Pariselle - Euzet 2009

Systematic revision of dactylogyridean parasites
(Monogenea) from cichlid fishes in Africa, the

Levant and Madagascar
Antoine PARISELLE
Institut de Recherche pour le Développement,
Laboratoire d’écologie et de systématique, Centre de Belair,
BP 1386, 18524 Dakar (Sénégal)
antoine.pariselle@ird.fr
Louis EUZET
Station méditerranéenne de l’Environnement littoral, Université Montpellier II,
1 Quai de la Daurade, F-34200 Sète (France)
euzet@univ-montp2.fr
Pariselle A. & Euzet L. 2009. — Systematic revision of dactylogyridean parasites (Monogenea)

from cichlid fishes in Africa, the Levant and Madagascar. Zoosystema 31 (4) : 849-898.
ABSTRACT
Systematic revision is made of the six dactylogyridean genera (Monogenea)
KEY WORDS described on African, Levantine and Malagasy cichlid fishes: Urogyrus Bilong
Monogenea, Bilong, Birgi & Euzet, 1994 (including one species); Enterogyrus Paperna, 1963
Dactylogyridea, (including eight species); Insulacleidus Rakotofiringa & Euzet, 1983 (including
Urogyridae,
Ancyrocephalidae, three species); Onchobdella Paperna, 1968 (including eight species); Scutogyrus
Cichlidae, Pariselle & Euzet, 1995 (including six species) and Cichlidogyrus Paperna,
Africa,
Levant, 1960 (including 71 species). Validities of species are discussed. Diagnoses and
Madagascar. illustrated determination keys are given for all taxa.
RÉSUMÉ
Révision systématique des Monogènes Dactylogyridea parasites de poissons Cichlidae
en Afrique, au Proche-Orient et à Madagascar.
Les auteurs font la révision systématique des six genres de Monogènes Dac-
tylogyridea connus pour être parasites des poissons Cichlidae en Afrique,
MOTS CLÉS au Proche-Orient et à Madagascar : Urogyrus Bilong Bilong, Birgi & Euzet,
Monogenea, 1994 (comprenant une espèce) ; Enterogyrus Paperna, 1963 (comprenant huit
Dactylogyridea, espèces) ; Insulacleidus Rakotofiringa & Euzet, 1983 (comprenant trois espèces) ;
Urogyridae,
Ancyrocephalidae, Onchobdella Paperna, 1968 (comprenant huit espèces) ; Scutogyrus Pariselle &
Cichlidae, Euzet, 1995 (comprenant six espèces) et Cichlidogyrus Paperna, 1960 (comprenant
Afrique,
Proche-Orient, 71 espèces). La validité de toutes les espèces est discutée. Les diagnoses et des
Madagascar. clés d’identification illustrées sont données pour tous les taxons.
ZOOSYSTEMA • 2009 • 31 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 849
Pariselle A. & Euzet L.
INTRODUCTION CAS Institute of Parasitology, Czechoslovak Acad-

emy of Sciences, České Budějovice, Czech
Republic;
Today six genera, belonging to Dactylogyridea MNHN Muséum national d’Histoire naturelle, Paris;
Bychowsky, 1937, have been proposed on Afri- MRAC Musée royal de l’Afrique centrale, Tervuren.
can, Levantine and Malagasy cichlids (due to the
specificity of Monogenea, the description of a new
Quadriacanthus species [genus originally described SYSTEMATICS
from Siluriformes] from a tilapiine host by Paperna
[1973] – i.e. Q. tilapiae from Oreochromis esculentus Family UROGYRIDAE
(Graham, 1928) – in the Victoria Lake is indubitably Bilong Bilong, Birgi & Euzet, 1994
the result of a sampling error [mix of samples or
contamination]. Oreochromogyrus mymensinghi Genus Urogyrus
described from Bangladesh by Ferdousi & Chandra Bilong Bilong, Birgi & Euzet, 1994
[2002] on introduced Oreochromis niloticus, has
to be considerer as nomen inquirenda because the TYPE AND ONLY SPECIES. — Urogyrus cichlidarum Bilong
Bilong, Birgi & Euzet, 1994, by original designation.
description of this new genus and new species is
based on the misidentification of Cichlidogyrus DIAGNOSIS. — Urogyridae. Four pairs of cephalic glands.
larvae): one Urogyridae Bilong Bilong, Birgi & Two pairs of ocellae with lenses. Intestinal caeca without
Euzet, 1994 (Urogyrus Bilong Bilong, Birgi & Euzet, diverticula, joined posteriorly. One pair of ventral anchors;
right anchor abortive. One transverse bar (ventral).
1994) and five Ancyrocephalidae Bychowsky, 1937 Fourteen ventral uncinuli. Very large testis, median,
(Enterogyrus Paperna, 1963; Insulacleidus Rakotofir- inter-caecal. Vas deferens encircling left intestinal caeca.
inga & Euzet, 1983; Onchobdella Paperna, 1968; Seminal vesicle present. One prostatic reservoir. Male
Scutogyrus Pariselle & Euzet, 1995 and Cichlidogyrus copulatory complex with penis and accessory piece in two
Paperna, 1960). The large number of species described parts forming pincer shape. Ovary on the anterior right
side of testis. Straight vagina with dextrolateral opening.
(especially in Cichlidogyrus), the lack of revision Parasite in the urinary bladder of African cichlids.
(except Price 1967; Paperna 1979 or Pariselle 1996)
or of determination keys, and newly acquired genetic
data (see Pouyaud et al. 2006), lead us to undertake Urogyrus cichlidarum
a systematic revision of dactylogyridean parasites Bilong Bilong, Birgi & Euzet, 1994
from cichlid fishes of these regions.
Urogyrus cichlidarum Bilong Bilong, Birgi & Euzet,
1994: 562-564, figs 1-3.
MATERIAL AND METHODS TYPE MATERIAL. — Holotype: MNHN 114 HF Tk11.
Paratype: MNHN 115 HF Tk12.
Terminology used is that of Pariselle & Euzet (1995c): TYPE HOST. — Benitochromis batesii (Boulenger, 1901)
uncinulus instead of hooklets; for large hooks, as (first species cited by Bilong Bilong et al. [1994]).
“gripus” has not been accepted by parasitologists, TYPE LOCALITY. — Mbila, Nyong Basin, Cameroon.
we will use “anchor” instead of “hamulus”, because
ADDITIONAL HOSTS. — Parananochromis caudifasciatus
this term is close to the Latin anchora, and describes (Boulenger, 1913); Pungu maclareni (Trewavas, 1962);
well the working of these haptoral sclerites (see Stomatepia pindu Trewavas, Green & Corbet, 1972;
Lim et al. 2001). and Tilapia sp.
Host species validities and spellings are given ac- SITE. — Urinary bladder.
cording to FishBase (Froese & Pauly 2009): www.
fishbase.net. ADDITIONAL LOCALITIES. — Batouri, Kadei Basin,
Cameroon on Parananochromis caudifasciatus; Barombi
M’Bo Lake, Cameroon on Pungu maclareni and Sto-
ABBREVIATIONS matepia pindu; Sangmelima, Lobo River, Cameroon
BMNH The Natural History Museum, London; on Tilapia sp.
850 ZOOSYSTEMA • 2009 • 31 (4)

Dactylogyridean parasites (Monogenea) from cichlid fishes
Family ANCYROCEPHALIDAE Bychowsky, 1937 TYPE MATERIAL. — Holotype: Hebrew University, Depart-
ment of Parasitology, no specified number.
Genus Enterogyrus Paperna, 1963 TYPE HOST. — Of E. cichlidarum: Tilapia zillii (Gervais,
1848) (first species cited by Paperna [1963]).
TYPE SPECIES. — Enterogyrus cichlidarum Paperna, 1963, Of E. niloticus: Oreochromis niloticus (Linnaeus, 1758).
by original designation.
OTHER SPECIES INCLUDED. — Enterogyrus amieti Bilong TYPE LOCALITY. — Of E. cichlidarum: Rubin River,
Bilong, Euzet & Birgi, 1996; E. barombiensis Bilong Israel.
Bilong, Birgi & Euzet, 1991; E. coronatus Pariselle, Of E. niloticus: Barher Mouise, Nile River, Egypt.
Lambert & Euzet, 1991; E. crassus Bilong Bilong, Birgi & ADDITIONAL HOSTS. — Sarotherodon galilaeus sanagaensis
Euzet, 1996; E. foratus Pariselle, Lambert & Euzet, 1991; (Thys van den Audenaerde, 1966) and Tilapia nyongana
E. malmbergi Bilong Bilong, 1988; E. melenensis Bilong Thys van den Audenaerde, 1971 (Bilong Bilong et al.
Bilong, Birgi & Lambert, 1989. 1996).
DIAGNOSIS. — Ancyrocephalidae. Pear-shaped body
enlarged posteriorly; thick tegument transversely striated SITE. — Stomach.
(except on haptor). Four ocellae, members of anterior pair ADDITIONAL LOCALITIES. — Jordan and coastal systems,
wider apart and smaller than those of posterior, anterior Israel (Paperna 1979); South of Cameroon on O. niloticus
with or without lenses. Pharynx muscular; short oesophagus (Bilong Bilong et al. 1989); Sakbayémé, Sanaga Basin,
followed by two lateral intestinal caeca without diverticula, Cameroon on Sarotherodon galilaeus sanagaensis and So’o,
joined posteriorly. Haptor cup-shaped, opening ventrally Nyong Basin, Cameroon on Tilapia nyongana (Bilong
or in two parts: one bulbous and one peduncular shaped. Bilong et al. 1996).
Two unequal pairs of anchors. One ventral transverse
bar, straight or V-shaped. Fourteen equal uncinuli, each
with points orientated ventrally (haptor cup-shaped) or REMARKS
anteriorly (haptor bulbous and peduncular shaped). Testis at The comparison by Khidr (1990) of Enterogyrus species
level of junction of intestinal caeca. Vas deferens encircling specimens from Tilapia zillii and Oreochromis niloticus
left intestinal caeca, widening to form a seminal vesicle, in Egypt revealed no differences and led this author
continuing as narrow duct to base of penis. Sub-median to synonymise E. niloticus with E. cichlidarum.
tubular and spirally coiled (clockwise) penis, enlarged at
base; accessory piece absent. Ovary median pre-testicular,
ovoid, sometimes U-shaped. Vagina not observed. Stomach
parasite of Levantine and African cichlid fishes. Enterogyrus amieti
Bilong Bilong, Euzet & Birgi, 1996
REMARKS
Two species are not considered in this study because Enterogyrus amieti Bilong Bilong, Euzet & Birgi, 1996:
38, 39, figs 4, 5.
they were described from the stomach of Etroplus
suratensis (Block, 1790) in Asia. These species (En- TYPE MATERIAL. — Holotype: MNHN 473 HF Tg72.
terogyrus globodiscus (Kulkarni, 1969) and E. papernai Paratype: MRAC 37.362.
Gussev & Fernando, 1973) are characterised by TYPE HOST. — Sarotherodon galilaeus sanagaensis (Thys
having two transverse bars (vs. only one for African van den Audenaerde, 1966).
and Levantine species), according to Paperna (1979)
we think that this difference will be sufficient to TYPE LOCALITY. — Sakbayémé, Sanaga Basin, Cam-
eroon.
justify splitting Enterogyrus into two genera.
SITE. — Stomach.
Enterogyrus cichlidarum Paperna, 1963

Enterogyrus barombiensis
Enterogyrus cichlidarum Paperna, 1963: 183-186, figs Bilong Bilong, Birgi & Euzet, 1991
1-3. — Bilong Bilong et al. 1989: 104, figs 4, 5.
Enterogyrus niloticus Eid & Negm, 1987: 79-84, Enterogyrus barombiensis Bilong Bilong, Birgi & Euzet,
figs 1-4. — Khidr 1990: 741, 742. 1991: 105, 106, figs 2, 3.
ZOOSYSTEMA • 2009 • 31 (4) 851

TYPE MATERIAL. — Holotype: MNHN 111HF Tk7. TYPE LOCALITY. — Mouth of the Casamance River,
Paratype: MNHN 112HF Tk8. Senegal.
TYPE HOST. — Pungu maclareni (Trewavas, 1962). ADDITIONAL HOST. — Sarotherodon melanotheron melan-
otheron Rüppel, 1852.
ADDITIONAL HOSTS. — Stomatepia pindu Trewavas,
1972, Konia eisentrauti (Trewavas, 1962). SITE. — Stomach.
TYPE LOCALITY. — Barombi Mbo Crater Lake, Cam- ADDITIONAL LOCALITY. — Layo station, Ebrié Lagoon,
eroon. Côte d’Ivoire.
SITE. — Stomach.
Enterogyrus malmbergi Bilong Bilong, 1988
Enterogyrus coronatus
Pariselle, Lambert & Euzet, 1991 Enterogyrus malmbergi Bilong Bilong, 1988: 52, 53,
figs 1, 2.
Enterogyrus coronatus Pariselle, Lambert & Euzet, 1991: TYPE MATERIAL. — No type material mentioned in the
213-216, figs 1D2, 4, 5A. original description.
TYPE MATERIAL. — Holotype: MNHN 89 HF Tk1. TYPE HOST. — Oreochromis niloticus (Linnaeus, 1758).
Paratypes: MNHN 89 HF Tk2; BMNH 1990.12.7.2.
TYPE LOCALITY. — Edebda, Sanaga River, Cameroon.
TYPE HOST. — Tilapia guineensis (Bleeker, 1862).
SITE. — Stomach.
TYPE LOCALITY. — Layo station, Ebrié Lagoon, Côte
d’Ivoire.
SITE. — Stomach. Enterogyrus melenensis
Bilong Bilong, Birgi & Lambert, 1989
Enterogyrus crassus
Bilong Bilong, Euzet & Birgi, 1996 Enterogyrus melenensis Bilong Bilong, Birgi & Lambert,
1989: 101-104, figs 2, 3.
Enterogyrus crassus Bilong Bilong, Euzet & Birgi, 1996: TYPE MATERIAL. — Holotype: MRAC 37.226.
38, figs 2, 3. Paratype: MRAC 37.227.
TYPE MATERIAL. — Holotype: MNHN 472 HF Tg72. TYPE HOST. — Hemichromis fasciatus Peters, 1857.
Paratype: MRAC 37.363.
TYPE LOCALITY. — Melen, Yaoundé, Cameroon.
TYPE HOST. — Tilapia nyongana Thys van den Audenaerde,
1971. SITE. — Stomach.
TYPE LOCALITY. — So’o, Nyong Basin, Cameroon. ADDITIONAL LOCALITIES. — Nyong, Sanaga and Lobé
Basins, Cameroun.
SITE. — Stomach.
“Enterogyrus hemihaplochromii”
Enterogyrus foratus
Pariselle, Lambert & Euzet, 1991 “Enterogyrus hemihaplochromii” – Bender 1979.
Enterogyrus foratus Pariselle, Lambert & Euzet, 1991: TYPE MATERIAL. — No type material mentioned in the
212, 213, figs 1A, 1D1, 2, 5B. original description.
TYPE MATERIAL. — Holotype: MNHN 90 HF Tk3. HOST. — Pseudocrenilabrus multicolor Schoeller, 1903.
Paratypes: MNHN 90 HF Tk4; BMNH 1990.12.7.1.
SITE. — Stomach.
TYPE HOST. — Sarotherodon melanotheron heudelotii
(Duméril, 1859). LOCALITY. — East Africa.
852 ZOOSYSTEMA • 2009 • 31 (4)

REMARKS TYPE HOST. — Paratilapia polleni Bleeker, 1868.

“Enterogyrus hemihaplochromii” was described in TYPE LOCALITY. — No type locality mentionned in the
an unpublished thesis; this name is then a nomen original description.
nudum.
SITE. — Gills.
Enterogyrus sp. ADDITIONAL LOCALITIES. — Near Majunga, West of

Madagascar; Katsepy and Ampizoroa Lakes, Madagascar;
Enterogyrus sp. – Cone et al. 1987: 312, 313, figs 1, Périmet and Ivoloina aquaculture stations, Tamatave
2, 3. Province, Madagascar.
HOST. — Captive Pomacanthus paru (Bloch, 1787).
LOCALITY. — Caribbean Sea. Insulacleidus paretropli
SITE. — Foregut. Rakotofiringa & Euzet, 1983
REMARKS Insulacleidus paretropli Rakotofiringa & Euzet, 1983:

111, 112, figs 8-10.
According to Pariselle et al. (1991) this species,
due to its morphology and anatomy, does not TYPE MATERIAL. — No type material mentioned in the
belong to Enterogyrus, but rather to Diplectanotrema original description.
Johnston & Tiegs, 1922 or Pseudempleurosoma TYPE HOST. — Paretroplus polyactis Bleeker, 1878.
Yamaguti, 1965 which individuals are located in
the pharynx of marine fishes. TYPE LOCALITY. — Ambila, Antananarivo, Madagas-
car.
Genus Insulacleidus Rakotofiringa & Euzet, 1983 SITE. — Gills.
TYPE SPECIES. — Insulacleidus paratilapiae Rakotofiringa &

Euzet, 1983, by original designation. Insulacleidus ptychochromidis
OTHER SPECIES INCLUDED. — Insulacleidus ptychochro- Rakotofiringa & Euzet, 1983
midis Rakotofiringa & Euzet, 1983 and I. paretropli
Rakotofiringa & Euzet, 1983. Insulacleidus ptychochromidis Rakotofiringa & Euzet,
1983: 110, 111, figs 5-7.
DIAGNOSIS. — Ancyrocephalidae. Three pairs of cephalic
glands. Two pairs of ocellae. Intestinal caeca without TYPE MATERIAL. — No type material mentioned in the
diverticula, joined posteriorly. Two pairs of anchor, one original description.
dorsal and one ventral. Two transverse bars, one dorsal
one ventral. Fourteen uncinuli. Testis median posterior TYPE HOST. — Ptychochromis oligacanthus (Bleeker,
to the body. Vas deferens encircling left intestinal caeca. 1868).
Seminal vesicle present. One prostatic reservoir. Male TYPE LOCALITY. — Ambila, Lemaitro, Tamatave Province,
copulatory complex with penis and accessory piece. Madagascar.
Pre-testicular median ovary. Vagina with lateral dextral
opening. Seminal receptacle present. Gill parasite on SITE. — Gills.
Malagasy cichlid fishes.
Insulacleidus paratilapiae Genus Onchobdella Paperna, 1968

Rakotofiringa & Euzet, 1983
TYPE SPECIES. — Onchobdella voltensis Paperna, 1968,
Insulacleidus paratilapiae Rakotofiringa & Euzet, 1983: by original designation.
107-110, figs 1-4.
OTHER SPECIES INCLUDED. — Onchobdella aframae
TYPE MATERIAL. — No type material mentioned in the Paperna, 1968; O. bopeleti Bilong Bilong & Euzet, 1995;
original description. O. krachii Paperna, 1968; O. melissa Pariselle & Euzet,
ZOOSYSTEMA • 2009 • 31 (4) 853

