Parasitol Res (2012) 110:1185–1200

DOI 10.1007/s00436-011-2612-0

ORIGINAL PAPER

An overview of the Gyrodactylus (Monogenea:

Gyrodactylidae) species parasitizing African catfishes,
and their morphological and molecular diversity
Iva Přikrylová & Radim Blažek & Maarten P. M. Vanhove

Received: 2 August 2011 / Accepted: 4 August 2011 / Published online: 18 August 2011
# Springer-Verlag 2011

Abstract An overview of Gyrodactylus infecting catfishes studied in Senegal, two more species, Gyrodactylus trans-
from the African continent is provided, including new data vaalensis and Gyrodactylus gelnari n. sp. were recognised.
from Sudan, Senegal, Kenya and Mozambique. Haptoral In addition, Gyrodactylus turkanaensis n. sp. from the gills
sclerite morphometry and nuclear ribosomal DNA sequen- of Kenyan C. gariepinus was described and an undescribed
ces revealed the presence of eight Gyrodactylus species. On Gyrodactylus sp. was recorded from Sudanese representa-
Senegalese Synodontis nigrita, Gyrodactylus synodonti n. tives of the same host. Detailed morphometrical and
sp. and Gyrodactylus nigritae n. sp. are described. These molecular comparisons of the species are presented and
are the first reports of gyrodactylid parasites from mocho- discussed. The study highlights the hitherto understudied
kid hosts. From the fins of North African catfish Clarias diversity of viviparous monogenean parasites throughout
gariepinus collected in Mozambique, Gyrodactylus alekosi Africa.
n. sp. and Gyrodactylus rysavyi were identified. G. rysavyi
was also reported from Kenyan C. gariepinus and Senegal-
ese Clarias anguillaris. From the fins of C. anguillaris Introduction

With over 3,100 valid species, the catfishes (order

Siluriformes) are among the most successful teleosts
I. Přikrylová (*) : R. Blažek
(Sabaj et al. 2003). The numerous recent discoveries
Department of Botany and Zoology, Faculty of Science,
Masaryk University, demonstrate that their biodiversity is yet far from
Kotlářská 2, understood (Lundberg et al. 2000). Several representatives
61137 Brno, Czech Republic of the order are of economic value in view of their
e-mail: ivaprik@sci.muni.cz
importance in aquaculture, with Clarias gariepinus
R. Blažek (Burchell, 1822) as prime African example (Haylor
Department of Fish Ecology, Institute of Vertebrate Biology, 1993; Chocha Manda 2010). Their biological and phylo-
Academy of Sciences of the Czech Republic, genetic diversity, as well as their predominantly freshwater
Květná 8,
habitat and global distribution, render siluriforms a key
603 65 Brno, Czech Republic
target group in ecological, evolutionary and biogeographic
M. P. M. Vanhove research (Lundberg et al. 2007). Another fruitful scientific
Laboratory of Animal Diversity and Systematics, approach can be catfish parasitology, certainly when
Department of Biology, Katholieke Universiteit Leuven,
applied to monogenean flatworm parasites, in view of
Charles Deberiotstraat 32,
3000 Leuven, Belgium their close relationship to their host species because of
their one-host lifecycle and high host specificity. Indeed,
M. P. M. Vanhove research on Monogenea from catfish has proven useful
Ichtyology Unit, African Zoology Department,
both in disentangling monogenean speciation mechanisms
Royal Museum for Central Africa,
Leuvensesteenweg 13, (Pariselle et al. 2003) and in reconstructing catfish
3080 Tervuren, Belgium biogeographical history (Barson et al. 2010). Moreover,
1186 Parasitol Res (2012) 110:1185–1200

several monogenean genera have proven to be pathogenic were collected from catfishes of the families Clariidae and
to economically important cultured catfish. Examples Mochokidae. Two flatworm species, parasitizing clariid
include Quadriacanthus Paperna, 1961 and Gyrodactylus hosts, could be assigned to G. transvaalensis and G. rysavyi
von Nordmann, 1832 (Kabata 1985; Obiekezie and Taege on the basis of haptoral hard part morphology. The
1991; Paperna 1996). remaining six represent new Gyrodactylus spp., the
Monogeneans belonging to Gyrodactylus are viviparous descriptions of which are presented here. The internal
skin and gill parasites of many freshwater and marine transcribed spacer (ITS) region of the nuclear rDNA is the
fishes. Only 28 Gyrodactylus species have been described molecular marker complementing morphological character-
from African fishes, of which only seven are known from isation of the species. Since its first application in
catfishes (Christison et al. 2005; Nack et al. 2005; Gyrodactylus taxonomy (Cunningham et al. 1995), the
Přikrylová et al. 2009a; Vaughan et al. 2010; Vanhove et ITS region has widely been used in combination with
al. 2011a; García-Vásquez et al. 2011). This is a limited species descriptions, because of the variability in its
number compared to the more than 400 valid Gyrodactylus subregions ITS-1 and ITS-2 (Cable et al. 2005; Rokicka
spp. worldwide (Harris et al. 2004), or to the 78 species of et al. 2009; Mullen et al. 2010; Vanhove et al. 2011a). The
Cichlidogyrus Paperna, 1960 in Africa, Madagascar and the present study provides new data about monogenean parasite
Levant (Pariselle and Euzet 2009; Vanhove et al. 2011b; diversity in Africa, and complements what little informa-
Gillardin et al. 2011). C. gariepinus is known to host five tion, if any, is available from the studied areas (Khalil and
African Gyrodactylus species. Two of them, Gyrodactylus Polling 1997).
alberti Paperna, 1973 and Gyrodactylus clarii Paperna,
1973 where described from Uganda (Paperna 1973);
another two, Gyrodactylus rysavyi Ergens, 1973 and Materials and methods
Gyrodactylus groschafti Ergens, 1973 were contemporane-
ously described from Egypt (Ergens 1973); and finally, Specimen collection and preparation
South Africa saw the description of Gyrodactylus trans-
vaalensis Prudhoe and Hussey, 1977. Another clariid host, Monogenean parasites belonging to Gyrodactylus were col-
Gyrodactylus camerunensis Lönnberg, 1895 yielded two lected from the catfishes Synodontis nigrita Valenciennes,
new gyrodactylid species in Cameroun: G. camerunensis 1840 (Teleostei: Siluriformes: Mochokidae), C. gariepinus
Nack, Bilong Bilong and Euzet, 2005 and Gyrodactylus and Clarias anguillaris (Linnaeus, 1758) (Teleostei: Siluri-
nyongensis Nack, Bilong Bilong and Euzet, 2005. Clarias formes: Clariidae) during parasitological investigations of
jaensis Boulenger, 1909 and Clarias pachynema Bou- freshwater fishes in four African countries carried out between
lenger, 1903 were mentioned as additional host fishes for November 2004 and January 2010. Details on the localities
these two Gyrodactylus species. So far, no other records and collection dates are provided in Table 1. Parasites were
exist of Gyrodactylus parasitizing African catfishes. removed from the hosts’ fins, body surface and gills. Their
During a survey of fish parasites on several localities in haptors were excised, fixed with ammonium picrate–glycerine
four different African countries, eight Gyrodactylus spp. (Malmberg 1970) and mounted on a slide for subsequent

