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135
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2, pp. 135-156, 2 figs., 2 tabs., January 2018
© 2018 by Verlag Dr. Friedrich Pfeil, München, Germany – ISSN 0936-9902
An updated checklist of the ichthyofauna
of the Mono River basin
(Benin and Togo: West Africa)
Djiman Lederoun*, ** Jos Snoeks***, ****, Philippe Lalèyè*,
Pierre Vandewalle** and Emmanuel Vreven***, ****
In order to evaluate the impact of anthropogenic changes, such as the future construction of a dam at Adjarala,
on the fish diversity of the Mono River basin, a list of the ichthyofauna of this basin has been compiled. This list
was established based on data obtained from collections in natural history museums and from the literature, and
updated following the most recent systematic revisions. A total of 60 native and one introduced species, belonging
to 40 genera and 23 families, are reported. The families Cyprinidae and Cichlidae, with nine species each, are best
represented in the list. Mormyridae, Alestidae and Clariidae account for six species each, while all other families
contribute three species or less. Of the 60 native species recorded, three are typically marine, while five others are
estuarine. Based on museum records, Raiamas senegalensis (Cyprinidae), previously not reported from the Mono
basin, has been added. Earlier attributions of Marcusenius brucii, M. cyprinoides, Petrocephalus simus (Mormyridae),
Labeo coubie (Cyprinidae), Brycinus leuciscus (Alestidae), Phractura ansorgii (Amphiliidae) and Synodontis melanopterus (Mochokidae) to the Mono basin proved to be based on misidentifications. The present study shows that
the fish species diversity of the Lower Mono is most probably underestimated, due to inadequate sampling and
the resulting lack of archived material from this portion of the basin.
Dans le but d’évaluer l’impact des activités anthropiques, telle que la construction prochaine d’un barrage à
Adjarala, sur la diversité des poissons du bassin du fleuve Mono, une liste de l’ichthyofaune de ce bassin a été
compilée. Cette liste a été établie à partir des collections des musées d’histoire naturelle et des données de la
littérature. Elle a été actualisée en suivant les dernières révisions systématiques. Un total de 60 espèces natives
et une espèce introduite, appartenant à 40 genres et 23 familles, sont rapportés. Les familles des Cyprinidae et
des Cichlidae, avec neuf espèces chacune, sont les mieux représentées. Les Mormyridae, Alestidae et Clariidae
comptent chacune six espèces alors que les autres familles contribuent avec trois espèces ou moins. Parmi les 60
espèces natives inventoriées, trois sont typiquement marines tandis que cinq autres sont estuariennes. Sur la base
des collections des musées, Raiamas senegalensis (Cyprinidae), non signalé du bassin du Mono, a été ajouté. Les citations antérieures de Marcusenius brucii, M. cyprinoides, Petrocephalus simus (Mormyridae), Labeo coubie (Cyprinidae),
Brycinus leuciscus (Alestidae), Phractura ansorgii (Amphiliidae) et Synodontis melanopterus (Mochokidae) du bassin
du Mono, sont basées sur des identifications erronées. La présente étude a montré que la diversité des espèces de
poissons du cours inférieur du Mono est probablement sous estimée, en raison d’un échantillonnage inadéquat
et de l’absence de collection provenant de cette partie du bassin.
*
Laboratory of Hydrobiology and Aquaculture, Faculty of Agricultural Sciences, University of Abomey-Calavi, 01 BP: 526 Cotonou, Benin. E-mail: (DL) ldjiman@yahoo.fr; (PL) laleyephilippe@gmail.com
** Laboratory of Functional and Evolutionary Morphology, University of Liège, Chemistry Institute B6, Sart
Tilman, 4000 Liege, Belgium. E-mail: (PV) p.vandewalle@ulg.ac.be
*** Royal Museum for Central Africa, Vertebrate Section, Ichthyology, Leuvensesteenweg 13, 3080 Tervuren,
Belgium. E-mail: (EV) emmanuel.vreven@africamuseum.be (corresponding author).
**** KU Leuven, Laboratory of Biodiversity and Evolutionary Genomics, Charles Deberiotstraat 32, 3000 Leuven,
Belgium. E-mail: (JS) jos.snoeks@africamuseum.be
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
136
Copyright © Verlag Dr. Friedrich Pfeil
Introduction
Western exploration and documentation of the
ichthyofauna of the Mono basin started only
quite recently. Indeed, Daget’s (1950) paper on
the freshwater fishes of the coastal regions of
Togo and Dahomey (now Benin), contained the
very first data on the Mono’s ichthyofauna. Two
cichlid species, Tilapia zilli (now Coptodon zillii;
see Dunz & Schliewen, 2013) and Tilapia heudeloti
macrocephala (now Sarotherodon melanotheron melanotheron; see Trewavas, 1983) were reported from
the Grand-Popo Lagoon (Benin) by Daget (1950),
although apparently no reference specimens were
deposited in any natural history museum. The first
major fish collections were made by the ‘Laboratoire d’Hydrobiologie du Service des Eaux, Forêts
et Chasses du Dahomey’ in the 1950s during the
colonial period (Gras, 1961; Lalèyè et al., 2004).
Based on the study of these collections, which
unfortunately have been lost, Gras (1961) reported
five freshwater and four marine fish species from
the lower reaches of the Mono basin in Benin. Five
years later, Thys van den Audenaerde was the first
to explore the ichthyofauna of the Upper Mono
in Togo. His small collection from that basin,
deposited at the Royal Museum for Central Africa
(RMCA), contained 129 specimens belonging to
nine widespread West African species, currently
identified as: Enteromius ablabes, Labeo parvus,
Malapterurus beninensis, Clarias gariepinus, Heterobranchus longifilis, H. isopterus, Epiplatys togolensis,
Hemichromis fasciatus and Coptodon guineensis.
From 1969 to 1970, other important ichthyological expeditions were undertaken. These expeditions, such as those of Loiselle in 1969, Verheyen,
Hulselmans and Puylaert in 1969, Stoffels in 1970
and Thys van den Audenaerde and Opdenbosch
in 1970, were largely organized by the RMCA, and
enabled further exploration of the Mono basin in
Togo. In addition, members of the 1969 expeditions also made collections for the J. L. B. Smith
Institute of Ichthyology [now the South African
Institute for Aquatic Biodiversity (SAIAB)], and
the United States National Museum (USNM).
However, none of these collections has since been
the subject of any particular publication.
Later collections from the Mono River basin,
made in the period 1975 to 1986, were deposited
in major natural history museums all over the
world. In 1975, several specimens of Enteromius
callipterus caught in the upper course of the Mono
in Togo were deposited in the fish collection of
the British Museum of Natural History (BMNH).
Another expedition undertaken by Kulo and
Kritsky in 1985-1986 around Kolokopé, i. e. the
upper course of the Mono basin in Togo, resulted
in the deposition of a small collection of fishes
from the Mono River in the American Museum
of Natural History (AMNH).
However, the most important collections,
comprising more than 1000 specimens from 24
localities, were assembled between 1981 and 1986
by Lévêque, Paugy and Bénech in the Togolese
part of the Mono basin. In the context of the
Onchocerciasis Control Programme (OCP) they
explored this section of the main river, as well
as numerous tributaries. Their collections were
all deposited at the Muséum National d’Histoire
Naturelle (MNHN) in Paris. This sampling effort
yielded much new data on the fish fauna of the
Mono River basin, some of which was published
in Paugy & Bénech (1989), including the first
checklist of this river’s ichthyofauna. These data
were also incorporated into a guide to the fresh
and brackish water fishes of West Africa (Lévêque
et al., 1990, 1992). Nevertheless, the ichthyofauna
of the Mono River basin remains underexplored.
Two ichthyological monitoring stations were
set up in the middle section of the Mono basin
to evaluate the effects of the insecticides used in
the context of the OCP, one at Atchinédji and
the other at Tététou (Paugy et al., 1988). The
construction of the Nangbéto dam between the
two monitoring stations in 1987, caused huge
disruptions, including floods, in the natural flow
regime. As a result, further sampling was abandoned. Thus, since 1987 no additional systematic
sampling has been undertaken in the Mono basin.
Nevertheless, after a short study visit to Benin in
1997, Vandewalle returned to the RMCA with a
single specimen of Labeo senegalensis [previously
reported and catalogued as L. coubie (see Lévêque,
2003)] from the Sazué River (Benin), a left-bank
tributary of the Lower Mono. Finally, Musschoot
& Lalèyè (2008), while studying the collections
of Synodontis schall from the Mono and Ouémé
basins, recognized and described a new species,
S. ouemeensis, currently considered endemic to
these two drainages and the Ogun basin (Nigeria).
In spite of the work briefly reviewed above,
much remains to be learned about the ichthyofauna of the Mono River basin. Indeed, the identifications of many specimens housed in natural history
collections were never subsequently reviewed,
and are now outdated. In addition, since the conLederoun et al.: Ichthyofauna of the Mono River basin
137
Copyright © Verlag Dr. Friedrich Pfeil
struction of the Nangbéto dam in 1987, no major
ichthyological studies have been undertaken to
complete and synthesize our current knowledge
of the fish fauna of the Mono. Thus, we feel that
an updated list of its ichthyofauna is not only
desirable but important. Such a list will become
even more significant if the planned construction
of a second dam at Adjarala on the lower course
of the Mono basin goes ahead.
1°E
ı
2°E
ı
9°N -
- 9°N
8°N -
- 8°N
7°N -
- 7°N
Material and methods
Study area. With its lower reaches forming the
border between Togo and Benin over a stretch of
about 100 km, the Mono is a transnational coastal
basin (Fig. 1). The river itself rises in the Koura
Hills at Alédjo (≈ 9°21' N 01°27' E) in northwestern Benin. It is approximately 360 km long and
drains a watershed of approximately 22 000 km2
(Paugy & Bénech, 1989) between latitudes 6°10'
and 9°00' North and longitudes 0°30' and 1°50'
East. Close to the Atlantic Ocean, the river splits
into two branches, one flowing towards the east
and entering the Beninese lagoon system (the
coastal lagoon of Grand-Popo and Lake Ahémé),
and the smaller segment meandering to the west
into the Togolese lagoon system (Lake Togo and
the Vogan Lagoon) (Fig. 1).
Two main climatic regions can be distinguished within the Mono watershed: (1) the
tropical zone, situated north of the 8th parallel
and characterized by two seasons, a dry (November to March) and a rainy one (April to October)
with an average total rainfall of between 1000 and
1300 mm/year; and (2) the sub-equatorial zone,
situated south of the 8th parallel and characterized
by four seasons, with two dry seasons (December
to March and July to September) alternating with
two rainy seasons (March to July and September
to November), and an average annual rainfall of
900 to 1100 mm (Paugy & Bénech, 1989; Amoussou, 2010; Laïbi et al., 2012).
Currently, the Nangbéto hydroelectric dam,
located approximately 180 km upstream of the
mouth of the Mono, is the only major hydrological
intervention within the watershed. Its reservoir,
which became operational in 1987, covers an area
of ≈ 180 km², has a maximum depth of ≈ 40 m
and a water storage capacity of approximately
1715 · 106 m3. Prior to 1987, the Mono River at
Athiémé (watershed ≈ 21 500 km²) (Fig. 1) was
characterized by a significant flow from June to
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
ı
1°E
ı
2°E
Fig. 1. Hydrographic map of the Mono basin: upper
course (dark grey); middle course (light grey); lower
course (very light grey). , sampling localities recorded
for specimens housed in the MNHN and the RMCA;
, sampling localities taken from the literature. Localities cited in the text: 1, Atchinédji; 2, Nangbéto dam;
3, Adjarala; 4, Athiémé; 5, Grand-Popo Lagoon.
November, with a maximum flow rate in September (423.1 m3/s) and a flow close to zero from
December to May (1.48 m3/s) (Amoussou, 2010).
The installation of the Nangbéto dam markedly
altered the flow regime; the flow is now permanent and floods are less intense (Oyédé, 1991). The
flow rate still reaches its peak in September, albeit
with a slight reduction of 3.2 % in the maximum
flow (409.7 m3/s), while the minimum flow rate
has increased by 97.2 % (to 52.6 m3/s) (Amoussou,
2010). The construction of a second hydroelectric
dam began in 2016 at the Adjarala Rapids (Fig. 1),
approximately 100 km downstream of Nangbéto
(Anonymous, 1992, 1997).
138
Copyright © Verlag Dr. Friedrich Pfeil
Data and methods. Data were obtained from
natural history museum collections and from
relevant publications (Daget, 1950; Gras, 1961;
Paugy & Bénech, 1989; Lévêque & Bigorne,
1985a-b; De Vos, 1995; Bigorne & Paugy, 1991;
Lévêque et al., 1991; Paugy et al., 1994; Paugy
et al., 2003a-b; Musschoot & Lalèyè, 2008). The
compiled species list has been updated using
the most recent systematic revisions. Specimens
were mainly identified with the keys in Paugy et
al. (2003a-b) except for those groups for which
more recent revisions were available (Musschoot
& Lalèyè, 2008; Decru et al., 2012, 2013).
All specimens originating from the Mono basin and housed at the RMCA were re-examined.
