Hydrobiologia (2018) 810:1–14

https://doi.org/10.1007/s10750-017-3486-7

FRESHWATER BIVALVES

Conservation of freshwater bivalves at the global

scale: diversity, threats and research needs
Manuel Lopes-Lima . Lyubov E. Burlakova . Alexander Y. Karatayev .
Knut Mehler . Mary Seddon . Ronaldo Sousa

Received: 11 December 2017 / Revised: 11 December 2017 / Accepted: 16 December 2017 / Published online: 25 January 2018
Ó Springer International Publishing AG, part of Springer Nature 2018

Abstract Bivalves are ubiquitous members of fresh- threats that vary across the globe; however, pollution
water ecosystems and responsible for important func- and natural system (habitat) modifications being
tions and services. The present paper revises consistently found as the most impacting. Freshwater
freshwater bivalve diversity, conservation status and bivalves are among the most threatened groups in the
threats at the global scale and discusses future research world with 40% of the species being near threatened,
needs and management actions. The diversity patterns threatened or extinct, and among them the order
are uneven across the globe with hotspots in the Unionida is the most endangered. We suggest that
interior basin in the United States of America (USA), global cooperation between scientists, managers,
Central America, Indian subcontinent and Southeast politicians and general public, and application of
Asia. Freshwater bivalves are affected by multiple new technologies (new generation sequencing and
remote sensing, among others) will strengthen the
quality of studies on the natural history and conser-
Guest editors: Manuel P. M. Lopes-Lima, Ronaldo G. Sousa,
Lyuba E. Burlakova, Alexander Y. Karatayev & Knut Mehler / vation of freshwater bivalves. Finally, we introduce
Ecology and Conservation of Freshwater Bivalves

M. Lopes-Lima (&) L. E. Burlakova
A. Y. Karatayev
K. Mehler

CIBIO/InBIO - Research Center in Biodiversity and Great Lakes Center, Buffalo State College, 1300
Genetic Resources, University of Porto, Campus Agrário Elmwood Ave, Buffalo, NY 14222, USA
de Vairão, Vairão, Portugal
e-mail: manuelpmlopeslima@gmail.com R. Sousa
CBMA - Centre of Molecular and Environmental
M. Lopes-Lima
R. Sousa Biology, Department of Biology, University of Minho,
CIIMAR/CIMAR—Interdisciplinary Centre of Marine Campus Gualtar, 4710-057 Braga, Portugal
and Environmental Research, University of Porto,
Terminal de Cruzeiros do Porto de Leixões, Avenida
General Norton de Matos, S/N, 4450-208 Matosinhos,
Portugal

M. Lopes-Lima
M. Seddon
SSC/IUCN—Mollusc Specialist Group, Species Survival
Commission, International Union for Conservation of
Nature, c/o The David Attenborough Building, Pembroke
Street, Cambridge CB2 3QZ, UK

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the articles published in this special issue of Hydro- needs and urgent management actions that may help
biologia under the scope of the Second International conserve these animals, and introduce the articles
Meeting on Biology and Conservation of Freshwater published in this special issue resulting from the
Bivalves held in 2015 in Buffalo, New York, USA. Second International Meeting on Biology and Con-
servation of Freshwater Bivalves held in 2015 in
Keywords Bivalvia
Unionida
Venerida
IUCN Buffalo, United States of America (USA).
Red List
Freshwater mussels
Conservation