Table 1 Localities, collection period and number of Gyrodactylus spp. analysed in the present study (N = number of specimens used for
morphometric study, n = number of specimens analysed by molecular methods)

Country Locality Host species Date of collection N/n

Senegal Niokolo Koba National Park S. nigrita March 2007 58/7

Niokolo Koba River (13°03.92′ N, 13°10.14′ W)
Niokolo Koba National Park C. anguillaris March 2006, 2007 and 2008 29/3
Mare Simenti (13°01.79′ N, 13°17.6′ W)
Kenya Lake Turkana, Kalokol C. gariepinus September 2008 21/0
(03°33.58′ N, 35°55.24′ E)
Lake Turkana, Todonyang village C. gariepinus September 2008 and 2009 11/0
(4°26.15′ N, 35°56.38′ E)
Mozambique temporal stream close to the village C. gariepinus February 2009 11/11
Bala Bala (24°19.34′ S, 33°02.36′ E)
Sudan Sennar, Blue Nile C. gariepinus January 2010 2/2
(13°32.81′ N, 33°38.17′ E)
Parasitol Res (2012) 110:1185–1200 1187

morphological analysis. The anterior ends of the parasite schemes. For trimming the resulting alignment, trimAl v.1.2
bodies were stored in SPECTRANAL absolute ethanol (Capella-Gutiérrez et al. 2009) was used. The optimal
(Allied-Signal, Riedel-de Haën). Specimens collected in model of molecular evolution was estimated by jModelTest
Sudan were directly fixed in absolute ethanol and slides were v.0.1.1 (Posada 2008; see also Guindon and Gascuel 2003,
prepared in the laboratory following Rokicka et al. (2007). Felsenstein 2005). Based on the corrected Akaike Informa-
tion Criterion (Hurvich and Tsai 1989), the TVM (Posada
Morphometric analysis 2003) + Γ model was selected. To allow subsequent
implementation in phylogenetic software, the model with
Morphological analysis of the collected parasite specimens the second best corrected Akaike score was chosen, namely
was performed in the Laboratory of Parasitology, Department the GTR (Tavaré 1986; Rodriguez et al. 1990) + Γ model,
of Botany and Zoology, Masaryk University, Brno, Czech with a gamma-shape parameter of 0.2. Reconstruction of a
Republic, based on morphology of the hard parts using a model-averaged phylogeny showed that tree topology did
phase-contrast microscope (Olympus BX51). Hard parts were not display phylogenetic uncertainty because of model
drawn with the aid of a drawing attachment. Measurements of selection. Hence, model choice should not influence the
hamuli and bars are those suggested by Přikrylová et al. results. In PhyML v.3.0 (Guindon and Gascuel 2003), a
(2008), and body size parameters and marginal hook maximum likelihood (ML) search was performed under the
measurements were taken based on Christison et al. (2005). optimised model. Nodal support was assessed through
All measurements are in micrometer, unless otherwise stated, 1,000 bootstrap samples using the nearest-neighbour
and are presented as the mean with the range in parentheses. interchange branch swapping algorithm. For the maximum
For comparative investigation of African Gyrodactylus parsimony (MP) method, PAUP* v.4.01b (Swofford 2001)
species, type material was obtained as reported in was used with the PaupUp interface (Calendini and Martin
Přikrylová et al. (2009b) and studied by the principal 2005). A heuristic search was carried out using 1,000
author. Principal Component Analysis (PCA) was carried replicates of nearest-neighbour interchange branch swapping;
out on the correlation matrix of haptor morphometrics gaps were treated as fifth state. Pairwise genetic distances,
using PAST (Hammer et al. 2001). according to the selected model, were calculated by the same
software. Bayesian inference (BI), also using the GTR + Γ
DNA extraction and amplification model, was implemented in MrBayes v.3 (Huelsenbeck and
Ronquist 2001, Ronquist and Huelsenbeck 2003). Posterior
Preservation ethanol was evaporated in a vacuum probabilities were calculated over 1.106 generations,
centrifuge, after which DNA was extracted using the sampling the Markov chain every 100 generations. One-
Qiagen Blood and Tissue Isolation kit, according to the fourth of the samples was discarded as “burn-in”. Conver-
manufacturer’s protocol. DNA was eluted in 50 μl. The sion of alignment files was carried out using ALTER v.1.2
ITS region of the rDNA was amplified with the primers (Glez-Peña et al. 2010) and a tree was drawn in FigTree v.1.3
ITS-1F (5′-GTTTCCGTAGGTGAACCT-3′; Rokicka et (http://tree.bio.ed.ac.uk/software/figtree).
al. 2007) and ITS-2R (5′-TCCTCCGCTTAGTGATA-3′;
Matějusová et al. 2001), in a Mastercycler eP gradient
thermocycler (Eppendorf). Each amplification reaction Results
contained 1 μl of template DNA, 1× PCR buffer,
1.25 mM MgCl2, 100 μM dNTPs, 0.1 mg/ml μl bovine Gyrodactylus parasites were found on three catfish species,
serum albumin, 0.5 μM of each primer (Generi Biotech) Synodontis nigrita, C. gariepinus and C. anguillaris. Eight
and 1.5 U of Taq polymerase in a total volume of 20 μl. Gyrodactylus spp. were identified. From Senegelase S.
Five microliters of PCR product was visualised on Gold nigritae, two new species, G. nigritae n. sp. and G.
View stained agarose gel (1%) and the remaining 15 μl was synodonti n. sp., were described. These two species infect
purified using the High Pure PCR Product Purification Kit different sites of the host’s body. While specimens of G.
(Roche). Sequencing, using identical primers as in initial nigritae n. sp. were only collected on barbels, G. synodonti
amplification, was carried out with the Big Dye Chemistry n. sp. specimens were consistently found on fins. These
Cycle Sequencing Kit v.3.1 and an ABI 3130 Genetic species are the first Gyrodactylus representatives recorded
Analyser automated sequencer (Applied Biosystems). on mochokid catfish. On Senegalese C. anguillaris, two
known gyrodactylid species were observed, namely G.
Sequence alignment and phylogenetic analyses rysavyi and G. transvaalensis. A third one appeared to be a
new species based on the morphology of its haptoral parts,
Sequences were aligned using MUSCLE v.3.8 (Edgar 2004) and is described as Gyrodactylus gelnari n. sp. The only
under default distance measures and sequence weighting Senegalese catfish Gyrodactylus representative from which
1188 Parasitol Res (2012) 110:1185–1200

