corse
Franche-Comt
le-de-France
le-de-France
le-de-France
le-de-France
Aube
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
le-de-France
LanguedocRoussillon
LanguedocRoussillon
LanguedocRoussillon
LanguedocRoussillon
LanguedocRoussillon
LanguedocRoussillon
Limousin
Limousin
Limousin
Lorraine
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Pays de la Loire
Corse-du-Sud
Haute-Corse
Jura
Essonne
Hauts-de-Seine
Seine-et-Marne
Seine-SaintDenis
Val-dOise
Val-dOise
Val-de-Marne
Val-de-Marne
Ville de Paris
Ville de Paris
Yvelines
Aude
Reference ID
No. entriesa
1280
1280
0
0
0
1280
1280
0
1280
10
6
4
0
1280
0
1280
0
1280
1280
0
Gard
Hrault
Hrault
1280
PyrnesOrientales
PyrnesOrientales
Creuse
Haute-Vienne
Haute-Vienne
Moselle
Arige
Aveyron
Aveyron
Gers
Gers
Haute-Garonne
Haute-Garonne
Hautes-Pyrnes
Lot
Tarn
Tarn
Tarn-et-Garonne
Loire-Atlantique
Loire-Atlantique
Maine-et-Loire
Maine-et-Loire
Mayenne
20
1280
0
0
1280
1280
0
0
1280
0
1280
0
1280
0
0
0
1280
1280
0
1280
0
1280
0
2
12
3
5
3
5
2
Continued
248
Appendix 1
Country
Administrative
region 1
Administrative
region 2
France
France
France
France
France
Pays de la Loire
Picardie
Poitou-Charentes
Poitou-Charentes
Poitou-Charentes
France
Poitou-Charentes
France
France
France
France
Poitou-Charentes
Poitou-Charentes
Poitou-Charentes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Baden-Wrttemberg
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Anatoliki Makedonia
kai Thraki
Attiki
Dytiki Makedonia
Dytiki Makedonia
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Germany
Greece
Greece
Greece
Greece
Greece
Greece
Reference ID
No. entriesa
Vende
Oise
Charente
Charente
CharenteMaritime
CharenteMaritime
Deux-Svres
Vienne
Vienne
Alpes-de-HauteProvence
Alpes-Maritimes
0
0
0
1280
0
Alpes-Maritimes
1280
Bouches-duRhne
Bouches-duRhne
Hautes-Alpes
2
4
1280
0
0
1280
0
26
15
4
2
20
1280
0
14
Var
21
Var
1280
Vaucluse
Ain
Ain
Ardche
Ardche
Drme
Drme
Haute-Savoie
Isre
Isre
Loire
Rhne
Rhne
Savoie
Drama
0
1280
0
1280
0
1280
0
0
1280
0
0
1280
0
948
0
24
18
23
3
34
4
2
3
2
Evros
Kavala
Attica
Florina
Grevena
0
0
0
2
9
Continued
249
Country
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Greece
Hungary
Hungary
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Italy
Macedonia
Macedonia
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Administrative
region 1
Administrative
region 2
Dytiki Makedonia
Dytiki Makedonia
Ipeiros
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Kentriki Makedonia
Peloponnisos
Stere Ellda
Stere Ellda
Stere Ellda
Borsod-AbajZempln
Budapest
Apulia
Campania
Campania
Campania
Emilia-Romagna
Emilia-Romagna
Lazio
Lazio
Lazio
Marche
Molise
Molise
Piemonte
Sardegna
Sicily
Sicily
Sicily
Sicily
Sicily
Sicily
Toscana
Toscana
Umbria
Skopje
Skopje
Anenii Noi
Balti
Basarabeasca
Bender
Briceni
Cahul
Calarasi
Cantemir
Causeni
Chisinau
Cimislia
Kastoria
Kozani
Preveza
Khalkidiki
Kilkis
Pella
Pieria
Serrai
Thessaloniki
Arcadia
Euboea
Evritania
Fthiotis
Foggia
Avellino
Caserta
Forl-Cesena
Ravenna
Latina
Rieti
Roma
Macerata
Campobasso
Isernia
Cuneo
Nuoro
Palermo
Palermo
Palermo
Palermo
Palermo
Palermo
Florence
Perugia
Centar
Zelenikovo
Reference ID
0
0
0
0
0
0
0
0
0
0
0
0
0
443
322
0
802
0
0
0
0
0
0
0
0
0
0
0
0
0
242
366
366
366
998
1331
0
0
0
0
667
667
667
667
667
667
667
667
667
667
667
No. entriesa
3
3
3
4
3
2
2
2
2
2
5
Continued
250
Appendix 1
Country
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Morocco
Morocco
Administrative
region 1
Morocco
Morocco
Portugal
Criuleni
Donduseni
Drochia
Dubasari
Edinet
Falesti
Floresti
Gagauzia
Glodeni
Hncesti
Ialoveni
Leova
Nisporeni
Ocnita
Orhei
Rezina
Rscani
Sngerei
Soldanesti
Soroca
Stefan Voda
Straseni
Taraclia
Telenesti
Transnistria
Ungheni
Mekns-Talalet
Rabat-SalZemmour-Zaer
Rabat-SalZemmour-Zaer
Tanger-Ttouan
Tanger-Ttouan
Aveiro
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Romania
Beja
Bragana
Castelo Branco
vora
Leiria
Lisboa
Portalegre
Santarm
Santarm
Santarm
Santarm
Setbal
Setbal
Setbal
Setbal
Setbal
Suceava
Morocco
Administrative
region 2
Reference ID
No. entriesa
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
667
0
0
2
4
Skhirate-Tmara
Larache
Tanger-Assilah
Oliveira de
Azemis
Mrtola
0
0
0
Ifrane
Khmisset
Castelo Branco
vora
Benavente
Coruche
Santarm
Alccer do Sal
Grndola
Grndola
Palmela
0
1143
0
0
1143
1143
1143
1143
0
0
0
1143
0
0
1143
0
460
10
6
Continued
251
Country
Romania
Romania
Romania
Romania
Romania
Serbia
Serbia
Serbia
Slovakia
Slovakia
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Spain
Switzerland
Switzerland
Switzerland
Tunisia
Tunisia
Tunisia
Tunisia
Tunisia
Administrative
region 1
Suceava
Teleorman
Teleorman
Timis
Tulcea
Macvanski
Niavski
Zlatiborski
Koick
Zilinsk
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Andaluca
Aragn
Aragn
Castilla y Len
Castilla y Len
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Castilla-La Mancha
Catalua
Catalua
Catalua
Comunidad de
Madrid
Comunidad Foral
de Navarra
Extremadura
Extremadura
Islas Baleares
La Rioja
Pas Vasco
Pas Vasco
Pas Vasco
Pas Vasco
Graubnden
Graubnden
Graubnden
Bja
Jendouba
Nabeul
Nabeul
Zaghouan
Administrative
region 2
Ljubovija
Aleksinac
Priboj
Cdiz
Crdoba
Granada
Huelva
Jan
Jan
Jan
Mlaga
Sevilla
Huesca
Zaragoza
Soria
Toledo
Albacete
Albacete
Cuenca
Guadalajara
Toledo
Toledo
Girona
Lleida
Tarragona
Madrid
Navarra
Badajoz
Cceres
Baleares
La Rioja
lava
Guipzcoa
Vizcaya
Nefza
An Draham
Haouaria
Takelsa
Zriba
Reference ID
462
460
462
177
460
0
0
0
1349
1349
0
0
0
0
0
611
611
0
0
0
0
0
821
0
821
821
821
989
989
0
0
0
0
No. entriesa
7
12
11
9
3
4
2
2
4
0
0
1345
0
77
0
0
0
435
435
435
0
0
0
0
0
8
19
3
4
5
3
2
Continued
252
Appendix 1
Country
Administrative
region 1
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Turkey
Aksaray
Ankara
Ankara
Ankara
Bolu
Burdur
Burdur
orum
Elzig
Giresun
Izmir
Kayseri
Konya
Nevsehir
Samsun
Sivas
Tokat
Van
Administrative
region 2
Reference ID
1353
0
1353
1353
0
0
1008
0
0
0
0
0
0
0
0
0
0
0
No. entriesa
20
3
9
6
3
3
3
If more than one.
Table A1.23. Geographic distribution data for Dermacentor reticulatus. See Appendix 2 (hosted on http://
Country
Administrative
region 1
Czech Republic
France
France
France
France
France
France
Jihomoravsky
Alsace
Aquitaine
Aquitaine
Aquitaine
Aquitaine
Aquitaine
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Auvergne
Auvergne
Bourgogne
Bourgogne
Bourgogne
Bourgogne
Bretagne
Bretagne
Bretagne
Centre
Centre
Haute-Normandie
le-de-France
le-de-France
le-de-France
le-de-France
Administrative
region 2
Bas-Rhin
Dordogne
Gironde
Landes
Lot-et-Garonne
PyrnesAtlantiques
Allier
Haute-Loire
Cte-dOr
Nivre
Sane-et-Loire
Yonne
Ctes-dArmor
Finistre
Ille-Et-Vilaine
Indre
Indre-et-Loire
Eure
Essonne
Hauts-de-Seine
Seine-et-Marne
Seine-Saint-Denis
Reference ID
No. entriesa
427
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
Continued
253
Country
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
France
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Germany
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Hungary
Moldova
Administrative
region 1
le-de-France
le-de-France
le-de-France
le-de-France
Limousin
Limousin
Lorraine
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Midi-Pyrnes
Nord-Pas-de-Calais
Pays de la Loire
Pays de la Loire
Pays de la Loire
Poitou-Charentes
Poitou-Charentes
Poitou-Charentes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Rhne-Alpes
Baden-Wrttemberg
Bayern
Bayern
Berlin
Brandenburg
Hessen
Sachsen
Sachsen-Anhalt
Thringen
Bcs-Kiskun
Baranya
Borsod-AbajZempln
Budapest
Csongrd
Fejr
Fejr
Gyor-Moson-Sopron
Gyor-Moson-Sopron
KomromEsztergom
Somogy
Szabolcs-SzatmrBereg
Vas
Veszprm
Anenii Noi
Administrative
region 2
Val-dOise
Val-de-Marne
Ville de Paris
Yvelines
Corrze
Haute-Vienne
Moselle
Arige
Aveyron
Gers
Haute-Garonne
Hautes-Pyrnes
Tarn-et-Garonne
Pas-de-Calais
Loire-Atlantique
Maine-et-Loire
Sarthe
Charente
Charente-Maritime
Vienne
Ain
Drme
Isre
Loire
Rhne
Oberpfalz
Reference ID
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
1280
576
236
1075
236
236
236
236
236
576
341
322
341
322
341
322
341
322
341
341
No. entriesa
3
3
4
3
4
4
2
2
3
2
341
341
341
341
667
Continued
254
Appendix 1
Country
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Moldova
Netherlands
Netherlands
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Poland
Portugal
Romania
Romania
Romania
Romania
Romania
Slovakia
Slovakia
Slovakia
Slovakia
Slovakia
Spain
Spain
If more than one.
Administrative
region 1
Calarasi
Chisinau
Criuleni
Hncesti
Ialoveni
Nisporeni
Orhei
Straseni
Ungheni
Zeeland
Zuid-Holland
Lublin
Lublin
Lublin
Masovian
Podlachian
Podlachian
Podlachian
Warmian-Masurian
Warmian-Masurian
Bragana
Buzau
Caras-Severin
Gorj
Mures
Tulcea
Banskobystrick
Koick
Trenciansk
Trnavsk
ilinsk
Pas Vasco
Principado de
Asturias
Administrative
region 2
Warsaw
Hajnwka
Gizycko
Gizycko
Reference ID
667
667
667
667
667
667
667
667
667
686
686
103
105
1282
1099
1282
464
385
118
120
1143
769
769
769
769
769
140
140
140
140
140
77
821
No. entriesa
12
29
5
5
4
Table A1.24. Data from serological surveillance of Crimean-Congo haemorrhagic fever (CCHF) in domestic animals.
Serology
Year/s
Reference
Albania
Goats
Serological
surveillance
20032005
Central African
Republic
Zebu cattle
Serological
surveillance
19921993
China
Sheep, camels
Serological
surveillance
20042005
China
Sheep
Serological
surveillance
2001
Egypt
Domestic buffaloes,
Serological
cattle, sheep, goats,
surveillance
donkeys, horses,
mules, pigs, dogs
Sheep
Serological
surveillance
1977
Iran
Dairy cattle
Serological
surveillance
20062008
Iran
Livestock
Serological
surveillance
20002007
Iran
Sheep, goats
Serological
surveillance
20032005
Papa, A., Velo, E., Papadimitriou, E., Cahani, G., Kota, M. and Bino, S.
(2009) Ecology of the Crimean-Congo hemorrhagic fever endemic area in
Albania. Vector-Borne and Zoonotic Diseases 9, 713716.
Guilherme, J.M., Gonella-Legall, C., Legall, F., Nakoume, E. and Vincent,
J. (1996) Seroprevalence of ve arboviruses in Zebu cattle in the Central
African Republic. Transactions of the Royal Society of Tropical Medicine
and Hygiene 90, 3133.
Sun, S., Dai, X., Ashian, M., Wang, X., Meng, W., Fend, C., Zhang, F., Hang, C.,
Hu, Z. and Zhang, Y. (2009) Epidemiology and phylogenetic analysis of
the Crimean-Congo hemorrhagic fever viruses in Xinjiang, China. Journal
of Clinical Microbiology 47, 25362543.
Qing, T., Saijo, M., Lei, H., Niikura, M., Maeda, A., Ikegami, T., Xinjung, W.,
Kurane, I. and Morikawa, S. (2003) Detection of immunoglobulin G to
Crimean-Congo hemorrhagic fever virus in sheep sera by recombinant
nucleoprotein-based enzyme linked immunosorbent and immunouorescence assays. Journal of Virological Methods 108, 111116.
Darwish, M.A., Imam, I.Z.E., Omar, F.M. and Hoogstraal, H. (1977) A
seroepidemiological survey for Crimean-Congo hemorrhagic fever virus
in humans and domestic animals in Egypt. Journal of the Egyptian Public
Health Association 52, 156163.
Avic-upanc, T. (2007) Epidemiology of Crimean-Congo hemorrhagic fever
in the Balkans. In: Ergnl, . and Whitehouse, C.A. (eds) Crimean Congo
Haemorrhagic Fever: A Global Perspective. Springer, Dordrecht, pp. 7588.
Lotfollahzadeh, S., Nikbakht Boroujeni, G.R., Mokhber Dezfouli, M.R. and
Bokaei, S. (2009) A serosurvey of Crimean-Congo haemorrhagic fever
virus in dairy cattle in Iran. Zoonoses and Public Health 58, 5459.
Chinikar, S., Goya, M.M., Shirzadi, M.R., Ghiasi, M.S., Mirahmadi, R., Haeri,
A., Moradi, M., Afzali, N., Rahpeyma, M., Zeinali, M. and Meshkat, M. (2008)
Surveillance and laboratory detection system of Crimean-Congo haemorrhagic fever in Iran. Transboundary and Emerging Diseases 55, 200204.
Bokaie, S., Mostafavi, E., Haghdoost, A.A., Keyvanfar, H., Gooya, M.M., Meshkat, M., Davari, A. and Chinikar, S. (2008) Crimean Congo hemorrhagic fever
in northeast of Iran. Journal of Animal and Veterinary Advances 7, 343350.
Continued
Greece
19711976
255
Species
Location
256
Table A1.24. Conitnued.