Diversity patterns at the global scale

Introduction FB are a polyphyletic group of animals restricted to

fresh waters with a little over 1,200 described species
Freshwater ecosystems are among the most threatened (Bogan, 2008; Bogan & Roe, 2008; Graf, 2013). The
on the planet facing unprecedented pressures related to main core of the group (99%) is composed of
the increase of human population and socioeconomic freshwater mussels of the order Unionida (strictly
development (Dudgeon et al., 2006; Vörösmarty et al., freshwater) (72%) and species belonging to 7 families
2010). Increasing anthropogenic pressure worldwide within the order Venerida (27%) (Fig. 1). The Vener-
results in habitat loss, habitat modification and frag- ida are composed mainly of families comprising 94%
mentation, overexploitation of natural resources (in- of the species the pea- or fingernail-clams Sphaeriidae
cluding water), pollution, introduction of invasive (67%) and the Cyrenidae (27%), which include, for
alien species (IAS) and climate change (Malmqvist & example, the invasive Asian clam Corbicula fluminea
Rundle, 2002; Strayer & Dudgeon, 2010). Biodiver- (Müller, 1774). The family Dreissenidae family (3%),
sity crisis is one of the major consequences of steeply well-known to contain important invasive alien
rising human demands, and among the animals with species (IAS) such as the quagga mussel Dreissena
high extinction rates are freshwater bivalves (FB) bugensis Andrusov, 1897 and the zebra mussel
(Strayer et al., 2004; Lydeard et al., 2004; Régnier Dreissena polymorpha (Pallas, 1771), is also included
et al., 2009; Lopes-Lima et al., 2014, 2017a). The in the order Venerida. The remaining handful of FB
future survival of FB is highly impaired and consid- species are scattered among other essentially marine
ering the large suite of ecosystem services they orders or families within the order Venerida (Fig. 1).
provide (Vaughn, 2017) scientists, managers, politi- FB are present in all continents except in glaciated
cians and the general public need to strengthen their (with the exception of few sphaeriid species) and
cooperation in order to conserve these species. desert areas, but the diversity patterns are not evenly
Whereas over the last years multiple studies have distributed (Fig. 2). The diversity is higher in the
been published concerning the biology, ecology and Nearctic (NA), Neotropics (NT) and Indotropics (IN)
conservation of FB, the majority of them were carried with & 25% species being found in each ecoregion.
out in North America and Europe (Lopes-Lima et al., The Palaearctic (PA) and Afrotropics (AF) have a
2014). Consequently, a great ignorance about basic lower diversity (& 10%) with Australasia (AU) being
aspects (e.g. distribution, diversity, abundance, pop- the poorest ecoregion (& 5%) (Fig. 2A). There are
ulation structure and life cycle) concerning species also distinct distribution patterns across the main
inhabiting South America, Africa and Asia still taxonomic groups. The Unionida is similar to the
persists and much more information is needed for general pattern for all FB, with 33% of the species
these continents. inhabiting the NA and 6% inhabiting the PA (Fig. 2B).
In the present paper, we compile data on FB The distribution of pea clams is completely distinct
diversity patterns, conservation status and threats from with the hotspots of diversity being the NT (31%) and
the International Union for Conservation of Nature the PA (22%), while the remaining diversity is
(IUCN) database using a species list adapted from scattered among the other continents (Fig. 2C).
Graf & Cummings (2017) and mapped them in Sphaeriids are also the only FB species that are
ecoregions adapted from Graf & Cummings (2007) capable of living at the higher latitudes of the Arctic,
and Haag (2010). We also briefly discuss research such as the islands of Iceland, Greenland, Baffin,

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Fig. 1 Global diversity of freshwater bivalves divided by families. Total number of species in brackets

Fig. 2 Diversity by ecoregions. A All freshwater bivalves; Cummings (2007) and Haag (2010): NA Nearctic, NT Neotrop-
B Unionida; C Sphaeriidae; D Cyrenidae ? remaining fresh- ical, PA Palaearctic, AF Afrotropical, IN Indotropical, AU
water bivalve groups. Ecoregions adapted from Graf & Australasian. Glaciated and desert areas void of mussels in grey

Svalbard and Novaya Zemlya (Schiøtte & Warén, diversity hotspot is in the IN that contains almost 70%
1992; Bespalaya et al., 2017). Finally, for Cyrenidae of such species, followed by the PA (18%), while other
and for a few other remaining species, the major ecoregions have a much lower diversity (Fig. 2D).