molecular data were successfully obtained was G. rysavyi. A single pair of cephalic lobes. Dimensions of the haptoral
different gyrodactylid parasite altogether was found on sclerites are given in Table 2. Hamuli sturdy, with flattened
Kenyan C. gariepinus. Morphological characters and metrics area on inner part of root (Fig. 1a). Simple dorsal bar.
suggest that it represents a new species, hereafter described Ventral bar ribbed at bar proper and bar membrane. Robust
as Gyrodactylus turkanaensis n. sp. On Mozambican C. shaft of marginal hook sickle rises straight from the base
gariepinus, G. rysavyi and another unknown Gyrodactylus and curves regularly (Figs. 1b, 2c). Sickle base with short
were identified. Haptoral morphology, supported by the blunt toe and pronounced heel slanted with rounded
molecular data, demonstrate the latter represents a new terminal edge.
species, described here as Gyrodactylus alekosi n. sp. Molecular characterisation. A 930-bp fragment covering
Sampling in Sudan revealed the presence of gyrodactylid ITS-1 (388 bp), 5.8S rDNA (156 bp), ITS-2 (383 bp) and
parasites on C. gariepinus fins, of which only two specimens 28S (3 bp) was successfully sequenced from four speci-
could be retrieved and sequenced. While molecular and mens and submitted to GenBank under accession number
morphological characterisations of these specimens strongly FR850682. The entire sequence was identical for the four
support their assignment to another new species, a formal specimens. A BlastN (Zhang et al. 2000) search in
species description could not be prepared due to lack of GenBank in September 2010 using the entire sequence
material. Morphological descriptions and (when applicable) revealed no identical or close hits.
molecular characterisation of all species analysed are Type host: C. gariepinus (Burchell, 1822) (Siluriformes:
provided, in alphabetical order, below. Clariidae)
Type locality: temporal stream close to Bala Bala village
(24°19.34′ S, 33°02.36′ E), southern Mozambique.
Descriptions Infection site: Fins.
Type material: Collected February 2009. Holotype and two
Gyrodactylus alekosi n. sp. paratypes deposited in the Institute of Parasitology, Biology
Description based on seven excised haptors. Hence, body Centre, Academy of Sciences of the Czech Republic in České
dimensions nor internal morphology could be investigated. Budějovice, Czech Republic (Accession number M-518).
The external morphology was partially observed. Body Etymology: The specific epithet honours Alekos, a special
elongate with clearly separate haptor, prohaptor with a friend of the first author.

Table 2 Comparison of the measurements (in μm, average with range in parentheses) of the haptoral hard parts of Gyrodactylus spp. collected
from S. nigrita and C. gariepinus from the current study with the measurements taken on paratype material of G. rysavyi

Measurement G. alekosi n. sp G. gelnari n. sp. G. nigritae n. sp. G. rysavyi (n=25) G. rysavyi (n=3)
(n=7) present study (n=22) present study (n=35) present study present study paratypes re-examination

HTL 51.1 (50.5–53)m 71.4 (69–75) 45.5 (43–51.5) 98.6 (89–106.5) 91.8 (90–93)
HPL 26.4 (25–27.5) 35.7(33.5–38) 26.3 (25–28.3) 48.9 (45.5–53) 46.4 (45–47.5)
HSL 29 (28.5–30) 44.4 (42.5–47) 28.5 (27–31) 59.7 (54.5–64) 57.8 (57.5–58.5)
HRL 26.3 (25–29)m 34.4 (31.5–37.5) 23.4 (19–30) 49 (38–57) 42.5 (40–44)
VBL 5.3 (4.5–5.5)n 6.6 (5.5–8)g 4.3 (3.5–5.5)a 8.3 (6–10.5)e 6 (5.5–6.5)
VBML 8.9 (8–10)n 17 (14.5–19)g 8.6 (7–12)a 21.8 (16–27)f 19.7 (19–21.5)
VBW 15.5 (13.5–17.5)n 19.5 (17.5–22)h 14.2 (13–15)a 33.4 (28.5–38.5)e 29.9 (28–31.5)
DBL 1.9 1.9 (1.5–2.5)h 1.3 (1–1.5)c 2.8 (1.8–3.5)h 2.3 (2–2.5)
DBW 14.6 19 (17.5–20.5)g 12.9 (10–16)d 23.7 (19–28)h 21.5 (19.5–22.5)
MHTL 33.3 (32–34.5)l 25.7 (24–28)b 32.6 (30.5–36.5)g 30.7
MHSL 4.7 (4.5–5)m 4.3 (3.5 -5)j 5.5 (5–6.5)d 4.2 (4–5)f 4.3 (4–4.5)
MHHL 18.5 (18–19)o 29 (27.5–31)i 20.3 (18.5–22)c 28.4 (26.2–32)f 26.8 (26.5–27)
MHSDW 3.6(3.5–4)m 4.3 (4–5)j 4 (3.5–4.5)d 3.5 (3–4)k
MHSPW 4 (4–4.5)m 3.5 (3–4)j 3.8 (3.5–4.5)d 3.6 (3–4)g 3.4 (3–4)
MHSAD 5.1 (5–5.5)m 5.7 (5.5–6)k 5.3 (5–6)d 4.5 (4–5)f 4.3 (4–5)

HTL hamulus total length; HPL hamulus point length; HSL hamulus shaft length; HRL hamulus root length; VBL ventral bar median length;
VBML ventral bar membrane length; VBW ventral bar width; DBL dorsal bar length; BDW dorsal bar width; MHTL marginal hook total length;
MHSL marginal hook sickle length; MHHL marginal hook handle length; MHSDW marginal hook sickle distal width; MHSPW marginal hook
sickle proximal width; MHSAD marginal hook sickle aperture distance
Number of specimens measured: a=33, b=32, c=30, d=28, e=23, f=22, g=21, h=20, i=17, j=13, k=12, l=11, m=5, n=3, o=2
Parasitol Res (2012) 110:1185–1200 1189

Fig. 1 Line drawings and

phase-contrast photomicro-
graphs of Gyrodactylus alekosi
n. sp. (a–b), Gyrodactylus
gelnari n. sp. (c–f) and
Gyrodactylus nigritae n. sp.
(g–j). a Hamulus. b Marginal
hook sickles. c Hamulus. d
Detail of the cirrus. e Marginal
hook sickles. f Hamuli complex
with detail of the ventral bar. g
Hamulus. h Detail of the cirrus.
i Marginal hook sickles. j
Hamuli complex with detail
of the ventral bar. Scale bars:
a, c, f, g, j 20 μm; b, d, e, h, i
10 μm

Remarks. Hamuli dimensions of G. alekosi n. sp. are similar present. Gut not extending beyond level of testes. Pharyn-
to those of G. camerunensis (total hamuli length: 50.5–53 vs. geal bulb 58 (47–69, n=10) long, 54 (43–54, n=10) wide
45–64; hamulus point length: 25–27.5 vs. 20–28), but both across the posterior bulb. Male copulatory organ (MCO,
species differ substantially in the shape of their marginal Fig. 1d) spherical, posterior to the pharyngeal bulb, 16 (13–
hook sickles. The shaft of the marginal hook sickle of 18, n=9) in diameter. MCO armed with one principal spine
G. alekosi n. sp. starts perpendicular to the foot, is regularly and a single row of 10–11 smaller spines. Measurements of
curved, and more sturdy than that of G. camerunensis which the haptoral sclerites are given in Table 2. Hamuli slender
has a thinner sickle proper rising in a slanted manner. with flattened area on the inner part of the root (Fig. 1c, f).
Dorsal bar simple. Surface of the ventral bar proper and its
G. gelnari n. sp. membrane ribbed (Fig. 1f). Narrow shaft of the marginal
Description based on 22 coverslip-flattened specimens. hook sickle distinctively points forward from the foot and
Elongated body of length 802 (688–925, n=6), width at curves slightly at a wide angle towards the point, the latter
level of uterus 128 (88–179, n=6). Excretory bladders extending beyond the toe (Fig. 1e). Heel slanted down-
1190 Parasitol Res (2012) 110:1185–1200