Species
Serology
Year/s
Reference
Iran
Sheep
Serological
surveillance
2008
Iran
Cattle, sheep, goats
Serological
surveillance
1975
Iraq
Serological
surveillance
1975
Kosovo
Sheep, cattle
1977
Kosovo
Cattle, sheep
Serological
surveillance
Serological
surveillance
Telmadarraiy, Z., Moradi, A.R., Vatandoost, H., Mostafavi, E., Oshaghi, M.A.,
Zahirnia, A.H., Haeri, A. and Chinikar, S. (2008) Crimean-Congo hemorrhagic fever: a seroepidemiological and molecular survey in Bahar, Hamadan Province of Iran. Asian Journal of Animal and Veterinary Advances 3,
321327.
Saidi, S., Casals, J. and Faghih, M.A. (1975) Crimean hemorrhagic feverCongo (CHF-C) virus antibodies in man, and in domestic and small
mammals, in Iran. American Journal of Tropical Medicine and Hygiene
24, 353357.
Tantawi, H.H., Shony, M.O. and Al-Tikriti, S.K. (1981) Antibodies to CrimeanCongo haemorrhagic fever virus in domestic animals in Iraq: a seroepidemiological survey. International Journal of Zoonoses 8, 115120.
Avic-upanc (2007); see Greece above for full reference.
Oman
Cattle, sheep, goat,