Specimens held in other museums, in particular
those from the Muséum National d’Histoire Naturelle (MNHN, Paris), were re-examined when
their identification seemed doubtful, i. e. for those
species for which their presence in the Mono basin
is unlikely based on our current knowledge of
their distribution. Species reported from the Mono
basin by Daget (1950), Gras (1961) and Paugy &
Bénech (1989) for which no collection records exist
were retained in our list, since the published reports were actually based on collected specimens,
most of which were identified by experts in the
field (see Table 1). Of the species listed by Paugy
et al. (1994), only those based on collection records
were accepted and included while species whose
provenance was inferred from distribution data
only were disregarded.
To obtain a more informative picture of the
distribution of its fish fauna, the Mono basin was
first divided into three major sections (lower, middle and upper) according to its longitudinal profile
reconstructed from data on elevation (Anonymous, 1962) and also into three other sections
based on the location of the Nangbéto dam (see
Fig. 1), i. e. (i) downstream of the dam, (ii) the
section occupied by Lake Nangbéto itself, and
(iii) upstream of the lake. Species lists for these
two different subdivisions are given in Table 1.
The sequence of families follows that of Nelson et
al. (2016), while the genera and species are listed
in alphabetical order.
The ecological character of the fish assemblages, i. e. freshwater, estuarine or marine species, was defined according to Albaret (1994).
All sampling localities within each section
were mapped (see Fig. 1). When coordinates of
the sampling localities were missing from the
museum labels, the relevant gazetteers for Benin
(USBGN, 1965) and Togo (USBGN, 1966) were
used.
Two empirical models published by Daget
& Iltis (1965) and Hugueny & Lévêque (2006)
were used to predict the species richness of the
Mono basin. This should enable us to evaluate if
the estimates of species diversity obtained here
are compatible with those expected for a basin
with its surface area. In addition, the log-linear
relationship between the surface areas of some
West African river basins and their currently
known species richness, as presented by Gourène
et al. (1999), was also applied to the data for the
Mono basin. The data on the species richness of
the basins included are from Hugueny & Lévêque
(2006).
Fish collection acronyms used are as follows:
AMNH, American Museum of Natural History, New York; BMNH, The Natural History
Museum, London; DPB, Direction des Pêches
du Bénin, Cotonou; MNHN, Muséum National
d’Histoire Naturelle, Paris; MRAC, Musée Royal
de l’Afrique Centrale, Tervuren; SAIAB, South
African Institute for Aquatic Biodiversity, Grahamstown; and USNM, National Museum of
Natural History, Smithsonian Institution, Washington, DC. Other abbreviations employed are:
HL, Head Length; MRACDL = New collections
deposited at MRAC by Djiman Lederoun; and
SL, Standard length. All locality data have been
translated into English.
Results
Sixty native and one introduced species, representing 40 genera and 23 families, have been found
in the Mono basin (Table 1). With nine species
each, Cyprinidae and Cichlidae are the most
species-rich families, followed by Mormyridae (6),
Alestidae (6), Clariidae (6). All the other families in
our list are represented by three species or less (see
Table 1). Oreochromis niloticus (Cichlidae) is the
only introduced species. Three species, i. e. Arius
latiscutatus, Drepane africana and Pseudotolithus
senegalensis, are typically marine, while five others,
i. e. Aplocheilichthys spilauchen, Awaous lateristriga,
Coptodon guineensis, Nematogobius maindroni and
Sarotherodon melanotheron are estuarine species
according to Albaret’s (1994) criteria.
The middle and upper courses of the Mono
exhibit the highest species richness with 45 and
44 species, respectively (73 and 72 % of the total
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
fish fauna); wheras the lower course harbors 29
species (47 %). However, these numbers may be
biased due to the fact that the lower course has
been poorly explored (Fig. 1). Sixteen species (26 %
of the fish fauna), belonging to 13 genera and
nine families, occur in all three parts of the basin.
Nevertheless, each section also hosts a number
of typical species. Thus, ten species (five marine
or estuarine species and five freshwater species)
are characteristic for the lower course, while four
(one estuarine species i.e. Awaous lateristriga and
three freshwater species) are only found in the
middle course, and six (all freshwater species)
in the upper course (Table 1).
Thirteen species (21 % of the fish fauna) were
collected from the Mono River at Nangbéto prior
to the construction of the dam and the creation
of its lake (Table 1). These are all widespread
species within the Mono basin. For the sections
downstream and upstream of Lake Nangbéto
itself, the species richness is respectively 53 (86 %
of the fish fauna) and 50 (81 %) species. Forty-two
species are common to downstream and upstream
sections of the lake. Furthermore, eleven (five
marine or estuarine and six freshwater species)
are characteristic for the downstream section,
while eight (all freshwater species) are restricted
to the upstream section. All species reported in the
present study were known from the Mono basin
before the construction of the Nangbéto dam.
The mean maximum species richness predicted by the formula developed by Daget & Iltis
(1965) is 61, while the Hugueny & Lévêque (2006)
model yields a figure of 55 and the linear model
of Gourène et al. (1999) results in 70 species (Table 2). Based on the average of the pooled means
(62 species), it can be assumed that approximately
98 % of the estimated species richness is currently
known. If one adopts the highest estimate (70
species), our current knowledge covers only 87 %
of the species richness; however, the number of
known species does fall within the lower bound
predicted by that model (Table 2).
Based on our current knowledge of West
African fish fauna, the taxonomic status of some
species reported from the Mono basin is uncertain. In the following, we discuss some special
issues: previous misidentifications, species cited
for the first time from the Mono basin, and junior
synonyms previously used in the literature.
Polypteridae. Only a single species of this family,
Polypterus senegalus, was reported from the Lower
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
139
Mono by Paugy & Bénech (1989), but unfortunately no specimen was preserved. However,
several specimens of this species were sampled
during our recent expeditions from 2011 to 2013
and have been deposited at the MRAC. All belong
to the nominal subspecies P. s. senegalus known
from West Africa, while the other subspecies,
P. s. meridionalis is only known from the Congo
River, specifically from the section extending
from the Upper Lualaba River downstream to
Yangambi (Banister & Bailey, 1979; Gosse, 1963,
1984, 1990).
Osteoglossidae. Lévêque et al. (1991) reported
Heterotis niloticus from the Mono as an introduced
species. A single specimen, collected in Lake
Toho (Lower Mono) in 1986, is housed at the
DPB. In addition, several specimens, which have
been deposited at the RMCA, were collected in a
number of small lakes located in the lower course
and in the main course of the Mono downstream
of the Nangbéto dam during our recent expeditions, indicating that the species is widespread
in the basin. The first transfers of this species in
Africa date from the 1950s when the species was
introduced from Cameroon in to the Democratic
Republic of the Congo and Gabon (see Welcomme,
1988). However, the species had already been
reported from Benin (Daget, 1950; Gras, 1961)
and Togo (Daget, 1950) on the basis of material
collected during the colonial period in the 1940s
and 1950s. That its presence in the area predates
the initial period of introductions within Africa
is further attested by a single specimen (MRAC
73190) collected at Lake Togo, Togoville in 1947.
Therefore, H. niloticus is herein considered native to the area, and has already been reported
as such by several authors in the past (see Daget
& Iltis, 1965; Micha & Frank, 1976; Daget, 1984;
Paugy, 2003a).
Mormyridae. Based on existing collections, six
valid species are currently known from the Mono
basin. Two of these, i. e. Brienomyrus brachyistius
and Mormyrus hasselquistii, are apparently confined to the tributaries, while the remaining four
species are present in both its main course and
its tributaries.
The evidence adduced for the presence of
Marcusenius brucii, M. cyprinoides and Petrocephalus levequei based on MNHN specimens and
Bigorne (2003) can now be rejected, as detailed in
the following. Two specimens (MNHN 1985-140
140
Copyright © Verlag Dr. Friedrich Pfeil
Table 1. Annotated checklist of the fishes of the Mono River basin, together with their currently known distribution within the basin based on museum records and published literature reports. The list is arranged by (a)
longitudinal profile and (b) relative to the position of the Nangbéto dam. D, downstream; i, introduced species;
LC, Lower course; mc, main course; MC, Middle course; Na, Nangbéto; t, tributaries; U, Upstream; UP, Upper
course; %, species present; *, marine species; #, estuarine species; +, species present but not reported by Paugy
& Bénech (1989). Acronyms of museums are mentioned in Collections column, when relevant (see Data and
methods). Major collections and bibliographical details are also given.
a
Species
Polypteridae (1)
Polypterus senegalus senegalus
%
Osteoglossidae (1)
Heterotis niloticus
+
Mormyridae (6)
Brienomyrus brachyistius
Marcusenius senegalensis
Mormyrops anguilloides
%
%
+
+
+
+
+
%
%
%
%
%
Mormyrus hasselquistii
Mormyrus rume
Petrocephalus bovei
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
Cyprinidae (9)
LC
t
UP
%
D Na Na U Na
%
+
%
Enteromius callipterus
%
%
%
Enteromius chlorotaenia
Enteromius macrops
Enteromius nigeriensis
Enteromius sublineatus
Labeo parvus
%
%
%
+
+
+
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
Brycinus longipinnis
%
%
%
%
%
%
Brycinus macrolepidotus
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
Brycinus nurse
Hydrocynus forskalii
Rhabdalestes septentrionalis
Amphilius atesuensis
Phractura clauseni
%
%
%
%
%
%
%
Synodontis ouemeensis
%
%
Malapterurus beninensis
%
Clariidae (6)
Clarias (Clarioides) agboyiensis
Clarias (Clarias) anguillaris
Clarias (Clarioides) buthupogon
Clarias (Clarias) gariepinus
%
%
%
%
%
%
%
Malapteruridae (1)
%
%
%
%
%
Synodontis cf. obesus
%
%
%
%
%
+
%
Amphiliidae (2)
%
%
+
%
Hepsetus odoe
%
%
Brycinus cf. imberi
Hepsetidae (1)
%
+
Enteromius ablabes
Distichodontidae (1) Distichodus rostratus
Mochokidae (2)
mc
UP
%
Labeo senegalensis
Raiamas senegalensis
Alestidae (6)
mc
t
MC MC
b
Family
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
%
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
and MNHN 1991-0941) previously catalogued as
M. brucii are herein identified as M. senegalensis.
Both specimens have conical teeth and hence
clearly differ from other West African Marcusenius
species, such as M. thomasi, M. mento, M. furci-
Collections
Literature
Paugy & Bénech, 1989
MRACDL, DPB
MNHN,
MNHN,
MNHN,
USNM
MNHN,
MNHN,
MNHN,
MRAC Paugy & Bénech, 1989; Bigorne, 2003
MRAC Bigorne, 2003
MRAC, Paugy & Bénech, 1989; Bigorne, 2003
MRAC Bigorne, 2003
MRAC Paugy & Bénech, 1989; Bigorne, 2003
USNM Paugy & Bénech, 1989; Bigorne, 2003
MNHN, MRAC,
USNM
BMNH, MNHN,
USNM
MNHN, USNM
MNHN
MRACDL
MNHN
MNHN, MRAC,
USNM
MNHN
MRAC
MNHN
Paugy & Bénech, 1989
Gras, 1961; Paugy & Bénech, 1989;
Lévêque, 2003
Paugy & Bénech, 1989; Lévêque, 2003
Paugy & Bénech, 1989; Lévêque, 2003
Gras, 1961
Paugy & Bénech, 1989; Lévêque, 2003
Gras, 1961; Paugy & Bénech, 1989;
Lévêque, 2003
Paugy & Bénech, 1989
Paugy & Bénech, 1989
MNHN, MRAC Gras, 1961; Paugy & Bénech, 1989;
Paugy, 2003c
AMNH, MNHN, Paugy & Bénech, 1989
MRAC, USNM
MNHN, MRAC, Paugy & Bénech, 1989; Paugy, 2003c
USNM
AMNH, MNHN, Paugy & Bénech, 1989; Paugy, 2003c
MRAC, USNM
MRAC
Paugy & Bénech, 1989; Paugy, 2003c
MNHN
Paugy & Bénech, 1989; Paugy, 2003c
MNHN, MRAC, Paugy & Bénech, 1989; Paugy, 2003b
USNM
MNHN, USNM Paugy & Bénech ,1989; Skeleton et al., 2003
MNHN
Paugy & Bénech, 1989; Skeleton et al., 2003
MNHN
MNHN
Paugy & Bénech, 1989; Paugy & Roberts,
2003
Musschoot & Lalèyè, 2008
MRAC
Paugy & Bénech, 1989; Norris, 2003
MRAC
Paugy
MRAC
Paugy
MNHN
Paugy
AMNH, MNHN, Paugy
MRAC
&
&
&
&
Bénech,
Bénech,
Bénech,
Bénech,
1989;
1989;
1989;
1989;
Teugels,
Teugels,
Teugels,
Teugels,
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
2003a
2003a
2003a
2003a
141
dens, M. ussheri, M. meronai and M. deboensi, all
of which have bicuspid teeth. We counted 12
scales around the caudal peduncle, indicating
that they are not conspecific with M. cyprinoides
or M. abadii either, as both of these species have 16
circumpeduncular scales. Therefore, the MNHN
specimens might belong either to M. senegalensis
or M. brucii, although the presence of the latter
species in Mono basin is questionable, as Jégu &
Lévêque (1984), who examined several specimens
of M. senegalensis from the Mono River, did not
report M. brucii from that basin. According to
the key to the species of the genus Marcusenius
published by Bigorne & Paugy (1990), M. brucii
is only known from the Ogun and Oshun rivers
in Nigeria. However, in the second edition of
the guide to the fresh and brackish water fishes
of West Africa, Bigorne (2003) reported both
M. brucii and M. senegalensis from the Mono. We
believe that this report of M. brucii was based
on the two MNHN specimens referred to here,
with which the diagnostic characters proposed by
Bigorne (2003) show some overlap. Indeed, the
body depth ranges from 3.1 to 4.6 times the SL
in M. senegalensis (vs. 2.9 to 3.3 times in M. brucii)
and the depth of the caudal peduncle varies from
2.0 to 3.6 times SL in M. senegalensis (vs. 1.9 to 2.0
times in M. brucii). For the two specimens from the
Mono in the MNHN, the body depth ranges from
3.1 to 3.2 times SL and the depth of the caudal
peduncle varies from 2.0 to 2.3 times in its length.