1995; O. pterigyalis Paperna, 1968; O. silverai Pariselle & REMARKS

Euzet, 1995; O. spirocirra Paperna, 1968. Dossou (1973), in his PhD Thesis on the fish parasites
DIAGNOSIS (according to Pariselle & Euzet [1995a]). — from South Benin, re-described O. aframae and
Ancyrocephalidae. Three pairs of cephalic glands. Two indicated a new host: Pelmatochromis caudifasciatus
posterior ocellae with lenses, two small anterior ocellae. Boulenger, 1913 (the species determination of this
Intestinal caeca without diverticula, joined posteriorly. new host remains uncertain; in fact Parananochromis
Two pairs of anchors: one large dorsal, one smaller
ventral. Three transverse bars: one dorsal, two ventral
caudifasciatus (Boulenger, 1913) is limited to South-
well separated. Fourteen uncinuli of same length. Testis ern Cameroon, the Northern part of the Congo and
posterior inter-caecal. Vas deferens encircling left intestinal the Rio Muni; this distribution excludes the Benin
caeca. Seminal vesicle present. One prostatic reservoir. rivers [see Froese & Pauly 2009]).
Male copulatory complex with penis and accessory
piece. Median pre-testicular ovary. Vaginal opening
lateral, dextral, sclerotised. Gill parasites on African Onchobdella bopeleti
cichlid fishes. Bilong Bilong & Euzet, 1995
Onchobdella bopeleti Bilong Bilong & Euzet, 1995:
Onchobdella voltensis Paperna, 1968 254-256, figs 2-4.
TYPE MATERIAL. — Holotype: MNHN 471 HF Tg71.
Onchobdella voltensis Paperna, 1968: 201, fig. 1. Paratypes: MNHN 471 HF Tg71; BMNH 1994.5.5.4;
TYPE MATERIAL. — Holotype: BMNH 1968.9.4.5. MRAC 37.367 and 37.368.
Paratype: BMNH 1968.9.4.6. TYPE HOST. — Hemichromis fasciatus Peters, 1857.
TYPE HOST. — Hemichromis fasciatus Peters, 1857. TYPE LOCALITY. — No type locality mentionned in the
original description.
TYPE LOCALITY. — Afram sector, Volta Lake, Ghana.
SITE. — Gills.
ADDITIONAL HOST. — Hemichromis bimaculatus Gill,
1862. ADDITIONAL LOCALITIES. — Kribi, Ejambwé River,
Cameroon; Bamako, Niger River, Mali; Senegal, Gambia,
SITE. — Gills. Mali and Côte d’Ivoire (Pariselle & Euzet 2004).
ADDITIONAL LOCALITIES. — Kete Krachi, Volta Lake
and Adutor Lagoon Volta River Delta, Ghana on H. Onchobdella krachii Paperna, 1968
bimaculatus; Benin and Cameroon on Hemichromis
fasciatus (Dossou & Birgi, 1984); Senegal, Gambia, Mali Onchobdella krachii Paperna, 1968: 205, figs 3d, 4c-e.
and Côte d’Ivoire on Hemichromis fasciatus (Pariselle &
Euzet 2004). TYPE MATERIAL. — BMNH 1968.9.4.4; MRAC MT
35.574 (Paperna 1979).
TYPE HOST. — Chromidotilapia guentheri (Sauvage,
Onchobdella aframae Paperna, 1968 1882).
TYPE LOCALITY. — Kpandu and Kete Krachi, Volta
Onchobdella aframae Paperna, 1968: 205: fig. 4f-h. Lake, Ghana.
TYPE MATERIAL. — Holotype: BMNH 1968.9.4.3. SITE. — Gills.
TYPE HOST. — Hemichromis fasciatus Peters, 1857.
TYPE LOCALITY. — Afram sector, Volta Lake, Ghana. Onchobdella melissa Pariselle & Euzet, 1995
SITE. — Gills. Onchobdella melissa Pariselle & Euzet, 1995a: 206,
fig. 4.
ADDITIONAL LOCALITIES. — Benin and Cameroon (Dos-
sou & Birgi 1984); Senegal, Gambia, Mali and Côte TYPE MATERIAL. — Holotype: MNHN 466 HF Tg65.
d’Ivoire (Pariselle & Euzet 2004). Paratypes: MNHN 467 HF Tg66; BMNH 1994 5.5.2;
854 ZOOSYSTEMA • 2009 • 31 (4)

MRAC 37.365 A and 37.366 B. Onchobdella spirocirra Paperna, 1968

TYPE HOST. — Pelmatochromis buettikoferi (Steindachner, Onchobdella spirocirra Paperna, 1968: 201-203, figs 2,
1894). 3a.
TYPE LOCALITY. — Koleya, Konkouré River, Guinea. TYPE MATERIAL. — Holotype: BMNH 1968.9.4.2.
SITE. — Gills. TYPE HOST. — Hemichromis bimaculatus Gill, 1862.
TYPE LOCALITY. — No type locality mentionned in the
Onchobdella pterigyalis Paperna, 1968 original description.
Onchobdella pterigyalis Paperna, 1968: 203-205, figs SITE. — Gills.
3c, e, 4a, b.
ADDITIONAL LOCALITY. — Afram sector, Volta Lake,
TYPE MATERIAL. — Holotype: BMNH 1968.9.4.1. Ghana.
TYPE HOST. — Hemichromis bimaculatus Gill, 1862.
Genus Scutogyrus Pariselle & Euzet, 1995
T YPE LOCALITY . — Nunguia dam, East of Accra,
Ghana. TYPE SPECIES. — Scutogyrus minus (Dossou, 1982), by
SITE. — Fins. subsequent designation (Pariselle & Euzet 1995b).
ADDITIONAL SITE. — Gills. OTHER SPECIES INCLUDED. — Scutogyrus bailloni Pariselle

& Euzet, 1995; S. chikhii Pariselle & Euzet, 1995; S.
ADDITIONAL LOCALITY. — Mamahuma stream, East of ecoutini Pariselle & Euzet, 1995; S. gravivaginus (Paperna
Accra, Ghana & Thurston, 1969); S. longicornis (Paperna & Thurston,
1969).
REMARKS
DIAGNOSIS. — Ancyrocephalidae. Three pairs of cephalic
Dossou (1973), in his PhD thesis on the fish parasites glands. Two posterior ocellae with lenses. Two small
from South Benin, re-described O. pterigyalis (as anterior ocellae, sometimes absent. Intestinal caeca
O. pterygialis) and indicated a new host: Pelmato- without diverticula, joined posteriorly. Two pairs of
chromis caudifasciatus Boulenger, 1913 (the species anchors, one dorsal and one ventral. Dorsal transverse
determination of this new host remains uncertain; in bar highly arched, enlarged laterally, winged, having in
its median portion two very long auricles hollow at their
fact, it has been synonymised with Parananochromis bases. Ventral transverse bar arched, rigid, supporting
caudifasciatus (Boulenger, 1913) which is limited one large, thin oval plate marked by fan-shaped median
to Southern Cameroon, the Northern part of the thickenings. Fourteen uncinuli. Testis median, posterior
Congo and the Rio Muni; this distribution excludes to the body. Vas deferens dextral, not encircling intestinal
the Benin Rivers). caeca. Seminal vesicle present. One prostatic reservoir.
Male copulatory complex with penis and accessory piece.
Median pre-testicular ovary. Vaginal opening sub-lateral,
Onchobdella silverai Pariselle & Euzet, 1995 dextral; vagina tubular, sclerotised. Seminal receptacle
present. Gill parasites of African cichlid fishes.
Onchobdella silverai Pariselle & Euzet, 1995a: 206-208,
fig. 5.
Scutogyrus minus (Dossou, 1982)
Paratypes: MNHN 467 HF Tg66; BMNH 1994 5.5.3; Cichlidogyrus longicornis minus Dossou, 1982: 309-312,
MRAC 37.365 B and 37.366 A. figs 19-23.
TYPE HOST. — Pelmatochromis buettikoferi (Steindachner, Cichlidogyrus minus – Pariselle & Euzet 1995b: 158-161,
1894). figs 2, 3.
TYPE LOCALITY. — Koleya, Konkouré River, Guinea. Scutogyrus minus – Pariselle & Euzet 1995b.
SITE. — Gills. TYPE MATERIAL. — Holotype: MNHN Tj141.
ZOOSYSTEMA • 2009 • 31 (4) 855

Paratype: MNHN Tj142. of a sampling error [mix of samples or contamination] or

of host identification); Oreochromis mortimeri (Trewavas,
OTHER MATERIAL. — MNHN 460 HF Tg55; MNHN 1966) (Douëllou 1993); O. mossambicus (Ferdousi &
460 HF Tg56 and Tg57; BMNH 1994.4.7.1; MRAC Chandra 2002).
37.357 (Pariselle & Euzet 1995b).
SITE. — Gills.
TYPE HOST. — Sarotherodon melanotheron Rüppel,
1852. ADDITIONAL LOCALITIES. — Kete Kratchi, Volta Lake,
Ghana (Paperna 1969); Cairo, Nile River, Egypt on
TYPE LOCALITY. — Ouémé and Couffo Rivers, Benin. O. niloticus (Ergens 1981); Kariba Lake, Zimbabwe on
O. mortimeri (Douëllou 1993). Senegal and Côte d’Ivoire
SITE. — Gills. on O. niloticus (Pariselle & Euzet 1995b). Bangladesh in
ADDITIONAL LOCALITIES. — Guinea and Côte d’Ivoire BAU and BFRI fish farm ponds on introduced O. niloticus
(Pariselle & Euzet 1995b). and O. mossambicus (Ferdousi & Chandra 2002).
REMARKS
REMARKS Dossou (1982) suggested that C. longicornis longi-
Dossou (1982) suggested that Cichlidogyrus longicornis could be raised to species level, this was done
cornis minus could be raised to species level. Douëllou by Douëllou (1993).
(1993) indicated that this sub-species is uncertain
and had to be invalidated. Pariselle & Euzet (1995b)
did not agree with Douëllou. Scutogyrus gravivaginus
(Paperna & Thurston, 1969)
Scutogyrus longicornis Cichlidogyrus longicornis gravivaginus Paperna & Thurston,

1969: 20-23, fig. 3c, e-g.
(Paperna & Thurston, 1969)
Cichlidogyrus gravivaginus – Douëllou 1993: 172, 173,
Cichlidogyrus longicornis longicornis Paperna & Thurston, fig. 73.
1969: 20-22, fig. 3a, b, d.
Scutogyrus gravivaginus – Pariselle & Euzet 1995b: 163-
Cichlidogyrus longicornis – Douëllou 1993: 170-172, 165, figs 6, 7.
fig. 71, 72. TYPE MATERIAL. — Holotype: MRAC M.T. 35.932
(Paperna 1979).
Scutogyrus longicornis – Pariselle & Euzet 1995b: 161-
Paratypes: private collection of Paperna.
163, figs 4, 5.
OTHER MATERIAL . — MNHN 129 HF (Douëllou
Actinocleidus muelleri Ferdousi & Chandra, 2002: 56, 1993).
57, fig. 4.
TYPE HOST. — Oreochromis leucostictus (Trewavas, 1933).
TYPE MATERIAL. — Holotype from Sarotherodon galilaeus:
MRAC M.T. 35.931 (Paperna 1979). TYPE LOCALITY. — Jinja and Albert Lake, Uganda.
Paratypes: private collection of Dr Paperna. ADDITIONAL HOSTS. — Oreochromis variabilis (Boulenger,
OTHER MATERIAL. — MNHN 128 HF (Douëllou 1993); 1906) (Paperna 1979); Oreochromis mortimeri (Trewavas,
MNHN 461 HF Tg58 and Tg59; BMNH 1994.4.7.2; 1966) (Douëllou 1993).
MRAC 37.358 (Pariselle & Euzet 1995b). SITE. — Gills.
T YPE HOST . — Sarotherodon galilaeus (Linnaeus, ADDITIONAL LOCALITY. — Kariba Lake, Zimbabwe
1758). on Oreochromis mortimeri (Trewavas, 1966) (Douëllou
1993).
TYPE LOCALITY. — Lakes Georges and Albert, Uganda.
REMARKS
ADDITIONAL HOSTS. — Oreochromis niloticus (Linnaeus,
1758) and Tilapia zillii Gervais, 1848 (Paperna 1979) (for Dossou (1982) suggested that C. longicornis gravivagi-
T. zillii: as species belonging to Scutogyrus are only found nus could be raised to species level, this was done
on mouthbreeder cichlid fishes, it is probably the result by Douëllou (1993).
856 ZOOSYSTEMA • 2009 • 31 (4)

Scutogyrus bailloni Pariselle & Euzet, 1995 C. arfii Pariselle & Euzet, 1995; C. berradae Pariselle &
Euzet, 2003; C. berrebii Pariselle & Euzet, 1994; C. bi-
Scutogyrus bailloni Pariselle & Euzet, 1995b: 165, 166, furcatus Paperna, 1960; C. bilongi Pariselle & Euzet,
figs 8, 9. 1996; C. bonhommei Pariselle & Euzet, 1998; C. bouvii
Pariselle & Euzet, 1997; C. bychowskii (Markevich, 1934);
TYPE MATERIAL. — Holotype: MNHN 462 HF Tg60. C. cirratus Paperna, 1964; C. cubitus Dossou, 1982;
Paratypes: MNHN 462 HF Tg61; BMNH 1994.4.7.3; C. dageti Dossou & Birgi, 1984; C. digitatus Dossou,
MRAC 37.359. 1982; C. dionchus Paperna, 1968; C. dossoui Douëllou,
1993; C. douellouae Pariselle, Bilong Bilong & Euzet,
TYPE HOST. — Sarotherodon galilaeus (Linnaeus, 1758). 2003; C. ergensi Dossou, 1982; C. euzeti Dossou & Birgi,
1984; C. falcifer Dossou & Birgi, 1984; C. flexicolpos
TYPE LOCALITY. — W National Park, Mékrou River, Pariselle & Euzet, 1995; C. fontanai Pariselle & Euzet,
Niger. 1997; C. gallus Pariselle & Euzet, 1995; C. giostrai Pariselle,
SITE. — Gills. Bilong Bilong & Euzet, 2003; C. guirali Pariselle &
Euzet, 1997; C. halinus Paperna, 1969; C. halli (Price &
Kirk, 1967); C. haplochromii Paperna & Thurston, 1969;
Scutogyrus ecoutini Pariselle & Euzet, 1995 C. hemi Pariselle & Euzet, 1998; C. inconsultans Birgi &
Lambert, 1986; C. karibae Douëllou, 1993; C. kothiasi
Scutogyrus ecoutini Pariselle & Euzet, 1995b: 167, 168, Pariselle & Euzet, 1994; C. kouassii N’Douba, Thys
figs 10, 11. van den Audenaerde & Pariselle, 1997; C. lagoonaris
Paperna, 1969; C. legendrei Pariselle & Euzet, 2003;
TYPE MATERIAL. — Holotype: MNHN 463 HF Tg62. C. lemoallei Pariselle & Euzet, 2003; C. levequei Pariselle &
Paratypes: MNHN 463 HF Tg63; BMNH 1994.4.7.4; Euzet, 1996; C. longicirrus Paperna, 1965; C. longipenis
MRAC 37.360. Paperna & Thurston, 1969; C. louipaysani Pariselle &
Euzet, 1994; C. microscutus Pariselle & Euzet, 1996;
TYPE HOST. — Sarotherodon occidentalis (Daget, 1962). C. nandidae Birgi & Lambert, 1986; C. njinei Pariselle,
Bilong Bilong & Euzet, 2003; C. nuniezi Pariselle & Euzet,
TYPE LOCALITY. — La Ramié, Bourouma River, Guinea. 1998; C. ornatus Pariselle & Euzet, 1995; C. ouedraogoi
Pariselle & Euzet, 1996; C. paganoi Pariselle & Euzet,
SITE. — Gills. 1997; C. papernastrema Price, Peebles & Bamford, 1969;
C. philander Douëllou, 1993; C. pouyaudi Pariselle &
Euzet, 1994; C. quaestio Douëllou, 1993; C. reversati
Scutogyrus chikhii Pariselle & Euzet, 1995 Pariselle & Euzet, 2003; C. rognoni Pariselle, Bilong
Bilong & Euzet, 2003; C. sanjeani Pariselle & Euzet,
Scutogyrus chikhii Pariselle & Euzet, 1995b: 171, 172, 1997; C. sanseoi Pariselle & Euzet, 2004; C. sclerosus
figs 12, 13. Paperna & Thurston, 1969; C. slembroucki Pariselle &
TYPE MATERIAL. — Holotype: MNHN 464 HF Tg64. Euzet, 1998; C. testificatus Dossou, 1982; C. teugelsi
Paratypes: BMNH 1994.4.7.5; MRAC 37.361. Pariselle & Euzet, 2004; C. thurstonae Ergens, 1981;
C. tiberianus Paperna, 1960; C. tilapiae Paperna, 1960;
TYPE HOST. — Oreochromis mossambicus (Peters, 1852). C. vexus Pariselle & Euzet, 1995; C. yanni Pariselle &
Euzet, 1996; C. zambezensis Douëllou, 1993.
TYPE LOCALITY. — Cayo Lake, Congo.
DIAGNOSIS (according to Pariselle & Euzet [1997]). —
SITE. — Gills. Ancyrocephalidae. Three pairs of cephalic glands. Two
posterior ocellae with crystalline lenses. Two small in-
consistent anterior ocellae. Intestinal caeca unbranched,
Genus Cichlidogyrus Paperna, 1960 joined posteriorly. Two pairs of anchors, one dorsal and
one ventral. Two transverse bars, dorsal with two auricles,
TYPE SPECIES. — Cichlidogyrus arthracanthus Paperna, ventral V-shaped. Fourteen uncinuli. Median posterior
1960, by original designation. testis. Vas deferens on right side, not encircling intestinal
caecum. Seminal vesicle present. One prostatic reservoir.
OTHER SPECIES INCLUDED. — Cichlidogyrus acerbus Male copulatory complex with penis and accessory piece.
Dossou, 1982; C. aegypticus Ergens, 1981; C. agnesi Median pre-testicular ovary. Sub-median vaginal dextral
Pariselle & Euzet, 1994; C. albareti Pariselle & Euzet, opening. Vagina sclerotised or not. Seminal receptacle
1998; C. amieti Birgi & Euzet, 1983; C. amphoratus present. Gill parasites of African Cichlidae, Nandidae
Pariselle & Euzet, 1995; C. anthemocolpos Dossou, 1982; and Cyprinodontidae.
ZOOSYSTEMA • 2009 • 31 (4) 857