camels
Serological
surveillance
19951996
Niger
Cattle, sheep, goats, Serological

camels
surveillance
19841988
Saudi Arabia
Sheep and goats
19931994
Senegal
Birds
Serological
surveillance
Serological
surveillance
19952009
19911992
Humolli, I., Dedushaj, I., Zupanac, T.A. and Muaj, S. (2010) Epidemiological, serological and herd immunity of Crimean-Congo haemorrhagic fever
in Kosovo. Medicinski Arhiv 64, 9193.
Williams, R.J., Al-Busaidy, S., Mehta, F.R., Maupin, G.O., Wagoner, K.D.,
Al-Awaidy, S., Suleiman, A.J., Khan, A.S., Peters, C.J. and Ksiazek, T.G.
(2000) Crimean-Congo haemorrhagic fever: a seroepidemiological and
tick survey in the Sultanate of Oman. Tropical Medicine and International
Health 5, 99106.
Mariner, J.C., Morrill, J. and Ksiazek, T.G. (1995) Antibodies to hemorrhagic
fever viruses in domestic livestock in Niger: Rift Valley fever and CrimeanCongo hemorrhagic fever. American Journal of Tropical Medicine and
Hygiene 53, 217221.
Agag, A.E. (1998) Rapid detection of Congo-Crimean haemorrhagic fever
antibodies. Assiut Veterinary Medical Journal 39, 280291.
Zeller, H.G., Cornet, J.P. and Camicas, J.L. (1994) Crimean-Congo haemorrhagic fever virus infection in birds: eld investigations in Senegal.
Research in Virology 145, 105109.
Appendix 1
Location
Sheep
Serological
surveillance
19861987
South Africa
Sheep and cattle,

wild vertebrates
Serological
surveillance
1993
South Africa
Cattle
Serological
surveillance
19811984
South Africa
Ostriches
Serological
surveillance
1984
Sudan
Ruminants
Serological
surveillance
1997
United Arab
Emirates
Serological
surveillance
19941995
Zimbabwe
Cattle
Serological
surveillance
19811984
Wilson, M.L., LeGuenno, B., Guillaud, M., Desoutter, D., Gonzalez, J.P.
and Camicas, J.L. (1990) Distribution of Crimean-Congo hemorrhagic
fever viral antibody in Senegal: environmental and vectorial correlates.
American Journal of Tropical Medicine and Hygiene 43, 557566.
Burt, F.J., Swanepoel, R. and Braack, L.E.O. (1993) Enzyme-linked immunosorbent assays for the detection of antibody to Crimean-Congo haemorrhagic fever virus in the sera of livestock and wild vertebrates. Epidemiology and Infection 111, 547558.
Swanepoel, R., Shepherd, A.J., Leman, P.A., Shepherd, S.P., McGillivray,
G.M., Erasmus, M.J., Searle, L.A. and Gill, D.E. (1987) Epidemiologic and
clinical features of Crimean-Congo hemorrhagic fever in southern Africa.
American Journal of Tropical Medicine and Hygiene 36, 120132.
Shepherd, A.J., Swanepoel, R., Leman, P.A. and Shepherd, S.P. (1987) Field
and laboratory investigation of Crimean-Congo haemorrhagic fever virus
(Nairovirus, family Bunyaviridae) infection in birds. Transactions of the
Royal Society of Tropical Medicine and Hygiene 81, 10041007.
Hassanein, K.M., El-Azazy, O.M., Yousef, H.M. (1997) Detection of CrimeanCongo haemorrhagic fever virus antibodies in humans and imported
livestock in Saudia Arabia. Transactions of the Royal Society of Tropical
Medicine and Hygiene 91, 356537.
Khan, A.S., Maupin, G.O., Rollin, P.E., Noor, A.M., Shurie, H.H., Shalabi,
A.G., Wasef, S., Haddad, Y.M., Sadek, R., Ijaz, K., Peters, C.J. and
Ksiazek, T.G. (1997). An outbreak of Crimean-Congo hemorrhagic fever
in the United Arab Emirates, 19941995. American Journal of Tropical
Swanepoel et al. (1987); see South Africa above for full reference.
Senegal
257
258
Appendix 1
Table A1.25. Geographic distribution data for Rickettsia spp. Data were extracted from the references
that are listed by country at the end of the table.
Country
Rickettsia species
Algeria
R. aeschlimannii
Region
Armenia
Northern Algeria (Ain

Ennoussour, Jijel,
Tafaraoui)
R. conorii Malish
Oran, Northern Algeria
(Algiers, El Tarf)
R. massiliae
Northern Algeria (Tizi
Ouzou)
R. monacensis
Northern Algeria (El Tarf)
R. sibirica mongoliti- Southern Algeria
monae
R. slovaca
Armenia
Austria
Austria
Bulgaria
R. helvetica
R. slovaca
R. conorii Malish
Bulgaria
Bulgaria
Bulgaria
Croatia
Croatia
Croatia
Croatia
R. helvetica
R. monacensis
R. slovaca
R. aeschlimannii
R. conorii Malish
R. helvetica
R. hoogstraali
Croatia
Croatia
Cyprus
Denmark
Egypt
Egypt
Egypt
R. rhipicephali
R. slovaca
R. conorii Malish
R. helvetica
R. aeschlimannii
R. conorii Malish
R. sibirica mongolitimonae
R. aeschlimannii
Algeria
Algeria
Algeria
Algeria
France
France
France
France
France
France
France
Germany
Germany
Germany
All mainland
Alpine region
Burgas, Haskovo,
Pazardzhik, Plovdiv,
Stara Zagora, Varna
Soa region
Soa region
NA
Southern Croatia
Dalmatia region, Split
North-western Croatia
Southern Croatia
NA
Jutland county
NA
NA
NA
Identied vector
Human disease
(reported cases)
Hyalomma (Hy.)
detritum, Hy.
marginatum
Rhipicephalus (Rh.)
sanguineus
Rh. turanicus
No
Ixodes ricinus
NAa
No
Yes
Dermacentor
marginatus
I. ricinus
NA
Rh. sanguineus
No
I. ricinus
I. ricinus
NA
Hy. marginatum
NA
D. reticulatus
Haemaphysalis (Ha.)
sulcata
Rh. turanicus
D. marginatus
Rh. sanguineus
I. ricinus
Hy. marginatum
NA
NA
Yes
Yes
No
Yes
Yes
No
No
Yes
No
Yes
No
No
No
Yes
NA
No
No
Yes
Yes
Corsica
Hy. marginatum, Hy. No

rupes (on migratory
birds)
R. conorii Malish
South of France, Corsica Rh. sanguineus
Yes
R. helvetica
Central France, Puy-de- I. ricinus
Yes
Dme
R. massiliae
Camargue, Marseille,
Rh. sanguineus, Rh. No
Sisteron
turanicus
R. raoulti
Marne region
D. reticulatus
No
R. sibirica mongoliti- Southern France,
NA
Yes
monae
Camargue, Marseille
R. slovaca
Southern France
D. marginatus
Yes
R. aeschlimannii
Saxony-Anhalt, Zerbst
D. reticulatus
No
R. helvetica
Southern Germany,
I. ricinus
No
Baden-Wrttemberg
and Bavaria
R. monacensis
Southern Germany,
I. ricinus
No
Bavaria
Continued
259
Country
Rickettsia species
Germany
Germany
R. raoulti
R. slovaca
Greece
Greece
Greece
Greece
Greece
Greece
Hungary
Hungary
Hungary
Hungary
Israel
Region
Southern Germany
Southern Germany,
Alpine region
R. aeschlimannii
Cephalonia Island
R. conorii Malish
All mainland, Cephalonia
Island, Crete Island
R. massiliae
Central Greece,
Cephalonia Island
R. rhipicephali
Cephalonia Island
R. sibirica mongoliti- Crete Island (Sitia)
monae
R. slovaca
Northern Greece
(Halkidiki)
R. helvetica
NA
R. monacensis
NA
R. raoulti
NA
R. slovaca
NA
Italy
R. conorii Israeli
Beersheva, Broil Hail,
spotted fever (ISF)
HaderaCaesarea,
strain
Haifa Bay, Kfar Saba,
Kibbutz, Reim
Tzeelim
R. massiliae
NA
R. sibirica mongoliti- South Israel (Or Haner)
monae
R. aeschlimannii
Sardinia, Sicily
R. conorii ISF
Sicily
R. conorii Malish
All mainland, Sardinia,
Sicily
R. helvetica
Central and northeastern Italy
R. massiliae
Sardinia, Sicily (Palermo)
Italy
R. monacensis
Italy
Italy
R. raoulti
R. slovaca
Central and northeastern Italy (Drena)

Tuscany
Sicily, Tuscany
Italy
Kosovo
Libya
Malta
Moldova
Rickettsia PoTi 169

R. conorii Astrakhan
R. conorii ISF
R. conorii Malish
R. helvetica
Sardinia
NA
NA
NA
NA
Moldova
R. monacensis
NA
Morocco
R. aeschlimannii
Morocco
R. conorii Malish
Taza region (between