While the specimen with a caudal peduncle depth
of 2.3 can be attributed to M. senegalensis, this is
not the case for the second. Indeed, both its values
(3.1 and 2.0) lie within the ranges given for both
nominal species, and we are therefore unable to
assign this latter specimen to either M. senegalensis
or M. brucii based on the diagnosis provided by
Bigorne (2003). Paugy & Bénech (1989) had also
mentioned that the taxonomic status of the two
nominal species was unclear. Indeed, they specifically stated that it was also difficult to distinguish
between the two species in the Ogun River, from
which M. brucii was originally described. In addition, they speculated that M. brucii replaces
M. senegalensis in the Mono basin. A revision of
the status of both nominal species is underway,
but preliminary results seem to indicate that
M. brucii is a junior synonym of M. senegalensis
(Boden, pers. comm.). Since we were unable to
allocate the examined specimens to one of the
two nominal species, we use the older name, i. e.
M. senegalensis.
142
Copyright © Verlag Dr. Friedrich Pfeil
As regards Marcusenius cyprinoides, its presence in the Mono basin is attested by a single
specimen (MNHN 2002-0799). However, according to Bigorne (2003), M. cyprinoides is only
known from the Chad basin, the Benue and the
Lower Niger. The specimen from the Mono has
conical teeth and 12 circumpeduncular scales and
is in fact conspecific with the two M. senegalensis
specimens cited above.
The re-examined specimens of Petrocephalus levequei (MNHN 1985-0782: five out of 19:
62.5-76.5 mm SL) are identified here as P. bovei.
Table 1. (continued).
a
Family
Species
LC
Heterobranchus isopterus
Heterobranchus longifilis
Claroteidae (2)
%
%
Chrysichthys (Chrysichthys) auratus
Chrysichthys (Melanodactylus) nigrodigitatus
Ariidae (1)
Arius latiscutatus
Schilbeidae (2)
Schilbe intermedius
Schilbe mystus
mc
t
MC MC
%
%
%
%
%
b
mc
UP
t
UP
%
%
%
%
%
%
%
+*
%
%
%
%
%
%
%
%
%
%
%
Nothobranchiidae (2) Epiplatys togolensis
%
%
%
#
%
%
%
Poeciliidae (1)
Aplocheilichthys spilauchen
#
#
Centropomidae (1)
Lates niloticus
%
Sciaenidae (1)
Pseudotolithus (Pseudotolithus) senegalensis
+*
+*
Drepaneidae (1)
Drepane africana
+*
+*
Cichlidae (9)
Chromidotilapia guntheri guntheri
%
%
%
%
Coptodon dageti
Coptodon guineensis
Gobiidae (2)
Hemichromis bimaculatus
%
Hemichromis fasciatus
%
%
Oreochromis niloticus
i
i
Sarotherodon galilaeus galilaeus
%
%
Sarotherodon melanotheron melanotheron
#
Awaous lateristriga
Nematogobius maindroni
Channidae (1)
Parachanna obscura
Protopteridae (1)
Total
Protopterus annectens annectens
%
%
%
%
%
#
#
%
%
%
%
+
%
#
%
%
%
%
%
%
%
%
%
i
%
%
%
%
i
%
%
#
#
#
%
#
#
#
#
#
%
%
%
%
%
%
%
%
%
%
%
29
%
%
+
%
Ctenopoma kingsleyae
%
#
Coptodon zillii
Anabantidae (1)
%
#
Fundulopanchax (Paludopanchax) filamentosus
%
%
+*
%
%
D Na Na U Na
#
#
%
31
34
45
39
33
53
13
50
44
Lederoun et al.: Ichthyofauna of the Mono River basin
143
Copyright © Verlag Dr. Friedrich Pfeil
According to Bigorne (2003), Petrocephalus occurs in two distinct areas in West Africa: (1) the
Sudano-Sahelian zone, which includes the Mono
basin; and (2) the Guinean zone, which extends
from the coastal basins of Guinea to the Ivory
Coast. Given that P. levequei is known from the
Collections
Literature
MNHN, MRAC Paugy & Bénech, 1989
MRAC
Paugy & Bénech, 1989; Teugels, 2003a
MNHN, MRAC, Paugy & Bénech, 1989; Risch, 2003
SAIAB
MRAC
Paugy & Bénech, 1989; Risch, 2003
DPB
Gras, 1961
AMNH, MRAC Paugy & Bénech, 1989; DeVos, 2003
MNHN
Paugy & Bénech, 1989; DeVos, 2003
MNHN
Paugy & Bénech, 1989; Wildekamp &
Van der Zee, 2003
Paugy & Bénech, 1989; Wildekamp &
Van der Zee, 2003
Paugy & Bénech, 1989; Wildekamp &
Van der Zee, 2003
MRACDL
Paugy & Bénech, 1989; Paugy, 2003d
Gras, 1961
Gras, 1961
MNHN, MRAC,
USNM
MRAC, USNM
MRAC
Paugy & Bénech, 1989; Teugels & Thys
van den Audenaerde, 2003
Teugels & Thys van den Audenaerde, 2003
Paugy & Bénech, 1989; Teugels & Thys
van den Audenaerde, 2003
MRAC
Daget, 1950; Paugy & Bénech, 1989; Teugels & Thys van den Audenaerde, 2003
MRACDL
Paugy & Bénech, 1989; Teugels & Thys
van den Audenaerde, 2003
MNHN, MRAC, Paugy & Bénech, 1989; Teugels & Thys
USNM
van den Audenaerde, 2003
MRACDL
Paugy & Bénech, 1989; Ahouansou Montcho & Lalèyè, 2008
MNHN, MRAC Gras, 1961; Paugy & Bénech, 1989; Teugels & Thys van den Audenaerde, 2003
MRACDL
Daget, 1950; Gras, 1961; Paugy & Bénech, 1989
MRAC, USNM
MNHN
Paugy & Bénech, 1989; Harrison et al., 2003
Paugy & Bénech, 1989; Harrison et al., 2003
MRACDL
Paugy & Bénech, 1989
MNHN
Paugy & Bénech, 1989; Teugels, 2003b
MRACDL
Paugy & Bénech, 1989
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
Guinean zone, it should not occur in the Mono
basin. Seven species are known from the Sudano-Sahelian zone: P. ansorgii, P. bane, P. bovei,
P. pallidomaculatus, P. pellegrini, P. sauvagii, and
P. soudanensis. The specimens examined have a
uniform silvery colour without the sub-dorsal spot
usually found in P. ansorgii, P. pallidomaculatus,
P. sauvagii and P. soudanensis (Bigorne, 2003). The
specimens examined have 23-26 dorsal-fin rays
and are therefore clearly different from P. bane,
which has at least 29 (Bigorne, 2003). Therefore,
the specimens might be conspecific with P. bovei
or P. pellegrini. As the length of the anal-fin base
varies from 3.5 to 3.6 times the SL (vs. 3.8-4.5 in
P. pellegrini) and P. pellegrini is known only from
the Niandan, a tributary of the Niger in Guinea
(Bigorne, 2003), the MNHN specimens are herein
identified as P. bovei, a species already known
from the Mono basin (Bigorne, 2003).
Cyprinidae. Together with the Cichlidae, Cyprinidae is the most species-rich family within the
basin, with nine reported species; i. e. including
six species of Enteromius. The genus Barbus sensu
lato is known to be a paraphyletic assemblage with
three different ploidy levels: diploid (2n = 48 or
50), tetraploid (2n = 100) and hexaploid (2n = 148150) (Golubtsov & Krysanov, 1993; Guégan et al.,
1995; Berrebi et al., 1990, 1996; Machordom &
Doadrio, 2001). Currently, the name Barbus sensu
stricto is only used for some tetraploid European
species and some species endemic to the Maghreb
and north-east Africa (Doadrio, 1990; Berrebi,
1998; Seegers et al., 2003). Nowadays, the West
African Barbus s. l. are divided into two main
groups: the large-sized hexaploid species and the
small diploid species (Berrebi et al., 1990). While
the former are assigned to the genus Labeobarbus
(Berrebi, 1998; Skelton, 2001), the latter have long
been referred to as ‘Barbus’ (Berrebi et al., 1996).
Table 2. Species richness of the Mono basin as predicted
by the different models used.
Model
Daget & Iltis (1965)
Hugueny & Lévêque (1999)
Linear relationship
Predicted Current species
species
richness
richness (N = 60) as %
of predicted
species richness
61
55
70
98.4
109.1
85.7
144
Copyright © Verlag Dr. Friedrich Pfeil
However, the genus name Enteromius, being the
oldest African generic-level name available, has
recently been proposed to accommodate all African diploid ‘Barbus’ species (Yang et al., 2015).
Therefore, and although paraphyletic in its current
delineation, this new nomenclature, as further
motivated by Skelton (2015, 2016), has been followed throughout the present paper and is used
here for all small-sized, diploid species found in
the Mono basin.
Raiamas senegalensis is herein reported for the
first time from the Upper Mono. Indeed, although
not reported by Lévêque & Bigorne (1983) in
their revision of the West African Leptocypris and
Raiamas, a single lot is available (MRAC 73-11P-674-684) registered as Barilius macrostoma (i. e.
one of junior synonyms of Raiamas senegalensis).
Paugy & Bénech (1989) reported the capture of
a few specimens of Labeo coubie in the lower and
middle reaches of the Mono basin. However,
although these authors deposited their fish collections at the MNHN, no L. coubie specimens
were found among this material. Nevertheless, a
single specimen (MRAC 97-007-P-0001), collected
at Houndjo-Houndji (≈ 06°18' N 01°50' E) on the
lower course of the Mono was also identified as
L. coubie (see Lévêque, 2003). The scale formula
for this specimen is: 6.5/36/6.5; 4.5; 16. The upper lip is damaged, preventing the examination
of its inner surface, but it is known to be smooth
in L. senegalensis (vs. with transverse folds in
L. coubie). However, the snout of this specimen
lacks tubercles (vs. snout with small, starred,
nuptial tubercles in L. coubie) (Lévêque, 2003). In
addition, the specimen (196 mm SL) has 55 gillrakers on the first gill arch [52-65 (size range 150250 mm SL) for L. senegalensis vs. 37-47 (size range
150-250 mm) for L. coubie] (Lévêque, 2003). This
feature clearly falls outside the range of L. coubie.
Finally, the general appearance of the specimen is
rather pale (as in L. senegalensis) while L. coubie is
dark, with bluish-grey to purplish-black dorsum
and lateral parts) (Lévêque, 2003). Although, both
L. coubie and L. senegalensis are widespread in
western Africa, the former has not been reported
from the coastal basins between the Volta and the
Niger, an area that includes the Mono basin. Taking all these criteria together, the MRAC specimen
was identified as L. senegalensis, a species also collected in the lower course of the Mono during our
recent expeditions (2011-2013). We therefore conclude that L. coubie is absent from the Mono basin.
Alestidae. Six valid species are recorded from the
Mono. However, the taxonomy of one of them,
Brycinus imberi, remains unclear. Preliminary
results of a systematic revision of B. imberi have
raised questions about the conspecificness of the
Mono specimens with the syntypes originating
from the Lower Zambezi River in Mozambique
(Musschoot, pers. comm.). Both populations differ
in the relative position of their fins and in body
depth. As such, the specimens from the Mono are
here referred to as B. cf. imberi.
Three specimens (MRAC 73-014-P-0086,
MRAC 73-005-P-880, and MRAC 73-11-P-56)
previously identified as Brycinus leuciscus are here
re-identified as B. nurse. These specimens lack a
parietal fontanel and have 8 teeth in the external
premaxillary row (vs. fontanel present and 6 teeth
on the external premaxillary row in B. leuciscus,
B. luteus, B. intermedius, B. longipinnis, and B. derhami), and have 5.5 scales above the lateral line (vs.
4.5 in B. carolinae, B. nigricauda, B. imberi, B. brevis,
and B. macrolepidotus). Lévêque et al. (1991) seem
to have reported B. leuciscus from the Mono on
the basis of these erroneous identifications. These
re-identifications thus confirm the absence of the
B. leuciscus in the area as reported by Paugy (1986).
Amphilidae. Two species, Amphilius atesuensis
and Phractura clauseni, are present in the Mono
River basin. An examination of five specimens
(MNHN 1987-0715) captured in the Amou River
at Amou oblo (≈ 07°23' N 00°52' E) and attributed
to P. ansorgii by Paugy & Bénech (1989), concluded
that they are conspecific with Phractura specimens
from the Oulé River at Ezimé (≈ 07°29' N 00°56' E)
(MNHN 1986-0242: 1 specimen) and the Amou
River (MNHN 1986-0243: 1 specimen), identified as P. clauseni by Paugy & Bénech (1989). In
all these re-examined MNHN specimens, the
posterior tip of the pectoral fin does not reach
the base of the ventral fin, which is considered
diagnostic for P. clauseni (vs. the posterior tip of
the pectoral fin does extend to the base of the
ventral fin in P. ansorgii) (Skelton et al., 2003).