Fig. 2 Drawings of the

marginal hook sickles of
Gyrodactylus spp. of African
catfish hosts. a G. synodonti n.
sp. b G. nigritae n. sp. c G.
alekosi n. sp. d G. turkanaensis
n. sp. e G. gelnari n. sp. f G.
camerunensis (paratype
specimen 263HG Ti148). g G.
transvaalensis (present study).
h G. transvaalensis (paratype
specimen 1978.11.3.1–12). i G.
rysavyi (Senegalese specimen).
j G. rysavyi (Kenyan specimen).
k G. rysavyi (Mozambican
specimen). l G. rysavyi
(paratype specimen m-134/1).
m Gyrodactylus sp. Scale bars
10 μm

wards with a slightly rounded edge. Muscular disc lateral to observed (Fig. 2e and d). Although the sickles proper of
the hamuli. both species point forward, the profile of the marginal hook
Type host: C. anguillaris (Linnaeus, 1758) (Siluriformes: sickle foot of G. turkanaensis n. sp. is regularly triangular,
Clariidae) smoothly joining into the foot of the sickle. In contrast, in
Type locality: Mare Simenti, Niokolo Koba National Park G. gelnari n. sp., the foot is more elongate and clearly
(13°01.79′ N, 13°17.6′ W), Senegal separated from the sickle proper. Regrettably, samples were
Infection site: Fins. not adequately preserved for molecular analysis, thus
Type material: Collected March 2007. Holotype and two genetic data on G. gelnari n. sp. cannot be presented.
paratypes deposited in the Institute of Parasitology, Biology
Centre, Academy of Sciences of the Czech Republic in České G. nigritae n. sp.
Budějovice, Czech Republic (Accession number M-520). Description based on 32 coverslip-flattened specimens and
Etymology: the specific epithet honours Prof. Milan Gelnar, three excised haptors of sequenced individuals. Body elongate,
the principal investigator supervising the first author, in 819 (589–1079, n=32) long, width at level of uterus 135 (98–
recognition of his support of her research ideas. 209, n=32). Prohaptor with single pair of cephalic lobes.
Remarks. The shape of the marginal hook sickle of G. Pharyngeal bulb 51 (37–67, n=32) long, 47 (31–53, n=32)
gelnari n. sp. is similar to that of G. camerunensis, its shaft wide across posterior bulb. Excretory bladders present. Gut
similarly rising shaft from the base, but the sickle shaft of extending beyond level of testes. MCO (Fig. 1h) posterior to
G. gelnari n. sp. is thinner than in G. camerunensis. The pharyngeal bulb, 18 (14–22, n=13) in diameter. MCO armed
heel of the marginal hook sickle of G. camerunensis is with one principal spine and a single row of seven to nine
more elongate and distinctively rounded than in G. gelnari smaller spines. Ribbed ventral bar proper and ventral bar
n. sp. (Fig. 2e and f). Moreover, the hamuli of G. gelnari n. membrane (Fig. 1j). Measurements of the haptoral sclerites
sp. (mean total length 71.4 μm) are larger than those of G. are given in Table 2. Hamuli sturdy because of widened joint
camerunensis (mean 54 μm, Nack et al. 2005). Based on of shaft and root; flattened area on inner part of root (Fig. 1g).
the dimensions of the haptoral sclerites, G. gelnari n. sp. Simple dorsal bar. Ventral bar without lateral processes,
resembles G. turkanaensis n. sp. (Tables 2 and 3), but surface of ventral bar proper ribbed, tongue-shaped ventral
differences in marginal hook sickle morphology can be bar membrane. Marginal hook sickle shaft broad, slightly
Parasitol Res (2012) 110:1185–1200 1191

Table 3 Comparison of the measurements (in μm, average with range in parentheses) of the haptoral hard parts of Gyrodactylus spp. collected
from C. gariepinus and C. anguillaris from the present study with the measurements made on paratype specimens of G. transvaalensis

Measurement G. synodonti n. sp. G. turkanaensis n. sp. G. transvaalensis G. transvaalensis Gyrodactylus sp.

(n=23) present study (n=21) present study (n=8) present study (n=2) re-examination (n=2) present study

HTL 80.3 (75–87) 82.3 (77–87.5) 43.5 (41.5–44.5) (41.5–45) 78

HPL 37.2 (34–40.5) 36.4 (34–38.5) 22.3 (20–23) (20.5–23) (36.5–39)
HSL 47.2 (43–54.5) 49.3 (47.5–51) 27.8 (25.5–30) (27.5 -28.5) 54
HRL 41.9 (37–46) 42.2 (34–44.5) 20 (20–22) (14.5–17) 39.5
VBL 8 (7–10) 7.4 (5.5–8.5)c 3.9 (3.5–4.5)l 4.5
VBML 17.7 (15.5–21.5) 19.3 (17–21)c 8.1 (7.5–9)m
VBW 25 (23–28)a 20.1 (18–22.5)c 11.4 (9.5–13.5)l 10.5
DBL 2.3 (2–3)a 2.2 (2–2.5)d 1.1 (1–1.5)o
DBW 19.9 (17–24.5)d 18.5 (16.5–21)f 13.2 (12–14.5)o
MHTL 27.9 (26.5–29.5)f 30.8 (29–32)n 21.5 (20.5–22)l 23
MHSL 6.3 (5.5–7)e 4 (3.5–5)i 5 (4.5–5.5)l (5–5.5) (4–4.5)
MHHL 21.7 (19.5–22.5)h 27.4 (26–28.5)f 16 (15.5–17)k
MHSDW 4.4 (3.5–5.5)e 3.9 (3–4.5)j 4 (4–4.5)l (3.1–3.3)
MHSPW 3.8 (3.5–4)e 3.4 (3–3.5)i 3.7 (3.5–4)l 4 (3.8–4.3)
MHSAD 6.6 (6–7)e 5.4 (4.5–6)j 5.7 (5.5–6)l (5.5–5.7) (5.3–5.7)