Babboudir and
Babezhare)
NA
Israel
Israel
Italy
Italy
Italy
Italy
Identied vector
Human disease
(reported cases)
D. reticulatus
D. marginatus
No
No
Hy. anatolicum
Rh. sanguineus
No
Yes
Rh. turanicus
No
Rh. sanguineus
Hy. anatolicum
No
Yes
NA
No
I. ricinus
I. ricinus
D. reticulatus
D. marginatus,
D. reticulatus
Rh. sanguineus,
Rh. turanicus
No
No
No
Yes
Yes
Rh. turanicus
Hyalomma sp.
No
No
Hy. marginatum
Rh. sanguineus
Rh. sanguineus, Rh.
turanicus
I. ricinus
No
Yes
Yes
Rh. sanguineus, Rh.

turanicus
I. ricinus
Yes
No
No
D. marginatus
D. marginatus, Ha.
punctata
Rh. turanicus
Rh. sanguineus
NA
NA
I. ricinus (on
migratory birds)
I. ricinus (on
migratory birds)
Hy. marginatum
Yes
Yes
Yes
Rh. sanguineus
Yes
No
NA
Yes
Yes
No
No
Continued
260
Appendix 1
Country
Rickettsia species
Region
Identied vector
Human disease
(reported cases)
Morocco
R. helvetica
I. ricinus
No
Morocco
Morocco
R. massiliae
R. monacensis
Rh. sanguineus
I. ricinus
No
No
Morocco
R. raoulti
D. marginatus
No
Morocco
R. slovaca
D. marginatus
No
Netherlands
R. helvetica

Babboudir and
Babezhare)
NA
Babboudir and
Babezhare)
Babboudir and
Babezhare)
Babboudir and
Babezhare)
NA
No
Netherlands
Poland
R. raoulti
R. helvetica
I. hexagonus,
I. ricinus
D. reticulatus
I. ricinus
Poland
Poland
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Portugal
Romania
Russia
(Astrakhan
region)
Russia
(Astrakhan
region)
Russia
(European
part)
NA
Central and southern
Poland
R. raoulti
Eastern, central and
southern Poland
(Biaowieza,
Radomsko, Warszawa)
R. slovaca
Central and southern
Poland
R. aeschlimannii
Alentejo
R. conorii ISF
All mainland
R. conorii Malish
All mainland
R. helvetica
Alentejo and Lisbon
regions
Alentejo and Trs-osR. massiliae
Montes regions
R. monacensis
Alentejo and Lisboa
regions, Madeira
Island
R. raoulti
Alentejo and Trs-osMontes regions
R. sibirica
Alentejo region, Coimbra
mongolitimonae
and Lisbon districts
R. slovaca
Alentejo and Trs-osMontes regions,
Coimbra district
Rickettsia PoTi 169 Alentejo region
R. conorii Malish
Southern Romania
R. conorii Astrakhan Astrakhan region
strain
No
No
D. reticulatus,
I. ricinus
No
I. ricinus
No
H. marginatum
Rh. sanguineus
Rh. sanguineus
No
Yes
Yes
No
Rh. sanguineus
No
I. ricinus
No
H. marginatum
No
Rh. pusillus
Yes
D. marginatus, D.
reticulatus
Yes
Rh. pusillus
NA
Rh. pumilio, Rh.
sanguineus
No
Yes
Yes
R. raoulti
Astrakhan region
Rh. pumilio
No
R. aeschlimannii
NA
Ha. punctata
No
Continued
261
Country
Rickettsia species
Region
Identied vector
Human disease
(reported cases)
Russia
(European
part)
Russia
(European
part)
Serbia
Serbia
Serbia
Slovakia
R. raoulti
Voronezh
D. reticulatus
No
R. slovaca
NA
D. marginatus
No
R. conorii
R. helvetica
R. monacensis
R. monacensis
NA
I. ricinus
I. ricinus
I. ricinus
NA
No
Slovakia
Slovakia
R. raoulti
R. slovaca
NA
NA
NA
North-eastern
(Becherov) and southwestern (Pounajske)
regions
Biaowieza
NA
Slovenia
Slovenia
Slovenia
Spain
R. conorii Malish
R. helvetica
R. monacensis
R. aeschlimannii
NA
NA
NA
Castilla y Lon, Madrid,
Rioja
Spain
R. conorii Malish
Spain
Spain
R. helvetica
R. massiliae
Andaluca, Canary
Islands, Castilla-La
Mancha, Castilla
y Lon, Catalonia,
Madrid, Mallorca
Island
Castilla y Lon, Jaen
Catalonia
Spain
Spain
Spain
Spain
Spain
Sweden
Switzerland
Switzerland
D. reticulatus
D. marginatus, D.
reticulatus, Ha.
inermis, Ha.
punctata, I. ricinus
Rh. sanguineus
I. ricinus
I. ricinus
Ha. inermis, Ha.
punctata, Hy.
marginatum,
Rh. bursa, Rh.
sanguineus, Rh.
turanicus
Rh. sanguineus
No
No
Yes
Yes
No
No
No
Yes
No
Rh. pusillus, Rh.
No
sanguineus, Rh.
turanicus
R. monacensis
Basque country, Castilla Ha. punctata, I. ricinus Yes
y Lon, Jaen, Rioja
R. raoulti
Basque country, Jaen,
D. marginatus, D.
Yes
Rioja
reticulatus, Ha.
punctata
R. rioja
NA
NA
Yes
R. sibirica mongoliti- NA
Rh. pusillus
Yes
monae
R. slovaca
Castilla-La Mancha, Cas- D. marginatus, D.
Yes
tilla y Lon, Granada,
reticulatus
Jaen, Madrid, Rioja
R. helvetica
NA
I. ricinus
Yes
R. helvetica
Alpine region
I. ricinus
No
R. massiliae
Canton Ticino
Rh. sanguineus
No
Continued
262
Appendix 1
Country
Rickettsia species
Region
Identied vector
Human disease
(reported cases)
Switzerland
Switzerland
Tunisia
Tunisia
Tunisia
Turkey
UK
Ukraine,
Crimea
Ukraine,
Crimea
Ukraine,
Crimea
Ukraine,
Crimea
R. monacensis
R. slovaca
R. conorii ISF
R. conorii Malish
R. monacensis
R. conorii Malish
R. helvetica
R. aeschlimannii
NA
Alpine region
Sfax
Monastir, Sousse, Tunis
NA
Trakya
NA
NA
I. ricinus
D. marginatus
NA
Rh. sanguineus
I. ricinus
NA
I. ricinus
Hy. marginatum
No
No
Yes
Yes
No
Yes
No
No
R. conorii
Rh. sanguineus
Yes
R. helvetica
Basin of the Black

and Caspian Seas
Eastern Ukraine
I. ricinus
No
R. monacensis
Eastern Ukraine
I. ricinus
No
NA, not available.
List of scientific papers from which data

were extracted for the table above.
Algeria
Bitam, I., Parola, P., Matsumoto, K., Rolain, J.M., Baziz, B., Boubidi, S.C., Harrat, Z., Belkaid, M. and Raoult,
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Mokrani, N., Parola, P., Tebbal, S., Dalichaouche, M., Aouati, A. and Raoult, D. (2008) Rickettsia aeschlimannii infection, Algeria. Emerging Infectious Diseases 14, 18141815.
Mouffok, N., Parola, P., Lepidi, H. and Raoult, D. (2009) Mediterranean spotted fever in Algeria new trends.
International Journal of Infectious Diseases 13, 227235.
Armenia
Balayeva, N.M., Eremeeva, M.E. and Raoult, D. (1994) Genomic identification of Rickettsia slovaca
among spotted fever group rickettsia isolates from Dermacentor marginatus in Armenia. Acta
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Austria
Rehcek, J. (1993) Rickettsiae and their ecology in the Alpine region. Acta Virologica 37, 290301.
Blaschitz, M., Narodoslavsky-Gfller, M., Kanzler, M., Walochnik, J. and Stanek, G. (2008) First detection of Rickettsia helvetica in Ixodes ricinus ticks in Austria. Vector-Borne Zoonotic Diseases 8,
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263
Bulgaria
Alexandrov, E., Alexandrova, D., Kamarinchev, B., Kantardjiev, T. and Dimitrova, Z. (2007) Marseilles fever
(Mediterranean spotted fever) in Bulgaria state and problems. In: Alexandrov, E., Kazar, J., Hechemy,
K. and Kantardjiev, T. (eds) Contemporary State of the Rickettsioses in the World and in Bulgaria. Prof.
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Christova, I., Van De Pol, J., Yazar, S., Velo, E. and Schouls, L. (2003) Identification of Borrelia burgdorferi
sensu lato, Anaplasma and Ehrlichia species, and spotted fever group rickettsiae in ticks from southeastern Europe. European Journal of Clinical Microbiology and Infection 22, 535542.
Komitova, R., Lakos, A., Aleksandrov, A., Christova, I. and Murdjeva, M. (2003) A case of tick-transmitted
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Croatia
Dobec, M., Golubic, D., Punda-Polic, V., Kaeppeli, F. and Sievers, M. (2009) Rickettsia helvetica in Dermacentor
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Duh, D., Punda-Polic, V., Avic-upanc, T., Bouyer, D., Walker, D.H., Popov, V.L., Jelovsek, M., Gracner, M.,
Trilar, T., Bradaric, N., Kurtti, T.J. and Strus, J. (2010) Rickettsia hoogstraalii sp. nov., isolated from hardand soft-bodied ticks. International Journal of Systematic and Evolutionary Microbiology 60, 977984.
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Psaroulaki, A., Loukaidis, F., Hadjichristodoulou, C. and Tselentis, Y. (1999) Detection and identification of
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Germany
Hildebrandt, A., Fritzsch, J., Franke, J., Sachse, S., Dorn, W. and Straube, E. (2010) Co-circulation of emerging
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Rumer, L., Graser, E., Hillebrand, T., Talaska, T., Dautel, H., Mediannikov, O., Roy-Chowdhury, P., Sheshukova,
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272
Appendix 2 and 3
Appendices 2 and 3
Appendix 2
List (By ID Number) of References from which Geographic Data Were Extracted (hosted on
http://www.cabi.org/resources/tatbd).
Appendix 3
List of Scientific Papers That Could Not Be Retrieved (hosted on http://www.cabi.org/
resources/tatbd).
Appendix 4 Keys for Genera