Cichlidogyrus arthracanthus Paperna, 1960 TYPE HOST. — Tilapia zillii (Gervais, 1848).
Cichlidogyrus arthracanthus Paperna, 1960: 3-5, figs 1, 2. TYPE LOCALITY. — Cairo, Nile River, Egypt.
TYPE MATERIAL. — No type material mentioned in the ADDITIONAL HOSTS. — Tilapia guineensis (Bleeker,
original description. 1862) (Pariselle & Euzet 1995c), Tilapia dageti Thys
van den Audenaerde, 1971, Tilapia louka Thys van den
TYPE HOST. — Tilapia zillii (Gervais, 1848). Audenaerde, 1969 and Tilapia walteri Thys van den
Audenaerde, 1968 (Pariselle & Euzet 1996).
TYPE LOCALITY. — Fish culture research station, Dor,
Israel. SITE. — Gills.
ADDITIONAL HOSTS. — Tristramella simonis (Günther, ADDITIONAL LOCALITIES. — Senegal, Guinea, and Congo
1864) and T. sacra (Günther, 1865); Tilapia guineensis on T. guineensis (Pariselle & Euzet 1995c); Côte d’Ivoire
(Bleeker, 1862) (Pariselle & Euzet 1995c). and Burkina Faso on T. zillii, Côte d’Ivoire on T. dageti;
Guinea on T. louka and T. walteri (Pariselle & Euzet
SITE. — Gills. 1996).
ADDITIONAL LOCALITIES. — Western shore, Sea of Galilee, REMARKS
Israel. Ghana (Paperna 1965, 1979), Uganda (Paperna & Dossou (1982) described this species in Benin, but
Thurston 1969), Egypt (Ergens 1981), Senegal, Guinea,
Ivory Coast and Congo (Pariselle & Euzet 1995c). confused the identification with C. gallus Pariselle &
Euzet, 1995 (see below under Cichlidogyrus gallus).
The specimens of C. aegypticus from T. louka and
T. walteri show a shape of the vagina slightly dif-
Cichlidogyrus acerbus Dossou, 1982 ferent from that described on T. zillii, which was
Cichlidogyrus acerbus Dossou, 1982: 314-316, figs 26-
not deemed sufficient for Pariselle & Euzet (1996)
28. to create a new species.
TYPE MATERIAL. — Holotype: MNHN Tj135.
Paratype: MNHN Tj136. Cichlidogyrus agnesi Pariselle & Euzet, 1995
TYPE HOST. — Sarotherodon melanotheron melanotheron Cichlidogyrus agnesi Pariselle & Euzet, 1995c: 189,
Rüppell, 1852. 190, fig. 2.
TYPE LOCALITY. — Ouémé and Couffo, Benin. TYPE MATERIAL. — Holotype: MNHN 220 HF Tk22.
Paratypes: MNHN 220 HF Tk23, BMNH 1993.5.14.7,
ADDITIONAL HOSTS. — Sarotherodon melanotheron MRAC 37.338.
heudelotii (Duméril, 1859) (Pariselle 1995).
SITE. — Gills.
TYPE LOCALITY. — Tiassalé, Bandama River, Côte
ADDITIONAL LOCALITIES. — Ebrié lagoon, Bakré Lake d’Ivoire.
and Samakro, Côte d’Ivoire on type host (Pariselle 1995).
Kogon River, Koba, Konkouré River, Forecariah, Guinea; SITE. — Gills.
Saint Louis, Retba, Somone Lagoon, Ziguinchor, Senegal;
Banjul, Gambia on S. m. heudelotii (Pariselle 1995). ADDITIONAL LOCALITIES. — Senegal, Guinea and Congo
on type host [Pariselle & Euzet 1996].
Cichlidogyrus aegypticus Ergens, 1981

Cichlidogyrus albareti Pariselle & Euzet, 1998
Cichlidogyrus aegypticus Ergens, 1981: 211, fig. 7.
Cichlidogyrus albareti Pariselle & Euzet, 1998: 275,
TYPE MATERIAL. — Originally deposited at the CAS, 276, fig. 2.
with no mention of reference number.
Voucher specimen: MNHN 484 HF Tg96 and Tg97 TYPE MATERIAL. — Holotype: MNHN 543 HF Tk73.
(Pariselle & Euzet 1996). Paratypes: MNHN 543 HF Tk74, BMNH 1997.1.30.1,
858 ZOOSYSTEMA • 2009 • 31 (4)

MRAC 37.408. Cichlidogyrus anthemocolpos Dossou, 1982

TYPE HOST. — Tilapia brevimanus Boulenger, 1911. Cichlidogyrus anthemocolpos Dossou, 1982: 307-309,
T YPE LOCALITY . — La Ramié, Bourouma River, figs 16-18.
Guinea. TYPE MATERIAL. — Holotype: MNHN Tj133.
SITE. — Gills. Paratype: MNHN Tj134.
ADDITIONAL LOCALITY. — Ndyarendi, Kogon River, TYPE HOST. — Tilapia zillii (Gervais, 1848).
Guinea. TYPE LOCALITY. — Ouémé, Benin.
SITE. — Gills.
Cichlidogyrus amieti Birgi & Euzet, 1983
Cichlidogyrus arfii Pariselle & Euzet, 1995
Cichlidogyrus amieti Birgi & Euzet, 1983: 101-104,
figs 1-3. Cichlidogyrus arfii Pariselle & Euzet, 1995a: 204, 205,
fig. 3.
TYPE MATERIAL. — No type material mentioned in the
original description. Cichlidogyrus dionchus Paperna, 1968: 91, pl. 1, fig. 7.
TYPE HOST. — Aphyosemion cameronense (Boulenger, TYPE MATERIAL. — Holotype: MNHN 470 HF Tg69.
1903) (first species cited by Birgi & Euzet [1983]) Paratypes: MNHN 470 HF Tg70, BMNH 1994.5.5.1,
(Nothobranchiinae Radda & Pürzl, 1981). MRAC 37.364.
TYPE LOCALITY. — Kala, Zamacoe, Cameroon. TYPE HOST. — Pelmatochromis buettikoferi (Steindachner,
1894).
ADDITIONAL HOST. — Aphyosemion obscurum (Ahl,
1924), junior synonym of A. cameronense. TYPE LOCALITY. — Koleya, Konkouré River, Guinea.
SITE. — Gills. SITE. — Gills.
REMARKS
REMARKS In the original description of C. dionchus, Paperna
Authors indicated in the original description that the (1968) gave three different illustrations for copulatory
presence of a Cichlidogyrus species on an Aphyosemion organs, the figure 7 pl. 1 p. 91 (without references
species is probably the result of a lateral transfer to host) resemble C. arfii, thus corresponding
from a cichlid fish. individuals have to be considered as C. arfii.
Cichlidogyrus amphoratus Cichlidogyrus berradae Pariselle & Euzet, 2003

Pariselle & Euzet, 1996
Cichlidogyrus berradae Pariselle & Euzet, 2003: 196,
Cichlidogyrus amphoratus Pariselle & Euzet, 1996: 115, fig. 3.
figs 6, 7. TYPE MATERIAL. — Holotype: MNHN 538 HF Tk63.
TYPE MATERIAL. — Holotype: MNHN 486 HF Tg100 Paratypes: MNHN 538 HF Tk64, BMNH 1997.1.29.1,
Paratypes: MNHN 486 HF Tg99, BMNH 1996.1.5.16, MRAC 37 409, CAS M-385.
MRAC 37.384. TYPE HOST. — Tilapia cabrae Boulenger, 1899.
TYPE HOST. — Tilapia louka Thys van den Audenaerde, TYPE LOCALITY. — Cayo Lake, Cabinda.
1969.
ADDITIONAL HOST . — Tilapia guineensis (Bleeker,
TYPE LOCALITY. — Laramié, Bourouma River, Guinea. 1862).
ZOOSYSTEMA • 2009 • 31 (4) 859

ADDITIONAL LOCALITY. — Loufoualéba Lake, Congo we do not agree: C. bifurcatus is a valid species, as
on T. guineensis. its description, illustrations and hosts never vary
in Paperna publications (1960, 1979).
Cichlidogyrus berrebii Pariselle & Euzet, 1994
Cichlidogyrus bilongi Pariselle & Euzet, 1995
Cichlidogyrus berrebii Pariselle & Euzet, 1994: 230, fig. 2.
TYPE MATERIAL. — Holotype and paratypes: MNHN Cichlidogyrus bilongi Pariselle & Euzet, 1995c: 190,
213 HF Tk14. 191, fig. 3.
Paratypes: BMNH 1993.3.4.1; MRAC 37.330, 37.331
TYPE MATERIAL. — Holotype: MNHN 218 HF Tk18.
and 37.333.
TYPE HOST. — Tylochromis jentinki (Steindachner, 1895) MRAC 37.337.
TYPE LOCALITY. — Ebrié Lagoon, Côte d’Ivoire. TYPE HOST. — Tilapia guineensis (Bleeker, 1862).
SITE. — Gills. TYPE LOCALITY. — Tiassalé, Bandama River, Guinea.
ADDITIONAL LOCALITIES. — Kogon River, Guinea and SITE. — Gills.

Sine Saloum Rivers, Senegal.
ADDITIONAL LOCALITY. — Layo research station, Ebrié
Lagoon, Côte d’Ivoire. Senegal and Congo on type
REMARKS host (nobis).
Pariselle & Euzet (1994) noticed the unusual mor-
phology of the dorsal transverse bar of Cichlidogyrus
from Tylochromis Regan, 1920 hosts (auricles are Cichlidogyrus bonhommei
continuous with convex face instead of attached on Pariselle & Euzet, 1998
this face); pending new morphological or molecular
Cichlidogyrus bonhommei Pariselle & Euzet, 1998: 279,
data from these or other parasites from Tylochromis, 280, fig. 5.
they left these species in Cichlidogyrus.
Cichlidogyrus bifurcatus Paperna, 1960 MRAC 37.403.
Cichlidogyrus bifurcatus Paperna, 1960: 13, 14, fig. 9; TYPE HOST. — Tilapia buttikoferi (Hubrecht, 1881).
1979: 9, pl. 1, figs 1-6.
TYPE LOCALITY. — Katonga, Little Scarcies River, Sierra
TYPE MATERIAL. — Paratype: MRAC M.T. 35.703 Leone.
(Paperna, 1979).
SITE. — Gills.
TYPE HOST. — Haplochromis flaviijosephi (Lortet, 1883).
TYPE LOCALITY. — Sea of Galilee, Israel. Cichlidogyrus bouvii Pariselle & Euzet, 1997
ADDITIONAL HOSTS. — Juveniles of Oreochromis au- Cichlidogyrus bouvii Pariselle & Euzet, 1997: 223, fig. 2.
reus (Steindachner, 1864); Haplochromis squamipinnis
Regan, 1921, H. elegans Trewavas, 1933, H. aeneocolor
Greenwood, 1973, H. limax Trewavas, 1933, H. sp. and
MRAC 37.395.
Pseudocrenilabrus multicolor (Schoeller, 1903) (Paperna
1979). TYPE HOST. — Sarotherodon occidentalis (Daget, 1962).
SITE. — Gills. TYPE LOCALITY. — La Ramié, Bourouma River, Guinea.
SITE. — Gills.
REMARKS ADDITIONAL LOCALITIES. — Kamouri, Batapon River,
Douëllou (1993) thought that C. bifurcatus was Guinea; Katonga, Little Scarcies River, Sierra Leone on
probably, at times, confused with C. haplochromii, type host.
860 ZOOSYSTEMA • 2009 • 31 (4)

Cichlidogyrus bychowskii (Markevich, 1934) REMARKS

Cichlidogyrus cirratus and C. nematocirrus have high
Ancyrocephalus bychowskii Markevich, 1934. morphological similarities and are both found on
Cichlidogyrus bychowskii – Paperna 1960: 14. O. niloticus. The only difference between these two
species is the length of the penis (320 vs. 145 μm)
TYPE MATERIAL. — No type material mentioned in the which is not easy to measure (very long, sinuous and
thin penis), thus we consider that C. nematocirrus
TYPE HOST. — Hemichromis bimaculatus Gill, 1862. is a junior synonym of C. cirratus.
TYPE LOCALITY. — From a fish reared in the Leningrad
aquarium.
Cichlidogyrus cubitus Dossou, 1982
ADDITIONAL HOSTS. — Hemichromis fasciatus Peters, 1857
and Sarotherodon galilaeus (Linnaeus, 1758) [Paperna Cichlidogyrus cubitus Dossou, 1982: 306, 307, figs 13-
1968, 1969]. 15.
SITE. — Gills. TYPE MATERIAL. — Holotype: MNHN Tj114.
Paratype: MNHN Tj115.
ADDITIONAL LOCALITIES. — Ghana on type host (Paperna
1965); Ghana on type host, Hemichromis fasciatus Peters, TYPE HOST. — Tilapia zillii (Gervais, 1848).
1857 and Sarotherodon galilaeus (Linnaeus, 1758) (Paperna
1968, 1969). TYPE LOCALITY. — Ouémé, Benin.
SITE. — Gills.
Cichlidogyrus cirratus Paperna, 1964
Cichlidogyrus cirratus Paperna, 1964: 4, 5, figs 10, 11. Cichlidogyrus dageti Dossou & Birgi, 1984
Cichlidogyrus nematocirrus Paperna, 1969: 863, figs 54, Cichlidogyrus dageti Dossou & Birgi, 1984: 106-108,
55; 1979: 13, pl. 3, figs 3-7. figs 9-11.
TYPE MATERIAL. — Of C. cirratus: Holotype and paratypes TYPE MATERIAL. — Holotype: MNHN 208 TC Tj134.
in the helminthological collection of the Department of
Parasitology Hebrew University, Hassad Medical School, TYPE HOST. — Hemichromis fasciatus Peters, 1857.
Israel, no number mentioned.
Of C. nematocirrus: Holotype: MRAC M.T. 35.583 TYPE LOCALITY. — No type locality mentionned in the
(Paperna 1979). original description.
TYPE HOST. — Of C. cirratus: Sarotherodon galilaeus SITE. — Gills.

(Linnaeus, 1758).
Of C. nematocirrus: Oreochromis niloticus (Linnaeus, ADDITIONAL LOCALITY. — Nyong Basin, Cameroon
1758) (Paperna 1979). on type host.
TYPE LOCALITY. — Of C. cirratus Tiberias Lake, Israel.

Of C. nematocirrus: Man-made Volta Lake, Ghana.
Cichlidogyrus digitatus Dossou, 1982
ADDITIONAL HOST. — Of C. cirratus: Oreochromis niloticus
(Ergens 1981). Cichlidogyrus digitatus Dossou, 1982: 303-306, figs 10-
Of C. nematocirrus: Tilapia zillii (Gervais, 1848), Oreo- 12.
chromis variabilis (Boulenger, 1906) and Oreochromis
esculentus (Graham, 1928) (Paperna 1979). Cichlidogyrus cf. brevicirrus – Paperna 1969: 864, figs 78,
79.
SITE. — Gills.
Cichlidogyrus halinus Paperna, 1969 (part): 860, 861,
ADDITIONAL LOCALITIES. — Cairo, Nile River, Egypt fig. 71 left drawing (individuals from Tilapia guineensis
(Ergens 1981) on O. niloticus. Senegal on type host in the original description) (cf. Pariselle & Euzet 1996:
(nobis). 117, 118, fig. 10).
ZOOSYSTEMA • 2009 • 31 (4) 861

TYPE MATERIAL. — C. digitatus: Holotype: MNHN TYPE LOCALITY. — Of C. dionchus: Mamahuma stream,
Tj116. Paratype: MNHN Tj117. Accra, Ghana.
Of C. brevicirrus: Jinja, Kazi, Victoria Lake, Uganda.
OTHER MATERIAL. — C. cf. brevicirrus: “paratype”:
MRAC M.T. 35.586 from Tilapia discolor (Günther, ADDITIONAL HOSTS. — Of C. dionchus: Chromidotilapia
1903) (Paperna 1979) (see Remarks under Cichlidogyrus guentheri (Sauvage, 1882) (Paperna 1979).
dionchus chapter below). Of C. brevicirrus: Haplochromis longirostris (Hilgendorf,
1888), Haplochromis obesus (Boulenger, 1906), Haplo-
TYPE HOST. — C. digitatus: Tilapia zillii (Gervais, 1848). chromis obliquidens (Hilgendorf, 1888), Haplochromis
sp., Hoplotilapia retrodens (Hilgendorf, 1888) and Tilapia
TYPE LOCALITY. — Ouémé, Benin. zillii (Gervais, 1848).
ADDITIONAL HOSTS. — Tilapia guineensis (Bleeker, 1862) SITE. — Gills.
(Paperna 1969). Tilapia dageti Thys van den Audenaerde,
1971, T. guineensis (Bleeker, 1862) and T. louka Thys van ADDITIONAL LOCALITIES. — Afram sector, Volta Lake and
den Audenaerde, 1969 (Pariselle & Euzet 1996). Tilapia Adutor lower reaches of the Volta River, Ghana on H. fasciatus;
brevimanus Boulenger, 1911 (Pariselle & Euzet 1998). Ghana on Chromidotilapia guentheri (Paperna 1979).
SITE. — Gills.
REMARKS
ADDITIONAL LOCALITIES. — Volta Lake, Ghana on Tilapia Paperna (1968) used for this species two spellings:
guineensis (original description of C. halinus in Paperna
1969). Senegal, Guinea, Côte d’Ivoire and Congo on C. dioncus and C. dionchus, as the first one was latter
Tilapia guineensis (Pariselle & Euzet 1995c); Côte d’Ivoire, use by Paperna (1979) and other authors, we accept
Guinea, Senegal, Mali and Gambia on Tilapia zillii, this spelling. Dossou & Birgi (1984) indicated that
T. dageti, T. guineensis and T. louka (Pariselle & Euzet1996); a proportion of individuals named C. dionchus by
Guinea on T. brevimanus (Pariselle & Euzet 1998). Paperna (1968: fig. 6, p. 91) correspond to C. falcifer
(individuals from Hemichromis fasciatus) and we agree
REMARKS with that. Dossou (1982) indicated that C. dionchus
In our opinion, measurements and figures given in the (on Sarotherodon galilaeus) is a valid species. Some
original description of C. cf. brevicirrus by Paperna hosts were added by Paperna (1979): Pseudocrenilabrus
(1969) are almost identical to those of C. digitatus multicolor (Schoeller, 1903), Tilapia rendalli (Boul-
Dossou, 1982, with which this species had to be enger, 1897) and Haplochromis wingatii (Boulenger,
synonymised, despite the opinion of Paperna (1979), 1902), but probably because of the presence of C. cf.
who synonymised it with C. dionchus. brevicirrus, which was synonymised by this author
Pariselle & Euzet 1996 synonymised a part of with C. dionchus, when we propose to synonymise
the individuals named C. halinus with C. digita- C. cf. brevicirrus with C. digitatus. In the same way,
tus (those found on Tilapia guineensis by Paperna Paperna (1979) deposits a paratype (MRAC M.T.
[1969: fig 71 left drawing]). 35.586, probably C. cf. brevicirrus) from Tilapia
discolor (Günther, 1903) for C. dionchus, we consider
this specimen as being a C. digitatus. In the original
Cichlidogyrus dionchus Paperna, 1968 description by Paperna (1968) the only true C. dionchus
was illustrated by the drawing 5 (pl. 1, p. 91) and by
Cichlidogyrus dionchus Paperna, 1968: 89, 90, pl. 1,
figs 4, 5.
the figures of C. brevicirrus in its original description
(Paperna & Thurston 1969: 18), this latter species
Cichlidogyrus brevicirrus Paperna & Thurston, 1969: 17, had been synonymised with C. dionchus by Paperna
18. — Paperna 1979: 9, 10. (1979); we agree with this conclusion.
TYPE MATERIAL. — Holotype: MRAC M.T. 35.585 from
type host (Paperna 1979).
TYPE HOST. — Of C. dionchus: Sarotherodon galilaeus Cichlidogyrus dossoui Douëllou, 1993
(Linnaeus, 1758).
Of C. brevicirrus: Haplochromis guiarti (Pellegrin, 1904). Cichlidogyrus dossoui Douëllou, 1993: 174, 175, fig. 9.
862 ZOOSYSTEMA • 2009 • 31 (4)