Number of specimens measured: a=21, b=20, c=19, d=17, e=16, f=15, g=13, h =12, i=11, j=10, k=7, l=6, m=5, n=4, o=3

pointing forwards, with regularly curved sickle and narrowing and its point terminates immediately after curving slightly.
point extending beyond the toe (Fig. 1i). Sickle foot ovate in The heel of G. transvaalensis is rounded, and elongate in
profile, rounded at the top and waved at the bottom. downward direction, while that of G. nigritae n. sp., despite
Molecular characterisation. A 870 bp fragment covering also being rounded at the top of the heel, is not that
ITS-1 (338 bp), 5.8S rDNA (156 bp) and ITS-2 (376 bp) elongate and more symmetrical to the toe.
was sequenced from three specimens and submitted to
GenBank under accession number FR850686. The whole G. rysavyi Ergens, 1973
sequence was identical for the three specimens. A BlastN Description based on 25 coverslip-flattened (11 Kenyan, 4
search in GenBank in September 2010 using the entire Mozambican, 10 Senegalese) and six sequenced (four
sequence revealed no identical or close hits. Mozambican, two Senegalese) specimens. Body elongate with
Type host: Synodontis nigrita Valenciennes, 1840 (Silur- clearly separate haptor, total body length 1,315 (1052–1674,
iformes: Mochokidae) n=7), width at level of uterus 171 (134–204, n=7). Cephalic
Type locality: Niokolo Koba River, Passage Koba (13°03.92′ region bilobed, each lobe containing a spike sensillum.
N, 13°10.14′ W), Niokolo Koba National Park, Senegal. Excretory bladders present. Pharyngeal bulb 81 (61–113,
Infection site: Barbels. n=10) long, 74 (56–105, n=10) wide across the posterior
Type material: Collected March 2007. Holotype and two part. MCO posterior to pharyngeal bulb, 23.5 (19–29, n=8)
paratypes deposited in the Institute of Parasitology, Biology in diameter, armed with one large principal spine and 11 thin
Centre, Academy of Sciences of the Czech Republic in České small spines in a single row. Intestinal crura not extending
Budějovice, Czech Republic (Accession number M-517). beyond anterior edge of testes. Hamuli slender because root
Etymology: The specific epithet is derived from the specific substantially narrows after its widened joint with the shaft
epithet of the host fish. (Fig. 3a). Flattened area on inner part of root. Simple dorsal
Remarks. G. nigritae n. sp. is similar to G. transvaalensis in bar. Ventral bar without lateral processes, surface of ventral
the overall size of the hamuli. However, G. nigritae n. sp. bar proper and ventral bar membrane ribbed (Fig. 3b).
differs from the latter species in hamulus point length Measurements of the haptoral sclerites are given in Table 2.
(mean 26.3 vs. 22.3 μm) and marginal hook total length Broad sickle shaft of marginal hook sickle points forwards
(mean 25.7 vs. 21.5 μm). In addition, these species’ from the sickle foot, and immediately curves regularly. Sickle
marginal hook sickles have a different shape. The sickle point downwardly directed, extending beyond the toe. Toe
proper of G. nigritae n. sp. is rather robust and curves and heel of the sickle foot slantedly rounded.
regularly, into a pronounced point with closed aperture. Molecular characterisation. A 943-bp PCR product cover-
Conversely, the sickle proper of G. transvaalensis is slender ing ITS-1 (383 bp), 5.8S (157 bp), ITS-2 (383 bp) and 28S
1192 Parasitol Res (2012) 110:1185–1200

Fig. 3 Line drawings and

phase-contrast photomicro-
graphs of Gyrodactylus rysavyi
Ergens, 1973 (a–c), Gyrodacty-
lus synodonti n. sp. (d–g). a
Hamuli. b Detail of the ventral
bar. c Marginal hook sickles. d
Hamulus. e Detail of the cirrus. f
Marginal hook sickles. g Hamuli
complex with detail of the
ventral bar. Scale bars: a, d, e, g
20 μm; b, c, f 10 μm

(20 bp) was sequenced from four Senegalese specimens and ITS-2 sequences differ in six bp. A BlastN search in
submitted to GenBank under accession number FR850679. GenBank in September 2010 using the entire sequence
The entire sequence was identical in these four specimens. An revealed no identical or close hits.
additional sequence of a 872 bp PCR product covering ITS-1 Type host: C. gariepinus (Burchell, 1822) (Siluriformes:
(342 bp), 5.8S (157 bp) and ITS-2 (373 bp) was obtained from Clariidae)
three specimens collected in Mozambique and submitted to Other host: C. anguillaris (Linnaeus, 1758) (Siluriformes:
GenBank under accession number FR850681. Pairwise Clariidae)
genetic distances are summarised in Table 4. The differences Type locality: Nile River, Cairo, Egypt
in ITS sequences of G. rysavyi of both countries are as Other localities: Mare Simenti, Niokolo Koba National Park
follows: the partial ITS-1 differs in seven individual (13°01.79′ N, 13°17.6′ W), Senegal; Kalokol (13°01.79′ N,
positions (four substitutions and three deletions), and the 13°17.6′ W), Todonyang village (4°26.15′ N, 35°56.38′ E),

Table 4 Gamma-corrected pair-

wise GTR distances (in %) be- 1 2 3 4 5
tween the species included in the
phylogenetic analysis, for a 1 Gyrodactylus synodonti n. sp.
881 bp dataset consisting of par- 2 G. nigritae n. sp. 1.9
tial first and second internal tran-
3 G. alekosi n. sp. 19.6 18.3
scribed spacers (ITS-1 and ITS-2)
and intervening 5.8S rDNA 4 Gyrodactylus sp. (Sudan) 20.6 19.3 13.8
5 Gyrodactylus rysavyi (Senegal) 20.9 19.6 13.8 1.7
6 Gyrodactylus rysavyi (Mozambique) 20.6 19.6 13.3 2.5 1.2
Parasitol Res (2012) 110:1185–1200 1193

Turkana Lake, Kenya; temporal stream close to Bala Bala elongate, 842 (602–1080, n=13) long and 113 (67–164,
village (24°19.34′ S, 33°02.36′ E), southern Mozambique. n=13) wide at level of uterus. Prohaptor with single pair of
Infection site: Fins. cephalic lobes. Excretory bladders present. Pharyngeal bulb
Remarks. The shape of the marginal hook sickles of G. 45 (35–61, n=11) long, 41 (25–58, n=11) wide. MCO
rysavyi specimens collected from three distant localities is (Fig. 3e) posterior to pharyngeal bulb, 19 (16–23, n=8)
identical to that of the re-examined paratypes (Fig. 2). The in diameter. MCO armed with one principal spine and a
length of hamulus shaft and point of G. rysavyi from the single row of seven to nine smaller spines. Intestinal crura
present study correspond with measurements on the paratype not extending beyond anterior edge of testes. Measure-
material. Other agreements in the measurements include all ments of the haptoral sclerites are given in Table 3.
analysed features of the marginal hooks. There were some Slender hamuli with well-defined long root. Hamuli root
small discrepancies in the total hamuli length, this charac- slightly curved outwards, inner part of root flattened.
teristic being larger (89–106.5 μm) in the present study than Dorsal bar simple. Ventral bar with small processes;
in the paratype material (90–93 μm). The species can be surface of ventral bar proper and its membrane ribbed
discriminated from all other Gyrodactylus spp. parasitizing (Fig. 3g). Sickle proper of marginal hook points slightly
clariids, by the size of its sclerites, the latter being larger than forward and is relatively slender, turns slightly into a quite
all studied species. PCA (Fig. 4) does show clustering open arch (Fig. 3f). Sickle point directed forwards,
according to region of origin. However, the types, originating terminating beyond toe. Toe elongate and blunt. Muscular
from Egypt, cluster together with the geographically distant disc lateral to hamuli.
Senegalese specimens. Moreover, the Mozambican speci- Molecular characterisation. An 868-bp fragment covering
mens are positioned amidst the other locations, all three of ITS-1 (338 bp), 5.8S rDNA (156 bp) and ITS-2 (374 bp)
which (Senegal, Lake Turkana and Nile) are part of the Nilo- was sequenced from four specimens and submitted to
Sudanese ichthyofaunal province (Lévêque 1997). Hence, GenBank under accession number FR850684. The entire
morphometric groups from various ichthyogeographical sequence was identical for all four specimens. A BlastN
regions seem to overlap. On top of the arguments mentioned, search in GenBank in September 2010 using the entire
it is also the current sample size that does not allow to raise sequence revealed no identical or close hits.
our G. rysavyi populations to species level. Type host: Synodontis nigrita Valenciennes, 1840 (Silur-
iformes: Mochokidae)
Gyrodactylus synodonti n. sp. Type locality: Niokolo Koba River, Passage Koba
Description based on 19 coverslip-flattened specimens and (13°03.92′ N, 13°10.14′ W), Niokolo Koba National Park,
four excised haptors of sequenced specimens. Body Senegal.
Infection site: Fins.
Type material: Collected March 2007. Holotype and two
paratypes deposited in the Institute of Parasitology, Biology
Centre, Academy of Sciences of the Czech Republic in České
Budějovice, Czech Republic (Accession number M-516).
Etymology: The specific epithet is derived from the generic
name of the host fish.
Remarks. G. synodonti n. sp. is similar to G. turkanaensis n.
sp. (see Table 3) in hamuli dimensions. The two species can
be discriminated by the marginal hook total length (mean:
25.7 vs. 30.8 μm, respectively). The difference in shapes of
their marginal hook sickles are distinctive: G. synodonti n. sp.
has a broader, regularly curved sickle proper, while in G.
turkanaensis n. sp. it is thinner, rising from the sickle foot at a
forward angle. The sickle foot of G. turkanaensis n. sp. has a
triangular profile and joins smoothly with the sickle proper,
whereas in G. synodonti n. sp. the sickle toe is more elongate,
and the foot is clearly separated from the sickle proper.