and Species of Ticks in the
Mediterranean Basin
Agustn Estrada-Pea,1 Alan R. Walker2 and Maxime Madder3

University of Zaragoza, Spain; 2University of Edinburgh, Scotland, UK;
and 3Institute of Tropical Medicine, Antwerp, Belgium
and University of Pretoria, South Africa
2
3
The reliable determination

of the genera and species of
1
ticks that are involved is the first step towards any
further knowledge on the ecology, distribution or
epidemiology of tick-borne pathogens. Current DNAbased methods are not yet accurate enough to be
systematically used for this purpose. The old-fashioned
methods using a stereomicroscope and systematic keys
are still the best5 approach to determining tick taxonomy.
We want to provide in this work an overview of the
most important features of interest that can be used to
identify the ticks present in the Mediterranean basin
and most of Europe. It must be understood, however,
that our work provides information only about the
species of ticks that have an impact on domestic animals,
as well as those which are similar and could be easily
confused. It must also be stressed that the investigation
of immature4 stages might provide more information
than that of adults. Immature stages are more abundant,
and their importance may be overlooked by focusing
only on adult stages.
273
274
Appendix 4
Keys for genera: Amblyomma (A) and Dermacentor (B)

2
A
The genus Amblyomma
5
6.5 mm
13
12
9
11
The basis capituli has straight

lateral margins (4). Legs usually
have pale rings (5) with enamel
ornamentation in the scutum (6).
Eyes and festoons are present (7).
Spiracular plates are large (9), and
ventral plates in males are indistinct
(10) as flat plates posterior to the
anus. An anal groove is posterior to
the anus (11), coxae IV are of
normal size (12) and coxae I have
unequal paired spurs (13).
Dermacentor ticks (Plate B) are
medium sized (see 1) with anterior
mouthparts (2) and broad palpal
articles (3) with straight lateral
margins (4). Legs have no pale rings
(5) and the conscutum has enamel
(6). Eyes are present (7) and
festoons are present in males (8).
Spiracular plates are large (9) and
ventral plates are absent in males
(10). The anal groove (11) is
posterior to the anus, coxae IV are
very large (12) and coxae I have
large and equal paired spurs.
10
The only ornate ticks that are in

Europe or regions in its proximity
are the adults of the genera
Amblyomma and Dermacentor. The
7
former is absent in Europe,
although isolated specimens have
been collected in northern Africa.
Two species of the genus
Dermacentor are very common in
Europe, namely D. marginatus and
8 D. reticulatus. The first is a common
parasite present in Mediterraneantype areas, while the second is a
serious concern because of
repeated reports of its spread into
areas that were formerly free of the
species.
The size of Amblyomma ticks (Plate
A) is large (see 1), the mouthparts
are anterior (2) and palpal articles II
are longer than articles I and III (3).
3
4
7
5
1
6
4.0 mm
B
The genus
Dermacentor
3
13
6
1
3.0 mm
12
9
11
10
Keys for Ticks in the Mediterranean Basin
275
Keys for genera: Rhipicephalus (A) and Haemaphysalis (B)

Adults of the genus
Rhipicephalus are some of the
most common ticks in the
Mediterranean region (Plate
A). They are medium sized
(see 1) and mouthparts are
anterior (2). The palp articles
are all small (3) and the basis
capituli has distinctly angular
lateral margins, making a
hexagonal shape of the entire
basis capituli (4), which is a
diagnostic character. Legs
have no pale rings (5) and the
scutum has no enamel (6).
Eyes are present (7) and
festoons are present in males
(8) and in females but unclear
if fed. Spiracular plates are
large (9) and ventral plates
are present in the males (10),
usually as two pairs of plates.
Anal groove is posterior to
anus (11), coxae IV are of
normal size (12) and coxae I
have large and equal paired
spurs (13).
Adults of the genus
Haemaphysalis (Plate B) are
small (see 1), with anterior
mouthparts (2) and broad
palpal articles (3). The basis
capituli has straight lateral
margins (4). Legs have no pale
rings (5) and the scutum lacks
ornamentation (6). Eyes are
always absent (7) and festoons
are always present (8) but may
be difficult to see in engorged
females. The spiracular plates
are large (9) and ventral plates
are absent from the males (10).
The anal groove is posterior to
the anus (11), coxae IV are of
normal size (12) and coxae I
have no paired spurs there is
only one single spur present (13).
A
The genus Rhipicephalus
3
2
4
7
1
5
6
4.0 mm
8
6
7
8
13
13
12
12
9
11
10
11
10
4
5
7
6
1
3.0 mm
8
13
B
The genus
Haemaphysalis
12
9
11
10
276
Appendix 4
Keys for genera: Hyalomma (A) and Ixodes (B)

2
Both Hyalomma and Ixodes are genera of ticks with long mouthparts
and which lack enamel in the
scutum.
4
5
Ticks in the genus Hyalomma (Plate

A) are large (see 1) with anterior
mouthparts (2) and palp articles II
longer than articles I and III (3). The
basis capituli has medium angular
lateral margins (4) and the legs have
pale rings (5). The scutum has no
ornamentation (6) but Hy. lusitanicum may be an exception to this in
some specimens. The eyes are
always very convex (7) and festoons
are present in both sexes (8), but
difficult to see in engorged females.
Spiracular plates are large (9) and
ventral plates are present only in
males, usually as three distinct pairs
(10). The anal groove is posterior to
the anus (11), coxae IV are of normal size (12) and coxae I have large
and equal paired spurs (13).
7
6
5.5 mm
8
13
A
The genus Hyalomma
12
9
11
10
3
4
3.5 mm
8
B
The genus Ixodes
13
12
Adults of the genus Ixodes (Plate B)

are medium sized (see 1) with anterior mouthparts (2) and palp arti7
cles II longer than articles I and III
(3). The basis capituli has straight
lateral margins (4). Legs never have
pale rings (5) and the scutum lacks
enamel (6). Eyes and festoons are
always absent (7 and 8). Spiracular
plates are large (9) and ventral
plates are present in males only (10)
in the form of large flat plates. The
anal groove forms a loop anterior to
the anus, and in some species forms
a circle around the anus.
9
10
11
277
Keys for species

1
2
1
1 mm
1 mm
4
A
The genus Argas
Three main species of soft ticks (family

Argasidae) are commonly reported in the
western Palaearctic. The genera Argas and
Ornithodoros are characterized by their
ventral mouthparts (Plate A, right, 1; Plate B,
right 3) and the lack of a scutum or
conscutum (Plate A, left, 2) in the adults.
B
The genus Ornithodoros
1
4
8
5
1 mm
7
Two of these species are part of the genus

Argas, a tick commonly found on birds,
which is illustrated in plate A. The genus
Argas can be separated from the genus
Ornithodoros by the presence of a lateral
margin, which is marked by rectangular
plates in A. persicus and by lateral fine ridges
in A. reflexus (Plate A, left, 3 and 4). Such a
lateral suture is absent in the genus
Ornithodoros (Plate B, left, 1). Both genera
have a mammillate pattern on the dorsum
(Plate B, 2), which in Argas is convoluted,
with raised discs present, smoothly flat and
well defined.
278
Appendix 4
4
7
6
6
5
1 mm
1 mm
1 mm
3
3
2
2
4
6
6
The females of the genus Rhipicephalus, including