Moreover, the anteriormost point of the pelvic-fin
insertion is situated behind the level of a vertical
line drawn through the base of the last dorsal-fin
ray (vs. pelvic-fin insertion located at that level
in P. ansorgii). Therefore, all examined specimens
are here identified as P. clauseni.
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
Mochokidae. One genus and two valid species
of this family are currently found in the basin:
Synodontis cf. obesus and S. ouemeensis. A specimen
of S. melanopterus (MNHN 1981-0923) from the
Mono River has been re-identified as S. cf. obesus during this study. It is characterized by the
following combination of characters: gill slit not
extending beyond pectoral-fin insertion (vs. gill
slit extending ventrally beyond pectoral-fin insertion in S. dekimpei, S. batensoda and S. membranaceus); maxillary barbel unbranched (vs. maxillary
barbel branched in S. resupinatus, S. annectens and
S. clarias); humeral process lacking backwardly
directed spines (vs. humeral process with backwardly directed spines in S. budgetti and S. omias);
maxillary barbel longer than head (vs. maxillary barbel shorter than head in S. vermiculatus,
S. sorex, S. voltae, S. thysi, S. violaceus, S. macrophthalmus, S. courteti, S. xiphias, S. gobroni and
S. guttatus); maxillary barbel with a clearly visible,
broad membrane (vs. maxillary barbel without
membrane or with a barely visible rudiment at its
base in S. punctifer, S. ocellifer, S. tourei, S. koensis,
S. arnoulti, S. schall, S. ouemeensis, S. kogonensis,
and S. levequei); lobes of the caudal-fin having
no black edges (vs. caudal-fin lobes with black
edges: S. filamentosus and S. bastiani); only the
first dorsal-fin ray prolonged into a filament (vs.
at least three dorsal-fin rays prolonged into filaments in S. melanopterus, S. eupterus and S. velifer);
34 mandibular teeth (vs. more than 48 mandibular
teeth in S. ansorgii and S. comoensis); body covered
with numerous small spots (vs. body with few
large spots that are sometimes fused in S. waterloti
and S. robbianus); interorbital distance 36.9 % of
head width (vs. interorbital distance over 50 %
of head width in S. frontosus); post-ocular length
36.8 % of HL and interorbital distance 77.9 % of
snout length (vs. post-ocular length 39.6 up to
43.6 % of HL and interorbital distance 89.3 up to
103.2 % of snout length in S. nigrita).
It should be noted that, owing to their larger
numbers of mandibular teeth [32-43 vs. 21-31
in S. obesus (De Weirdt, pers. comm.)], the identification of several MNHN specimens (MNHN
1981-0923; 1982-0990 and 1986-0321) as well as
RMCA specimens (MRAC B1-026-P-0084-0087)
from the Mono basin as S. obesus (Paugy & Bénech,
1989; Paugy & Roberts, 2003) remains uncertain.
The status of these specimens is currently under
further study (De Weirdt, pers. comm.) as they
might represent a new species endemic to the
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
145
Mono basin and possibly to the Ouémé basin as
well. Pending the results of this study, the Mono
specimens are here attributed to S. cf. obesus.
Musschoot & Lalèyè (2008) described two
new West African Synodontis species, of which
only S. ouemeensis is present in our study area.
These authors concluded that, although S. schall
is widespread in West Africa, it is replaced by
S. ouemeensis in the Mono River. Hence, S. ouemeensis is currently regarded as being endemic to the
Ogun, Mono and Ouémé rivers (Nigeria, Benin
and Togo).
Malapteruridae. Only one representative, Malapterurus beninensis, is present in the Mono River.
The family was only known from MRAC specimens previously identified as M. electricus, which
was thought to be a widespread almost Panafrican
species. Roberts (2000) revalidated M. beninensis
which had been synonymized with M. electricus
by Gosse (1986). This distinction was subsequently
confirmed by Norris (2002). Apart from the Mono
River, M. beninensis is known from the Lower
Volta River in Ghana to the Chiloango River
system in Cabinda (Angola) and the Congo, and
from the island of Fernando Poo (Roberts, 2000;
Norris, 2002).
Ariidae. One specimen identified as Arius latiscutatus and captured in Grand-Popo Lagoon
(Fig. 1) in 1956, was examined in the collections
of the DPB and its identification confirmed. The
species has previously been reported from the
basin by Gras (1961).
Schilbeidae. Two valid species are currently
known from the Mono basin, i.e. Schilbe mystus, reported by Paugy & Bénech (1989) as its
junior synonym S. niloticus (see De Vos, 1995),
and S. intermedius. One specimen (MNHN 19810921) previously identified as S. mystus was
re-identified as S. intermedius following De Vos
(1995, 2003). This specimen lacks an adipose fin
and is therefore not conspecific with S. mystus, nor
with S. micropogon, S. brevianalis or S. mandibularis (see De Vos, 2003). The two remaining West
African species to which this specimen might be
attributed are S. intermedius and S. uranoscopus.
As it has nine gill rakers on the lower limb of the
first gill arch (8-13 in S. intermedius vs. 13-16 in
S. uranoscopus), the MNHN specimen is identified
here as S. intermedius.
146
Copyright © Verlag Dr. Friedrich Pfeil
Nothobranchiidae. Epiplatys togolensis is currently the only species known from the Mono
basin. Originally described as a subspecies of
E. sexfasciatus and elevated to species rank by
Wildekamp (1996), the only currently available
specimen is MNHN 1987-1440.
Poeciliidae. Paugy & Bénech (1989) reported
Aplocheilichthys keilhacki (as Micropanchax keilhacki)
from the Mono basin. In addition, some specimens
of A. keilhacki from the basin are housed at the
RMCA (MRAC 91-52-P-4-7: 16.0-22.3 mm SL).
However, the only valid species currently known
from the basin is Aplocheilichthys spilauchen.
Furthermore, in all four MRAC specimens the
pectoral fins insert below the mid-lateral line
(vs. pectoral fins on or above mid-lateral line in
all other West African species of Poeciliidae, i. e.
Procatopus aberrans, P. similis, Poropanchax normani,
P. rancureli, P. luxophthalmus, Rhexipanchax nimbaensis, R. lamberti, R. kabae, R. schioetzi, Micropanchax
scheeli [= M. keilhacki (see below)], M. bracheti,
M. ehrichi, M. kingie and M. pfaffi). In addition,
these specimens have 8 dorsal-fin rays, 11 or 12
anal-fin rays and 25-28 scales on the mid-lateral
row, formulae that agree well with the description of A. spilauchen as provided by Wildekamp
& Van der Zee (2003).
Moreover, Micropanchax keilhacki has been
described based on two specimens from the Togo
Lagoon near Djeta, southeastern Togo. However,
the species does not appear in the overview of the
West African species by Wildekamp & Van der
Zee (2003) and the genus does not occur in the
Mono (DL, unpublished data).
Sciaenidae. This family primarily consists of
marine species and is represented by a single
species in the Mono basin, i. e. Pseudotolithus (Pseudotolithus) senegalensis, which is known from the
Grand-Popo Lagoon in Benin only. Pseudotolithus
senegalensis has previously been reported from
the Mono basin as Otolithus brachygnathus by
Gras (1961). Although the species is widespread
along the West African coast (Bauchot, 2003a), its
presence in the Mono basin is currently not supported by any natural history museum specimen.
In addition, Vreven & Snoeks (2007) reported that
previous data on Pseudotolithus species in West
and Central Africa should be treated with caution, as there has been much confusion between
P. senegallus, P. senegalensis and P. typus. Here,
we accept the report of P. senegalensis from the
study area by Bauchot (2003a) pending the collection of fresh specimens. The species, however,
is not represented among the material collected
during our recent surveys (2011-2013) in the
Grand-Popo Lagoon.
Drepanidae. The only species from this family
reported from the Mono basin is Drepane africana. This species was erected for the subspecies
D. punctata africana, known from Mauritania to
Angola, while the subspecies D. punctata punctata
(now D. punctata) occurs along the Indo-Pacific
coast (Daget & Iltis, 1965). Gras (1961) was the
first to report this species (as D. punctata) from
the Grand-Popo Lagoon.
Although Drepane africana is well known
and widespread along the West African coast
(Bauchot, 2003b), its presence in the Mono basin
is not supported by any museum specimen. Furthermore, the species was not found during our
recent expeditions (2011-2013) to the Grand-Popo
Lagoon.
Cichlidae. With nine valid species currently
known from the Mono basin, this family is, together with the Cyprinidae, the most species-rich
in the study area. We found MRAC specimens
identified as Tilapia galilaea multifasciata [= Sarotherodon galilaeus multifasciatus] and as T. g. galilaea [= S. g. galilaeus], which would indicate that
both subspecies are found sympatrically in the
Mono. According to Trewavas (1983), these two
subspecies differ from each other in the median
number of dorsal spines: 15 in S. g. multifasciatus
vs. 16 in S. g. galilaeus. Most of the specimens
from the Mono basin have 16 dorsal spines. As
such, they are considered as S. g. galilaeus and not
S. g. multifasciatus, which conforms to the reported
occurrence of the former in the area (see Trewavas,
1983; Teugels & Thys van den Audenaerde, 2003).
The brackish water subspecies Sarotherodon
melanotheron melanotheron was reported from the
Mono at the Grand-Popo Lagoon by Daget (1950)
as its junior synonym Tilapia heudeloti macrocephala
(see Trewavas, 1983). Paugy & Bénech (1989)
and Paugy et al. (1994) reported the species from
the Mono by referring to Daget’s (1950) report.
Although the species is lacking in the existing
collections from the basin, several specimens were
recently collected from the Lower Mono (small
lakes and Grand-Popo Lagoon) and have been
deposited at the RMCA.
Oreochromis niloticus was introduced into
Lederoun et al.: Ichthyofauna of the Mono River basin
147
Copyright © Verlag Dr. Friedrich Pfeil
220 –
200 –
SR = 60.683 · LogSUF - 193.83
R2 = 0.8068
Ng
180 –
Species richness
160 –
Vo
140 –
120 –
Cr
100 –
80 –
60 –
Ou
Kn Sp
Jo
Ni
Mono
Ag
Og Ca
Se
Ko
40 –
Ma
Ba
Ga Co
Sa
Mo
Lo
20 –ı
3.5
ı
ı
ı
ı
ı
4.0
4.5
5.0
5.5
6.0
Log surface area
Fig. 2. Scatterplot of species richness (SR) against the logarithm of the surface area (LogSUF) of twenty West African
coastal river basins. , basins used to obtain the regression line; , position of the Mono basin on the regression
line based on its known surface area (22 000 km2) with an inferred species richness of 70 species; Ag, Agnébi;
Ba, Bandama; Ca, Cavally; Co, Comoé; Cr, Cross; Ga, Gambie; Jo, Jong; Kn, Konkouré; Ko, Kolenté; Lo, Lofa;
Ma, Mano; Mo, Moa; Ni, Nipoué; Ng, Niger; Og, Ogun; Ou, Ouémé; Sa, Sassandra; Se, Sewa; SP, St Paul;
Vo, Volta.
Benin and Togo for the purposes of aquaculture
in 1979 (Lazard, 1990). However, specimens
of O. niloticus escaped from the hatcheries and
quickly established themselves in the nearby
ponds and streams (Lalèyè et al., 2004). The only
specimen reported (but not preserved) from the
main course of the Mono was collected at Atchinédji (≈ 07°34' N 01°21' E), upstream of the Nangbéto dam reservoir, by Paugy & Bénech (1989).
Moreover, Ahouansou Montcho & Lalèyè (2008)
reported the species from Lake Toho (≈ 06°37' N
01°46' E: Lower Mono basin). The Monseigneur
Robert Sastre Aquaculture Station (≈ 06°37' N
01°46' E), from where O. niloticus specimens might
have escaped, is situated in the neighbourhood
of the lake.
Discussion
Paugy & Bénech (1989), based on both a literature review and a study of museum collections,
reported a total of 61 native and one introduced
species (Oreochromis niloticus) for the Mono basin.
Lévêque et al. (1991) reported only 59 native
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
species for the Mono basin, based on a survey of
museum collections only. A compilation of the
species occurrence information, as provided in
the faunal guides to the fresh- and brackish water
fishes of West Africa (Lévêque et al., 1990, 1992;
Paugy et al., 2003a-b) for the Mono basin, gives
a total of 52 native and one introduced species
for the basin.
The present study reports a total of 60 native
and one introduced species. Although this number differs very little from that obtained by Paugy
& Bénech (1989), our list does not include ten of
the species reported by those authors, while incorporating nine others. The species not included
in our list are: Marcusenius brucii, Petrocephalus
sp., Pollimyrus isidori (Mormyridae), Labeo coubie
(Cyprinidae), Phractura ansorgii (Amphiliidae),
Clarias camerunensis (Clariidae), Epiplatys bifasciatus, Foerschichthys flavipinnis (= F. nigeriensis),
Fundulosoma thierryi (Nothobranchiidae) and
Aplocheilichthys keilhacki (= Micropanchax keilhacki)
(Poeciliidae). The species of Clariidae and Nothobranchiidae were omitted because the localities
previously reported for them do not lie within
the Mono basin. Indeed, although they are known
148
Copyright © Verlag Dr. Friedrich Pfeil
from Togo [C. camerunensis from Missahohe
(≈ 06°57' N 00°35' E) and Kousountou (≈ 06°56' N
00°37' E); E. bifasciatus from Agalopé (≈ 06°26' N
01°16' E); F. nigeriensis from Agalopé and from
a tributary of the Lili River between Tsevié and
Aguatopé (≈ 06°26' N 01°15' E); and F. thierryi from
a pond at Assahoun near the railroad (≈ 06°27' N
00°55' E)], their presence in the Mono basin is not
substantiated by any museum specimen.