G. transvaalensis Prudhoe and Hussey, 1977

Fig. 4 Plot of principal component analysis on measurements of the
haptoral hard parts of Gyrodactylus rysavyi, for 11 Kenyan (plus sign),
Description based on eight coverslip-flattened specimens.
four Mozambican (ex symbol), ten Senegalese (filled square) and three Elongate body of total length 687 (547–856, n=3), width at
type specimens (open circle) level of uterus 100 (84–124, n=3). Cephalic region bilobed,
1194 Parasitol Res (2012) 110:1185–1200

each lobe containing a spike sensillum. Presence of Type host: C. gariepinus (Burchell, 1822) (Siluriformes:
excretory bladders observed. Pharyngeal bulb 46 (39–51, Clariidae)
n=3) long, 42 (38–45, n=3) wide. MCO posterior to Type locality: Kalokol, Lake Turkana (13°01.79′ N,
pharyngeal bulb, 14.5 (n=1) in diameter. Arrangement of 13°17.6′ W), Kenya.
MCO spines not discernible on whole mounts. Intestinal Infection site: Gills.
crura not extending beyond anterior edge of testes. Type material: Collected September 2008. Holotype and
Measurements of the haptoral sclerites are presented in two paratypes deposited in the Institute of Parasitology,
Table 3. Hamuli rather sturdy as root widens after its joint Biology Centre, Academy of Sciences of the Czech
with shaft (Fig. 5a, d). Root gently narrowing towards the Republic in České Budějovice, Czech Republic (Accession
proximal end. Inner part of root flattened. Dorsal bar number M-519).
simple. Ventral bar with without lateral processes; ventral Etymology: The specific epithet is derived from the name of
bar proper ribbed. Ventral bar membrane tongue-shaped the type locality.
(Fig. 5b). Marginal hook sickle foot ovate in profile, with Remarks. Although G. turkanaensis n. sp. resembles G.
slightly triangular toe and blunt heel pointing downwards. synodonti n. sp. in hamuli size, clear differences between
Sickle proper rises at slightly forward angle; sickle point these two species are observed in the marginal hook sickle
narrow and angled almost perpendicularly, just extending shape: G. turkanaensis n. sp. has a thinner sickle proper,
beyond toe (Figs. 5c, 2g). rising at a forward angle, while G. synodonti n. sp. has a
Type host: C. gariepinus (Burchell, 1822) (Siluriformes: broad, regularly curved sickle proper. The sickle foot of G.
Clariidae) turkanaensis n. sp. is of triangular profile with the heel
Other host: C. anguillaris (Linnaeus, 1758) (Siluriformes: continuing into the sickle proper, while in G. synodonti n.
Clariidae) sp. the sickle foot has a more elongate toe and the sickle
Type locality: Confluence of Olifants and Elands Rivers, proper is clearly separated from the foot. The marginal
South Africa. hook sickle shape of G. turkanaensis n. sp. resembles that
Other localities: Mare Simenti, Niokolo Koba National of G. groschafti the most, but the two species differ
Park (13°01.79′ N, 13°17.6′ W), Senegal. significantly by the size of their hamuli complex, with the
Infection site: Fins. hamuli total length of G. turkanaensis n. sp. exceeding that
Remarks. While the Senegalese G. transvaalensis speci- of G. groschafti (82.3 vs. 34.2 μm, respectively).
mens from C. anguillaris were collected at a large
geographic distance from the species’ type locality, their Gyrodactylus sp.
general morphology and hamuli dimensions correspond Only two excised haptors of sequenced specimens were
well to the re-examined G. transvaalensis type material available. Therefore, body dimensions nor internal morphol-
from South African C. gariepinus. This also goes for the ogy can be presented. Measurements of the haptoral sclerites
marginal hook sickle shape (Fig. 2g and h). are given in Table 3. Hamuli slender because of long, well-
defined root with flattened area on inner part (Fig. 5j). Dorsal
G. turkanaensis n. sp. bar simple. Ventral bar without lateral processes. Marginal
Description based on 21 coverslip-flattened specimens. Elon- hook sickle foot of triangular profile; toe and heel blunt
gate body of total length 900 (739–1088, n=6), width at level (Fig. 5k). Very narrow sickle shaft sharply angles forwards,
of uterus 140 (98–225, n=6). Prohaptor with single pair of turns into a very open, narrow, short point extending beyond
cephalic lobes. Excretory bladders present. Pharyngeal bulb the toe. Heel continuous with shaft.
64 (54–76, n=12) long, 56 (41–71, n=12) wide. MCO Molecular characterisation. A 910 bp PCR product
posterior to pharyngeal bulb, 19 (15–25, n=7) in diameter. covering ITS-1 (380 bp), 5.8S (157 bp), ITS-2 (373 bp)
Detailed arrangement of MCO spines not discernible on was sequenced from two specimens and submitted to
whole mounts. Intestinal crura extending beyond anterior GenBank under accession number FR850688. The entire
edge of testes. Measurements of the haptoral sclerites are sequence was identical for both specimens. A BlastN search
given in Table 3. Hamuli slender, with root narrowing in GenBank in September 2010 using the entire sequence
substantially after joining shaft (Fig. 5e,h). Hamuli with revealed no identical or close hits.
flattened area on inner part of root. Dorsal bar simple. Ventral Host: C. gariepinus (Burchell, 1822) (Siluriformes:
bar without lateral processes, surface of ventral bar proper Clariidae)
and ventral bar membrane ribbed (Fig. 5b). Marginal hook Locality: Sennar, Blue Nile (13°32.81′ N, 33°38.17′ E),
sickle foot of triangular profile; heel continuous with sickle Sudan.
shaft, the latter positioned at a forward angle. Sickle point Infection site: Fins.
short and thin, extending beyond the toe. Toe slant, slightly Remarks. Due to the limited number of studied specimens,
rounded. Muscular disc lateral to hamuli. the formal description of this unknown species is not
Parasitol Res (2012) 110:1185–1200 1195