Rh. bursa (left), Rh. sanguineus (centre) and
Rh. turanicus (right). The female of Rh. annulatus is
illustrated below this legend.
1 mm
It is difficult to distinguish between

females of different species of the
genus Rhipicephalus. A first separation can be made by the use of the
relative size of the spurs in coxae I
(not illustrated). They are very short
and indistinct in Rh. annulatus and
very large and well visible (even
with the naked eye) for the rest of
the species. The separation of
the females of the other species can
be achieved by investigating the
details of the genital opening, the
spiracular plate, the size and position of the eyes, and the dorsal punctations in the scutum, as illustrated
herein. Females of Rh. bursa are commonly easily separated because the
large size of their dorsal punctations.
More difficult is an adequate and
reliable separation of Rh. sanguineus
and Rh. turanicus. If difficulties are
found, it is preferably to report these
species as Rh. sanguineus complex
to avoid confusion.
The males of the

genus
Rhipicephalus
279
2
5
5
1
10
6
1
B
12
12
11
10
11
10
7
7
As with the females, differentiating between males of species of the genus Rhipicephalus in
the western Palaearctic may represent a considerable challenge. Special attention must to
be paid details such as the size and distribution of the conscutum punctations (Plates A, C,
E, 1), the observation of coxae I anterior spurs from a top position (Plates A, C, F, 3) the size
and shape of the adanal plates (Plates B, D, 7, 8), the presence or absence of a caudal
appendage (Plates B, D, E, F, 9) and the aspect of the spiracular plates (Plates B, D, E, F, 10).
All these features are illustrated herein. The final determination of species depends not
only on the presence or absence of a given character, but also on the combination of the
different features of each species. Illustrated are the dorsal and ventral views of the males
of Rh. bursa (Plates A, B), Rh. sanguineus (Plates C, D), Rh. turanicus (Plates E, F) and
Rh. annulatus (Plates G, H).
2
5
9
1
6
12
5
11
3
9
1 mm
10
The males of the

genus
Rhipicephalus
280
Appendix 4
The females of the genus Hyalomma

1
1
2
5
3
4
3
4
1 mm
1 mm
1
5
6
4
5
1 mm
1 mm
2
1
1
5
E
2
3
3 F
4
1 mm
5
1 mm
281
The previous page illustrates the females of the genus Hyalomma. As for the genus Rhipicephalus,
species in the genus Hyalomma are very difficult to determine to a high degree of reliability. We have
included herein the drawings of six species of the genus, which may be not present in the whole
European Mediterranean, but are common in the African coast. It is important to obtain adequate
material for differentiation of species, i.e. avoid engorged specimens. If engorged immature stages
are available, the most useful are to left to moult and the newly moulted adults are examined.
The drawings on the previous page refer to the females of Hy. anatolicum (Plate A),
Hy. excavatum (Plate B), Hy. scupense (Plate C), Hy. dromedarii (Plate D), Hy. lusitanicum (Plate E)
and Hy. marginatum (Plate F). Similar species, such as Hy. rufipes and Hy. turanicum are not
illustrated, although they may be found in places like northern Africa (i.e. Egypt) or Turkey.
The precise distribution of all these species is not yet well known. Hy. dromedarii is an introduced species, and a parasite of the one-humped camel, with morphological similarities to
other species in the genus.
In Hy. anatolicum (Plate A on the previous page), the scapular grooves profile is shallow (1) and
the grooves reach the posterior margin of scutum, which is pale coloured (2). The scutum posterior
margin is smooth (3) and the leg colouration has pale rings (4), but the legs are also pale, so the rings
may be difficult to observe. Punctation size is small and distribution of punctation is sparse (5).
In Hy. excavatum (Plate B on the previous page) the scapular grooves profile is steep (1)
and the grooves reach close the posterior margin of the scutum, which is dark coloured (2). The
scutum posterior margin is slightly sinuous (3), leg colouration is with pale rings (4) and the
legs are mainly dark. The distribution of punctation is localized, with medium numbers mainly
on the anterior of the scutum (5).
In Hy. scupense (Plate C on the previous page), the scapular grooves profile is shallow (1)
and the grooves reach the posterior margin of the scutum, which is dark coloured (2). The
scutum posterior margin is slightly sinuous (3). The leg colouration is without pale rings (5)
although there may be pale patches on the dorsal surface of leg segments. The size of punctations is size is small, and their distribution is localized (6) with small numbers on the anterior
of the scutum, giving it a smooth shiny appearance.
In Hy. dromedarii (Plate D on the previous page), the scapular grooves profile is steep (1)
and the grooves reach the posterior margin of the scutum, which is dark coloured (2).
The scutum posterior margin is slightly sinuous (3). The leg colouration is with pale rings (4).
The size of punctations is large (5) and their distribution is sparse.
The female of Hy. lusitanicum (Plate E on the previous page) has a steep scapular grooves
profile (1) and the grooves are obscured by lines or large punctations, but they reach the posterior margin of the scutum. The scutum is dark coloured but has faint white enamel ornamentation on the scapulae (2). The scutum posterior margin is distinctly sinuous, with distinctly
concave outlines posterior to the eyes (3). The leg colouration is with white enamel (4) as irregular patterns on the dorsal and lateral leg surfaces. Punctation size is large (5) but there are some
small ones and some large ones. These large punctations are shallow, often joined together and
with a rough internal surface. The distribution of punctation is localized, with the large ones in
the cervical fields and central areas.
The female of H. marginatum (Plate F on the previous page) has a steep scapular grooves
profile (1) and the grooves do not reach the posterior margin of the scutum. The scutum is dark
coloured (2). The scutum posterior margin is distinctly sinuous (3). The leg colouration is with
pale rings (5) and there are also patches of pale colour along the dorsal surfaces of the central
segments of all legs. The size of punctations is small (6) and their distribution is sparse.
The illustrations in the following two pages include males of the genus Hyalomma.
282
Appendix 4
The males of genus Hyalomma
1
2
1
10
10
39
The male of Hy. excavatum (Plate B)

has an apparent cervical depression
(1) with the conscutum dark and
heavily sclerotized (2). Lateral
grooves are short (3) but distinct
with a rough surface. There are two
posterior ridges (4) and a caudal
depression
is present. The central
5
4
festoon is pale (5), paracentral
festoons are joined anteriorly (6)
and a posteromedian groove is present (7). The paramedian
grooves are small or indistinct (8). Legs are mainly dark and
the pale rings are well visible (9). Punctations are large (10).
87
The male of Hy. anatolicum (Plate A,

this page) has cervical fields with
evident depression (1), a pale
coloured conscutum (2) and short
lateral grooves (3). There are two
posterior ridges (4) and a caudal
depression is present (5). The
paracentral festoons are separated
anteriorly (6) with a posteromedian
groove that is long and narrow (7).
The paramedian grooves are small
(8). The legs are pale in a patchy or
marbled pattern (9) and punctations
are small and their distribution,
sparse (10).
The male of Hy. scupense is pictured in Plate C. Its cervical

fields depression is apparent (1), with a dark coloured
conscutum (2) and long lateral grooves (3). Posterior ridges
number four (4) and a caudal depression is present. The central
festoon is pale coloured
(but may be dark) and
paracentral festoons are
separated anteriorly. There
is a posteromedian groove
(6). Paramedian grooves
2
are small (7). Leg
colouration is without pale
rings (8). Punctations are
3
small (9).
C
1
7
6
283
The males of genus Hyalomma
9
9
3
8
3
4
4
5
6
4
5
5
4
In the male of Hy. dromedarii (Plate A, this page) the cervical field depression is apparent (1)
and the conscutum is dark coloured (2). Lateral grooves are short (3) and the posterior ridges
number four (4). A caudal depression is present but partially obscured by posterior ridges. The
central festoon is pale coloured (5). The paracentral festoons are separate anteriorly. The
posteromedian groove is present (6), the paramedian grooves are large (7). The leg colouration
is with pale rings (8). Punctations are large and sparse (9).
The male of Hy. lusitanicum is pictured in Plate B. There is an apparent depression in
the cervical fields (1) and a dark coloured conscutum (2) which may have faint white
enamel ornamentation on the scapulae. Lateral grooves are short (3). Posterior ridges
number two and a caudal depression is present (4). The central festoon is pale coloured
(but may be dark coloured). Paracentral festoons are separate anteriorly (5). A posteromedian groove is present. Paramedian grooves are absent (6). Leg colouration is with enamel
ornamentation, as irregular patterns of marbling on the dorsal and lateral leg surfaces (7).
Punctations are large (although there are also small and very large punctations) and their
distribution is localized (8).
The male of Hy. marginatum is shown in Plate C. There is an apparent cervical depression
(1) and the conscutum is dark coloured (2). Lateral grooves are long and they continue
towards the eyes as lines of punctations (3). Posterior ridges number two and a caudal depression is present but shallow (4). The central festoon is dark coloured (5). Paracentral festoons are
separate anteriorly (5) and a posteromedian groove is present (6). Paramedian grooves are
small (7), but all the posterior grooves are shallow and may be indistinct. Leg colouration is
with pale rings; there are also patches of pale colour along the dorsal surfaces of the legs (8).
Punctations are small and their distribution is sparse (9).
284
Appendix 4
The males of the genus Dermacentor
There are only two species of ticks in