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Diversity at an ecoregion scale is also not dis- 8 families in the NT (Fig. 4). Even within the most
tributed evenly (Fig. 3). Within NA, the species species-rich FB family, the Unionidae, the IN exhibits
diversity is generally higher in the interior basins, a much higher taxonomic diversity than all of the other
while in the NT the diversity is higher in Central ecoregions, with representatives of all subfamilies of
America and in the Orinoco, Amazon and Paraguay Unionidae occurring there, except for the NA
River basins (Fig. 3A). In the AF, the Congo River Ambleminae.
basin is richer in Unionida species (Fig. 3B), and the We would like to stress that diversity patterns
Nile and Eastern African River basins have a higher described above may be underestimated and may
sphaeriid diversity (Fig. 3C). While the Western change substantially as a result of ongoing and future
Palaearctic is quite diverse in sphaeriids and dreis- surveys in the less studied regions of Southeast Asia,
senids, Laurasia has a higher diversity in the IN, from Africa, NT and AU. For example, Bolotov et al. (2017)
the Hindu to the Amur River basin (Figs. 3A, C and studying the FB of a poorly known and remote basin
D). Within IN, the diversity of sphaeriids is higher in (Sittaung) in Myanmar described two new genera and
the Indian subcontinent, while in the Unionida and the seven new species. Also, even in Europe and NA,
remaining groups the diversity is higher in Indochina which are the most well-studied continents, the
and Sundaland (Fig. 3). In AU, a higher number of knowledge of the diversity of Unionida is still
species is found in the East (Fig. 3). undergoing considerable changes (e.g. Froufe et al.,
Although specific diversity of FB is similar in NA, 2016a, b, 2017; Araujo et al., 2017; Lopes-Lima et al.,
NT and IN, there is a higher taxonomic diversity in the 2017a; Williams et al., 2017; Smith et al., 2018).
IN than in NA and NT. In the IN there are represen-
tative species of 5 orders and 10 families compared to
the 2 orders and 4 families in the NA and 3 orders and

Fig. 3 Diversity by ecoregions. A All freshwater bivalves; Neotropical, PA Palaearctic, AF Afrotropical, IN Indotropical,
B Unionida; C Sphaeriidae; D Cyrenidae ? remaining fresh- AU Australasian. Glaciated and desert areas lacking FB are in
water bivalve groups. Ecoregion subdivisions adapted from grey
Graf & Cummings (2007) and Haag (2010): NA Nearctic, NT

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Fig. 4 Taxonomic composition and diversity of freshwater bivalves in each ecoregion. Total number of species in brackets

Conservation status and major threats threatened and extinct species (including 25 [2.8%]
extinct or probably extinct species) is in Unionida,
FB are among the most threatened taxonomic groups while only 14.5% of Sphaeriidae and 8.8% of
in the world, with almost 40% of the species being Cyrenidae (plus all the other remaining species) have
near threatened, threatened or extinct (Fig. 5). How- a near-threatened or threatened status (Fig. 5, top
ever, this high imperilment is mainly due to the part). However, IUCN assessments are not evenly
contribution of Unionida since not all groups are distributed across taxa and countries and FB are a good
evenly threatened or assessed for conservation status example of this situation (Fig. 5). Thus, a higher
(Fig. 5, top). Based on the number of assessed species, percentage of large and more conspicuous unionids
the highest percentage (45%) of near-threatened, has been assessed compared to other FB groups

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Fig. 5 Map of IUCN Red List conservation status for Unionida Afrotropical, IN Indotropical, AU Australasian. On the scale bar:
freshwater mussels by ecoregions (bottom of the figure) and NE Not evaluated by the IUCN Red List; and the IUCN Red List
global conservation status for freshwater bivalves and each categories: DD data deficient, LC least concern, NT near
major freshwater bivalve group (top of the figure). Ecoregion threatened, VU vulnerable, EN endangered, CR critically
subdivisions adapted from Graf & Cummings (2007) and Haag endangered, CR (PE) critically endangered probably extinct,
(2010): NA Nearctic, NT Neotropical, PA Palaearctic, AF EX extinct

(Fig. 5, top). Some ecoregions (e.g. NA, AF, PA and and (E) a quantitative analysis of extinction probabil-
IN) have a high percentage of species evaluated, while ity (IUCN, 2012). Most of the near-threatened and
species from AU and especially NT have a very low threatened FB species have been assessed using
Red List coverage (Fig. 5, bottom part). The percent- criteria A and B and to a much lesser extent using
age of threatened and near-threatened Unionida criteria C and D (Fig. 6). Since criterion E needs
species is higher in NA (67%) and PA (52%) than in comprehensive data on a wide range of features (e.g.
other ecoregions, with the lowest percentage (19%) in demography, life history, habitat requirements, threats
the IN (Fig. 5, bottom). This does not necessarily and management options), no FB species was ever
mean that less species are threatened in the IN, since evaluated using this criterion (Fig. 6). Most FB
this ecoregion has a much higher percentage of data- species have been assessed based on their population
deficient species, reflecting the lower level of knowl- size reduction and geographic range contraction
edge and data on the threats available for IN species. compared to a few species with very small distribution
On the other hand, almost half of the species have been ranges. In fact, it is difficult to assign a threatened
assessed as of ‘‘least concern’’ in the AF, which might status using criterion D for most FB species due to
indicate a more favourable status of freshwater their generally large distribution ranges.
mussels in this ecoregion. The global pattern is more or less similar in all
The IUCN Red List assessments are based on a set ecoregions, with the exception of the NT and AF
of five criteria: (A) population size reduction, (Fig. 6). While the NT pattern may not be very
(B) small geographic range, (C) small population size representative of the ecoregion due to the few assessed
plus decline, (D) very small or restricted populations species, in AF it reflects the poor knowledge about the