Furthermore, only a single Marcusenius species, M. senegalensis, has been retained, as M. brucii, which was previously reported from the
Mono basin, is most probably a junior synonym
of M. senegalensis (Boden, pers. comm.). All Petrocephalus specimens examined belong to P. bovei, a
species also reported by Paugy & Bénech (1989).
The sole report of Pollimyrus isidori is based on
misidentifications of P. bovei (see Paugy & Bénech,
1989). Similarly, the earlier reports of Phractura
ansorgii and Aplocheilichthys keilhacki (= Micropanchax keilhacki) are shown here to rest on the
misidentification of P. clauseni and A. spilauchen,
respectively (see Results). The nine species added
to our list (see Table 1) are either based on comparisons with museum specimens (six species) or
on published records (three species).
Some reports of additional species, such as
Brycinus leuciscus, Marcusenius cyprinoides, Petrocephalus levequei and Synodontis melanopterus,
are based on misidentifications or mislocalizations (see Results). Parailia pellucida, previously
reported for the Mono basin by De Vos (2003),
was also removed from the list. That attribution
was presumably based on De Vos’ (1995) citation of Loiselle (1971), who reported the presence of P. pellucida in the Zio River at Toblekope
(≈ 06°17' N 01°13' E: Togo). It is clear, however,
that the Zio River belongs to the Lake Togo basin
(see Paugy & Bénech, 1989). This explains why no
specimens of this species from the Mono basin can
be found in natural history collections. Similarly,
Hippopotamyrus pictus was also not included in
our list as the specimens examined by Lévêque
& Bigorne (1985a) and Bigorne (2003) [MNHN
1981-736, MRAC 73-13-P-43-44 from the Oti River
(≈ 10°40' N 00°47' E) and MNHN 1982-964 from
the Kara River (≈ 10°01' N 00°25' E)] were actually
collected from tributaries of the Volta River basin
in northern Togo. Furthermore, the supposed
presence of Enteromius atakorensis in the Mono
basin mentioned by Lévêque et al. (1991) also
seems to be in error. Indeed, to our knowledge,
the only known specimen from Togo (AMNH
57314), also originates from the Kara River
(≈ 10°01' N 00 °25' E). Finally, two species names
are replaced in our list: Malapterurus beninensis
replaces M. electricus and Synodontis ouemeensis
replaces S. schall in the Mono basin.
Species occurrences reported in this study
were not compared to those cited in Paugy et
al. (1994). These authors established a list of 73
native taxa by using the distribution maps of all
West African species published by Lévêque et al.
(1990, 1992), rather than point sampling, to infer
the presence of species in particular catchment
areas. If a species was reported to be present in
the neighbouring basins to the left and the right,
Paugy et al. (1994) considered it to be present
in the intermediate basin, which is certainly not
always the case (see Gourène et al., 1999). For
example, recent studies have shown that Enteromius atakorensis (see Lévêque, 2003), Pollimyrus
adspersus (see Bigorne, 2003), Poropanchax normani,
Epiplatys spilargyreius (see Wildekamp & Van der
Zee, 2003) and Mastacembelus nigromarginatus (see
Vreven, 2003), reported from the Mono basin by
Paugy et al. (1994), are in fact not present.
The number of species actually present in
the basin almost certainly exceeds the total of 60
native species listed here. The lower course of
the basin has so far been sampled only cursorily
(Fig. 1) and there is reason to believe that further
sampling campaigns in this area will significantly
increase its currently known species richness.
However, using a variety of predictive models,
an average species richness of 62 ± 8 species was
obtained for the whole basin. The overall mean
therefore differs by only two species from our
current estimate of the total species richness of
the entire basin.
Welcomme (1985), Hugueny (1990), Oberdorff
et al. (1993), Tito de Morais & Lauzanne (1994),
Thiel et al. (1995), Koné et al. (2003) and Lalèyè
et al. (2004) showed that the number of species
increases as one proceeds downstream in the river.
According to the present inventory, the upper
and middle courses of the Mono each host more
species than the lower part of the basin. This disparity is likely to be due to a sampling bias, and
supports the hypothesis that particular attention
to sampling in the lower course of the basin will
reveal additional species. Indeed, Lévêque et al.
(1990, 1992) and Da Costa et al. (2000) noted that
the migration of marine and estuarine taxa into
rivers enriches their fish communities, especially
in their lower courses. Strikingly, such taxa are
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
poorly represented (≈ 13 %) in our inventory, and
additional marine and estuarine species, such as
Caranx spp., Eleotris spp., Cynoglossus senegalensis,
Elops spp., are to be expected within the extensive
estuarine area. For example, these taxa represent
≈ 25 % of the fish fauna of Ouémé River (Lalèyè
et al., 2004).
Although several species are characteristic
for each of the different sectors of the basin, this
finding is also likely to result, at least in part,
from sampling biases. The available data on the
ichthyofauna of the upper and middle courses are
mainly based on the collections made by Lévêque,
Paugy and Bénech (see Paugy & Bénech, 1989),
who used multiple fishing techniques. However,
our current knowledge of the ichthyofauna of the
lower course is based on the catches from artisanal
fisheries only.
The species richness of the Mono basin (60
species, 22 000 km2) is broadly comparable to that
of the well-known neighbouring coastal basins,
such as the Ouémé basin to the east [122 species for
50 000 km2 (Lalèyè et al. 2004)] and the Volta basin
to the west [147 species for 398 371 km2 (Hugueny
& Lévêque, 2006)], taking into account the differences in surface area (see Fig. 2). However, 20 species reported from the Mono basin are not known
from the Ouémé basin despite recent sampling undertaken by Lalèyè et al. (2004). The species missing in the Ouémé basin according to this list are:
Amphilius atesuensis, Aplocheilichthys spilauchen,
Arius latiscutatus, Awaous lateristriga, Enteromius
ablabes, E. nigeriensis, E. sublineatus, Brycinus cf.
imberi, Clarias anguillaris, C. buthupogon, Drepane
africana, Epiplatys togolensis, Fundulopanchax (Paludopanchax) filamentosus, Heterobranchus isopterus,
Mormyrus hasselquistii, Nematogobius maindroni,
Phractura clauseni, Pseudotolithus (P.) senegalensis,
Coptodon dageti and Synodontis cf. obesus. However,
only five species, i. e. A. lateristriga (MNHN 19840503), E. sublineatus (MNHN 1982-1349, MNHN
1982-1164, MNHN 1982-1351, MNHN 1982-1352),
E. togolensis (MRAC 73-5-P-3134-148), F. filamentosus (FB 2743415: not seen) and H. isopterus (MNHN
1982-0944), are represented by museum vouchers
from the Ouémé River, while three more species,
i. e. A. spilauchen (see Wildekamp & Van der Zee,
2003), C. anguillaris (see Teugels, 2003a) and
N. maindroni (see Harrisson et al., 2003), are only
listed in the literature, without preserved voucher
specimens. The absence of the remaining 12 species (excluding the marine species A. latiscutatus,
D. africana, and P. senegalensis), from the Ouémé is
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
149
most likely due to limited sampling of the lagoon
area. On the other hand, 80 species (46 marine
or estuarine species and 34 freshwater species)
reported from the Ouémé basin have not been
found in the Mono basin. In the case of marine
and estuarine species, this may be partly due to
the undersampling of the lower regions of the
Mono River (see also above).
Ten species, including the three aforementioned marine species, currently known from the
Mono basin, i. e. Arius latiscutatus, Brienomyrus
brachyistius, Brycinus cf. imberi, Clarias buthupogon,
Drepane africana, Epiplatys togolensis, Fundulopanchax filamentosus, Pseudotolithus senegalensis,
Synodontis cf. obesus and S. ouemeensis, are absent
from the Volta basin. Conversely, 102 species (10
marine or estuarine species and 92 freshwater
species) identified from the Volta basin are absent
from the Mono. Based on these data, it seems that,
despite its modest size, the Mono River harbors
its own particular ichthyofauna, which is not just
a depauperate version of the species diversity
found in its larger neighbours, the Ouémé and
the Volta. Therefore, further detailed documentation of these apparently complex patterns of
species distribution might shed new light on the
hydrographic history of the region.
In evaluating the decline and extinction of
fishes, at least five causes are typically listed
(Helfman, 2007; Tyus, 2011). Helfman (2007) refers
to these as the HIPPO factors: (1) Habitat loss;
(2) Introduced species; (3) Pollution; (4) (human)
Population and consumption; and (5) Overexploitation. Montgomery (2003) had previously
referred to history, i. e. our inability to learn from
past mistakes, as yet another important cause. According to Helfman (2007), for freshwater fishes,
the principal cause of decline and extinction is
habitat degradation, including disruption of the
bottom, removal of structure, water withdrawal,
hydrological alterations (including impoundments) eutrophication, and sediment deposition.
However, generally, a combination of several
of these HIPPO factors together with aspects of
the basin’s history is usually at work (Helfman,
2007; Tyus, 2011). Overall, the importance of each
factor in the erosion of fish diversity has already
been quite well documented and discussed for
other continents (see Miller et al., 1989) but is,
with the exception of Southern African species
(see Skelton, 1990) and the Lake Victoria cichlid
species flock (Harrison & Stiassny, 1999), largely
lacking in Africa as a whole. Indeed, in Southern
150
Copyright © Verlag Dr. Friedrich Pfeil
Africa, the two major, direct causes of decline
among freshwater fishes are habitat destruction
(H) and introduced species (I) (see Skelton, 1990),
whereas for Lake Victoria, the primary factors are
usually habitat alteration (H), competition and
predation by introduced species (Lates niloticus)
(I), overfishing (O) and pollution (P) (see Harrison
& Stiassny, 1999).
In the Mono River, all above-mentioned factors
are present, although in different degrees. The
dam built between 1984 and 1987 at the level of
Nangbéto village in Togo resulted in the formation
of a lake, which covers an area of 180 km2. This
changed the hydrological regime downstream
(see above). Oreochromis niloticus was introduced
into the newly created Lake Nangbéto in 1986 to
support the fisheries (Paugy & Bénech, 1989). This
man-made lake is now the main center of fishing
activity for the entire basin (DL, unpublished
data). The fish fauna of the small lakes in the lower
basin, i. e. mainly cichlids such as Sarotherodon
galilaeus and S. melanotheron, is currently subject
to intense and largely uncontrolled exploitation
(Lederoun et al., 2015, 2016). In addition, many
cotton fields (intensively treated with pesticides)
lie along the banks of the Mono, and their runoffs
drain into this river. Finally, the use of pesticides
and other toxic chemicals for fishing is a common
practice in the Lower Mono basin during the lowwater period, approximately between December
to March (DL, unpublished data). The combination of all these anthropogenic changes has most
probably impacted the fish fauna of the basin;
hence the pressing need to continue monitoring
and documenting its diversity.
In conclusion, it should be noted that, while
the present paper provides an assessment of our
current knowledge of the fish fauna of the Mono,
it is obvious that the existing collections do not
allow us to realistically evaluate the species richness of the river basin prior to the advent of the
Nangbéto dam. Sampling campaigns carried out
before dam construction did not cover the entire
basin, and the area downstream of the dam, in
particular, has been largely neglected. However,
with 60 native species currently known for its
watershed of about 22 000 km2, the fish species
diversity of the Mono basin is relatively similar
to that found in other basins of the West African
ichthyofaunal province. The list presented herein
will undoubtedly be an indispensable asset in assessing the possible impacts of the Nangbéto dam
on the fish diversity of the basin. In addition, it
will supply baseline data for further exploration
of the fish diversity of the basin. This is, among
others, especially true for the Mono downstream
of Adjarala rapids i. e. the lower course of the
Mono and this before the construction of a future
dam at this site.
Acknowledgements
We would like to thank Patrice Pruvost, Romain Causse,
Gaby Zora and Claude Ferrara (MNHN) for the loan of
specimens under their care; Tobias Musschoot (FishBase,
RMCA) for the map of the study area; Gert Boden (FishBase, RMCA) for the digital copy of the MRAC holdings
of the Mono and the surrounding region; Miguël Parrent
(RMCA) for curatorial assistance; Rainer Sonnenberg
(Zoologisches Forschungmuseum Alexander Koenig,
Bonn, Germany) for confirming the identification of
Epiplatys togolensis; and David De Weirdt for sharing his
knowledge of Synodontis. This work is part of the Ph.D.
dissertation of the first author at the University of Liège
funded by a scholarship of the Agence Universitaire de
la Francophonie (AUF) (2010-2013) and supplemented
by a research grant of the International Foundation for
Science (IFS, A/5327-1) (2013-2015).
Material examined. Polypterus senegalus senegalus.
MRAC 2011-026-P-0060-0068, 9, 224.8-282.8 mm TL;
Benin: Codjohoué, Mono River.
Heterotis niloticus. MRAC 2011-026-P-0052, 1,
262.6 mm SL; Bénin: Sazué River at Houndjo-Houndji,
affluent of Mono River.