the genus Dermacentor in the western Palaearctic, namely D. marginatus (Plate A) and D. reticulatus (Plate
B). Both are large, very ornate ticks
whose
identities
are
easily
determined.
The males of D. marginatus (Plate A)
have short cornua (1). The posterior
spur on palpal articles II is short on
the dorsal surface (2). In coxae I, the
gap between internal and external
spurs is medium, and the external
spur is slightly shorter than the
internal spur (3). The lateral groove
is indistinct, only more conspicuous
than the punctations (4). The
posterior spur on trochanter I is
short on the dorsal surface (5).
The male of D. reticulatus (Plate B)

has long cornua (1). The posterior
spur on palpal articles II is long on
the dorsal surface (2). In coxae I, the
gap between external and internal
spurs is narrow, and the external
spur is as long as the internal spur
(3). The lateral groove is in the form
of punctations only (there is no
groove visible) (4). The posterior
spur on trochanter I is long on the
dorsal surface (5).
As with the males, differentiating

between females of the two species
of the genus Dermacentor in the western Palaearctic is easy because they
are the only ornate ticks found. Both
species are included in this page:
D. marginatus (Plate A) and D. reticulatus (Plate B).
In the females of D. marginatus, the
shape of the porose areas is a narrow
oval, and they are slanting (1). The
posterior spur of palp articles II is
absent from the dorsal surface (2). In
coxae I, the gap between the external and internal spurs is medium,
and the external spur is slightly
shorter than the internal spur (3).
The posterior lips of the genital
aperture have a narrow V shape (4)
and are not truncated at the posterior margin.
285
The females of the genus Dermacentor

2
A
1
1 mm
The females of D. reticulatus have

porose areas shaped in a broad oval,
nearly circular, pattern (1). The posterior spur of palp articles II is
present on the dorsal surface (2). The
gap between coxae I external and
internal spurs is narrow, and the
external spur is as long as the internal spur, as displayed in (3). The
posterior lips of the genital aperture
have a broad U shape, which is truncated posteriorly (4).
2
B
1
3
4
1 mm
286
Appendix 4
There are two prominent species of

the genus Haemaphysalis in the
western Palaearctic, namely Ha.
punctata (Plate A) and Ha. sulcata
(Plate B). Other species, such as Ha.
numidiana, Ha. erinacei, Ha. concinna,
etc. are not so common or of
importance to the health of domestic
animals or humans, as far as we
know currently. New studies and
additional collections of these
species will most likely provide
interesting data about the importance
and distribution of this tick genus.
The male of the genus Haemaphysalis
2
1
The males of Ha. punctata have a

short lateral extension (1) on palp
articles II, and the dorsal spur is
absent from these articles (2). The
ventral spur is also absent from palp
segments II (3). The spurs of coxae
IV are long, extending to the level of
the posterior margin of the anus and
curving inwards (4). There are 11
festoons (5), and two of these are
enclosed by a lateral groove (6).
Distribution of the punctuations is
dense (7). The cornua are short (8).
4
6
3
1
4
6
5
In the males of Ha. sulcata, the lateral

extension of palp segments II is
longer than in Ha. punctata (1) and
the dorsal spur is absent from these
articles (2). Palp segments 2 The
ventral spur is also absent from palp
segments II (3). The spurs of coxae
IV are long, extending to the level of
the posterior margin of the anus,
and curving outwards (4). There are
nine festoons (5), and only one is
enclosed by a lateral groove (6).
Distribution of the punctuations is
sparse (7). The cornua are long (8).
In the females of Ha. punctata (Plate

A, this page), the lateral extension of
palp articles II is small (1), and the
dorsal spur is absent (2). The ventral
spur of palp segments III is also
absent (3). The spurs of coxae I to II
are of medium length, and those of
coxae IV are distinct (4). There are 11
festoons (5) and three of these are
enclosed by each lateral groove (6).
The distribution of punctuation is
dense (7).
In the females of Ha. sulcata (Plate B,
this page), the lateral extension to
palp articles II is small (1) and the
dorsal spur is absent from these
articles (2). The ventral spur is
present on palp articles III (3). The
spurs of coxae I to III are short and
those of coxae IV are indistinct (4).
There are nine festoons (5), and only
one festoon is enclosed by each
lateral groove (6). The distribution
of punctation is sparse (7).
287
The females of the genus Haemaphysalis

3
1
A
2
4
1 mm
3
B
There are several species of the

genus Ixodes in the western
Palaearctic. They are rare in northern
Africa, and much more common in
Europe. Details are given here for
female ticks only, and for the species
I. ricinus, I. gibbosus and I. hexagonus.
Illustrations of the females of the
genus Ixodes are included on the
next page (Plates A to F) in both
dorsal (A, C, E) and ventral views
(B, D, F).
2
4
6
1 mm
288
Appendix 4
The females of the genus Ixodes

3
2
4 1
4
5
7
1 mm
6
A
The female of I. gibbosus (Plates C,

D) is very similar to that of I. ricinus,
4 but there are important differences
in the tarsi, which are very tapered
Plate C, 3), the auriculae are short
(Plate D, 2) and the internal spurs of
coxae I are shorter. The posterior
alignment of the anal groove is long
and converging (D, 5).
1
1
4
56
7
1 mm
C
4 1
5
7
6
1 mm
In female I. ricinus (Plate A), scapular grooves are present (1), palp
alignment slopes inward (2), the
tarsi are slightly tapered towards the
claws (3), punctations are distinct
(4), setae are present on the scutum
(5), setae on the alloscutum are thin
and colourless (6), and the scutum
posterior margin is slightly sinuous
(7). On its ventral surface (Plate B),
I. ricinus has coxae of normal type
(1), auriculae are distinct (2), coxae I
internal spurs are very long (3),
coxae II to IV external spurs are distinct (4), alignment of the posterior
anal groove is long and diverging
(5), and the genital aperture is
between coxae IV (6).
6
F
I. hexagonus is a very different spe3 cies (Plates E, F). The dorsal surface
(Plate E) scapular grooves are absent
(1), palp alignment slopes inward
(2), the tarsi are blunt stepped
towards the taris (3), punctations are
4
distinct (4), setae are present on the
scutum but small and sparse (5),
setae on the alloscutum are thin and
colourless (6). On the ventral surface
(Plate F), coxae are of normal type
(1), auriculae are indistinct (2), internal spurs of coxae I are short (3),
external spurs of coxae II to IV are
indistinct (4), posterior alignment of
the anal groove is long and diverging (5), and the genital aperture is
between coxae III (6).
Index
African horse sickness (AHS), 5758

African swine fever (ASF) virus (ASFV)
Asfarviridae family, 33
CCHF, 3637
domestic pigs, 3334
genetic diversification, 3536
infected warthogs, 33
Mediterranean basin, 7
no treatment/effective vaccine, 3334
non-infected pig population, 34
occurrence, 77
Ornithodoros, 911
Ornithodoros moubata, 34
Ornithodoros porcinus, 34
swine haemorrhagic diseases, 34
tonsils and mandibular lymph node, 34
viral swine disease, 33
virus-tick interaction, 35
wild pigs, 34
wild pigs and soft ticks, 34
Anaplasmoses
alphaproteobacteria, 39
Anaplasma bovis, 42
Anaplasma centrale, 4142
Anaplasma marginale, 41
Anaplasma ovis, 42
Anaplasma phagocytophilum, 3941
Anaplasma platys, 4243
hard ticks transmitted, 3940
intraerythrocytic bacteria, 39
Argasidae
Argas
Ar. persicus, 8
Ar. polonicus, 9
Ar. reflexus, 89
eight-legged nymph instar, 78
larvae, 8
nocturnal and endophilic, 7
oval shaped and dorsoventrally
flattened, 7
spiracular plates, 7
unfed adults, 7
economic and medical impact, 7
gnathosoma, 7
leathery cuticle and nymphal stages, 7
Ornithodoros
ASFV, 1011
discrepancies concerning infection, 10
identification, 11
Malawi strain, 10
occurrence, 8990
O. erraticus, 910
O. moubata group, 10
O. moubata/porcinus ticks, 10
sexual dimorphism, 7
unsclerotized, 7
xerophilic arthropods, 7
Astrakhan fever (AF), 48
babesioses, 5354
basic biology and geographical distribution
Argasidae (see argasidae)
Ixodidae (see Ixodidae)
Nutalliellidae, 6
sac-like appearance, 6
Boophilus ticks, 20