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of the NA and the PA, habitat modifications seem to

negatively affect more species in these ecoregions
than in the AF and IN. Exploitation is a much more
detrimental threat in the IN than elsewhere (Fig. 7). In
fact, harvesting of mussels for human consumption in
East and Southeast Asia is a major economic activity;
for example, in Vietnam it may reach up to 50,000 tons
per year in each major basin (Köhler et al., 2012).
Furthermore, the ratio of agriculture related threats is
higher in AU and PA, mainly due to water diversion
and extraction.

Research and conservation actions needs

Many species of FB are still poorly understood,

especially in Central America, Southeast Asia and
Sundaland (Lopes-Lima et al., 2014, 2017b). This lack
Fig. 6 IUCN Red List criteria used for the assessment of
freshwater bivalve species by ecoregions. Ecoregion subdivi-
of knowledge hampers their status assessment.
sions adapted from Graf & Cummings (2007) and Haag (2010): The IUCN database indicates that research needs
AF Afrotropical, AU Australasian, IN Indotropical, NA Nearctic, are generally lower for the NA and PA compared to the
NT Neotropical, PA Palaearctic. IUCN Red List criteria: other ecoregions, especially for the three top research
A population size reduction, B geographic range, C small
population size and decline, D very small or restricted
needs, i.e. population size and distribution, identifica-
populations tion of threats and life history (Fig. 8A). This may be
explained by the stronger research effort and higher
financial support available for North American and
population size and trends. This is due to the lack of European studies. However, even in these ecoregions,
research that is being done in the AF, where survey basic data on distribution, population size, accurate
and monitoring studies are almost non-existent identification of threats and basic life history traits are
(Lopes-Lima et al., 2014; Sousa et al., 2016, 2017). still lacking for many species. Taxonomical data and
FB are affected by multiple threats that range from knowledge on life history traits are particularly needed
natural system modifications to degradation, pollution, for AF species. The same general trends in research
introduction of IAS, exploitation and human distur- needs can be seen for all species assessed by IUCN as
bance. Within the assessed FB species for the IUCN well as for data-deficient species (Fig. 8B).
Red List, pollution is still the most recorded global Due to the high risk of extinction, many species
threat comprising 42% of all threats (Fig. 7). Natural urgently need worldwide conservation actions. Land
system (habitat) modifications such as the construction and water protection was found to be the top conser-
of dams and channels are the second most cited threat vation measure globally and throughout all ecore-
(20%), followed by urban development, exploitation, gions, but especially for IN species (Fig. 9). Land and
agriculture, climate change, mining and IAS, together water management is also shown to be one of the top
representing less than 10%. Other disturbances such as priorities for FB conservation, particularly in the PA
transport, recreational activities and geological events and AF ecoregions (Fig. 9). Other types of conserva-
only play a minor role. tion actions showed quite distinct patterns among
The relative percentage of recorded threats is ecoregions. For example, a stronger legislation is
generally similar across the main ecoregions with a likely required for the AF, PA, NA and AU, but law
few notable exceptions. For instance, in the NA and enforcement needs to be enhanced only in the AF and
PA species seem to be less threatened by climate PA ecoregions. Moreover, increasing awareness of the
change than the tropical and southern hemisphere general public about the importance of conserving FB
ecoregions. Conversely, in the more developed areas is quite essential for the PA and particularly in the IN.