TYPE MATERIAL. — Holotype: MNHN 139 HF. Cichlidogyrus euzeti Dossou & Birgi, 1984
TYPE HOST. — Tilapia rendalli (Boulenger, 1897). Cichlidogyrus euzeti Dossou & Birgi, 1984: 108, 109,
TYPE LOCALITY. — Kariba Lake, Zimbabwe. figs 12-14.
ADDITIONAL HOSTS. — Oreochromis mortimeri (Trewavas, TYPE MATERIAL. — Holotype: MNHN 209 TC Tj135.
1966) and Serranochromis macrocephalus (Boulenger, TYPE HOST. — Hemichromis fasciatus Peters, 1857.
1899).
TYPE LOCALITY. — Benin and Cameroon.
SITE. — Gills.
SITE. — Gills.
Cichlidogyrus douellouae ADDITIONAL LOCALITY. — Congo on type host (Pariselle &
Pariselle, Bilong Bilong & Euzet, 2003 Euzet 2004).
Cichlidogyrus douellouae Pariselle, Bilong Bilong & Euzet,

2003: 203-205, fig. 3. Cichlidogyrus falcifer Dossou & Birgi, 1984
Paratypes: MNHN 535 HF Tk59, BMNH 1997.2.3.1, Cichlidogyrus falcifer Dossou & Birgi, 1984: 104-106,
MRAC 37.410. figs 5-8.
TYPE HOST. — Sarotherodon galilaeus (Linnaeus, 1758). Cichlidogyrus dionchus Paperna, 1968 (part): 89-91, pl.
1, fig. 6 (individuals coming from Hemichromis fasciatus)
TYPE LOCALITY. — W National Park, Mékrou River, (cf. Dossou & Birgi 1984: 104-106).
Niger.
TYPE MATERIAL. — Holotype: MNHN 207 TC Tj133.
ADDITIONAL HOST. — Sarotherodon galilaeus sanagaensis
(Thys van den Audenaerde, 1966). TYPE HOST. — Hemichromis fasciatus Peters, 1857.
SITE. — Gills. TYPE LOCALITY. — Benin and Cameroon.
ADDITIONAL LOCALITIES. — Bama, Kou River, Burkina SITE. — Gills.

Faso on type host, Edéa, Sanaga River, Cameroon on
S. g. sanagaensis. ADDITIONAL LOCALITIES. — Congo, Senegal, Mali,
Gambia and Côte d’Ivoire on type host (Pariselle &
Euzet 2004).
Cichlidogyrus ergensi Dossou, 1982
REMARKS
Cichlidogyrus ergensi Dossou, 1982: 301-303, figs 7-9. Dossou & Birgi (1984) indicated that C. falcifer
Cichlidogyrus sp. — Ergens 1981: 212, 213. corresponds to a proportion of specimens named
C. dionchus by Paperna in 1968; we agree with that:
TYPE MATERIAL. — Holotype: MNHN Tj113. among the three figures given by Paperna (1968:
TYPE HOST. — Tilapia zillii (Gervais, 1848). 91), we consider number 6 (pl. I) to be a C. falcifer
and probably was found on H. fasciatus.
TYPE LOCALITY. — Ouémé, Benin.
ADDITIONAL HOST. — Tilapia guineensis (Bleeker, 1862)
(Pariselle & Euzet 1995c). Cichlidogyrus flexicolpos Pariselle & Euzet, 1995
SITE. — Gills. Cichlidogyrus flexicolpos Pariselle & Euzet, 1995c: 193,
194, fig. 5.
ADDITIONAL LOCALITIES. — Cairo, Nile River, Egypt
(description of Cichlidogyrus sp. by Ergens [1981]); TYPE MATERIAL. — Holotype: MNHN 217 HF Tk16.
Senegal, Guinea, Côte d’Ivoire and Congo on Tilapia Paratypes: MNHN 217 HF Tk17, BMNH 1993.5.14.9,
guineensis (Bleeker, 1862) (Pariselle & Euzet 1995c). MRAC 37.340.
ZOOSYSTEMA • 2009 • 31 (4) 863

TYPE HOST. — Tilapia guineensis (Bleeker, 1862). lii (Dossou 1982). Senegal, Guinea, and Congo on
T. guineensis and T. walteri (Pariselle & Euzet 1996).
TYPE LOCALITY. — Tiassalé, Bandama River, Côte
d’Ivoire.
ADDITIONAL HOST. — Tilapia dageti Thys van den Cichlidogyrus giostrai
Audenaerde, 1971 (Pariselle & Euzet 1996). Pariselle, Bilong Bilong & Euzet, 2003
SITE. — Gills. Cichlidogyrus giostrai Pariselle, Bilong Bilong & Euzet,
2003: 205, 206, fig. 4.
ADDITIONAL LOCALITIES. — Layo research station, Ebrié
Lagoon, Côte d’Ivoire; Congo River, Congo on type TYPE MATERIAL. — Holotype: MNHN 529 HF Tk46.
host. Senegal and Guinea on T. guineensis and T. dageti Paratypes: MNHN 529 HF Tk47, BMNH 1997.2.3.2,
(Pariselle & Euzet 1996). MRAC 37.406.
TYPE HOST. — Sarotherodon caudomarginatus (Boulenger,
1916).
Cichlidogyrus fontanai Pariselle & Euzet, 1997
TYPE LOCALITY. — Tondon Road, Badi River, Guinea.
Cichlidogyrus fontanai Pariselle & Euzet, 1997: 223,
224, fig. 3. Site. — Gills.
Paratypes: MNHN 532 HF Tk 53, BMNH 1997.1.28.3, Cichlidogyrus guirali Pariselle & Euzet, 1997
MRAC 37.396.
Cichlidogyrus guirali Pariselle & Euzet, 1997: 224-226,
TYPE HOST. — Sarotherodon occidentalis (Daget, 1962). fig. 4.
TYPE LOCALITY. — La Ramié, Bourouma River, Guinea. TYPE MATERIAL. — Holotype: MNHN 533 HF Tk54.
SITE. — Gills. Paratypes: MNHN 533 HF Tk55, BMNH 1996.1.5.22-
25, MRAC 37.386.
ADDITIONAL LOCALITY. — Kamouri, Batapon River,
Guinea on type host. TYPE HOST. — Sarotherodon occidentalis (Daget, 1962).
TYPE LOCALITY. — La Ramié, Bourouma River, Guinea.
Cichlidogyrus gallus Pariselle & Euzet, 1995 SITE. — Gills.

ADDITIONAL LOCALITIES. — Kamouri, Batapon River, Guinea
Cichlidogyrus gallus Pariselle & Euzet, 1995c: 191-193, and Little Scarcies River, Sierra Leone on type host.
fig. 4.
Cichlidogyrus aegypticus – Dossou 1982: 296-299, figs Cichlidogyrus halinus Paperna, 1969
1-3 (cf. Pariselle & Euzet 1995c: 191). Not C. aegypicus
Ergens, 1981. Cichlidogyrus halinus Paperna, 1969: 860, 861, figs 70,
TYPE MATERIAL. — Holotype: MNHN 221 HF Tk24. 71 right drawing, 72.
Paratypes: MNHN 221 HF Tk25, BMNH 1993.5.14.10, Cichlidogyrus erectus Dossou, 1982: 316-318, fig. 29-30
MRAC 37.336. (cf. Pariselle & Euzet 1996: 117).
TYPE HOST. — Tilapia guineensis (Bleeker, 1862). TYPE MATERIAL. — Of C. halinus: Holotype: MRAC
TYPE LOCALITY. — Tiassalé, Bandama River, Côte M.T. 35.928 (Paperna 1979).
d’Ivoire. Of C. erectus: Holotype: MNHN Tj 143.
ADDITIONAL HOSTS. — Tilapia zillii (Gervais, 1848) TYPE HOST. — Of C. halinus: Sarotherodon melanotheron
(Dossou 1982). Tilapia walteri Thys van den Audenaerde, heudelotii (Duméril, 1861).
1968 (Pariselle & Euzet 1996). Of C. erectus: Sarotherodon melanotheron melanotheron
Rüppell, 1852.
SITE. — Gills.
TYPE LOCALITY. — Of C. halinus: Peshs lagoon, east of
ADDITIONAL LOCALITIES. — Layo research station, Ebrié Accra, South Ghana.
Lagoon, Côte d’Ivoire. Ouémé River, Benin on T. zil- Of C. erectus: Ouémé and Couffo, Benin.
864 ZOOSYSTEMA • 2009 • 31 (4)

SITE. — Gills. 1933) and Oreochromis spilurus spilurus (Günther, 1894).

C. halli victorianus: Oreochromis esculentus (Graham,
1928) (Paperna 1979).
Cichlidogyrus halli (Price & Kirk, 1967) C. tubicirrus magnus: Sarotherodon galilaeus (Linnaeus,
1758), Oreochromis leucostictus (Trewavas, 1933).
Cleidodiscus halli Price & Kirk, 1967: 139-141, figs
1-12. — Paperna 1979: 10. SITE. — Gills.
Cichlidogyrus halli typicus Paperna, 1979: 11, pl. 2, ADDITIONAL LOCALITIES. — Of C. halli: Kariba Lake,
fig. 1. — Douëllou 1993: 160-164, fig. 3. Zimbabwe on Oreochromis mortimeri and Serranochromis
Cichlidogyrus tubicirrus magnus Paperna & Thurston, macrocephalus (Douëllou 1993). Guinea and Sierra Leone
1969: 26, fig. 6a, e. — Paperna 1979: 10. on Sarotherodon occidentalis (Pariselle & Euzet 1997).
Of C. halli typicus: lakes Georges, Albert and Edward,
Cichlidogyrus magnus – Dossou 1982: 312-314, figs 24, Kajansi fish ponds, Uganda on O. niloticus; Lake Albert,
25. — Douëllou 1993: 160-164. Uganda and Volta Lake and lower Volta River, Ghana
Cichlidogyrus halli victorianus Paperna, 1979: 11, 12, on S. galilaeus; lakes Georges and Albert; Uganda on
pl. 2, figs 2, 3. O. leucostictus; Kajansi fish ponds, Uganda on introduced
O. niloticus vulcani and O. spilurus spilurus.
TYPE MATERIAL. — Of C. halli: Holotype: Albany
Museum, Grahamstown, Republic of South Africa. REMARKS
Paratypes: Helminthological Collection, United States
National Museum, Washington, DC; MRAC, no numbers Price & Kirk (1967) synonymised Cichlidogyrus with
mentioned. Voucher specimen from type host: MNHN Cleidodiscus Mueller, 1934; this was confirmed by Price
126 HF (Douëllou 1993). (1967). Price et al. (1969) rehabilitated Cichlidogyrus
Of C. halli typicus: Paratype: M.T. 35.511 from Oreo- with five species (C. arthracanthus, C. bifurcatus,
chromis niloticus (only type specimen mentioned by C. bychowskii, C. longicirrus and C. tiberianus),
Paperna 1979).
Of C. tubicirrus magnus: “Types”: private collection of but kept in Cleidodiscus (valid genus) two species
Paperna, no number mentioned. (C. halli and C. tilapiae). Paperna (1979) move back
Of C. halli victorianus: Holotype: MRAC M.T. 35.941 to Cichlidogyrus the two Cleidodiscus species found
from Oreochromis variabilis (Paperna 1979). on African cichlid fishes (C. halli and C. tilapiae);
TYPE HOST. — Of C. halli and C. halli typicus: Oreochromis this was confirmed by Douëllou (1993).
shiranus shiranus (Boulanger, 1897) (original host in Considering measurements and figures by Paperna
Price & Kirk 1967). (1979: 11, pl. 2 figs 2, 3) and after the examination of
Of C. tubicirrus magnus: Oreochromis niloticus (Linnaeus, type material, we did not find significant differences
1758).
Of C. magnus: Sarotherodon melanotheron Rüppel, 1852 between C. halli victorianus and C. halli. So, despite
(in Dossou 1982). the opinion of Douëllou (1993), we consider that C. h.
Of C. halli victorianus: Oreochromis variabilis (Boulenger, victorianus had to be synonymised with C. halli.
1906). The determination by Ergens (1981) of Tilapia
TYPE LOCALITY. — Of C. halli and C. halli typicus: Fort zillii (Gervais, 1848) as host for C. halli in Egypt
Johnston, Upper Shire River, Malawi. remains for us doubtful, in fact, this species is only
Of C. tubicirrus magnus: Albert and Georges Lakes, found on mouthbreeders Cichlidae (i.e. genera
Uganda. Oreochromis and Sarotherodon).
Of C. magnus: Ouémé and Couffo, Bénin.
Of C. halli victorianus: Victoria Lake, Entebbe and
Jinja, Uganda. Cichlidogyrus haplochromii
ADDITIONAL HOSTS. — C. halli: Oreochromis mortimeri
Paperna & Thurston, 1969
(Trewavas, 1966) and Serranochromis macrocephalus Cichlidogyrus haplochromii Paperna & Thurston, 1969:
(Boulenger, 1899) (Douëllou 1993). Sarotherodon oc- 18-20, fig. 2. — Paperna 1979: 12, pl. 3, figs 1, 2. —
cidentalis (Daget, 1962) (Pariselle & Euzet 1997). Douëllou 1993: 168-170, fig. 6.
C. halli typicus: Oreochromis niloticus (Linnaeus, 1758), Sa-
rotherodon galilaeus (Linnaeus, 1758), Oreochromis leucostictus Cichlidogyrus dionchus Paperna, 1968 (see Remarks
(Trewavas, 1933), Oreochromis niloticus vulcani (Trewavas, below). — Paperna 1979: 9, 10, pl. 1, fig. 7.
ZOOSYSTEMA • 2009 • 31 (4) 865

Cichlidogyrus tubicirrus minutus Paperna & Thurston, 1969: Hoplotilapia retrodens, Oreochromis leucostictus, Tilapia sp.,
26-28, fig. 6b, f (individuals from haplochromiine hosts). Albert Lake, Nabugabo, Bunyoni and Mulehe, Uganda
and Ruaha River, Tanzania on Haplochromis spp.
Cichlidogyrus tubicirrus – Paperna 1979: 14 (individuals
from haplochromiine hosts).
REMARKS
TYPE MATERIAL. — “Types”: private collection of Dr
Paperna, no number mentioned.
Paperna (1979) admitted he probably confused this
Of C. tubicirrus: “Types”: private collection of Paperna, no species, according to hosts and locations with C. tu-
number mentioned (original description of C. tubicirrus bicirrus and C. bifurcatus. Douëllou (1993) indicated
minutus); “Holotype”: MRAC M.T. 34.291 from Haplo- that C. haplochromii represents probably a species
chromis schubotzi Boulenger, 1914 (Paperna 1979). complex. Paperna (1979: pl. 1, fig. 7) gave drawings
OTHER MATERIAL. — MNHN 134 HF (Douëllou of male copulatory organs as being C. dionchus; we
1993). think that it is in fact an illustration of C. haplochromii
genitalia; Paperna in the original description specified
TYPE HOST. — Haplochromis guiarti (Pellegrin, 1904). neither hosts nor measurements and provided no
Of C. tubicirrus: Haplochromis schubotzi (Paperna
1979). drawings of haptoral sclerotised parts. The validity
of type host, type locality, holotype and paratypes
TYPE LOCALITY. — Jinja, Uganda for C. haplochromii cited by Paperna (1979) for Cichlidogyrus tubicirrus is
and C. tubicirrus.
dubious. In the original description of Cichlidogyrus
ADDITIONAL HOSTS. — C. haplochromii: Haplochromis tubicirrus minutus by Paperna & Thurston (1969),
macrognathus Regan, 1922, Haplochromis nubilus (Boul- the figure and measurements correspond exactly to
enger, 1906), Haplochromis obliquidens (Hilgendorf, 1888), C. haplochromii (cf. Douëllou 1993: 165, 169), at least
Haplochromis wingatii (Boulenger, 1902), Haplochromis
sp., Hoplotilapia retrodens (Hilgendorf, 1888), Macropleu- for specimens from hosts of the genus Haplochromis
rodus bicolor (Boulenger, 1906) and Platytaeniodus degeni (like C. haplochromii), which have to be synonymised
Boulenger, 1906. Haplochromis petronius Greenwood, 1973 with C. haplochromii (this concurs with remarks in
(Paperna 1979). Pharyngochromis darlingi (Boulenger, Douëllou 1993: 169, 170).
1911) (Douëllou 1993).
C. tubicirrus: Haplochromis angustifrons Boulenger, 1914,
Haplochromis guiarti (Pellegrin, 1904), Haplochromis longi-
rostris (Hilgendorf, 1888), Haplochromis macrops (Boulenger, Cichlidogyrus hemi Pariselle & Euzet, 1998
1911), Haplochromis obesus (Boulenger, 1906), Haplochromis
obliquidens (Hilgendorf, 1888), Haplochromis squamipin- Cichlidogyrus hemi Pariselle & Euzet, 1998: 276, fig. 3.
nis Regan, 1921, Haplochromis sp., Hoplotilapia retrodens TYPE MATERIAL. — Holotype: MNHN 544 HF Tk75.
(Hilgendorf, 1888) (original description of C. tubicirrus Paratypes: MNHN 544 HF Tk76, MRAC 37.411.
minutus); Haplochromis aeneocolor Greenwood, 1973,
Haplochromis limax Trewavas, 1933, Haplochromis nubilus TYPE HOST. — Tilapia brevimanus Boulenger, 1911.
(Boulenger, 1906), Haplochromis nigripinnis Regan, 1921, TYPE LOCALITY. — La Ramié, Bourouma River, Guinea.
Haplochromis elegans Trewavas, 1933, Haplochromis petronius
Greenwood, 1973, Tilapia sp. (Paperna 1979). SITE. — Gills.
SITE. — Gills. ADDITIONAL LOCALITIES. — Kogon and Ndyarendi,
Kogon River, Guinea.
ADDITIONAL LOCALITIES. — C. haplochromii: Kazi, Uganda
on type host; George Lake, Uganda on Haplochromis
wingatii and Haplochromis sp., Kariba Lake, Zimbabwe Cichlidogyrus inconsultans
on Pharyngochromis darlingi (Douëllou 1993). Birgi & Lambert, 1987
C. tubicirrus: Georges Lake, Uganda on Haplochromis
angustifrons, H. elegans, H. limax, H. macrops, H. nigripinnis, Cichlidogyrus inconsultans Birgi & Lambert, 1987: 103.
H. petronius, H. schubotzi, H. squamipinnis, Haplochromis New name for Cichlidogyrus euzeti Birgi & Lambert,
sp. (original description of C. tubicirrus minutus); Kazinga 1986: 525-527, figs 4-6.
Channel, Uganda on Haplochromis squamipinnis, H. ae-
neocolor, North Victoria Lake, Uganda on Haplochromis TYPE MATERIAL. — No type material mentioned in the
guiarti, H. longirostris, H. obesus, H. obliquidens, H. nubilus, original description.
866 ZOOSYSTEMA • 2009 • 31 (4)