virus (CCHFV)
crop rotation and land fallowing, 4
289
290
Index

virus (CCHFV) (continued)
distribution map, 38
eco-epidemiology, 3
epidemic of, 3
Hyalomma anatolicum, 17
Hyalomma marginatum, 16
IgM and IgG positivity rates, 3
land use changes, 28
pH inactivation, 37
potential vertebrate host reservoirs, 36
Thrace region, 3
tick-borne zoonotic infection, 36
S-, M-and L-RNA segments, 36
Ehrlichioses
Ehrlichia canis, 43
Ehrlichia ruminantium, 43
Europe and Mediterranean basin, tick-borne
infections
African horse sickness (AHS), 5758
anaplasmoses (see anaplasmoses)
ASF (see African swine fever (ASF) )
Bartonella infections, 58
Crimean-Congo haemorrhagic fever (CCHF),
3637
Ehrlichioses (see Ehrlichioses)
hepatozoonosis, 56
Lyme borrelioses (LB), 5152
piroplasmoses
babesioses, 5354
theilerioses, 5456
Q fever, 5859
Recurrent (relapsing) fever, 52
rickettsioses
Rickettsia aeschlimannii, 4950
Rickettsia conorii (see Rickettsia conorii)
Rickettsia helvetica, 50
Rickettsia massiliae, 49
Rickettsia monacensis, 50
Rickettsia sibirica mongolitimonae strain, 48
Rickettsia slovaca, 4849
Rickettsia species, 4445
TBE group (see Tick-borne encephalitis (TBE) )
tularaemia, 5960
Flaviviridae, 37
Francisella tularensis infection, 86
Greek goat encephalitis virus (GGEV), 39
Integrated Consortium on Ticks and Tick-borne

Diseases, 76
Israeli spotted fever (ISF), 47

Ixodes ricinus
Anaplasma phagocitophylum, A. marginale,
A. centrale, 41
castor bean/sheep tick, 12
endophilic species, 12
Hepatozoon transmission, 56
louping ill virus (LIV), 38
louping ill virus primary vector and
virus reservoir, 39
Lyme disease, 2
occurence, 9091
Palaearctic region, 2
Rickettsia monacensis, 50
TBEV and TBE, 2
three-host tick, 12
vegetation, 2
zoonoses, 12
Ixodidae
Amblyomma
Am. americanum, 1819
Am. hebraeum, 18
Am. maculatum, 18
Am. variegatum, 18
ornate eyes and coxal spurs, 18
parasitize reptiles, 18
Dermacentor
D. marginatus, 2021, 96
D. reticulatus, 2122, 9697
ornate patterning, 20
Haemaphysalis
capitulum, 14
Ha. concinna, 14, 92
Ha. inermis, 15, 92, 93
Ha. punctata, 14, 9192
Ha. sulcata, 15
festoons, 14
parasitize livestock, 13
three-host ticks, 14
Hyalomma
capituli, 16
genetic instability, 1516
Hy. anatolicum, 17
Hy. excavatum, 17
Hy. marginatum, 16, 93
Hy. scupense, 1718
immature stage identification, 16
morphological characteristics, 16
pale rings, 15
parasites of livestock, 16
Ixodes
host-seeking behaviour, 12
I. canisuga, 13
I. frontalis, 13
I. gibbosus, 13
I. hexagonus, 13
I. persulcatus, 12
Index
I. ricinus (see Ixodes ricinus)

I. trianguliceps, 13
I. uriae, 13
prostriate hard ticks, 12
morphology, 11
one nymphal stage, 11
Rhipicephalus
brown ticks, 19
hexagonal basis capituli, 19
parasitize wild and domestic
artiodactyls, 19
Rh. annulatus, 20, 9596
Rh. bursa, 20, 9495
Rh. camicasi, 1920
Rh. sanguineus, 1920, 94
Rh. turanicus, 1920

Lyme borrelioses (LB), 5152
Mediterranean spotted fever (MSF), 4546

monitoring and surveillance
system (MoSS), 99
data collection methods, 101
disease identification, 99100
disease-positive animal, 100
goals and objectives identification, 100
hazards, 100
public health and socio-economic
impact, 100
random sampling, 101
sample populations, 101102
serological testing, 100
storage and tracking methods, 101
Nomenclature of territorial units for statistics

(NUTS), 76
Nutalliella namaqua, 6
Ornate sheep tick, 2021

Ovine encephalitis/encephalomyelitis
virus, 3839
Q fever, 5859
recurrent (relapsing) fever, 52

Rickettsia conorii
Astrakhan fever (AF), 48
Israeli spotted fever (ISF), 47, 81
Mediterranean spotted fever (MSF), 4546
spotted fever group (SFG) rickettsiae, 46
291
soft ticks (see Argasidae)

spread and distribution
animals movement
domestic animals movement, 2930
exotic and wildlife species, 29
migratory birds, 29
climate changes, 2728
land use changes, 28
surveillance and control measures
action-oriented programme, 99
argasidae, 110
data collection methods, 103104
definition, 9899
flag/drag cloth, 98
hard ticks control options
acaricides, 104
biological control, 106107
chemical control of ticks on animals,
104106
genetic resistance, 107108
habitat modification, 109
integrated tick management, 109110
personal protection, 108
pheromone/hormone-assisted control, 106
tick prophylaxis, 104
vaccine, 108
mark-and-recapture method, 98
MoSS (see monitoring and surveillance
system (MoSS) )
ornithodoros, 110
predefined threshold, 99
scratching technique, 98
tick specimens collection, 99
theilerioses, 5456
tick-borne encephalitis (TBE) virus (TBEV)
aetiological agent, 37
Apodemus flavicollis, 38
Clethrionomys glareolus, 38
Flavivirus, 37
Greek goat encephalitis virus (GGEV), 39
human infections, 37
I. ricinus, 3738
Kumlinge virus, 37
neurological syndrome, 38
viral subtypes, 37, 78
tick-borne pathogens occurrence
anaplasmoses and ehrlichioses, 80
African swine fever (ASF), 77
babesioses, 8283
bartonelloses, 8586
borrelioses, 8182
Crimean-Congo haemorrhagic fever
(CCHF), 7778
equine piroplasmosis, 8485
292
tick-borne pathogens occurrence (continued)

Integrated Consortium on Ticks and
Tick-borne Diseases, 76
NUTS, 76
Q fever, 8687
Index
rickettsioses, 8081
theilerioses, 83, 84
tick-borne encephalitis group viruses, 7880
tularaemia, 86
tularaemia, 5960

Tick and Tickborne Disease

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Ticks and Tick-borne Diseases

Geographical Distribution and Control Strategies

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Ticks and Tick-borne Diseases

Mo Salman and Jordi Tarrs-Call

Agustn Estrada-Pea, Rbert Farkas, Thomas G.T. Jaenson,

CABI is a trading name of CAB International

Tel: +1 800 552 3083 (toll free)

CAB International 2013. All rights reserved. No part of this

Emerging (Re-emerging) Tick-borne Infections and the

Tick-borne Infections (Including Zoonoses) in Europe

Table A1.3. Geographic distribution data for the tick-borne encephalitis

Appendix 2 List (By ID Number) of References from which Geographic Data

Appendix 3 List of Scientific Papers That Could Not Be Retrieved

The colour plate section can be found following p. 8

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Agustn Estrada-Pea, Professor of Parasitology, Department of Animal Pathology

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Mo Salman1 and Jordi Tarrs-Call2

Significance of Tick-borne Diseases

Mo Salman and Jordi Tarrs-Call

Methods Used for the Systematic

Tick-borne African Swine Fever virus OR ASF

AND (search terms related to the

Aland OR Albania OR Andorra OR

Concerns occurrence of a tick species or a tick-borne

Mo Salman and Jordi Tarrs-Call

Tick genus and species, recorded as in the original paper.

Table I.3. Criteria used in the second screening for relevance.

English abstract and text Abstract not available

A more detailed description of the systematic literature review process is given in

Identification Methods for Tick Species, Including the Most

Mo Salman and Jordi Tarrs-Call

External structure of adult argasid ticks (the example is Ornithodoros)

four pairs of legs

External structure of adult ixodid ticks (the example is Hyalomma)

MALE, DORSAL VIEW

FEMALE, DORSAL VIEW

FEMALE, VENTRAL VIEW

MALE, VENTRAL VIEW

EndNote X1.0.1 [Bld 2682], Thomson. http://www.endnote.com

Size (unfed Large

Dermacentor Haemaphysalis Amblyomma

Mo Salman and Jordi Tarrs-Call

Emerging (Re-emerging) Tick-borne

Mo Salman1 and Agustn Estrada-Pea2

Ticks are arthropods and therefore they are

example, limited variations in temperature

Mo Salman and Agustn Estrada-Pea

(Talleklint and Jaenson, 1998) and colonize a

(Rogers and Randolph, 2006). It is, however,

Emerging Tick-borne Infections and Dissemination

sites with active foci of the infection and sites

amplification foci, with consequently higher

Mo Salman and Agustn Estrada-Pea

active immune protection (Gargili et al.,

the enzootic amplification cycle in Turkey