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Fig. 7 Main threats for

freshwater bivalves
recorded from the IUCN
Red List database by
ecoregions. Ecoregion
subdivisions adapted from
Graf & Cummings (2007)
and Haag (2010): NA
Nearctic, NT Neotropical,
PA Palaearctic, AF
Afrotropical, IN
Indotropical, AU
Australasian

Fig. 8 Research needs for freshwater bivalves recorded from Cummings (2007) and Haag (2010): NA Nearctic, NT Neotrop-
the IUCN Red List database by ecoregions. A All assessed ical, PA Palaearctic, AF Afrotropical, IN Indotropical, AU
species in the IUCN Red List; B data-deficient species in the Australasian. DD data-deficient species in the IUCN Red List
IUCN Red List. Ecoregion subdivisions adapted from Graf &

A special interest in species ex situ propagation and example, new remote sensing techniques like under-
reintroduction programs is exhibited for the NA, water video and side-scan sonars may help survey FB
emphasising the vast knowledge already accumulated populations, and identify more favourable habitats
for many species in the ecoregion (Fig. 9). (Powers et al., 2014; Mehler et al., 2016). Use of
Although many research gaps and conservation drones in semi-arid regions can aid in tracking and
needs have been identified in the last years, many identifying the remaining pools after droughts where
recent technological advances can provide us with mussels take refuge. These technologies and also the
new insights that are needed for FB research. For use of environmental DNA analyses may help

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Fig. 9 Conservation needs

for freshwater bivalves
extracted from the IUCN
Red List database by
ecoregions. Ecoregion
subdivisions adapted from
Graf & Cummings (2007)
and Haag (2010): NA
Nearctic, NT Neotropical,
PA Palaearctic, AF
Afrotropical, IN
Indotropical, AU
Australasian

gathering basic biological and ecological data on et al., 2015) may help to promote a better FB
distribution and abundance, which are still missing for representation within protected area networks.
many species (Stoeckle et al., 2016). More powerful
genetics and morphometric tools are also increasingly
available, for instance, new statistical tools for species The proceedings of the Second International
delimitation using molecular and/or other types of data Meeting on Biology and Conservation
such as morphometry and anatomical traits (e.g. of Freshwater Bivalves
Froufe et al., 2016b; Pfeiffer et al., 2016). These tools
are particularly important owing to the fact that many All the research and conservation needs summarised
species present hidden cryptic diversity (Froufe et al., above make the facilitation of cooperation among
2016b; Pfeiffer et al., 2016). Additionally, next- scientists from different countries and continents
generation sequencing is now allowing for quicker particularly important. For example, recent reviews
and less expensive robust phylogenies using methods published by multinational teams of scientists pro-
like whole-transcriptome and whole-mitogenome vided vital baseline information about FB on different
analyses with a wide range of markers (Guerra et al., continents (e.g. Pereira et al., 2014 for South America,
2017; Lopes-Lima et al., 2017c). Furthermore, using Walker et al., 2014 for Australia; Lopes-Lima et al.,
reduced genome representations or snip analyses, it is 2017a for Europe; Williams et al., 2017 for North
now possible to get more information on the phylo- America, and Zieritz et al., 2017 for East and
geographic patterns of species and on the definition of Southeast Asia). Additionally, intercontinental coop-
conservation units (Catchen et al., 2017; Desalle & erative research is also becoming increasingly com-
Amato, 2017). mon (see for example Zieritz et al., 2016; Lopes-Lima
While most of the global protected areas network is et al., 2017b). In order to discuss the current and future
aimed at protecting essentially terrestrial vertebrates, research challenges and needs, the Second Interna-
the identification of sites to conserve freshwater tional Meeting on Biology and Conservation of
vertebrates and invertebrates such as FB is also of Freshwater Bivalves was hosted by the Great Lakes
crucial importance (Darwall et al., 2011; Maceda- Centre at SUNY Buffalo State College in Buffalo,
Veiga et al., 2017). Using the IUCN Key Biodiversity New York, USA, from 4 to 8 October 2015, bringing
Areas (KBAs) network (IUCN, 2016) or new system- together over 80 scientists from 19 countries and four
atic conservation planning approaches (e.g. Hermoso continents (Europe, North America, South America
and Australia) (Burlakova et al., 2017).