TYPE HOST. — Polycentropsis abbreviata Boulenger, 1901 new morphological or molecular data from these
(Nandidae). or other parasites of Tylochromis, they left these
TYPE LOCALITY. — Mouanko, Cameroon. species in Cichlidogyrus.
SITE. — Gills.
Cichlidogyrus kouassii N’Douba,
REMARKS Thys van den Audenaerde & Pariselle, 1997
This species was re-named C. inconsultans because
the name C. euzeti Dossou & Birgi, 1984 already Cichlidogyrus kouassii N’Douba, Thys van den Aude-
naerde & Pariselle, 1997: 432, 433, fig. 2.
exists for a parasite from Hemichromis fasciatus.
Cichlidogyrus karibae Douëllou, 1993 MRAC 37.394.
Cichlidogyrus karibae Douëllou, 1993: 175-177, fig. 9.
TYPE LOCALITY. — Bakro, Ayamé Lake, Côte d’Ivoire.
TYPE MATERIAL. — Holoype (?): MNHN 136 HF.
SITE. — Gills.
TYPE HOST. — Sargochromis codringtonii (Boulenger,
1908).
TYPE LOCALITY. — Kariba Lake, Zimbabwe. Cichlidogyrus lagoonaris Paperna, 1969
ADDITIONAL HOST. — Oreochromis mortimeri (Trewavas, Cichlidogyrus lagoonaris Paperna, 1969: 861, figs 73-
1966). 77.
SITE. — Gills. Cichlidogyrus gibbus Dossou, 1982: 318, 319, figs 31-33.
TYPE MATERIAL. — C. lagoonaris: Holotype: MRAC
M.T. 35.929 (Paperna 1979).
Cichlidogyrus kothiasi Pariselle & Euzet, 1994 C. gibbus: Holotype: MNHN Tj137; paratype: MNHN
138.
Cichlidogyrus kothiasi Pariselle & Euzet, 1994: 232-234,
fig. 4. TYPE HOST. — Sarotherodon melanotheron melanotheron
Rüppell, 1852.
TYPE MATERIAL. — Holotype and paratypes: MNHN
212 HF Tk13. ADDITIONAL HOST. — Tilapia guineensis (Bleeker, 1862).
Paratypes: BMNH 1993.3.4.2, MRAC 37.332 and
37.334. TYPE LOCALITY. — C. lagoonaris: Lake Volta, Ghana.
C. gibbus: Ouémé and Couffo, Benin.
TYPE HOST. — Tylochromis jentinki (Steindachner, 1895).
SITE. — Gills.
TYPE LOCALITY. — Ebrié Lagoon, Côte d’Ivoire.
SITE. — Gills. REMARKS
We consider that C. gibbus, which male apparatus
ADDITIONAL LOCALITY. — Kogon River, Guinea on
type host.
is closely related to that of C. lagoonaris, is a junior
synonym. Paperna gives Sarotherodon melanotheron
heudelotii (Duméril, 1861) as type host for C. lagoon-
REMARKS aris coming from Ghana, and yet this fish species
Pariselle & Euzet (1994) noticed the unusual mor- is not present in this area (see Paugy et al. 2003),
phology of the dorsal transverse bar of Cichlidogyrus the sub-species from Ghana is S. m. melanotheron,
from Tylochromis (auricles are continuous with which is considered as the type host (as in Dossou
convex face instead of attached on this face); pending 1982 for C. gibbus).
ZOOSYSTEMA • 2009 • 31 (4) 867

Cichlidogyrus legendrei Pariselle & Euzet, 2003 TYPE HOST. — Hemichromis fasciatus Peters, 1857 (Paperna
1979).
Cichlidogyrus legendrei Pariselle & Euzet, 2003: 198,
fig. 5. TYPE LOCALITY. — New Tafo, Ghana.
TYPE MATERIAL. — Holotype: MNHN 539 HF Tk65. ADDITIONAL HOST. — Chromidotilapia guentheri (Sauvage,
Paratypes: MNHN 539 HF Tk66, BMNH 1997.1.29.2, 1882) (Paperna 1979).
MRAC 37.400, CAS M-387. SITE. — Gills.
TYPE HOST. — Tilapia cabrae Boulenger, 1899. ADDITIONAL LOCALITIES. — Benin and Cameroon on
TYPE LOCALITY. — Cayo Lake, Congo. type host (Dossou & Birgi 1984); Congo on type host
(Pariselle & Euzet 2004).
SITE. — Gills.
Cichlidogyrus longipenis
Cichlidogyrus lemoallei Pariselle & Euzet, 2003 Paperna & Thurston, 1969
Cichlidogyrus lemoallei Pariselle & Euzet, 2003: 198-200, Cichlidogyrus tubicirrus longipenis Paperna & Thurston,
fig. 6. 1969: 28, fig. 6d, h.
TYPE MATERIAL. — Holotype: MNHN 540 HF Tk67. Cichlidogyrus longipenis – Paperna 1979: 13.
MRAC 37.401, CAS M-388. TYPE MATERIAL. — “Holotype”: private collection of Dr
Paperna, no number mentioned.
TYPE HOST. — Tilapia cabrae Boulenger, 1899. “Syntype”: MRAC M.T. 35.921 (Paperna 1979).
TYPE LOCALITY. — Mouth of Bas Kouilou River, Congo.
TYPE HOST. — Astatoreochromis alluaudi Pellegrin,
SITE. — Gills. 1904.
ADDITIONAL LOCALITIES. — Loukoula River and Cayo TYPE LOCALITY. — Jinja, Uganda.
Lake, Congo.
SITE. — Gills.
Cichlidogyrus levequei Pariselle & Euzet, 1996

Cichlidogyrus louipaysani Pariselle & Euzet, 1995
Cichlidogyrus levequei Pariselle & Euzet, 1996: 120,
121, figs 12, 13. Cichlidogyrus louipaysani Pariselle & Euzet, 1995c:
195, fig. 6.
Paratypes: MNHN 482 HF Tg93, BMNH 1996.1.5.15, TYPE MATERIAL. — Holotype: MNHN 222 HF Tk26.
MRAC 37.388. Paratypes: MNHN 222 HF Tk27, BMNH 1993.5.14.11,
MRAC 37.341.
TYPE HOST. — Tilapia coffea Thys van den Audenaerde,
1970. TYPE HOST. — Tilapia guineensis (Bleeker, 1862).
TYPE LOCALITY. — Niambala, Oulé River, Guinea. TYPE LOCALITY. — Rice field, Koba, Guinea.
ADDITIONAL LOCALITY. — Koulé, Guinea.
ADDITIONAL LOCALITIES. — Somone Lagoon, Senegal
River, Sine Saloum River, Senegal and Banjul, Gambia
Cichlidogyrus longicirrus Paperna, 1965 River, Gambia.
Cichlidogyrus longicirrus Paperna, 1965: 108, 109, 111. —

Dossou & Birgi 1984: 102-104, figs 1-4. Cichlidogyrus microscutus Pariselle & Euzet, 1996
TYPE MATERIAL. — “Paratype”: MRAC M.T. 35.933 Cichlidogyrus microscutus Pariselle & Euzet, 1996: 113,
(Paperna 1979). 114, figs 4, 5.
868 ZOOSYSTEMA • 2009 • 31 (4)

TYPE MATERIAL. — Holotype: MNHN 488 HF Tg103. TYPE HOST. — Tilapia cessiana Thys van den Audenaerde,
Paratypes: MNHN 488 HF Tg104, BMNH 1996.1.5.10- 1968.
11, MRAC 37.385.
TYPE LOCALITY. — Toyebli, Nipoué River, Guinea.
ADDITIONAL HOST. — Tilapia buttikoferi (Hubrecht,
TYPE LOCALITY. — Loufoualéba Lake, Congo. 1881).
ADDITIONAL HOST. — Tilapia dageti Thys van den SITE. — Gills.
Audenaerde, 1971.
ADDITIONAL LOCALITIES. — Kandiafara, Kogon River,
SITE. — Gills. Guinea and Katonga, Little Scarcies River, Sierra Leone
on T. buttikoferi.
ADDITIONAL LOCALITY. — Niokolo-Koba National Park,
Gambia River, Senegal on T. dageti.
Cichlidogyrus ornatus Pariselle & Euzet, 1996
Cichlidogyrus nandidae Birgi & Lambert, 1986
Cichlidogyrus ornatus Pariselle & Euzet, 1996: 115-117,
figs 8, 9.
Cichlidogyrus nandidae Birgi & Lambert, 1986: 522-524,
figs 1-3. TYPE MATERIAL. — Holotype: MNHN 487HF Tg102.
Paratypes: MNHN 487 HF Tg101, BMNH 1996.1.5.1-2,
TYPE MATERIAL. — No type material mentioned in the MRAC 37.383.
original description..
TYPE HOST. — Tilapia zillii (Gervais, 1848).
TYPE HOST. — Polycentropsis abbreviata Boulenger, 1901
(Nandidae). TYPE LOCALITY. — Baoulé River in Côte d’Ivoire.
TYPE LOCALITY. — Mouanko, Cameroon. ADDITIONAL HOST. — Tilapia dageti Thys van den
Audenaerde, 1971.
SITE. — Gills.
SITE. — Gills.
Cichlidogyrus njinei ADDITIONAL LOCALITIES. — Bagoué River, Côte d’Ivoire on

type host and Comoé River, Côte d’Ivoire on T. dageti.
Pariselle, Bilong Bilong & Euzet, 2003
Cichlidogyrus njinei Pariselle, Bilong Bilong & Euzet, Cichlidogyrus ouedraogoi Pariselle & Euzet, 1996
2003: 207, fig. 5.
TYPE MATERIAL. — Holotype: MNHN 478 HF Tg77. Cichlidogyrus ouedraogoi Pariselle & Euzet, 1996: 121,
Paratypes: MNHN 478 HF Tg77, MRAC 37.370. 122, figs 14, 15.
TYPE HOST. — Sarotherodon galilaeus sanagaensis (Thys TYPE MATERIAL. — Holotype: MNHN 483 HF Tg94.
van den Audenaerde, 1966). Paratypes: MNHN 483 HF Tg95, BMNH 1996.1.5.21,
MRAC 37.387.
TYPE LOCALITY. — Edéa, Sanaga River, Cameroon.
TYPE HOST. — Tilapia coffea Thys van den Audenaerde,
SITE. — Gills. 1970.
TYPE LOCALITY. — Niambala, Oulé River, Guinea.
Cichlidogyrus nuniezi Pariselle & Euzet, 1998 SITE. — Gills.
Cichlidogyrus nuniezi Pariselle & Euzet, 1998: 276-278,

fig. 4.
Cichlidogyrus paganoi Pariselle & Euzet, 1997
Paratypes: MNHN 542 HF Tk70, BMNH 1997.1.30.3, Cichlidogyrus paganoi Pariselle & Euzet, 1997: 226,
MRAC 37.404. 227, fig. 5.
ZOOSYSTEMA • 2009 • 31 (4) 869

TYPE MATERIAL. — Holotype: MNHN 531 HF Tk51. ADDITIONAL LOCALITY. — Kogon River, Guinea.
MRAC 37.398.
REMARKS
TYPE HOST. — Sarotherodon occidentalis (Daget, 1962). Pariselle & Euzet (1994) noticed the unusual mor-
phology of the dorsal transverse bar of Cichlidogyrus
TYPE LOCALITY. — La Ramié, Bourouma River, Guinea.
from Tylochromis (auricles are continuous with
SITE. — Gills. convex face instead of attached on this face); pending
new morphological or molecular data from these
ADDITIONAL LOCALITIES. — Kamouri, Batapon River, or other parasites of Tylochromis, they left these
Guinea and Katonga, Little Scarcies River, Sierra Leone.
species in Cichlidogyrus.
Cichlidogyrus papernastrema Cichlidogyrus quaestio Douëllou, 1993

Price, Peebles & Bamford, 1969
Cichlidogyrus quaestio Douëllou, 1993: 181, 182, fig. 12.
Cichlidogyrus papernastrema Price, Peebles & Bamford,
1969: 120-122, figs 11-20. TYPE MATERIAL. — Holotype: MNHN 137 HF.
TYPE MATERIAL. — No type material mentioned in the TYPE HOST. — Tilapia rendalli (Boulenger, 1897).
TYPE LOCALITY. — Kariba Lake, Zimbabwe.
TYPE HOST. — Tilapia sparrmanii Smith, 1840.
A DDITIONAL HOSTS . — Sargochromis codringtonii
TYPE LOCALITY. — Natal, South Africa. (Boulenger, 1908) and Serranochromis macrocephalus
(Boulenger, 1899).
SITE. — Gills.
SITE. — Gills.
Cichlidogyrus philander Douëllou, 1993 REMARKS

Douëllou (1993) thought that this species is close to
Cichlidogyrus philander Douëllou, 1993: 183, 184, C. brevicirrus, and that Thurston (1970) probably
fig. 13. confused these two species in Uganda on Tilapia
TYPE MATERIAL. — Holotype: MNHN 137 HF.
rendalli (Boulenger, 1897).
TYPE HOST. — Pseudocrenilabrus philander philander

(Weber, 1897). Cichlidogyrus reversati Pariselle & Euzet, 2003
TYPE LOCALITY. — Kariba Lake, Zimbabwe. Cichlidogyrus reversati Pariselle & Euzet, 2003: 196-198,
fig. 4.
SITE. — Gills.
Cichlidogyrus pouyaudi Pariselle & Euzet, 1994 MRAC 37.402, CAS M-386.
TYPE HOST. — Tilapia cabrae Boulenger, 1899.
Cichlidogyrus pouyaudi Pariselle & Euzet, 1994: 230-232,
fig. 3. TYPE LOCALITY. — Mouth of Bas Kouilou River, Congo.
TYPE MATERIAL. — Holotype and paratypes: MNHN SITE. — Gills.
214 HF Tk15.
Paratypes: BMNH 1993.3.4.3, MRAC 37.335.
Cichlidogyrus rognoni
TYPE HOST. — Tylochromis jentinki (Steindachner, 1895) Pariselle, Bilong Bilong & Euzet, 2003
TYPE LOCALITY. — Ebrié Lagoon, Côte d’Ivoire.
Cichlidogyrus rognoni Pariselle, Bilong Bilong & Euzet,
SITE. — Gills: 2003: 202, 203, fig. 2.
870 ZOOSYSTEMA • 2009 • 31 (4)

TYPE MATERIAL. — Holotype: MNHN 536 HF Tk60. TYPE MATERIAL. — Holotype: MRAC M.T. 34.288
Paratypes: MNHN 536 HF Tk61, BMNH 1997.2.3.3, from type host.
MRAC 37.402. Paratype: M.T. 35.576 from hybrid (Paperna 1979);
Voucher specimen: MNHN 127 HF (Douëllou
TYPE HOST. — Oreochromis niloticus (Linnaeus, 1758). 1993).
TYPE LOCALITY. — Senegal River, Senegal. TYPE HOST. — Oreochromis mossambicus (Peters, 1852)
(Paperna 1979).
SITE. — Gills.
TYPE LOCALITY. — Kajansi, Uganda.
Cichlidogyrus sanjeani Pariselle & Euzet, 1997 ADDITIONAL HOSTS. — Oreochromis niloticus (Linnaeus,
1758), O. leucostictus (Trewavas, 1933), Haplochromis sp.
Cichlidogyrus sanjeani Pariselle & Euzet, 1997: 227, and Tilapia zillii (Gervais, 1848) (experimental infesta-
228, fig. 6. tion). Hybrid between O. mossambicus and O. niloticus,
O. spilurus niger (Günther, 1894) and O. aureus (Stein-
TYPE MATERIAL. — Holotype: MNHN 534 HF Tk56. dachner, 1864) (Paperna 1979). Oreochromis mortimeri
Paratypes: MNHN 534 HF Tk57, BMNH 1997.1.28.6, (Trewavas, 1966) and Serranochromis macrocephalus
MRAC 37.399. (Boulenger, 1899) (Douëllou 1993).
TYPE HOST. — Sarotherodon occidentalis (Daget, 1962). SITE. — Gills.
TYPE LOCALITY. — La Ramié, Bourouma River, Guinea. ADDITIONAL LOCALITIES. — George Lake, Uganda on
Oreochromis niloticus and Haplochromis sp.; Jinja, Uganda
SITE. — Gills. on O. leucostictus; Kajansi ponds, Uganda on Oreochromis
ADDITIONAL LOCALITIES. — Kamouri, Batapon River, spilurus niger, O. niloticus and Tilapia zillii; Jinja and Vic-
Guinea and Katonga, Little Scarcies River, Sierra toria Lake, Uganda on O. leucostictus; Fish pounds, Coastal
Leone. Plain, Israel on O. aureus. Kariba Lake, Zimbabwe on O.
mortimeri and Serranochromis macrocephalus (Douëllou
1993). BFRI fish farm ponds, Bangladesh (on introduced
Cichlidogyrus sanseoi Pariselle & Euzet, 2004 O. mossambicus; Ferdousi & Chandra 2002).
Cichlidogyrus sanseoi Pariselle & Euzet, 2004: 360-362,

figs 2, 3. Cichlidogyrus slembroucki
Pariselle & Euzet, 1998
TYPE MATERIAL. — Holotype: MNHN 236HG TI
233.
Paratypes: MNHN 236HG TI 233 bis, BMNH Cichlidogyrus slembroucki Pariselle & Euzet, 1998: 280-
2004.9.15.14-15. 281, fig. 6.
TYPE HOST. — Hemichromis fasciatus Peters, 1858. TYPE MATERIAL. — Holotype: MNHN 546 HF Tk79.
TYPE LOCALITY. — Kounougou River, Côte d’Ivoire. MRAC 37.405.
SITE. — Gills. TYPE HOST. — Tilapia buttikoferi (Hubrecht, 1881).
ADDITIONAL LOCALITIES. — Grand Blahoua Lagoon, TYPE LOCALITY. — Kandiafara, Kogon River, Guinea.
Côte d’Ivoire; Niokolo-Koba National Park, Gambia
River, Senegal; Sélingué, Niger River, Mali. SITE. — Gills.
Cichlidogyrus sclerosus Paperna & Thurston, 1969 Cichlidogyrus testificatus Dossou, 1982
Cichlidogyrus sclerosus Paperna & Thurston, 1969: 23, Cichlidogyrus testificatus Dossou, 1982: 319-321, figs 34-
24, fig. 4. — Douëllou, 1993: 164-166, fig. 4. 36.
Cichlidogyrus bangladeshi Ferdousi & Chandra, 2002: TYPE MATERIAL. — Holotype: MNHN Tj 139.
50-52, fig. 1. Paratype: MNHN Tj 140.
ZOOSYSTEMA • 2009 • 31 (4) 871

TYPE LOCALITY. — Ouémé, Benin. SITE. — Gills.