Brienomyrus brachyistius. MRAC 73-11-P-31-46, 16,
71.6-101.2 mm SL; Togo: Amoutchou River at Ebéva.
Marcusenius senegalensis. MNHN 1985-0140, 2,
130.1-133.1 mm SL; Togo: Amoutchou River at Ebeva.
– MNHN 1991-0941, 1, 89.4 mm SL; Togo: Na River
at Paratao. – MNHN 2002-0799, 1, 81.1 mm SL; Togo:
Ogou at Tchamba. – MRAC 2013-004-P-0114-0123, 10,
110.5-142.9 mm SL; Benin: Togodo, Mono River.
Mormyrops anguilloides. MRAC 2012-021-P-01030110, 8, 166.3-235.5 mm SL; Benin: Togodo, Mono River.
Mormyrus hasselquistii. MRAC 73-11-P-52, 1,
247.1 mm SL; Benin: Alédjo, Mono River.
Mormyrus rume. MRAC 2012-021-P-0111-0113, 3,
160.2-277.9 mm SL; Benin: Togodo, Mono River.
Petrocephalus bovei. MNHN 1985-0755, 1, 82.5 mm
SL; Benin: Atchinedji, Mono River. – MNHN 1985-0756,
1, 67.7 mm SL; Benin: Tététou, Mono River. – MRAC
2012-021-P-0098-0099, 2, 57.9-58.2 mm SL; Benin: Adjarala, Mono River.
Enteromius ablabes. MNHN 1988-1803, 2, 43.255.1 mm SL; Togo: Amou River at Oblo. – MRAC 73-5P-2413-468, 50, 17.8-43.0 mm SL; Togo: Agbofon near
Atakpamè, Mono River. – MRAC 76-32-P-4077-4078,
2, 19.0-30.3 mm SL; Togo: 5-10 miles for d’Atakpamé,
Mono River.
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
Enteromius callipterus. MNHN 1989-0540, 2, 46.253.9 mm SL; Togo: Amou River at Amou oblo. – MRAC
2013-004-P-0038-0062, 25, 40.6-43.3 mm SL; Benin:
Lomon River at Hontomè.
Enteromius chlorotaenia. MNHN 1981-992, 2, 71.873 mm SL; Togo: Atakpame, Mono River.
Enteromius nigeriensis. MRAC 2012-021-P-00010016, 16, 36.8-49.0 mm SL; Benin: Djonnougui, Mono
River. – MRAC 2013-004-P-0001-0032, 32, 33.9-48.8 mm
SL; Benin: Djonnougui, Mono River.
Enteromius sublineatus. MNHN 1982-1353, 1,
69.7 mm SL; Togo: Tététou, Mono River. – MNHN
1982-1165, 1, 78.6 mm SL; Togo: Kpessi, Mono River.
– MRAC 2013-004-P-0063-0089, 27, 14.0-45.9 mm SL;
Benin: Paratago to the right of the bridge on the way
to Aledjo, Mono River.
Labeo parvus. MRAC 73-5-P-1891, 1, 159.7 mm SL;
Togo: Agbofon near Atakpamè, Mono River. – MRAC
73-14-P-229-234, 6, 100.5-143.7 mm SL; Togo: Agbofon
near Atakpamè, Mono River. – MRAC 73-14-P-236242, 7, 79.9-165.3 mm SL; Togo: Ogbone, Mono River.
– MRAC 2013-004-P-0104-0108, 5, 84.9-239.4 mm SL;
Benin, Togodo, Mono River.
Labeo senegalensis. MNHN 1981-0913, 1, 141.5 mm
SL; Togo: Tététou, Mono River. – MRAC 97-007-P-0001,
1, 196 mm SL; Benin: Sazué River at Houndjo-Houndji,
affluent of Mono River. – MRAC 2012-021-P-0045-0048,
4, 110.7-134.8 mm SL; Benin: Togodo, Mono River.
Raiamas senegalensis. MRAC 73-11-P-674-684, 11,
91.8-163.5 mm SL; Togo: Fazao, Mono River.
Distichodus rostratus. MNHN 1981-0888, 1, 152.3 mm
SL; Togo: Tététou, Mono River. – MRAC 2012-021-P0102, 1, 188.8 mm SL; Benin: Djossouhé, Mono River.
Brycinus cf. imberi. MRAC 73-11-P-0059, 1, 73.2 mm
SL; Togo: Tététou, Mono River. – MRAC 2012-021-P0022-0028, 7, 80.2-91.9 mm SL; Togo: Nangbéto dam
at Atakè, Mono River.
Brycinus longipinnis. MRAC 73-014-P-0191-0228,
38, 61.0-70.0 mm SL; Togo: Dotékopé, Mono River.
– MRAC 2012-021-P-0029-0041, 13, 50.6-58.0 mm SL;
Togo: Nangbéto dam at Akodéseva, Mono River.
Brycinus macrolepidotus. MRAC 73-05-P-619-621,
3, 96.7-117.5 mm SL; Togo: Corrékopé, Mono River.
– MRAC 73-14-P-55-59, 3, 92.7-114.5 mm SL; Togo:
Kolékopé, Mono River. – MRAC 73-14-P-60-81, 22, 84.0130.4 mm SL; Togo: Dotékopé, Mono River. – MRAC
2011-026-P-0015-0019, 5, 130.9-178.6 mm SL; Benin:
Adjarala, Mono River.
Brycinus nurse. MRAC 73.005-P-880, 1, 149.2 mm
SL; Togo: Corrokopé, Mono River. – MRAC 73-11-P-56;
1, 107.3 mm SL; Togo: Ebeva, Mono River. – MRAC
73-014-P-0086, 1, 86.0 mm SL; Dotékopé, Mono River.
– MRAC 2011-026-P-0020-0021, 2, 140.5-142.5 mm SL;
Togo: Nangbéto dam at the end of the dike, Mono River.
Hydrocynus forskalii. MRAC 73-14-P-53-54, 2, 143.5208.5 mm SL; Togo: Ogbone, Mono River.
Rhabdalestes septentrionalis. MNHN 2000-0650, 2,
35.9-40.9 mm SL; Togo: Kpessi, Mono River. – MNHN
2000-0649, 2, 28.4-36.7 mm SL; Togo: Tététou, Mono
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
151
River. – MRAC 2012-021-P-0049-0088, 40, 47.4-54.7 mm
SL; Togo: Nangbéto dam at Akodéseva, Mono River.
Hepsetus odoe. MRAC 2012-021-P-0043-0044, 2,
160.5-166.5 mm SL; Benin: Togodo, Mono River.
Amphilius atesuensis. MNHN 1987-0711, 5, 49.164.8 mm SL; Togo: Amou River at Amou oblo.
Phractura clauseni. MNHN 1986-0242, 1, 68.8 mm
SL; Togo: Oule River at Ezime. – MNHN 1986-0243; 1,
46,7 mm SL; Togo: Amou River at Amou oblo. – MNHN
1987-0715, 5, 39.5-61.4 mm SL; Togo: Amou River at
Amou oblo. – MNHN 1987-716, 1, 64.6 mm SL; Togo:
Oule River at Ezime.
Synodontis cf. obesus. MRAC 17-25-P-37-38, 2, 52.856.0 mm SL; Togo: Tchamba, Mono River. – MRAC
73-14-P-359-362, 4, 104.5-144.0 mm SL; Togo: Kolékopé,
Mono River. – MRAC 73-14-P-363-368, 6, 89.5-159.5 mm
SL; Togo: Ogbone, Mono River. – MNHN 1981-0923, 1,
133.3 mm SL; Togo: Tététou, Mono River. – MNHN 19820990, 1, 97.2 mm SL; Togo: Tététou, Mono River. – MNHN
1986-0321, 1, 114.3 mm SL; Togo: Atchinedji, Mono River.
Synodontis ouemeensis. MNHN 1981-927, 1, 104 mm
SL; Togo: Atchinedji, Mono River. – MNHN 1981-928,
1, 149 mm SL; Togo: Tététou, Mono River. – MNHN
1982-995, 1, 55 mm SL; Kpessi, Mono River. – MNHN
2002-0783, 1, 107 mm SL; Togo: Atchinedji, Mono River.
Malapterurus beninensis. MRAC 73-11-P-797-807,
11, 61.5-168.0 mm SL; Togo: Ebeva, Mono River.
Clarias (Clarioides) agboyiensis. MRAC P-73072.0115,
1, 102.7 mm SL; Togo: Botike-Zogue, river between
Aufouin and Atlekogou. – MNHN 2002-783, 2, 66.980.2 mm SL; Togo: Anié River at Sotouboua.
Clarias (Clarias) anguillaris. MRAC 2013-004-P0109, 1, 203.0 mm SL; Benin: Grand-Popo Lagoon at
Onkuhoué, Mono River.
Clarias (Clarioides) buthupogon. MNHN 1986-0404, 3,
63.7-65.7 mm SL; Togo: Aou losso, Mono River. – MNHN
1986-0406, 1, 78.5 mm SL; Togo: Kri-kri, Mono River.
Clarias (Clarias) gariepinus. MRAC 2013-004-P-0110,
1, 203.0 mm SL; Benin: Grand-Popo Lagoon at Houssoukoé, Mono River.
Heterobranchus isopterus. MRAC 73-05-P-3021-3022,
2, 83.3-103.3 mm SL; Togo: Dotékopé, Mono River. –
MRAC 2013-004-P-0111-0112, 2, 133.0-155.7 mm SL;
Benin: Lomon River at Hounssahoué.
Heterobranchus longifilis. MRAC 2011-026-P-0051,
1, 221.8 mm SL; Benin: Djonnougui, Mono River.
Chrysichthys (Chrysichthys) auratus. MRAC 2011026-P-0035-0037, 3, 102.3-122.7 mm SL; Benin: Vodomey,
Mono River.
Chrysichthys (Melanodactylus) nigrodigitatus. MRAC
2011-026-P-0034, 1, 105.6 mm SL; Togo: Nangbéto dam
at Akodéséva, Mono River.
Schilbe intermedius. MNHN 1981-0921, 1, 181.3 mm
SL; Togo: Tététou, Mono River. – MRAC 2011-026-P-00710073, 3, 129.5-171.8 mm SL; Benin: Adjarala, Mono River.
Schilbe mystus. MRAC 2011-026-P-0074-0083, 10,
143.5-259.8 mm SL; Benin: Adjarala, Mono River.
Epiplatys togolensis. MNHN 1987-1440, 1, 34.1 mm
SL; Togo: Amou River at Amou oblo.
152
Copyright © Verlag Dr. Friedrich Pfeil
Fundulopanchax (Paludopanchax) filamentosus. MRAC
73-72-P-201-218, 18, 14.0-27.9 mm SL; Togo: AklakouMolokou, Mono River.
Aplocheilichthys spilauchen. MRAC 91-52-P-4-7, 4,
16.0-22.3 mm SL; Togo: Agamè, Mono River.
Lates niloticus. MRAC 2011-026-P-0053-0054, 2,
202.7-232.7 mm SL; Togo: Nangbéto dam at Akodéseva,
Mono River.
Chromidotilapia guntheri guntheri. MRAC 2011-026P-0032-0033, 2, 82.5-83.6 mm SL; Togo: Nangbéto dam
at Djatokopé, Mono River.
Coptodon dageti. MRAC 73-61-P-1-3, 3, 93.8120.7 mm SL; Togo: 36 km E of Ayengré, Mono River.
Coptodon guineensis. MRAC 2011-026-P-0089, 1,
193.4 mm SL; Benin: Gbagan Lagoon at Zogbédji, Mono
River.
Coptodon zillii. MRAC 73-14-P-448, 1, 149.5 mm SL;
Togo: Kolékopé, Mono River. – MRAC 2011-026-P-0090,
1, 129.0 mm SL; Benin: Codjohoué, Mono River. – MRAC
2013-004-P-0132, 1, 79.9 mm SL; Benin: Lomon River
at Hounssahoué under the bridge on the way to the
Tohoun border.
Hemichromis bimaculatus. MRAC 2011-026-P-00390046, 8, 43.0-67.5 mm SL; Benin: Lac Loké at Agbodo,
Mono River.
Hemichromis fasciatus. MRAC 2011-026-P-00470050, 4, 66.1-117.5 mm SL; Togo: Nangbéto dam at the
end of the dike, Mono River.
Oreochromis niloticus. MRAC 2011-026-P-0055-0059,
5, 80.1-126.4 mm SL; Togo: Nangbéto dam at Akodéséva, Mono River.
Sarotherodon galilaeus galilaeus. MRAC 2012-021-P0118, 1, 99.4 mm SL; Benin: Lake Doukon at Doukonta,
Mono River.
Sarotherodon melanotheron melanotheron. MRAC
73-14-P-396-397, 2, 136.0-178.2 mm SL; Togo: Korrékopé, Mono River. – MRAC 73-14-P-441-447, 7, 125.5177.0 mm SL; Togo: Dotékopé, Mono River. – MRAC
73-14-P-449-472, 24, 41.5-58.7 mm SL; Togo: Dotékopé,
Mono River. – MRAC 2011-026-P-0069-0070, 2, 118.9129.3 mm SL; Benin: Lac Toho at Logbo, Mono River.
Awaous lateristriga. MRAC 73-14-P-498-500, 3,
102.0-106.2 mm SL; Togo: Kolékopé, Mono River. –
MRAC 73-14-P-501-502, 2, 96.0-104.0 mm SL; Togo:
Dotékopé, Mono River. – MRAC 2011-026-P-0001-0005,
5, 65.0-89.5 mm SL; Benin: Djonnougui, Mono River.