Fig. 5 Line drawings and phase-contrast photomicrographs of sickle. d Hamuli complex. e Hamulus. f Marginal hook. g Detail of
Gyrodactylus transvaalensis Prudhoe and Hussey 1977 (Senegal) (a– the ventral bar. h Hamuli complex. i Detail of the cirrus. j Hamulus. k
d), Gyrodactylus turkanaeansis n. sp. (e–h) and Gyrodactylus sp. Marginal hook sickle. l Marginal hook. Scale bars: a, d, e, h, j 20 μm;
(Sudan) (j–l). a Hamulus. b Detail of the ventral bar. c Marginal hook b, c, f, g, i, k, l 10 μm

presented here. The size of the haptoral sclerites is similar more narrow sickle shaft than G. turkanaensis n. sp.
to that of G. turkanaensis n. sp., but the species differ in Furthermore, the sickle shaft of Gyrodactylus sp. is slightly
their marginal hook sickle shape: Gyrodactylus sp. has a curved resulting in a more open sickle aperture, and its
1196 Parasitol Res (2012) 110:1185–1200

sickle point is very short, whereas G. turkanaensis n. sp. five new species brings the total number of Gyrodactylus
has a more distinctively curved shaft and a longer point. species described from African fishes to 33 (Christison et
al. 2005; Nack et al. 2005; Přikrylová et al. 2009b; Vaughan
Sequences and phylogenetic analyses et al. 2010; Vanhove et al. 2011a; García-Vásquez et al.
2011).
After trimming of the aligned sequences, 881 bp were
retained in the combined dataset of ITS-1, 5.8S rDNA and Gyrodactylus spp. on mochokid catfishes
ITS-2. The matrix contained 170 variable sites of which
121 were parsimony-informative. There were no differences The finding of Gyrodactylus species on Senegalese S.
in topology between the BI, ML and MP trees (Fig. 6). nigrita represents the first record of these parasites on a
Three lineages are apparent. Apart from G. alekosi n. sp., mochokid host. Two new species identified from this host,
there are two highly supported clades: one grouping both G. nigritae n. sp. and G. synodonti n. sp., are clearly
Synodontis parasites, namely G. synodonti n. sp. and G. distinguished by the size of their haptoral sclerites and also
nigritae n. sp., and another one clustering G. rysavyi with by the shape of their marginal hook sickle (Tables 1 and 2
the yet undescribed species parasitizing Sudanese C. and Fig. 2). Moreover, different infection sites were
gariepinus. Gamma-corrected pairwise genetic distances observed for the two species. G. synodonti n. sp., bearing
(Table 4) are 1.9% between both Gyrodactylus spp. from larger hamuli, was only collected from the hosts’ fins, while
Synodontis, and 1.2–2.5% in the clade containing G. G. nigritae n. sp., with smaller hamuli, was found on
rysavyi. barbels only. A difference in hamulus size according to
The 5.8S rDNA gene was identical in all species under infection site is often observed in Gyrodactylus (Malmberg
study except for G. alekosi n. sp. In terms of overall genetic 1970; Huyse and Malmberg 2004).
distances, this species is more similar to the other Clarias
parasites (Gyrodactylus sp. and G. rysavyi) than to Comparisons between Gyrodactylus spp. on clariid
Synodontis parasite clade (G. nigritae n. sp. and G. catfishes
synodonti n. sp.).
Our finding of two Gyrodactylus spp. on C. gariepinus
from Mozambique represents the first record of gyrodac-
Discussion tylid parasites from this country. One of those Mozambican
species is G. rysavyi, originally described from the Nile
The comparison of gyrodactylids of catfishes from four River in Egypt (Ergens 1973) and also retrieved in Senegal
distant African countries provides an opportunity to and Kenya (current study). Its type host is C. gariepinus,
observe morphological and molecular diversity of these which has an almost pan-African distribution (Teugels
parasites on a continental scale. Furthermore, the present 1986). C. gariepinus largely overlaps in its West-African
study delivers original data on Gyrodactylus species distribution with the mudfish C. anguillaris, another
distribution on catfishes from Africa. The description of possible host to G. rysavyi, hence the wide geographical

Fig. 6 Midpoint rooted

phylogram for the Gyrodactylus
spp. under study, constructed
from an 881-bp dataset consist-
ing of partial first and second
internal transcribed spacers
(ITS-1 and ITS-2) and interven-
ing 5.8S rDNA. Statistical node
support is consistently shown as
follows: Bayesian posterior
probability/maximum likelihood
bootstrap/maximum parsimony
bootstrap. Branch lengths corre-
spond to the expected number of
substitutions per site under
Bayesian inference
Parasitol Res (2012) 110:1185–1200 1197