TYPE HOST. — Tilapia mariae Boulenger, 1899. ADDITIONAL LOCALITIES. — Ghana on Sarotherodon
galilaeus (Paperna 1968, 1969). Uganda on O. niloticus
SITE. — Gills. and Haplochromis longirostris (Paperna & Thurston 1969)
and on Oreochromis variabilis, O. esculentus and O. niloticus
(Paperna 1979). Senegal on type host (Pariselle et al.
2003). BFRI fish farm ponds, Bangladesh (on introduced
Cichlidogyrus teugelsi Pariselle & Euzet, 2004 O. mossambicus; Ferdousi & Chandra 2002).
Cichlidogyrus teugelsi Pariselle & Euzet, 2004: 362,
363, fig. 4.
REMARKS
TYPE MATERIAL. — Holotype: MNHN 237HG TI This species was re-described by Pariselle et al. (2003)
234. on the type host in Senegal as Ergens (1981) did not
Paratypes: MNHN 237HG TI 234 bis, BMNH mention, in the original description, the auxiliary
2004.9.15.12-13.
plate associated with the male apparatus.
TYPE HOST. — Hemichromis fasciatus Peters, 1857.
TYPE LOCALITY. — Kounougou River, Côte d’Ivoire.
Cichlidogyrus tiberianus Paperna, 1960
SITE. — Gills.
Cichlidogyrus tiberianus Paperna, 1960: 10-12, figs 7,
ADDITIONAL LOCALITIES. — Adiopodoumé, Côte d’Ivoire; 8. — Dossou 1982: 299-301, figs 4-6. — Douëllou,
Niokolo-Koba National Park, Gambia River, Senegal 1993: 173, 174, fig. 8.
and Sélingué, Niger River, Mali.
TYPE MATERIAL. — No type material mentioned in the
Cichlidogyrus thurstonae Ergens, 1981 TYPE HOST. — Tilapia zillii (Gervais, 1848).
Cichlidogyrus thurstonae Ergens, 1981: 211, 212, fig. 8. — TYPE LOCALITY. — Sea of Galilee, Israel.
Pariselle et al. 2003: 207-209, fig. 6.
ADDITIONAL HOSTS. — Tristramella simonis simonis
Cichlidogyrus tiberianus Paperna, 1960 individuals from: (Günther, 1864) and T. sacra (Günther, 1865) (experi-
Sarotherodon galilaeus (Linnaeus, 1758) (in Paperna 1968: mental infection only), Haplochromis flaviijosephi (Lortet,
92; 1969: 863); Oreochromis niloticus and Haplochromis 1883). Tilapia rendalli (Boulenger, 1897) (Paperna 1979;
longirostris (Hilgendorf, 1888) (in Paperna & Thurston Douëllou 1993). Tilapia coffea Thys van den Audenaerde,
1969: 24, 25, fig. 5d left drawing); from Oreochromis 1970, Tilapia dageti Thys van den Audenaerde, 1971,
variabilis (Boulenger, 1906), O. esculentus (Graham, T. guineensis (Bleeker, 1862), T. walteri Thys van den
1928) and O. niloticus (Linnaeus, 1758) (in Paperna Audenaerde, 1968 (Pariselle & Euzet 1995c, 1996).
1979: 14); see Remarks under C. tiberianus.
SITE. — Gills.
Cichlidogyrus gilli Ferdousi & Chandra, 2002: 52-54,
fig. 2. ADDITIONAL LOCALITIES. — Southern Ghana on type
host (Paperna 1965). Uganda on type host (Paperna &
TYPE MATERIAL. — Holotype: CAS, no number men- Thurston 1969). Uganda on T. rendalli (Paperna 1979).
tioned. Egypt on type host (Ergens 1981). Benin on type host
(Dossou 1982). Zimbabwe on T. rendalli (Douëllou
OTHER MATERIAL. — MNHN 537 HF Tk62 (Pariselle 1993). Senegal, Guinea, Côte d’Ivoire and Congo on
et al. 2003). Tilapia guineensis, T. coffea, T. dageti, T. walteri and type
TYPE HOST. — Oreochromis niloticus (Linnaeus, 1758). host (Pariselle & Euzet 1995c, 1996).
TYPE LOCALITY. — Cairo, Nile River, Egypt. REMARKS

ADDITIONAL HOST. — Haplochromis longirostris (Hilgen- Ergens (1981) in Egypt found and re-described
dorf, 1888) (Paperna & Thurston 1969; Ergens 1981). C. tiberianus on Tilapia zillii; he indicated that
Oreochromis mossambicus (Ferdousi & Chandra, 2002). C. tiberianus on Oreochromis niloticus and Haplo-
872 ZOOSYSTEMA • 2009 • 31 (4)

chromis longirostris found in Uganda by Paperna & ADDITIONAL HOSTS. — Oreochromis niloticus (Linnaeus,
Thurston (1969) (with some differences with original 1758), Tristramella sacra (Günther, 1865) and Tristramella
simonis (Günther, 1864) (experimental infestation). Tilapia
descriptions from Ghana or Israel) had to be con- busumana (Günther, 1903) and Hemichromis fasciatus
sidered as C. thurstonae. Dossou (1982) in Benin Peters, 1857 (Paperna 1968). Oreochromis leucostictus
found C. tiberianus on Tilapia zillii, he agreed (Trewavas, 1933), Oreochromis mossambicus (Peters, 1852),
with Ergens (1981) about the confusion made by Tilapia zillii (Gervais, 1848) (experimental infestation)
Paperna & Thurston (1969) between C. tiberianus and Haplochromis macrognathus Regan, 1922 (Paperna &
and C. thurstonae. Douëllou (1993), according to Thurston 1969). Chromidotilapia guentheri (Sauvage,
1882) (Paperna 1969). Oreochromis aureus (Steindachner,
Ergens (1981), considered that Paperna & Thurston 1864), Oreochromis niloticus vulcani (Trewavas, 1933),
(1969) confused C. tiberianus with C. thurstonae; Oreochromis spilurus niger (Günther, 1894), Oreochromis
she added that Paperna (1968, 1969 and 1979) may variabilis (Boulenger, 1906) and Oreochromis urolepis
have made the same error. Pariselle & Euzet (1995c, urolepis (Norman, 1922) (Paperna 1979). Oreochromis
1996) found C. tiberianus only from hosts of the mortimeri (Trewavas, 1966) (Douëllou 1993).
sub-genus Coptodon (namely: Tilapia coffea, T. dageti, SITE. — Gills.
T. guineensis, T. walteri and T. zillii). All these remarks
are leading us to the same conclusion: all individuals ADDITIONAL LOCALITIES. — Dor, Fish Culture Research
Station, Israel on Oreochromis niloticus; Ginossar, Fisheries
described as C. tiberianus from mouthbreeders hosts Field Station, Israel on Sarotherodon galilaeus; Western
(genus Oreochromis and Sarotherodon) have been shores, Sea of Galilee, Israel on Tristramella sacra. Southern
confused with C. thurstonae by Paperna & Thurston Ghana on O. niloticus and S. galilaeus (Paperna 1965).
(1969) or Paperna (1968, 1969 and 1979). Ghana on Tilapia busumana and Hemichromis fasciatus
(Paperna 1968). Uganda on Oreochromis leucostictus,
O. mossambicus, Tilapia zillii (experimental infestation)
and Haplochromis macrognathus (Paperna & Thurston
Cichlidogyrus tilapiae Paperna, 1960 1969). Volta Basin, South Ghana on Sarotherodon galilaeus,
Oreochromis niloticus and Chromidotilapia guentheri
Cichlidogyrus tilapiae Paperna, 1960: 5-9, figs 3-6. — (Paperna 1979). Egypt on Oreochromis niloticus and
Douëllou 1993: 167, 168, fig. 5. Tilpia zillii (Ergens 1981). Kariba Lake, Zimbabwe on
Oreochromis mortimeri (Douëllou 1993). Bangladesh in
Cleidodiscus tilapiae – Price 1967: 381. — Paperna BFRI and BAU fish farm ponds on introduced O. niloticus
1979: 14. and O. mossambicus (Ferdousi & Chandra 2002).
Cichlidogyrus tubicirrus minutus Paperna & Thurston, REMARKS
1969: 26-28, fig. 6c, g. — Paperna 1979: 14 (individuals
from Oreochromis leucostictus). Paperna (1979) synonymised Cichlidogyrus tubicirrus
minutus with Cichlidogyrus tubicirrus, the validity of
Cichlidogyrus tubicirrus – Paperna 1979: 14 (individuals type host, type locality, holotype and paratypes given
from Oreochromis leucostictus). in this paper is dubious; in the original description
Cichlidogyrus chandrai Ferdousi & Chandra, 2002: by Paperna & Thurston (1969), the figure and
54-56, fig. 3. measurements of specimens from Oreochromis
leucostictus, are almost identical to those of C. tilapiae
TYPE MATERIAL. — Of C. tilapiae: no type material
mentioned in the original description. (cf. Ergens 1981: 210), with which C. tubicirrus
Of C. tubicirrus (from Oreochromis leucostictus): Paratype: (from O. leucostictus) has to be synonymised.
MRAC M.T. 35932 (Paperna 1979).
TYPE HOST. — Sarotherodon galilaeus (Linnaeus, 1758) Cichlidogyrus vexus Pariselle & Euzet, 1995
(Paperna 1979).
Cichlidogyrus vexus Pariselle & Euzet, 1995: 195-197,
TYPE LOCALITY. — Jordan and coastal system, Israel fig. 7.
(Paperna 1979).
OTHER MATERIAL. — C. tilapiae: MNHN 130 HF Paratypes: MNHN 219 HF Tk21, BMNH 1993.5.14.12,
(Douëllou 1993). MRAC 37.339.
ZOOSYSTEMA • 2009 • 31 (4) 873

A Cichlidogyrus yanni Pariselle & Euzet, 1996

Cichlidogyrus yanni Pariselle & Euzet, 1996: 118-120,
2.9 fig. 11.
1.9 Paratypes: MNHN 481 HF Tg91, BMNH 1996.1.5.22-
25, MRAC 37.386.
TYPE HOST. — Tilapia zillii (Gervais, 1848).
TYPE LOCALITY. — Kogon River, Guinea.
0
ADDITIONAL HOSTS. — Tilapia dageti Thys van den
Audenaerde, 1971, T. guineensis (Bleeker, 1862), T. louka
B Thys van den Audenaerde, 1969, and T. walteri Thys van
den Audenaerde, 1968.
SITE. — Gills.
ADDITIONAL LOCALITIES. — Volta Noire River, Burkina
Faso on type host; Niokolo-Koba National Park, Gambia
2 River, Senegal on T. dageti; Layo Research Station, Ebrié
Lagoon, Côte d’Ivoire and Senegal River, Senegal on
T. guineensis; Bourouma River, Guinea on T. louka; Cavally
and Nipoué Rivers, Côte d’Ivoire on T. walteri.
1
C Cichlidogyrus zambezensis Douëllou, 1993

Cichlidogyrus zambezensis Douëllou, 1993: 178-181,
1.7 fig. 11.
TYPE MATERIAL. — Holotype: MNHN 138 HF.
TYPE HOST. — Serranochromis macrocephalus.
0
ADDITIONAL HOST. — Oreochromis mortimeri (Trewavas,
FIG. 1. — Definition of characters used in the identification keys of
1966).
Cichlidogyrus Paperna, 1960 species (from 54 species studied):
A, value repartition of the standardised length of dorsal transverse TYPE LOCALITY. — Kariba Lake, Zimbabwe.
bar auricles; B, value repartition of the standardised length of
uncinuli pairs III-VII; C, value repartition of the standardised length SITE. — Gills.
of uncinuli pair I.
TYPE LOCALITY. — Tiassalé, Bandama River, Côte IDENTIFICATION KEY FOR STUDIED
d’Ivoire. GENERA AND SPECIES
Following Dossou (1985), Pariselle & Euzet
ADDITIONAL HOST. — Tilapia zillii (Gervais, 1848). (1995c) and Pariselle (1996) the 71 species forming
SITE. — Gills. Cichlidogyrus, recorded from 66 different host
species (64 Cichlidae, one Cyprinodontidae, one
ADDITIONAL LOCALITIES. — Layo Research Station, Ebrié
Lagoon, Côte d’Ivoire on type host; Kossou Lake, Côte Nandidae), may be clustered in different groups ac-
d’Ivoire on type host and T. zillii. Senegal, Guinea and cording to the relative size of their haptoral sclerites.
Congo on type host (Pariselle & Euzet 1996). This has been confirmed by Pouyaud et al. (2006)
874 ZOOSYSTEMA • 2009 • 31 (4)

using both genetic and morphometrical data: “This be “standardised” by dividing its total length by
division into four groups based on morphological the total length of uncinuli pair II, which retain
criterions is also confirmed by the genetic distances”. its size from larval to adult stages of development
These groups can be easily characterised by morpho- in ancyrocephalid Monogenea.
metrical characters: length of the transverse dorsal – Length of transverse dorsal bar auricles: the
bar auricles, of the uncinulus pairs III-VII, and of distribution of this character values (Fig. 1A) shows
uncinuli pairs I. a clear gap between species belonging to Scutogyrus
The following may introduced morphometrical (> 2.9) (Fig. 2E) and Cichlidogyrus (< 1.9) (Fig. 2F).
differences into the size range for these characters: This unequivocal character, therefore, is relevant to
developmental stage of the examined parasite; in- diagnose Scutogyrus.
fluence of temperature on the size of the sclerites – Size of uncinuli pairs III-VII: the gap between
(Ergens & Gelnar 1985; Appelby 1996); equipment large and small standardized sizes of uncinuli pairs
used; method by which the measurements are taken, III-VII is located near 2 (Fig. 1B).
or other constraints (see Caltran et al. 1995). To – Size of uncinuli pair I: the gap between large
avoid these problems, the length of each examined and small standardized sizes of uncinuli pairs I is
sclerite, according to Pariselle & Euzet (2003), will located near 1.7 (Fig. 1C).
1. One transverse bar (ventral) ....................................................................................... 2

— More than one transverse bar (ventral and dorsal) ...................................................... 3
2. One asymmetrical pair of anchors (ventral) (Fig. 2A) ..................................... Urogyrus
— Two pairs of symmetrical anchors (Fig. 2B) ............................................. 6 Enterogyrus
3. Two transverse bars (one dorsal and one ventral) ........................................................ 4
— Three transverse bars (one dorsal, two ventral) (Fig. 2C) ...................... 15 Onchobdella
4. Dorsal bar with wings (= auricles) (Fig. 2D, a) ........................................................... 5
— Dorsal bar lacking wings (Fig. 2D, b) ................................................... 13 Insulacleidus
5. Dorsal bar winged with very long auricles, ventral bar associated with a plate (Fig. 2E)
................................................................................................................ 22 Scutogyrus
— Dorsal bar with short auricles ventral bar not associated with a plate (Fig. 2F) ..............
........................................................................................................... 27 Cichlidogyrus
6. Dorsal anchor total length more than 50 μm, ventral bar straight, more than 40 μm
(Fig. 3A, a) ................................................................................................................. 7
— Dorsal anchor total length less than 50 μm, ventral bar V-shaped, less than 40 μm (Fig. 3A,
b) ............................................................................................................................... 8
7. Ventral anchor total length less than 27 μm .............................. Enterogyrus malmbergi
— Ventral anchor total length more than 34 μm ................................... Enterogyrus crassus
8. Dorsal anchor with shaft longer than blade (Fig. 3B, a) .............................................. 9
— Dorsal anchor with shaft shorter than blade (Fig. 3B, b) .......................................... 11
9. Haptor cup-shaped .................................................................. Enterogyrus cichlidarum
— Haptor in two parts: one bulbous and one peduncular shaped (Fig. 3C) .................. 10
10. Penis with distal turns well separated (Fig. 3D, a) ........................ Enterogyrus coronatus
— Penis with distal turns adjoining (Fig. 3D, b) .................................. Enterogyrus foratus
ZOOSYSTEMA • 2009 • 31 (4) 875

11. Ventral bar total length less than 14 μm ........................................... Enterogyrus amieti
— Ventral bar total length more than 16 μm ................................................................ 12
12. Penis with 3-2-3 turns (Fig. 3E, a) .............................................. Enterogyrus melenensis
— Penis with 6-2-4 turns (Fig. 3E, b) ......................................... Enterogyrus barombiensis
13. Looped penis (Fig. 4B, a) ....................................................... Insulacleidus paratilapiae
— Straight penis (Fig. 4B, b) ........................................................................................ 14
14. Simple accessory piece (Fig. 4C, a) ............................................. Insulacleidus paretropli
— Accessory piece complex (Fig. 4C, b) ............................... Insulacleidus ptychochromidis
15. Dorsal bar clearly arched (M-shaped) (Fig. 5A, a) ..................................................... 16
— Dorsal bar straight or slightly V-shaped (Fig. 5A, b) ................................................. 19
16. Ventral bars longer than dorsal one (Fig. 5B, a) ......................... Onchobdella pterigyalis
— Ventral bars shorter than dorsal one (Fig. 5B, b) ....................................................... 17
17. Penis J-shaped (Fig. 5C, a) ........................................................... Onchobdella voltensis
— Penis long and sinuous or spirally coiled (Fig. 5C, b) ............................................... 18
18. Penis sinuous, no visible vagina (Fig. 5D, a) ............................... Onchobdella spirocirra
— Penis spirally coiled (4 to 5 turns), long and sinuous vagina (Fig. 5D, b) .......................
...................................................................................................... Onchobdella bopeleti
19. Dorsal bar slightly V-shaped, length more than 43 μm (Fig. 5E, a) ... Onchobdella silverai
— Dorsal bar straight, length less than 40 μm (Fig. 5E, b) ............................................ 20
20. Ventral anchor total length less than 11 μm (Fig. 5F, a) .................. Onchobdella melissa
— Ventral anchor total length more than 20 μm (Fig. 5F, b) ......................................... 21
21. Penis thin, G-shaped, total length 80 μm (Fig. 5G, a) .................... Onchobdella krachii
— Penis large, J-shaped, total length less than 60 μm (Fig. 5G, b) .... Onchobdella aframae
22. Penis and vagina very long and thin (Fig. 6A, a) .............................. Scutogyrus ecoutini
— Penis and vagina shorter (Fig. 6A, b) ........................................................................ 23
23. Penis length more than 70 μm (Fig. 6B, a) ............................................................... 24
— Penis length less than 70 μm (Fig. 6B, b) .................................................................. 25
24. Vagina tubular, thin-walled (Fig. 6C, a) ............................................ Scutogyrus bailloni
— Vagina large, thick-walled (Fig. 6C, b) ...................................... Scutogyrus gravivaginus
25. Accessory piece ending in two curved pincer-like hooks (Fig. 6D, a) ... Scutogyrus chikhii
— Accessory piece ending in pincers with one component hooked, the other blunt (Fig. 6D,
b) ............................................................................................................................. 26
26. Accessory piece basal extension well marked (Fig. 6E, a) .................... Scutogyrus minus
— No accessory piece extension (Fig. 6E, b) .................................... Scutogyrus longicornis
27. Uncinuli pairs III-VII long (“standardised length” more than 2) (Fig. 7A, a) ............ 28
— Uncinuli pairs III-VII short (“standardised length” less than 2) (Fig. 7A, b) .............. 53
28. Uncinuli pairs I large (“standardised length” more than 1.7) (Fig. 7B, a) .................. 29
— Uncinuli pairs I small (“standardised length” less than 1.7) (Fig. 7B, b) .................... 30
876 ZOOSYSTEMA • 2009 • 31 (4)