Nematogobius maindroni. MNHN 2000-0640, 1,
57.5 mm SL; Togo: Kpessi, Mono River. – MNHN 19880480, 2, 49.4-51.4 mm SL; Togo: Tététou, Mono River.
Ctenopoma kingsleyae. MRAC 2011-026-P-0038, 1,
95.0 mm SL; Benin: Togodo, Mono River. – MRAC
2012-021-P-0100-0101, 2, 89.7-100.0 mm SL; Benin: Lac
Toho at Logbo, Mono River.
Parachanna obscura. MNHN 1988-0478, 1, 52.7 mm
SL; Togo: Amou River at Amou oblo. – MRAC 2012021-P-0114-0117, 4, 143.9-178.0 mm SL; Benin: Lake
Toho at Logbo, Mono River.
Protopterus annectens annectens. MRAC 2013-004-P0131, 1, 239.8 mm TL; Benin: Djonnougui, Mono River.
Literature cited
Ahouansou Montcho, S. & P. A. Lalèyè. 2008. Some
aspects of biology of Oreochromis niloticus L. (Perciformes: Cichlidae) recently introduced in Lake
Toho (Benin, West Africa). International Journal of
Biological and Chemical Science, 2: 114-122.
Albaret, J.-J. 1994. Les poissons: biologie et peuplements.
Pp. 239-279 in: J. R. Durand, P. Dufour, D. Guiral
& S. G. Zabi (eds.), Environnement et ressources
aquatiques de Côte d’Ivoire. Tome 2: les milieux
saumâtres. L’exemple de la lagune Ebrié. ORSTOM, Paris.
Amoussou, E. 2010. Variabilité pluviométrique et dynamique hydro-sédimentaire du bassin-versant du
complexe fluvio-lagunaire Mono-Ahémé-Couffo
(Afrique de l’Ouest). Thèse présentée pour l’obtention du doctorat en Géographie-Spécialité:
Géographie physique appliquée. Université de
Bourgogne, Paris, 315 pp.
Anonymous. 1962. Données connues du bassin du Mono
in: Possibilité d’Aménagement Hydroagricole dans
le bassin du Mono. Office de la Recherche Scientifique et Technique Outre-Mer: 1-19.
— 1992. Barrage d’Adjarala (Togo-Bénin). Étude d’impacts hydrauliques et sédimentologiques. Rapport
d’études préparé par Coyne & Bellier. Université de
Bordeaux III, Institut de Géographie, Paris, 105 pp.
— 1997. Étude d’environnement de l’aménagement
hydroélectrique d’Adjarala (Togo-Bénin): Impacts
hydrobiologiques sur le système fluvio-lagunaire.
Résumé préparé par Coyne & Bellier. Communauté
Electrique du Bénin-Banque Mondiale, 59 pp.
Banister, K. E. & R. G. Bailey. 1979. Fishes collected
by the Zaïre River Expedition, 1974-75. Zoological
Journal of the Linnean Society, 66: 205-249.
Bauchot M. L. 2003a. Sciaenidae. Pp. 504-511 in: D.
Paugy, C. Lévêque & G. G. Teugels (eds.), Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 2. Collection Faune et Flore
Tropicales 40. MRAC & IRD, Tervuren & Paris.
— 2003b. Drepanidae. Pp. 514-515 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 2. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
Berrebi, P. 1998. Genetic divergence among morphotypes of Lake Tana (Ethiopia) barbs. Biological
Journal of the Linnean Society, 64: 369-384.
Berrebi, P., M. Kottelat, P. Skelton & P. Rab. 1996.
Systematics of Barbus: state of the art and heuristic
comments. Folia Zoologica, 45: 5-12.
Berrebi, P., C. Lévêque, G. Cattaneo-Berrebi, J. F. Agnèse,
J. F. Guégan & A. Machordom. 1990. Diploid and
tetraploid African Barbus (Osteichthyes, Cyprinidae): on the coding of differential gene expression.
Aquatic Living Resources, 3: 313-323.
Bigorne, R. 2003. Mormyridae. Pp. 155-222 in: D. Paugy,
C. Lévêque & G. G. Teugels (eds.), Faune des
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 1. Collection Faune et Flore
Tropicales 40. MRAC & IRD, Tervuren & Paris.
Bigorne, R. & D. Paugy. 1990. Description de Marcusenius
meronai, espèce nouvelle de Mormyridae (Teleostei)
de Sierra Leone. Ichthyological Exploration of
Freshwaters, 1: 33-38.
Bigorne, R. & D. Paugy. 1991. Note sur la systématique
des Petrocephalus (Teleostei: Mormyridae) d’Afrique
de l’Ouest. Ichthyological Exploration of Freshwaters, 2: 1-30.
Da Costa, K. S., G. Gourène, L. Tito De Morais & D. F.
E. Thys van den Audenaerde. 2000. Caractérisation
des peuplements ichtyologiques de deux fleuves
côtiers ouest-africains soumis à des aménagements
hydroagricoles et hydroélectriques. Vie et Milieu,
50: 65-77.
Daget, J. 1950. Poissons d’eau douce de la région côtière
du Togo et du Dahomey. Notes Africaines, 46: 57-59.
— 1984. Osteoglossidae. Pp. 57-58 in: J. Daget, J.
P. Gosse & D. F. E. Thys van den Audenaerde
(eds.), Check-list of the Freshwater Fishes of Africa
(CLOFFA), Volume I. ORSTOM & MRAC, Paris &
Tervuren.
Daget, J. & A. Iltis. 1965. Poissons de Côte d’Ivoire
eaux douces et saumâtres. Mémoire de l’Institut
Fondamental d’Afrique Noire, 74, 385 pp.
Decru, E., J. Snoeks & E. Vreven. 2013. The true identity
of the holotype of Hepsetus odoe and the names of
the two West African species of Hepsetus (Teleostei:
Hepsetidae). Ichthyological Exploration of Freshwaters, 24: 187-192.
Decru, E., E. Vreven & J. Snoeks. 2012. A revision of the
West African Hepsetus (Characiformes; Hepsetidae)
with a description of H. akawo sp. nov. and a redescription of H. odoe (Bloch, 1794). Journal of Natural
History, 46: 1-23.
De Vos, L. 1995. A systematic revision of the African
Schilbeidae (Teleostei, Siluriformes) with an annotated bibliography. Annales du Musée Royal de
l’Afrique Centrale, Sciences Zoologiques, 271: 1-450.
— 2003. Schilbeidae. Pp. 98-116 in: D. Paugy, C. Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 2. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
Doadrio, I. 1990. Phylogenetic relationships and classification of western palearctic species of the genus
Barbus (Osteichthyes, Cyprinidae). Aquatic Living
Resources, 3: 265-282.
Dunz, A. R. & U. K. Schliewen. 2013. Molecular phylogeny and revised classification of the haplotilapiine
cichlid fishes formerly referred to as “Tilapia”.
Molecular Phylogenetics and Evolution, 68: 64-80.
Golubtsov, A. S. & E. Y. U. Krysanov. 1993. Karyological
study of some cyprinid species from Ethiopia. The
ploidy differences between large and small Barbus
of Africa. Journal of Fish Biology, 42: 445-455.
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
153
Gosse, J. P. 1963. Le milieu aquatique et l’écologie des
poissons dans la région de Yangambi. Annales
du Musée Royal de l’Afrique Centrale, Sciences
Zoologiques, 116: 113-270.
— 1984. Polypteridae. Pp. 18-29 in: J. Daget, J. P. Gosse
& D. F. E. Thys van den Audenaerde (eds.), Checklist of the Freshwater Fishes of Africa (CLOFFA)
Volume I. ORSTOM & MRAC, Paris & Tervuren.
Gosse, J. P. 1986. Malapteruridae. Pp. 102-104 in: J.
Daget, J. P. Gosse & D. F. E. Thys van den Audenaerde (eds.), Check-list of the Freshwater Fishes
of Africa (CLOFFA), Volume 2. ISNB, MRAC &
ORSTOM, Bruxelles, Tervuren & Paris.
Gosse, J. P. 1990. Polypteridae. Pp. 79-87 in: C. Lévêque,
D. Paugy & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Tome I. Faune tropicale XXVIII. MRAC & ORSTOM,
Tervuren & Paris.
Gourène, G., G. G. Teugels, B. Hugueny, F. E. Dard
& D. F. E Thys van den Audenaerde. 1999. Evaluation de la diversité ichtyologique d’un bassin
ouest-africain après la construction d’un barrage.
Cybium, 23: 147-160.
Gras, R. 1961. Liste des poissons du bas-Dahomey
faisant partie de la collection du laboratoire d’hydrobiologie du service des eaux, forêts et chasses
du Dahomey. Bulletin IFAN, 23, Ser. A, 2: 573-586.
Guégan, J. F., P. Rab, A. Machordom & I. Dadrio. 1995.
New evidence of hexaploidy in ‘large’ African
Barbus with some consideration on the origin of
hexaploidy. Journal of Fish Biology, 47: 192-198.
Harrisson, I. J. & M. J. Stiassny. 1999. The quiet crises:
a preliminary listing of the freshwater fishes of the
world that are extinct or “missing in action”. Pp.
271-332 in: R. D. E. MacPhee & H.-D. Sues (eds.),
Extinctions in near time: cause, contexts, and consequences. Kluwer Academic/Plenum Publishing,
New York.
Harrisson, I. J., P. J. Miller & F. Pezold. 2003. Gobiidae.
Pp. 626-666 in: D. Paugy, C. Lévêque & G. G.
Teugels (eds.), Faune des poissons d’eaux douces
et saumâtres de l’Afrique de l’Ouest. Volume 2.
Collection Faune et Flore Tropicales 40. MRAC &
IRD, Tervuren & Paris.
Helfman, G. S. 2007. Fish conservation: a guide to understanding and restoring global aquatic biodiversity
and fishery resources. Washington, D. C., Island
Press, 584 pp.
Hugueny, B. 1990. Richesse des peuplements de poissons dans le Niandan (haut Niger, Afrique) en
fonction de la taille de la rivière et de la diversité
du milieu. Revue d’Hydrobiologie Tropicale, 23:
351-364.
Hugueny, B. & C. Lévêque. 2006. Richesse en espèce des
peuplements de poissons. Pp. 237-249 in: Lévêque,
C. & D. Paugy (eds.), Les poissons des eaux continentales africaines: diversité, écologie, utilisation
par l’homme. IRD, Paris.
154
Copyright © Verlag Dr. Friedrich Pfeil
Jégu, M. & C. Lévêque. 1984. Le genre Marcusenius
(Pisces, Mormyridae) en Afrique de l’Ouest. Revue
d’Hydrobiologie Tropicale, 17: 335-358.
Koné, T., G. G. Teugels, V. N’Douba, G. Gooré Bi &
E. P. Kouamelan. 2003. Premières données sur
l’inventaire et la distribution de l’ichtyofaune d’un
petit bassin côtier ouest africain: rivière Gô (Côte
d’Ivoire). Cybium, 27: 101-106.
Laïbi, R. A., A. Gardel, E. J. Anthony & L. M. Oyédé. 2012.
Apport des séries d’images landsat dans l’étude de
la dynamique spatio-temporelle de l’embouchure
de l’estuaire des fleuves Mono et Couffo au Bénin,
avant et après la construction du barrage de Nangbéto sur le Mono. Revue Télédétection, 10: 179-188.
Lalèyè, P., A. Chikou, J.-C. Philippart, G. Teugels & P.
Vandewalle. 2004. Étude de la diversité ichtyologique du bassin du fleuve Ouémé au Bénin (Afrique
de l’Ouest). Cybium, 28: 329-339.
Lazard, J. 1990. Transferts de poissons et développement
de la production piscicole. Exemple de trois pays
d’Afrique subsaharienne. Revue d’Hydrobiologie
Tropicale, 23: 251-265.
Lederoun, D., A. Chikou, E. Vreven, J. Snoeks, J. Moreau, P. Vandewalle & P. Lalèyè. 2015. Population
parameters and exploitation rate of Sarotherodon
melanotheron melanotheron Rüppell, 1852, Cichlidae,
in Lake Toho (Benin-West-Africa). Journal of Biodiversity and Environmental Sciences, 66: 259-271.
Lederoun, D., P. Vandewalle, A. A. Brahim, J. Moreau
& P. Lalèyè. 2016. Population parameters and
exploitation rate of Sarotherodon galilaeus galilaeus
(Cichlidae) in Lakes Doukon and Togbadji, Benin.
African Journal of Aquatic Science, 41: 151-160.
Lévêque, C. 2003. Cyprinidae. Pp. 322-436 in: D. Paugy,
C. Lévêque & G. G. Teugels (eds.), Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 1. Collection Faune et Flore
Tropicales 40. MRAC & IRD, Tervuren & Paris.
Lévêque, C. & R. Bigorne. 1983. Révision des Leptocypris et Raiamas (Pisces, Cyprinidae) de l’Afrique
de l’Ouest. Revue d’Hydrobiologie Tropicale, 16:
373-393.
Lévêque, C. & R. Bigorne. 1985a. Le genre Hippopotamyrus (Pisces, Mormyridae) en Afrique de l’Ouest,
avec la description d’Hippopotamyrus paugyi n. sp.
Cybium, 9: 175-192.