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The present special issue in Hydrobiologia com- historical range and population size of the threatened
prises a total of 34 papers (including this introductory species Popenaias popeii (Karatayev et al., 2015). On
note) summarising some of the information presented a smaller scale, a study on the longitudinal variation in
in this meeting. These papers cover a wide variety of freshwater mussel assemblages within two rivers is
topics, from a review of ecosystem services provided presented by Chambers & Woolnough (2016), while
by freshwater mussels (Vaughn, 2017) to papers Dittman et al. (2017) evaluate the microhabitat and
describing the diversity patterns and conservation of biology of the poorly studied pea clam Sphaerium
Unionida in East and Southeast Asia (Zieritz et al., striatinum. Two papers assess the growth of M.
2017) as a result of international collaboration. Seven auricularia (Nakamura et al., 2017) and of juvenile
papers focus on different biological aspects of inva- freshwater pearl mussels M. margaritifera at the river
sive bivalve species, including diversity changes by scale (Černá et al., 2017). One paper assesses the shell
species substitution (Karatayev et al., 2017), physio- phenotypic plasticity of Unio crassus (Zaja˛c et al.,
logical aspects (Labecka & Domagala, 2016), disper- 2017). The influence of the flood pulses and of near-
sion (Collas et al., 2016), ecological effects on native bed hydrodynamics on freshwater mussels is evalu-
bivalve species (Ferreira-Rodrı́guez et al., 2016), low ated by Callil et al. (2017) and Sansom et al. (2017),
palatability to distinct predators (Castro et al., 2017), respectively. Finally, toxicology and archaeology are
metabolite emission suppression in zebra mussels represented by a study of the effects of polycyclic
exposed to predation stress (Antoł et al., 2017) and the aromatic hydrocarbons on unionid mussels (Archam-
use of a new sonar technology and underwater bault et al., 2017) and the conservation implications of
imagery analysis for the survey of FB in rivers freshwater mussel remains in a Texan river (Popejoy
(Mehler et al., 2016). Propagation as a conservation et al., 2016).
tool was the subject of three studies: one about an Conservation of FB requires urgent collaboration
improved method of in vitro culture of glochidia (Ma between scientists, managers, politicians and the
et al., 2016), one introducing short-term breeding of general public, in order to share knowledge and
the Endangered freshwater pearl mussel Margaritifera efforts. An example of this collaboration is the
margaritifera (Linnaeus, 1758) as a new technique for International Meeting on Biology and Conservation
the augmentation of declining populations (Moorkens, of Freshwater Bivalves, but more efforts are necessary
2017) and one revising the challenges in the conser- for the transfer of knowledge between scientists and
vation progress of Margaritifera auricularia (Spen- the general public in order to raise awareness about the
gler, 1793) (Prié et al., 2017). Six papers used importance of FB conservation. These efforts can
molecular tools to describe genetic structure or include, but not be limited to the increase in visibility
phylogeographic patterns of European (Feind et al., of FB conservation issues in the media, better
2017), North American (Hewitt et al., 2016; Mathias engagement with local communities and stakeholders
et al., 2016) and South American species (da Cruz (e.g. providing training and lifelong learning oppor-
Santos-Neto et al., 2017); to reveal the uncommon tunities like workshops for public, better information
doubly uniparental inheritance of mitochondria in a dissemination and accessibility of collaborative
European species (Soroka and Burzynski, 2017) and research even integrating participants from civil
the sequencing of transcriptomic resources for an society into surveys and research projects), publica-
invasive species (Soroka et al., 2017). The interaction tions and additions to national collections.
between mussels and their host fishes was addressed in
three papers that evaluate the effects of stress (Douda Acknowledgements We thank the Editor-in-Chief Prof. Dr.
Koen Martens for his help and support and the coordinator of
et al., 2016), cross-immunity (Chowdhury et al., 2017)
Springer Journals Editorial Office, Deepan Selvaraj, for his hard
and temperature (Schneider et al., 2017) on the work and assistance in the editorial process of this Special Issue.
reproduction of freshwater mussels. Three papers This work was supported by FCT—Foundation for Science and
describe distribution patterns with distinct spatial and Technology, Project 3599—Promote the Scientific Production
and Technological Development and Thematic 3599-PPCDT by
temporal scales: the population trends of Unionidae in
FEDER as part of the project FRESHCO: multiple implications
Romania (Sı̂rbu and Benedek, 2017), the distribution of invasive species on Freshwater Mussel co-extinction
of freshwater mussels and their host fishes in Texas processes (Contract: PTDC/AGRFOR/1627/2014). FCT also
(Dascher et al., 2017) and a study that reconstructs the supported MLL under Grant (SFRH/BD/115728/2016).

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