A B C
VB
VB
VA VA
VB
DB
VA DA DA
a b
E F
VB
VA VB
VA
DB
DB
DA
DA
FIG. 2. — Illustration of the identification key for studied genera: A, haptoral morphology of Urogyrus Bilong Bilong, Birgi & Euzet,
1994 species; B, haptoral morphology of Enterogyrus Paperna, 1963 species; C, haptoral morphology of Onchobdella Paperna,
1968 species; D, shape of dorsal bars (not winged [a], winged [b]); E, haptoral morphology of Scutogyrus Pariselle & Euzet, 1995
species; F, haptoral morphology of Cichlidogyrus Paperna, 1960 species. Abbreviations: DA, dorsal anchor; DB, dorsal transverse
bar; VA, ventral anchor; VB, ventral transverse bar.
ZOOSYSTEMA • 2009 • 31 (4) 877

s
s
bl
bl
a
b
a b
C E
4
3
2
2
D
3 6
a b a b
FIG. 3. — Illustration of identification key for Enterogyrus Paperna, 1963 species: A, size of dorsal anchor (large [a], small [b]) and shape
of dorsal transverse bar (straight [a], V-shaped [b]); B, relative length of dorsal anchor blade and shaft (long shaft [a], short shaft [b]);
C, morphology of haptor with bulbous and peduncular parts; D, penis distal turns (separated [a] or adjoining [b]); E, number of turns
in the different parts of the penis. Abbreviations: bl, blade; s, shaft; 2, 3, 4 and 6, number of turns in the parts of the penis.
878 ZOOSYSTEMA • 2009 • 31 (4)

a b
a b
FIG. 4. — Illustration of identification key for Insulacleidus Rakotofiringa & Euzet, 1983 species: A, haptor morphology; B, shape of the
penis (looped [a], straight [b]); C, shape of the accessory piece (simple [a], complex [b]).
ZOOSYSTEMA • 2009 • 31 (4) 879

A
B
a b
a b
a b
G
E F
a
a
a
b b
b
FIG. 5. — Illustration of identification key for Onchobdella Paperna, 1968 species: A, shape of dorsal transverse bar (arched [a], straight
or V-shaped [b]); B, relative length of dorsal and ventral bars (ventral > dorsal [a], ventral < dorsal [b]); C, shape of penis (J-shaped [a],
sinuous or spirally coiled [b]); D, shape of penis (sinuous [a], spirally coiled [b]); E, shape and size of dorsal bar (long and V-shaped
[a], short and straight [b]); F, size of ventral anchor (small [a], large [b]); G, shape of the penis (G-shaped [a], J-shaped [b]).
880 ZOOSYSTEMA • 2009 • 31 (4)

A
B
a b
a b
C D
a b
a b
a b
FIG. 6. — Illustration of identification key for Scutogyrus Pariselle & Euzet, 1995 species: A, size of the penis (long [a], short [b]); B, size
of the penis (long [a], short [b]); C, shape of the vagina (tubular and thin-walled [a], large and thick-walled [b]); D, shape of the accessory
piece ending (two curved pincer-like hooks [a], one component hooked, the other blunt [b]); E, shape of the accessory piece (basal
extension well marked [a], no extension [b]). Scale bar: 30 μm.
ZOOSYSTEMA • 2009 • 31 (4) 881

29. Vagina not visible, single-looped penis (Fig. 7C, a) ............. Cichlidogyrus arthracanthus
— Sclerotised vagina, curved penis (Fig. 7C, b) ........................ Cichlidogyrus inconsultans
30. Very large body and sclerite size, large and S-shaped penis (Fig. 7D, a) ... Cichlidogyrus halli
— Other penis shapes (Fig. 7D, b) ................................................................................ 31
31. Male apparatus auxiliary plate(s) present (Fig. 8A, a) ................................................ 32
— Male apparatus without auxiliary plate (Fig. 8A, b) .................................................. 39
32. Presence of two small auxiliary plates (Fig. 8B, a) .......................... Cichlidogyrus guirali
— Presence of one auxiliary plate (Fig. 8B, b) ............................................................... 33
33. Reduced auxiliary plate (smallest axis < 10 μm) (Fig. 8C, a) ... Cichlidogyrus microscutus
— Large auxiliary plate (smallest axis > 10μm) (Fig. 8C, b) .......................................... 34
34. Accessory piece slightly S-shaped, ending bottle-opener-like (Fig. 8D, a) ......................
................................................................................................ Cichlidogyrus thurstonae
— Accessory piece bent at right angle (Fig. 8D, a) ........................................................ 35
35. Penis length less than 80 μm .................................................................................... 36
— Penis length more than 95 μm .................................................................................. 38
36. Extremity of accessory piece with large perpendicular diverticles, vagina thin walled and
amphora-shaped (Fig. 8E, a) .................................................... Cichlidogyrus aegypticus
— Extremity of accessory piece without large perpendicular diverticles (Fig. 8E, b) ...... 37
37. Accessory piece convex face with large semi-circular extension (Fig. 8F, a) .....................
....................................................................................................... Cichlidogyrus gallus
— Accessory piece convex face without large semi-circular extension (Fig. 8F, b) ...............
.................................................................................................... Cichlidogyrus paganoi
38. Penis length less than 105 μm, vagina length less than 50 μm ....................... C. bilongi
— Penis length more than 130 μm, vagina length more than 60 μm ................... C. agnesi
39. Long and narrow vagina (Fig. 9A, a) ........................................................................ 40
— Large and/or short vagina (Fig. 9A, b) ...................................................................... 42
40. Penis length more than 85 μm, ................................................. Cichlidogyrus flexicolpos
— Penis length less than 60 μm ................................................................................... 41
41. Accessory piece T-shaped (Fig. 9B, a) ...................................... Cichlidogyrus testificatus
— Accessory piece ending in a hook (Fig. 9B, b) ............................ Cichlidogyrus lemoallei
42. Short conical and right angle folded vagina (Fig. 9C, a) ........................................... 43
— Tubular vagina (almost constant diameter) (Fig. 9C, b) ............................................ 46
44. Accessory piece ending with smooth convex face (Fig. 9D, a) ........ Cichlidogyrus bouvii
— Accessory piece ending in a bifurcated hook with crenulated convex face (Fig. 9D, b) ...
..................................................................................................... Cichlidogyrus dossoui
45. Accessory piece ending folded back (Fig. 9E, a) ............................... Cichlidogyrus vexus
— Accessory piece ending in a hook (Fig. 9E, b) .......................... Cichlidogyrus douellouae
882 ZOOSYSTEMA • 2009 • 31 (4)

A B
III-VII III-VII I I
II
b II b
a
a
C D
a b
FIG. 7. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, length of uncinuli III to VII (long [a], short [b], pair
II is given as a reference); B, size of uncinuli I (large [a], small [b], pair II is given as a reference); C, C. arthracanthus Paperna, 1960
penis (a); C. inconsultans (Birgi & Lambert, 1986) sclerotised genitalia (b); D, shape of penis (C. halli (Price & Kirk, 1967) [a], other
species [b]). Abbreviations: I, II and III-VII, uncinuli numbering.
ZOOSYSTEMA • 2009 • 31 (4) 883

46. Short annulated vagina (Fig. 9F, a) .................................................. Cichlidogyrus hemi

— Vagina with smooth wall (Fig. 9F, b) ........................................................................ 47
47. Straight or slightly curved vagina (Fig. 10A, a) .......................... Cichlidogyrus legendrei
— Well curved or looped vagina (Fig. 10A, b) .............................................................. 48
48. Looped vagina (Fig. 10B, a) ..................................................................................... 49
— Curved vagina (not looped) (Fig. 10B, b) ................................................................. 51
49. Vagina forming a single loop (Fig. 10C, a) ............................... Cichlidogyrus tiberianus
— Vagina forming two semi-loops (Fig. 10C, b) ........................................................... 50
50. Vagina semi-loops in the same plane, crenulated distal accessory piece extremity (Fig. 10D,
a) ................................................................................................. Cichlidogyrus kouassii
— Vagina semi-loops in perpendicular planes, accessory piece distal extremity with a single
hook (Fig. 10D, b) ................................................................ Cichlidogyrus bonhommei
51. Curved accessory piece ending in two points, vagina with characteristic enlargement
(Fig. 10E, a) ...................................................................... Cichlidogyrus anthemocolpos
— Accessory piece bent at right angle, ending in a single point, vagina without enlargement
(Fig. 10E, b) ............................................................................................................. 52
52. Accessory piece with triangular extension at proximal third (Fig. 10F, a) .......................
...................................................................................................... Cichlidogyrus ergensi
— Accessory piece without triangular extension at proximal third (Fig. 10F, b) ..................
................................................................................................ Cichlidogyrus ouedraogoi
53. Auricles reduced, in continuity with dorsal bar anterior face (Fig. 11A, a) ................ 54
— Auricles developed (Fig. 11A, b) ............................................................................... 56
54. Uncinulus I “standardised length” more than 2 (Fig. 11B, a) ....... Cichlidogyrus kothiasi
— Uncinulus I “standardised length” less than 2 (Fig. 11B, b) ...................................... 55
55. Accessory piece with narrow base, penis with large heel (Fig. 11C, a) ... Cichlidogyrus berrebii
— Accessory piece with wide base, penis with thin heel (Fig. 11C, b) ... Cichlidogyrus pouyaudi
56. Uncinulus I “standardised length” more than 2 (Fig. 11D, a) ................................... 57
— Uncinulus I “standardised length” less than 2 (Fig. 11D, b) ...................................... 76
57. Penis with swollen portion (Fig. 11E, a) ................................................................... 58
— Penis without swollen portion (thin and tubular all along) (Fig. 11E, b) .................. 59
58. Penis length less than 35 μm ............................................. Cichlidogyrus papernastrema
— Penis length more than 45 μm .................................................. Cichlidogyrus philander
60. Sclerotised vagina ..................................................................................................... 61
— Vagina not visible ..................................................................................................... 62
61. Penis with developed heel, accessory piece S-shaped (Fig. 11F, a) ... Cichlidogyrus dageti
— Penis with poorly developed heel, accessory piece C-shaped (Fig. 11F, b) ......................
..................................................................................................... Cichlidogyrus falcifer
884 ZOOSYSTEMA • 2009 • 31 (4)

A B
a b a b
C D
a
a b
b
E F
a b
a b
FIG. 8. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, presence (a) or absence (b) of an auxiliary plate
associated with the male apparatus; B, presence of two (a) or one (b) auxiliary plate associated with the male apparatus; C, size of
the auxiliary plate associated with the male apparatus (small [a], large [b]); D, shape of the accessory piece (S-shaped [a], bent at right
angle [b]); E, shape of the extremity of the accessory piece (presence [a] or absence [b] of large perpendicular diverticles); F, shape
of the accessory piece convex face (presence [a] or absence [b] of a large semi-circular extension).
ZOOSYSTEMA • 2009 • 31 (4) 885

A B
a b
C D
a b
a b
a b
a b
FIG. 9. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, shape of the sclerotised vagina (long and narrow
[a], large and/or short [b]); B, shape of the accessory piece (T-shaped [a], ending in a hook [b]); C, shape of the vagina (short conical
and right angle folded [a], tubular [b]); D, shape of the accessory piece ending convex face (smooth [a], crenulated [b]); E, shape of
the accessory piece ending (folded back [a], hook shaped [b]); F, shape of vagina wall (annulated [a], smooth [b]).
886 ZOOSYSTEMA • 2009 • 31 (4)

A B C
b a b
a b
D E
a b
a b b
FIG. 10. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, shape of the vagina (straight or slightly curved
[a], well curved or looped [b]); B, shape of the vagina (looped [a], curved [b]); C, shape of vagina (one [a] or two semi [b] loops);
D, shape of vagina and accessory piece extremity (semi-loops in the same plane and crenulated [a], semi-loops in perpendicular
planes and single-hooked [b]); E, shape of accessory piece extremity and vagina (ending in two points and characteristic enlargement
[a], ending in a single point and no enlargement [b]); F, presence (a) or absence (b) of a triangular extension at proximal third of the
accessory piece.
ZOOSYSTEMA • 2009 • 31 (4) 887

62. Accessory piece forming a large gutter-like portion at its base (Fig. 12A, a) ...................
.................................................................................................... Cichlidogyrus nuniezi
— Narrow accessory piece (Fig. 12A, b) ........................................................................ 63
63. Accessory piece with a rounded bulge at distal third (Fig. 12B, a) ..................................
................................................................................................... Cichlidogyrus dionchus
— Accessory piece without a rounded bulge at distal third (Fig. 12B, b) ....................... 64
64. Accessory piece with double ending (Fig. 12C, a) ....................... Cichlidogyrus digitatus
— Accessory piece with single ending (Fig. 12C, b) ...................................................... 65
65. Accessory piece ending in a thin and long hook (Fig. 12D, a) .... Cichlidogyrus berradae
— Accessory piece ending in a wide and short hook (Fig. 12D, b) ................................ 66
66. Penis with large and rounded heel (Fig. 12E, a) ........................................................ 67
— Penis with narrow heel (Fig. 12E, b) ......................................................................... 68
67. Penis extremity folded back, accessory piece closed C-shaped (Fig. 12F, a) ....................
................................................................................................... Cichlidogyrus reversati
— Penis and accessory piece open C-shaped (Fig. 12F, b) ................. Cichlidogyrus halinus
68. Accessory piece > 27 μm (Fig. 12G, a) ........................................... Cichlidogyrus yanni
— Accessory piece < 26 μm (Fig. 12G, b) ........................................ Cichlidogyrus quaestio
69. Spirally coiled penis, length more than 350 μm (Fig. 13A, a) ................................... 70
— Not spirally coiled penis, length less than 150 μm (Fig. 13A, b) ............................... 72
70. Penis with 4 or 5 turns, not spirally coiled vagina (Fig. 13B, a) ...... Cichlidogyrus euzeti
— Penis with more than 4 or 5 turns, spirally coiled vagina (Fig. 13B, b) ..................... 71
71. Penis with 8 or 9 turns, length about 450 μm (Fig. 13C, a) ..... Cichlidogyrus longicirrus
— Penis with 14 or 15 turns, length about 1500 μm (Fig. 13C, b) ... Cichlidogyrus sanseoi
72. Penis with large diameter, slightly sinuous, without heel (Fig. 13D, a) ..........................
......................................................................................................... Cichlidogyrus arfii
— Filiform penis (Fig. 13D, b) ..................................................................................... 73
73. Nearly straight penis (Fig. 13E, a) ............................................... Cichlidogyrus albareti
— Curved penis (Fig. 13E, b) ....................................................................................... 74
74. G-shaped penis (Fig. 13F, a) ......................................................... Cichlidogyrus teugelsi
— Single-looped penis (Fig. 13F, b) .............................................................................. 75
75. Long vagina, accessory piece round ended (Fig. 14A, a) ............ Cichlidogyrus nandidae
— Short vagina, accessory piece ending in a hook (Fig. 14A, b) ... Cichlidogyrus bychowskii
76. Penis with swollen portion (Fig. 14B, a) ................................................................... 77
— Tubular penis without swollen portion (Fig. 14B, b) ................................................ 84
77. Well marked swollen portion (Fig. 14C, a) ............................................................... 78
— “Diffuse” swollen portion (Fig. 14C, b) .................................................................... 82
888 ZOOSYSTEMA • 2009 • 31 (4)

A B
a b
b
a
C D
b
a
b
a
a b
b
a
FIG. 11. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species; A, size of auricle on the dorsal bar (reduced [a],
developed [b]); B, relative size of uncinulus pair I (large [a], small [b]); C, shape of the accessory piece base and penis heel (narrow
and large [a], wide and thin [b]); D, relative size of uncinulus pair I (large [a], small [b]); E, presence (a) or absence (b) of a swollen
portion in the penis; F, size of the penis heel and of the accessory piece (well developed and S-shaped [a], poorly developed and
c-shaped [b]).
ZOOSYSTEMA • 2009 • 31 (4) 889

A B
a a b
b
C D E
a b a b a b
F G
a b a
FIG. 12. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, shape of the accessory piece base (gutter-like
[a], narrow [b]); B, presence (a) or absence (b) of a round bulge at distal third of the accessory piece; C, shape of the accessory piece
ending (double [a], single [b]); D, shape of the accessory piece ending hook (thin and long [a], wide and short [b]); E, shape of penis
heel (wide [a], narrow [b]); F, shape of penis extremity and accessory piece (folded back and close C [a], straight and open C [b]);
G, size of accessory piece (long [a], short [b]).
890 ZOOSYSTEMA • 2009 • 31 (4)

A B
a b
C
D
a b
E F
a
b
a b
FIG. 13. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, shape of the penis (spirally coiled [a], not spirally
coiled [b]); B, number of turns in spirally coiled penis and shape of the sclerotised vagina (4-5 and sinuous [a], more than 4-5 and
spirally coiled [b]); C, number of turns in spirally coiled penis (8-9 [a], 14-15 [b]); D, penis diameter (large [a], filiform [b]); E, shape of
the penis (straight [a], curved [b]); F, shape of the penis (G [a], single looped [b]).
ZOOSYSTEMA • 2009 • 31 (4) 891