Lévêque, C. & R. Bigorne. 1985b. Répartition et variabilité des caractères méristiques et métriques des
espèces du genre Mormyrus (Pisces – Mormyridae)
en Afrique de l’Ouest. Cybium, 9: 325-340.
Lévêque, C., D. Paugy & G. G. Teugels. 1990. Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Tome I. Faune tropicale XXVIII. MRAC
& ORSTOM, Tervuren & Paris, 384 pp.
Lévêque, C., D. Paugy & G. G. Teugels. 1991. Annotated
check-list of the freshwater fishes of the Nilo-Sudan
river basins, in Africa. Revue d’Hydrobiologie
Tropicale, 24: 131-154.
Lévêque, C., D. Paugy & G. G. Teugels. 1992. Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Tome II. Faune tropicale XXVIII. MRAC
& ORSTOM, Tervuren & Paris, 526 pp.
Loiselle, P. V. 1971. The fishes of the Zio River at
Togbkelope. Biology data on Schilbe mystus (err.
for S. intermedius) and Physailia pellucida = Parailia
pellucida). Part 2. Aquarium, 4: 50-60.
Micha, J. C. & V. Frank. 1976. Biologie des principales
espèces utilisées en pisciculture africaine. CIFA
Technical Paper, 4: 292-331.
Machordom, A. & I. Doadrio. 2001. Evolutionnary history and speciation modes in the cyprinid genus
Barbus. Proceedings of the Royal Society, Biological
Sciences, 268: 1297-1306.
Miller, R. R., J. D. Williams & J. E. Williams. 1989. Extinctions of North American fishes during the past
century. Fisheries, 14: 22-38.
Montgomery, D. R. 2003. King of fish: the thousand-year
run of salmon. Westview, Boulder, 290 pp.
Musschoot, T. & P. Lalèyè. 2008. Designation of a neotype for Synodontis schall (Bloch and Schneider, 1801)
and description of two new species of Synodontis
(Siluriformes: Mochokidae). Journal of Natural
History, 42: 1303-1331.
Nelson, J. S., T. C. Grande, & M. V. H. Wilson. 2016.
Fishes of the World. Fifth edition. John Wiley.
Hoboken, 752 pp.
Norris, S. M. 2002. A revision of the African electric
catfishes, family Malapteruridae (Teleostei, Siluriformes), with erection of a new genus and descriptions of fourteen new species and an annotated
bibliography. Musée Royale de l’Afrique Centrale,
Tervuren, Annales Sciences Zoologiques, 289: 1-155.
— 2003. Malapteruridae. Pp. 174-194 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 1. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
Oberdorff, T., E. Guilbert & J. C. Lucchetta. 1993. Patterns of fish species richness in the Seine River basin,
France. Hydrobiologia, 259: 157-167.
Oyédé, L. M. 1991. Dynamique sédimentaire actuelle
et messages enregistrés dans les séquences quartenaires et néogènes du domaine margino littoral du
Bénin (l’Afrique de l’Ouest). Thèse présentée pour
l’obtention du doctorat en géologie sédimentaire,
nouveau régime. Université de Bourgogne, Paris,
302 pp.
Paugy, D. 1986. Révision systématique des Alestes et
Brycinus africains (Pisces, Characidae). ORSTOM,
Études et thèses, 295 pp.
— 2003a. Osteoglossidae. Pp.145-147 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 1. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
— 2003b. Hepsetidae. Pp. 232-234 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
Lederoun et al.: Ichthyofauna of the Mono River basin
Copyright © Verlag Dr. Friedrich Pfeil
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 1. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
— 2003c. Alestidae. Pp. 236-282 in: D. Paugy, C. Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 1. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
— 2003d. Centropomidae. Pp. 451-453 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 2. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
Paugy, D. & V. Bénech. 1989. Les poissons d’eau douce
des bassins côtiers du Togo (Afrique de l’Ouest).
Revue d’Hydrobiologie Tropicale, 22: 295-316.
Paugy, D., V Bénech & K. Etou. 1988. La faune ichtyologique des bassins du Mono et du lac Togo (Togo).
Convention OMS-ORSTOM, 23: 1-110.
Paugy, D., C. Lévêque & G. G. Teugels. 2003a. Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 1. Collection Faune et Flore Tropicales 40. MRAC & IRD, Tervuren & Paris, 457 pp.
Paugy, D., C. Lévêque & G. G. Teugels. 2003b. Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 2. Collection Faune et Flore Tropicales 40. MRAC & IRD, Tervuren & Paris, 815 pp.
Paugy, D. & T. R. Roberts. 2003. Mochokidae. Pp. 196268 in: D. Paugy, C. Lévêque & G. G. Teugels (eds.),
Faune des poissons d’eaux douces et saumâtres de
l’Afrique de l’Ouest. Volume 2. Collection Faune et
Flore Tropicales 40. MRAC & IRD, Tervuren & Paris.
Paugy, D., K. Traoré & P. S. Diouf. 1994. Faune ichtyologique des eaux douces d’Afrique de l’Ouest. Pp.
35-66 in: G. G. Teugels, J. F. Gueguan & J. J. Albaret
(eds.), Diversité biologique des poissons des eaux
douces et saumâtres d’Afrique: synthèses géographiques: symposium PARADI. Annales du Musée
Royal de l’Afrique Centrale.
Risch, L. M. 2003. Claroteidae. Pp. 60-96 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 1. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
Roberts, T. R. 2000. A review of the African electric catfish family Malapteruridae, with descriptions of new
species. Occasional Papers in Ichthyology, 1: 1-15.
Seegers, L., L. De Vos & D. O. Okeyo. 2003. Annotated
checklist of the freshwater fishes of Kenya (exluding
the lacustrine haplochromines from Lake Victoria).
Journal of East African Natural History, 92: 11-47.
Skelton, P. H. 1990. The conservation and status of
threatened fishes in southern Africa. Journal of Fish
Biology, 37, supplement A: 87-95.
— 2001. A complete guide to the freshwater fishes of
southern Africa. Second edition. Stuik Publishers,
395 pp.
— 2015. Fishes. Pp. 96-123 in: C. Griffiths, J. Day &
M. Picker (eds.), Freshwater life: a field guide to
Ichthyol. Explor. Freshwaters, Vol. 28, No. 2
155
the plants and animals of southern Africa. Struik
Nature & Penguin Random House, Cape Town.
— 2016. Name changes and additions to the southern
African freshwater fish fauna. African Journal of
Aquatic Science, 41: 1-7.
Skelton, P. H., L. M. Risch & D. Paugy. 2003. Amphiliidae. Pp. 117-143 in: D. Paugy, C. Lévêque & G. G.
Teugels (eds.), Faune des poissons d’eaux douces
et saumâtres de l’Afrique de l’Ouest. Volume 2.
Collection Faune et Flore Tropicales 40. MRAC &
IRD, Tervuren & Paris.
Teugels, G. G. 2003a. Clariidae. Pp. 144-173 in: D.
Paugy, C. Lévêque & G. G. Teugels (eds.), Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 2. Collection Faune et Flore
Tropicales 40. MRAC & IRD, Tervuren & Paris.
— 2003b. Channidae. Pp. 443-446 in: D. Paugy, C.
Lévêque & G. G. Teugels (eds.), Faune des poissons
d’eaux douces et saumâtres de l’Afrique de l’Ouest.
Volume 2. Collection Faune et Flore Tropicales 40.
MRAC & IRD, Tervuren & Paris.
Teugels, G. G. & D. F. E. Thys van den Audenaerde.
2003. Cichlidae Pp. 520-600 in: D. Paugy, C. Lévêque
& G. G. Teugels (eds.), Faune des poissons d’eaux
douces et saumâtres de l’Afrique de l’Ouest. Volume
2. Collection Faune et Flore Tropicales 40. MRAC
& IRD, Tervuren & Paris.
Thiel, R., A. Sepulveda, R. Kafemann & W. Nellen.
1995. Environmental factors as forces structuring
the fish community of the Elbe Estuary. Journal of
Fish Biology, 46: 47-69.
Tito De Morais, L. & L. Lauzanne. 1994. Zonation longitudinale des peuplements ichtyologiques avant
la mise en eau de la retenue de Petit-Saut (Guyane
française). Revue d’Hydrobiologie Tropicale, 27:
467-483.
Trewavas, E. 1983. Tilapiine fishes of the genera Sarotherodon, Oreochromis and Danakilia. British Museum
(Natural History), London, 583 pp.
Tyus, H. M. 2011. Ecology and Conservation of Fishes.
CRC Press. Boca Raton, London, New York, 573 pp.
USBGN, 1965. Dahomey: official standard names approved by the United States Board on Geographic
Names. Washington, District of Columbia: Office of
Geography, Department of the Interior, Gazetteer,
91, iv + 89 pp.
— 1966. Togo: official standard names approved by
the United States Board on Geographic Names.
Washington, District of Columbia: Office of Geography, Department of the Interior, Gazetteer, 98,
iii + 100 pp.
Vreven, E. 2003. Mastacembelidae. Pp. 708-737 in: D.
Paugy, C. Lévêque & G. G. Teugels (eds.), Faune des
poissons d’eaux douces et saumâtres de l’Afrique
de l’Ouest. Volume 1. Collection Faune et Flore
Tropicales 40. MRAC & IRD, Tervuren & Paris.
Vreven, E. & J. Snoeks. 2007. Sciaenidae. Pp. 432-440
in: M. L. J. Stiassny, G. G. Teugels & C. D. Hopkins
(eds.), Poissons d’eaux douces et saumâtres de
156
Copyright © Verlag Dr. Friedrich Pfeil
basse Guinée, ouest de l’Afrique centrale, Volume 2.
MRAC & IRD, Tervuren & Paris.
Welcomme, R. L. 1985. River fisheries. FAO Fisheries
Technical Paper, 262. FAO, Rome, 330 pp.
— 1988. International introductions of inland aquatic
species. FAO Fisheries Technical Paper 294. FAO,
Rome, 318 pp.
Wildekamp, R. H. 1996. A world of killies. Atlas of the
oviparous cyprinodontiform fishes of the World.
American Killifish Association, Inc. Volume 3: 1-330.
Wildekamp, R. H. & J. R. Van der Zee. 2003. Cyprinodontiformes. Pp. 298-442 in: D. Paugy, C. Lévêque
& G. G. Teugels (eds.), Faune des poissons d’eaux
douces et saumâtres de l’Afrique de l’Ouest. Volume
2. Collection Faune et Flore Tropicales 40. MRAC
& IRD, Tervuren & Paris.
Yang L., T. Sado, M. V. Hirt, E. Pasco-Viel, M. Arunachalam, J. Li, X. Wang, J. Freyhof, K. Saitoh, A. M.
Simons, M. Miya, S. He & R. L. Mayden. 2015.
Phylogeny and polyploidy: resolving the classification of cyprinine fishes (Teleostei: Cypriniformes).
Molecular Phylogenetics and Evolution, 85: 97-116.
Received 14 March 2017
Revised 16 November 2017
Accepted 6 December 2017
Lederoun et al.: Ichthyofauna of the Mono River basin
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Ichthyological Exploration of Freshwaters
An international journal for field-orientated ichthyology
Volume 28 • Number 2 • January 2018
CONTENTS
Costa e Silva, Thiago, Frank V. R. Ribeiro, Carlos A. S. Lucena and Paulo H. F. Lucinda: Pimelodus speciosus (Teleostei: Pimelodidae), a new catfish species from the rio Tocantins
drainage, Brazil ...........................................................................................................................
97
Lalramliana, Beihrosa Solo, Samuel Lalronunga and Lalnuntluanga: Hemimyzon indicus, a
new species of balitorid fish from the Kaladan basin, Mizoram, northeast India (Teleostei:
Balitoridae)...................................................................................................................................
107
Nagy, Béla: Nothobranchius ditte, a new species of annual killifish from the Lake Mweru
basin in the Democratic Republic of the Congo (Teleostei: Nothobranchiidae) ..............
115
Lederoun, Djiman, Jos Snoeks, Philippe Lalèyè, Pierre Vandewalle and Emmanuel Vreven:
An updated checklist of the ichthyofauna of the Mono River basin (Benin and Togo:
West Africa) .................................................................................................................................
135
Nagy, Béla and Emmanuel Vreven: Micropanchax petnehazyi, a new species of lampeye
cyprinodontiform from the Lufira drainage, Democratic Republic of Congo (Cyprinodontiformes: Poeciliidae) ...........................................................................................................
157
Iqbal, Muhammad, Doni Setiawan and Ajiman: New data on the distribution of the endangered white-edge freshwater whipray Fluvitrygon signifer (Chondrichthyes: Dasyatidae)
171
Moelants, Tuur, Jos Snoeks and Emmanuel Vreven: Distichodus kasaiensis and D. ingae, two
new distichodontid species (Characiformes: Distichodontidae) from the Congo basin .
177
Cover photograph
Nothobranchius ditte Nagy (Photograph by Béla Nagy)
Béla Nagy
(this volume pp. 115-134)
Articles appearing in this journal are indexed in:
AQUATIC SCIENCES and FISHERIES ABSTRACTS
BIOLIS - BIOLOGISCHE LITERATUR INFORMATION SENCKENBERG
CAMBRIDGE SCIENTIFIC ABSTRACTS
CURRENT CONTENTS/AGRICULTURE, BIOLOGY & ENVIRONMENTAL SCIENCES and SCIE
FISHLIT
ZOOLOGICAL RECORD