range of G. rysavyi or its occurrence on other clariid hosts the discovery of more unusual haptoral structures e.g.
is not surprising. accessory bars in Mormyrogyrodactylus gemini Luus-
For G. gelnari n. sp. and G. turkanaensis n. sp., only Powell, Mashego and Khalil, 2003 (Luus-Powell et al.
morphometrical data were obtained. Differences in system- 2003); additional haptoral plates in Gyrodactylus thysi
atically important characters between them and with known Vanhove, Snoeks, Volckaert and Huyse, 2011 (Vanhove et al.
African congeners are deemed sufficient for their designa- 2011a), it is expected that this continent hosts a remarkable
tion as new species. While the haptoral sclerites of G. diversity in haptoral organisation among gyrodactylids.
gelnari n. sp. and G. turkanaensis n. sp. are morphomet- Most Gyrodactylus spp. parasitising catfishes can appar-
rically similar, the shape of their marginal hook sickles ently be divided into two morphotypes based on their
differ, enabling to clearly distinguish between them. hamulus shape. Furthermore, most of them have the same
New data on distribution and morphometrics are presented type of ventral bar, with ribbed ventral bar proper and ventral
for G. transvaalensis. In the original description (Prudhoe bar membrane (Figs. 3 and 5). Another character shared by
and Hussey 1977), the number of studied specimens is the same species is the flattened area on the inner part of the
unclear. Moreover, the associated drawings do not allow hamulus root. On this basis, the following species can be
morphological comparison, especially as regards details of regarded as belonging to a first morphotype: G. clarii, G.
the marginal hook sickle. Our reanalysis of paratype speci- groschafti, G. rysavyi, G. transvaalensis, G. camerunensis
mens provides more precise information, facilitating recog- and all newly described species. Gyrodactylus fusci Ky, 1968,
nition of G. transvaalensis. Hamuli total length, shaft length a parasite of Hong Kong catfish, Clarias fuscus Lacepède,
and point length of the new G. transvaalensis specimens 1803, should be included, too. From the original description
correspond with the paratype material and, as far as shaft and (Ky 1968, drawing 1B), it is evident that this species’ hamuli
point length are concerned, with the original measurements and ventral bar are of the same morphological type as the
by Prudhoe and Hussey (1977). However, there were some above-mentioned African Gyrodactylus spp. Asian G. fusci
discrepancies in root length, explaining the difference in the displays a similar hamulus total length (70–81 μm: Ky 1968)
reported hamulus total length. The values reported here for to G. synodonti n. sp. and G. turkanaensis n. sp. (75–87 μm
the root (20–22 μm) exceed those in both Prudhoe and and 77–87.5 μm, respectively), but these species differ
Hussey (1977; 15–20 μm) and in the paratype specimens substantially in the size of their marginal hooks. Marginal
(14.5–17 μm). The agreement between marginal hook hook total length in G. fusci (42–47 μm) considerably
metrics of our specimens and re-examined paratype speci- exceeds the 26.5–29.5 μm and 29–32 μm of, respectively,
mens is shown in Table 3. G. synodonti n. sp. and G. turkanaensis n. sp. The second
Only two Gyrodactylus sp. specimens from Sudan were morphotype can be characterised by large slender hamuli,
studied. Nevertheless, the morphological differences in their point considerably narrowing after emerging from the
marginal hook sickle shape between them and African shaft. As representatives of this group, G. alberti and G.
congeners demonstrate that they represent another hitherto nyongensis can be mentioned. These two species have a
unknown species. The shape of the marginal hook sickle of proportionally short marginal hook handle (MHHL) in
Gyrodactylus sp. is most reminiscent to G. turkanaensis sp. comparison with the length of their marginal hook sickle
n., but its sickle proper is thinner and its point less curved (MHSL): G. alberti 14.8 μm and 6.5 μm respectively, and G.
in Gyrodactylus sp. Due to the limited number of studied nyongensis 17 μm/11.6 μm (measured during the re-
specimens, the species can at present not be formally examination of type material). In contrast, in the species
described, and more material should be morphometrically from the first morphotype, the proportion MHHL/MHSL is
studied. However, molecular data on Gyrodactylus sp. are substantially higher: mean values 28 μm and 4.2 μm for G.
already presented. rysavyi and 29 μm and 4.3 μm for G. gelnari n. sp.
Some of the new species, namely G. synodonti n. sp., G.
gelnari n. sp. and G. turkanaensis n. sp., share an additional Molecular analyses
haptoral feature: a pair of muscular discs situated on the side
of the hamuli. These three species have quite large hamuli The first genetic data on Gyrodactylus parasitising African
(of mean total length 80.3, 71.4 and 82.3 μm, respectively), catfishes are presented, and phylogenetic tree reconstruc-
and this feature is likely to support the parasite’s attachment tion was performed. The molecular data concur with
to its host. However, such discs have not been observed species distinctions suggested on a morphological basis.
in G. rysavyi, another species of considerable hamuli size Over the region comprising ITS-1 and -2 and 5.8S rDNA,
(mean total length 101.2 μm). Quite similar haptoral the overall gamma-corrected pairwise genetic distance is
position and structures were observed in Diplogyrodacty- 1.2% between Senegalese and Mozambican G. rysavyi.
lus martini Přikrylová, Matějusová, Musilová, Gelnar and This approaches the value of 1% suggested to concur with
Harris, 2009 (Přikrylová et al. 2009a). As Africa has seen species delineation by Ziętara and Lumme (2002). However,
1198 Parasitol Res (2012) 110:1185–1200

the same authors state that a variety of other criteria would rysavyi. Of course, molecular analysis of the other
be needed to ascertain species distinction (Ziętara and Gyrodactylus species mentioned is necessary to investigate
Lumme 2003), e.g. regarding host specificity and consistent whether the haptor-based phenotypic affinities are mirrored
morphological differences. As these do not seem to be by genetic data, and whether the two morphotypes of
fulfilled for the two G. rysavyi populations (both infest C. Gyrodactylus parasites of catfish, suggested above, hold.
gariepinus, and morphometrics showed an overlap between Both (clariid) catfishes and their gyrodactylid parasites
sampled ichthyogeographical regions), we see no reason to often display a wide geographic range (cfr. the occurrence
rise any of them to species status. The genetic and of G. rysavyi in areas and drainage basins as far apart as
morphological differences observed are, in view of their Senegal and Mozambique). In this respect, additional
distant geographic origin, most likely a result of incipient molecular characterisation can also fruitfully assist in
speciation because of isolation-by-distance. inferring the (common) phylogeographic history of host
Phylogenetic analyses consistently revealed the existence of and parasite. This was exemplified in a study by Barson et
three lineages in the catfish Gyrodactylus spp.: (1) a clade of al. (2010) on Macrogyrodactylus Malmberg, 1957 of clariid
both Synodontis parasites included, (2) G. alekosi n. sp. and catfishes, revealing ancient connections between presently
(3) the Sudanese Gyrodactylus sp. together with G. rysavyi. distant host populations.
Several authors suggest the 5.8S rDNA gene to be useful in
distinguishing between Gyrodactylus subgenera as defined, Acknowledgements Collection of material was supported by a grant
from the Ministry of Education, Youth and Sports of the Czech
based on the excretory system, by Malmberg (1970), (Ziętara
Republic (Ichthyoparasitology Research Centre—Project no. LC522).
et al. 2002; Huyse et al. 2003). Hence, it is applied to give a The sampling in Kenya was supported by KMRI (Kenyan Marine and
clue about subgenus affiliation or the recognition of new Freshwater Research Institute, Lake Turkana Station, Kalokol),
subgenera (García-Vásquez et al. 2007; Vanhove et al. 2011a). namely by Dr. John Malala and David Lotuliakou. The sampling in
Sudan was organised with the help of Prof. Zuheir N. Mahmoud,
Remarkably, the 5.8S sequence of all genetically studied Faculty of Science, University of Khartoum. I.P. received financial
catfish Gyrodactylus spp. is identical, or, in the case of G. support from the research project of Masaryk University (MSM
alekosi n. sp., near-identical (differing in one deletion and one 0021622416) and from the grant P505/11/P470 from the Grant
transversion only). As previously mentioned, BLAST Agency of the Academy of Sciences of the Czech Republic. Dr. Tine
Huyse and Prof. Filip Volckaert (K.U.Leuven) are acknowledged for
searches using the entire amplified fragment did not reveal
useful suggestions and for hosting and scientifically supporting the
any close hits. When specifically comparing the 5.8S portion phylogenetic analyses. M.P.M.V. is recipient of a Ph.D. fellowship of
to a range of gyrodactylids (spanning the various Gyrodacty- the Research Foundation—Flanders (FWO-Vlaanderen).
lus subgenera, all African and South-American species for
which molecular data are available, and other gyrodactylid
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