79. No visible vagina, developed swollen portion (more than half penis total length) (Fig. 14D,
a) ................................................................................................ Cichlidogyrus sanjeani
— Visible S-shaped vagina, reduced swollen portion (less than one fifth of penis total length)
(Fig. 14D, b) ............................................................................................................ 80
80. Accessory piece ending in a hook (Fig. 14E, a) ...................... Cichlidogyrus amphoratus
— Accessory piece bifurcated, lined along convex face with tubercles (Fig. 14E, b) ............
.................................................................................................... Cichlidogyrus ornatus
81. Accessory piece with heavily sclerotised portion L-shaped, uncinuli pairs II and III to VII
(11 and 13 μm) (Fig. 14F, a) ........................................................ Cichlidogyrus giostrai
— Accessory piece with heavily sclerotised portion C-shaped, uncinuli pairs II and III to VII
(16 and 20 μm) (Fig. 14F, b) .......................................................... Cichlidogyrus njinei
82. Penis length less than 40 μm .................................................... Cichlidogyrus lagoonaris
83. Short, large and conical vagina (Fig. 14G, a) ......................... Cichlidogyrus zambezensis
— Short, thin and S-shaped vagina (Fig. 14G, b) ............................. Cichlidogyrus karibae
85. Sclerotised vagina ..................................................................................................... 86
— Vagina not visible ..................................................................................................... 88
86. Single C-shaped accessory piece with a thin extension at its base (Fig. 15A, a) ..............
.................................................................................................... Cichlidogyrus levequei
— Complex accessory piece with well-sclerotised C-shaped portion (Fig. 15A, b) ......... 87
87. Accessory piece with a C-shaped portion sharply arched, with thin plates at each extremity
(Fig. 15B, a) .......................................................................... Cichlidogyrus slembroucki
— Accessory piece with C-shaped portion slightly arched, without plates at its extremities,
and a digitations at its middle (Fig. 15B, b) .................................. Cichlidogyrus acerbus
88. Penis with heel (Fig. 15C, a) .................................................................................... 89
— Penis without heel (Fig. 15C, b) ............................................................................... 91
89. Very thin heel, accessory piece ending in claws (Fig. 15D, a) ...... Cichlidogyrus fontanai
— Developed heel, accessory piece with forked ending (Fig. 15D, b) ............................ 90
90. Penis length less than 40 μm, large dorsal bar auricles (Fig. 15E, a) ...............................
.................................................................................................... Cichlidogyrus rognoni
— Penis length more than 40 μm, dorsal bar auricles smaller (Fig. 15E, b) ........................
................................................................................................. Cichlidogyrus bifurcatus
91. Penis length more than 40 μm, accessory piece thin at level of insertion on basal bulb
(Fig. 15F, a) ......................................................................... Cichlidogyrus haplochromii
— Penis length less than 37 μm, accessory piece large at level of insertion (Fig. 15F, b) ......
.................................................................................................... Cichlidogyrus tilapiae
92. Accessory piece straight or slightly curved (Fig. 16A, a) ............................................ 93
— Accessory piece sharply curved (Fig. 16A, b) ............................................................ 95
892 ZOOSYSTEMA • 2009 • 31 (4)

A B
a b a
C D
b a b
a b
F G
a b
FIG. 14. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, size of the sclerotised vagina and shape of
accessory piece ending (long and rounded [a], short and hooked [b]); B, presence (a) or absence (b) of a swollen portion in the penis;
C, well marked (a) or diffuse (b) swollen portion in the penis; D, relative size of the swollen portion in the penis (developed [a], reduced [b]);
E, shape of accessory piece ending (hooked [a], bifurcated [b]) and presence of tubercles along the convex face; F, shape of accessory
piece heavily sclerotised portion (L [a], C [b]); G, shape of the sclerotised vagina (large and conical [a], thin and S-shaped [b]).
ZOOSYSTEMA • 2009 • 31 (4) 893

A B
a b
a b
D
a
a b
E F
a b
FIG. 15. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, shape of the accessory piece (simple with
extension [a], complex [b]); B, presence (a) or absence (b) of thin plates at the extremity of the accessory piece; C, presence (a) or
absence (b) of a penis heel; D, size of the penis heel and shape of the accessory piece ending (thin and presence of claws [a], developed
and forked [b]); E, size dorsal bar auricles (large [a], small [b]); F, size of the penis and width of accessory piece insertion at the level
of the penis (more than 40 μm and thin [a], less than 37 μm and large [b]).
894 ZOOSYSTEMA • 2009 • 31 (4)

A B
a b
a b
C D
a
a b b
a b
FIG. 16. — Illustration of identification key for Cichlidogyrus Paperna, 1960 species: A, shape of accessory piece (straight [a], sharply
curved [b]); B, absence (a) or presence (b) of a penis heel; C, shape of the sclerotised vagina (straight and large [a], sinuous and thin
[b]); D, shape of the accessory piece ending and of the penis (rounded and looped [a], forked and C [b]); E, accessory piece extremities
shape and length (smooth and equal [a], rough and unequal [b]).
ZOOSYSTEMA • 2009 • 31 (4) 895

93. Penis with no visible heel (Fig. 16B, a) ..................................... Cichlidogyrus longipenis
— Penis with developed heel (Fig. 16B, b) .................................................................... 94
94. Straight and large vagina, penis length 70 μm (Fig. 16C, a) ......... Cichlidogyrus sclerosus
— Long, thin and sinuous vagina, penis length 145 μm (Fig. 16C, b) ... Cichlidogyrus cirratus
95. Accessory piece with rounded ending, looped penis (Fig. 16D, a) ... Cichlidogyrus amieti
— Accessory piece forked ending, C-shaped penis (Fig. 16D, b) ................................... 96
96. Extremities of accessory piece smooth and about equal in length (Fig. 16E, a) ..............
..................................................................................................... Cichlidogyrus cubitus
— Extremities of accessory piece highly unequal in length, the longer one with rough patch
on its convex face (Fig. 16E, b) .............................................. Cichlidogyrus louipaysani
REFERENCES Urogyrus cichlidarum gen.nov., sp.nov., Urogyridae

fam.nov., monogène parasite de la vessie urinaire de
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morhua L. in the Oslo Fjord, Norway. Systematic poissons des eaux douces du Cameroun. Présence des
Parasitology 33: 199-207. genre Cichlidogyrus et Dactylogyrus chez Aphyosemion
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von Enterogyrus hemihaplochromii n. sp. (Trematodes, de France 108: 101-106.
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Zoologisches Institut der Universität Hohenheim, Monopisthocotylea, Ancyrocephalidae). Description
Hohenheim, Germany. de Cichlidogyrus nandidae n. sp. Annales de Parasitologie
BILONG BILONG C. F. 1988. — Enterogyrus malmbergi humaine et comparée 61: 521-528.
n. sp. (Monogenea-Ancyrocephalidae) parasite de BIRGI E. & LAMBERT A. 1987. — À propos de Cichlidogyrus
l’estomac du Cichlidae Tilapia nilotica Linné, 1757 euzeti Birgi & Lambert, 1986 décrit au Cameroun
au Sud-Cameroun. Annales de la Faculté des Sciences chez un Nandidae Polycentropsis abbreviata Boulenger,
de Yaoundé, Biologie-Biochimie 5: 51-58. 1901. Annales de Parasitologie humaine et comparée
BILONG BILONG C. F., BIRGI E. & LAMBERT A. 1989. — 62: 103.
Enterogyrus melenensis n. sp. (Monogenea, Ancyro- CALTRAN H., SILAN P. & ROUX M. 1995. — Ligophorus
cephalidae), parasite stomacal de Hemichromis fasciatus imitans (Monogenea) ectoparasite de Liza ramada
Peters, 1857 (Téléostéen, Cichlidae) du Sud-Cameroun. (Teleostei) I. Populations naturelles et variabilité
Revue de Zoologie africaine – Journal of African Zoology morphologique. Écologie 26: 95-104.
103: 99-105. CONE D. K., GRATZEK J. B. & HOFFMAN G. L. 1987. — A
BILONG BILONG C. F. & EUZET L. 1995. — Onchobdella study of Enterogyrus sp. (Monogenea) parasiting the
bopeleti n. sp. (Monogenea, Ancyrocephalidae) parasite foregut of captive Pomacanthus paru (Pomacanthidae) in
branchial de Hemichromis fasciatus (Peters, 1857) Georgia. Canadian Journal of Zoology 65: 312-316.
(Cichlidae). Journal of African Zoology 109: 253-258. DOSSOU C. 1973. — Recherches sur les Monogènes para-
BILONG BILONG C. F., EUZET L. & BIRGI E. 1996. — sites des poissons d’eau douce du Sud Dahomey. Ph.D.
Monogenean stomach parasites of cichlid fishes from Thesis, Université des Sciences et Techniques du
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BILONG BILONG C. F., BIRGI E. & EUZET L. 1991. — du Bénin III. Espèces nouvelles du genre Cichlidogyrus
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BILONG BILONG C. F., BIRGI E. & EUZET L. 1994. — 116 p.
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DOSSOU C. & BIRGI E. 1984. — Monogènes parasites PAPERNA I. 1968. — Monogenetic Trematodes col-
d’Hemichromis fasciatus Peters, 1857 (Teleostei, lected from fresh water fish in Ghana. Second report.
Cichlidae). Annales des Sciences naturelles, Zoologie Bamidgeh, Bulletin of Fish Culture in Israel 20: 80-
6: 101-109. 100.
DOUËLLOU L. 1993. — Monogeneans of the genus PAPERNA I. 1969. — Monogenetic Trematodes of the
Cichlidogyrus Paperna, 1960 (Dactylogyridae: An- fish of the Volta basin and South Ghana. Bulletin de
cyrocephalinae) from cichlid fishes of Lake Kariba l’I.F.A.N. 31: 840-880.
(Zimbabwe) with descriptions of five new species. PAPERNA I. 1973. — New species of Monogenea (Vermes)
Systematic Parasitology 25: 159-186. from African freshwater fish. A preliminary report.
EID N. & NEGM M. 1987. — Some morphological Revue de Zoologie et de Botanique africaines 87: 505-
study on a new species of endoparasitic monogenetic 518.
Trematode Enterogyrus niloticus in the intestine of PAPERNA I. 1979. — Monogenea of inland water fish
Tilapia nilotica. Journal of the Egyptian Veterinary in Africa. Annales du Musée royal d’Afrique centrale,
Medical Association 47: 79-86. sér in-8° (Zool.) 226: 1-131.
ERGENS R. 1981. — Nine species of the genus Cichlidogyrus PAPERNA I. & THURSTON J. P. 1969. — Monogenetic
Paperna, 1960 (Monogenea: Ancyrocephalinae) from Trematodes collected from cichlid fish in Uganda;
Egyptian fishes. Folia Parasitologica 28: 205-214. including the description of five new species of Cich-
ERGENS R. & GELNAR M. 1985. — Experimental verifica- lidogyrus. Revue de Zoologie et de Botanique africaines
tion of the effect of temperature on the size of hard parts 79: 15-33.
of opistohaptor of Gyrodactylus katharineri Malmberg, PARISELLE A. 1995. — Études des parasites de Cichlidae
1964. Folia Parasitologica 32: 377-380. en Afrique de l’Ouest, in AGNÈSE J.-F. (ed.), Comptes
FROESE R. & PAULY D. (eds) 2009. — FishBase. World rendus de l’atelier biodiversité et aquaculture. CRO
Wide Web electronic publication. http://www.fishbase. Abidjan: 44-52.
org (September 2009). PARISELLE A. 1996. — Diversité, spéciation et évolution
GUSSEV A. V. & FERNANDO C. H. 1973. — Dactylogy- des Monogènes branchiaux de Cichlidae en Afrique de
ridae (Monogenoidea) from stomach of Fishes. Folia l’Ouest. PhD Thesis, Université de Montpellier II,
Parasitologica (Praha) 20: 207-212. Montpellier, France, 199 p.
KHIDR A. A. 1990. — Population dynamics of Enterogyrus PARISELLE A., BILONG BILONG C. F. & EUZET L. 2003. —
cichlidarum (Monogenea: Ancyrocephalinae) from Four new species of Cichlidogyrus Paperna, 1960
the stomach of Tilapia spp. in Egypt. International (Monogenea, Ancyrocephalidae) all gill parasites
Journal for Parasitology 20: 741-747. from African mouthbreeder tilapias of the genera
LIM L. H. S., TIMOFEEVA T. A. & GIBSON D. I. 2001. — Sarotherodon and Oreochromis (Pisces, Cichlidae),
Dactylogyridean monogeneans of the siluriform with a re-description of C. thurstonae Ergens, 1981.
fishes of the Old World. Systematic Parasitology 50: Systematic Parasitology 56: 201-210.
159-197. PARISELLE A. & EUZET L. 1994. — Three new spe-
MARKEVICH A. P. 1934. — [Parasitic diseases of fish cies of Cichlidogyrus Paperna, 1960 (Monogenea,
and their control]. Publ. Koiz., Leningrad, 100 p. Ancyrocephalidae) parasitic on Tylochromis jentinki
(in Russian). (Steindachner, 1895) (Pisces, Cichlidae) in West
N’DOUBA V., THYS VAN DEN AUDENAERDE D.F.E. & Africa. Systematic Parasitology 29: 229-234.
PARISELLE A. 1997. — Description d’une espèce PARISELLE A. & EUZET L. 1995a. — Trois Monogènes
nouvelle de Monogène ectoparasite branchial de Tilapia nouveaux parasites branchiaux de Pelmatochromis buet-
guineensis (Bleeker, 1862) (Cichlidae) en Côte d’Ivoire. tikoferi (Steindachner, 1895) (Cichlidae) en Guinée.
Journal of African Zoology 111: 429-433. Parasite 2: 203-209.
PAPERNA I. 1960. — Studies on Monogenetic Trematodes PARISELLE A. & EUZET L. 1995b. — Scutogyrus n. g.
in Israel. 2 Monogenetic Trematodes of Cichlids. (Monogenea, Ancyrocephalidae) for Cichlidogyrus
Bamidgeh, Bulletin of Fish Culture in Israel 12: 20- longicornis minus Dossou, 1982, C. longicornis and
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n. sp., a monogenetic Trematode parasitic in the Cichlids. Journal of the Helminthological Society of
intestine of Fish. The Bulletin of the Research Council Washington 62: 157-173.
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Submitted on 5 August 2008;

accepted on 17 December 2008.
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Systematic revision of dactylogyridean parasites

(Monogenea) from cichlid fishes in Africa, the

Pariselle A. & Euzet L. 2009. — Systematic revision of dactylogyridean parasites (Monogenea)

INTRODUCTION CAS Institute of Parasitology, Czechoslovak Acad-

850 ZOOSYSTEMA • 2009 • 31 (4)

Enterogyrus cichlidarum Paperna, 1963

ZOOSYSTEMA • 2009 • 31 (4) 851

852 ZOOSYSTEMA • 2009 • 31 (4)

REMARKS TYPE HOST. — Paratilapia polleni Bleeker, 1868.

Enterogyrus sp. ADDITIONAL LOCALITIES. — Near Majunga, West of

REMARKS Insulacleidus paretropli Rakotoﬁringa & Euzet, 1983:

Genus Insulacleidus Rakotoﬁringa & Euzet, 1983 SITE. — Gills.

TYPE SPECIES. — Insulacleidus paratilapiae Rakotoﬁringa &

Insulacleidus paratilapiae Genus Onchobdella Paperna, 1968

ZOOSYSTEMA • 2009 • 31 (4) 853

1995; O. pterigyalis Paperna, 1968; O. silverai Pariselle & REMARKS

854 ZOOSYSTEMA • 2009 • 31 (4)

MRAC 37.365 A and 37.366 B. Onchobdella spirocirra Paperna, 1968

ADDITIONAL SITE. — Gills. OTHER SPECIES INCLUDED. — Scutogyrus bailloni Pariselle

ZOOSYSTEMA • 2009 • 31 (4) 855

Paratype: MNHN Tj142. of a sampling error [mix of samples or contamination] or

Scutogyrus longicornis Cichlidogyrus longicornis gravivaginus Paperna & Thurston,

856 ZOOSYSTEMA • 2009 • 31 (4)

ZOOSYSTEMA • 2009 • 31 (4) 857

Cichlidogyrus aegypticus Ergens, 1981

858 ZOOSYSTEMA • 2009 • 31 (4)

MRAC 37.408. Cichlidogyrus anthemocolpos Dossou, 1982

Cichlidogyrus amphoratus Cichlidogyrus berradae Pariselle & Euzet, 2003

ZOOSYSTEMA • 2009 • 31 (4) 859

SITE. — Gills. TYPE LOCALITY. — Tiassalé, Bandama River, Guinea.

ADDITIONAL LOCALITIES. — Kogon River, Guinea and SITE. — Gills.

860 ZOOSYSTEMA • 2009 • 31 (4)

Cichlidogyrus bychowskii (Markevich, 1934) REMARKS

TYPE HOST. — Of C. cirratus: Sarotherodon galilaeus SITE. — Gills.

TYPE LOCALITY. — Of C. cirratus Tiberias Lake, Israel.

ZOOSYSTEMA • 2009 • 31 (4) 861

862 ZOOSYSTEMA • 2009 • 31 (4)

Cichlidogyrus douellouae Pariselle, Bilong Bilong & Euzet,

SITE. — Gills. TYPE LOCALITY. — Benin and Cameroon.

ADDITIONAL LOCALITIES. — Bama, Kou River, Burkina SITE. — Gills.

ZOOSYSTEMA • 2009 • 31 (4) 863

Cichlidogyrus gallus Pariselle & Euzet, 1995 SITE. — Gills.

864 ZOOSYSTEMA • 2009 • 31 (4)

SITE. — Gills. 1933) and Oreochromis spilurus spilurus (Günther, 1894).

ZOOSYSTEMA • 2009 • 31 (4) 865

866 ZOOSYSTEMA • 2009 • 31 (4)

ZOOSYSTEMA • 2009 • 31 (4) 867

Cichlidogyrus levequei Pariselle & Euzet, 1996

Cichlidogyrus longicirrus Paperna, 1965: 108, 109, 111. —

868 ZOOSYSTEMA • 2009 • 31 (4)

Cichlidogyrus njinei ADDITIONAL LOCALITIES. — Bagoué River, Côte d’Ivoire on

Cichlidogyrus nuniezi Pariselle & Euzet, 1998: 276-278,

ZOOSYSTEMA • 2009 • 31 (4) 869

Cichlidogyrus papernastrema Cichlidogyrus quaestio Douëllou, 1993

Cichlidogyrus philander Douëllou, 1993 REMARKS

TYPE HOST. — Pseudocrenilabrus philander philander

870 ZOOSYSTEMA • 2009 • 31 (4)

Cichlidogyrus sanseoi Pariselle & Euzet, 2004: 360-362,

SITE. — Gills. TYPE HOST. — Tilapia buttikoferi (Hubrecht, 1881).