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39
Iclitliyol.
Explor. Freshwaters, Vol. 1,
No.
1,
pp. 39-48,2
figs.,
1
tab., January
1990
O
1990
by
Verlag
Dr.
Friedrich Pfeil, München,
FRG
-
ISSN
0936-9902
Relict tropical fish fauna
in
Central Sahara
Christian
Lévêque
*
Following a taxonomic revision of freshwater fishes recorded from the Sahara, an updated species list is given. It
appears that most species collected in small water bodies are relict populations of widespread species occuring in
surrounding river basins, except for
Barbus
apleurogramina
which might be a relict of a more ancient fish fauna.
Distribution patterns are discussed in view of the supposed extent of river catchments during the Holocene.
Une révision taxinomique des espèces de poissons récoltées dans le Sahara a permis de dresser une liste actualisée
prenant en compte les connaissances récentes. I1 apparaît que la plupart des espèces collectées dans les petites
collections d'eau sont des populations reliques d'espèces largement répandues dans les bassins hydrographiques
voisins, àl'exception de
Barbus
npleurogranana
qui pourrait être un vestige d'une faune plus ancienne. La distribution
actuelle est discutée,
à
la lumière des données sur l'extension supposée des bassins hydrographiques durant
l'Holocène.
Introduction
Since the begining of this century, numerous
Saharan expeditions were able to collect several
fish species in the widely scattered and isolated
small patches of water in this area (Daget, 1959a,
1968; Dumont, 1979, 1987; Estève. 1949, 1952;
Fowler, 1949; Le Berre, 1989; Monod, 1951,1954;
Pellegrin, 1913,1919a-b, 1931,1934,1936).
If
the
occurence of species known from Sudanese ri-
vers far south was quite surprising for the first
discoverers, it is nowadays obvious that their
occurence results from the dynamics
of
changing
climates over the area (Dumont, 1982; Maley,
1983).
As a result of the improvement
of
our
know-
ledge on African fishes, and following recent
taxonomic revisions, it turned out that confusion
existed in the identification of some species, often
based on small specimens, sometimes poorly
preserved. The aim
of
this paper is to give an
updated taxonomic review
of
the relict fish
populations in the Sahara, and to discuss their
significance. This paper only deals with fish
of
afrotropical
origin,
bearing
in
mind that northern
Africa is nowadays mainly colonized by Palae-
arctic or Mediterranean species. Many of the
specimens collected are housed in the Muséum
national d'Histoire naturelle, Paris
(MNHN),
the
British Museum of natural History, London
(BMNH)
and the Musée royal de l'Afrique cen-
trale, Tervuren (MRAC), and part of those col-
lections were examined.
*
Chercheur ORSTOM, Laboratoire d'Ichtyologie, Muséum national d'Histoire naturelle,
43
rue Cuvier, 75005
Paris, France
ORSTOM
Fonds
Doeumentaire
Ichthyol. Explor. Freshwaters,
Vol.
1,
No.
1
40
\
a
L.
parvus
A
B.
deserti
A
Lévêque:
Relict
fishes
in
Sahara
AE.
spilargyreins
D
*A.
defontainesi
8
H.
bimaculatus
41
Fig.
1.
Distribution of relict tropical fish species in Sahara and surrounding river basins. Data on
Clarias
from Teugels
(1986)
and on
S.
galilaeus
from Trewavas
(1983).
List
of
species collected in Central Sahara
OnFigure
1
allknownSaharanrecordsarerepor-
ted. For surrounding river basins, scattered
sym-
bols give an idea of the distribution range for
each species. Detailed distribution record for
these basins are published by Lévêque et al. (in
press), Paugy
&
Benech (in press) and Teugels et
al. (1988).
Cvprinidae
Barbus
ayleurogramma
Boulenger, 1911
(Fig. 1A)
Reported by Daget (1968) from Aoué, on the
western side of the Ennedi
(MNHN
no 1967-647).
The specimens examined exhibit the following
characters: mouth terminal, at the level of the
middle of the eye; only one barbel on each side,
its tip reaching the anterior margin of the eye; last
simple dorsal ray bony and slightly denticulated
behind (clearly marked only in large specimens);
Ichthyol.
Explor.
Freshwaters,
Vol.
1,
No.
1
42
7 branched dorsal rays; 22 to 23 scales in longi-
tudinal series, 8 around caudal peduncle;
no
lateral line; a black spot at the base of the caudal
and anal fins; scales edged with dark brown. This
description is diagnostic for
B. aplezirogramma
as
described by Boulenger
(1911).
The presence of
B. apleuvogramma
on
the
south-western side of the Ennedi is surprising
(see discussion) as the species was only known
from Lake Victoria and related streams.
Barbus bynni occidentalis
Boulenger,
191
1
(Fig.
1B)
Daget (1959a) and Blache et al. (1964)
reported
B. batesi
from Totous (Tibesti). For
Trewavas (1974) the identification is doubtful
and Lévêque
&
Guégan (in press) reidentified the
three juvenile specimens as
B. bynni occidentalis.
The latter is present in the Chad basin, whereas
B.
batesi
is known from Cameroun.
’
Barbus deserti
Pellegrin, 1909 (Fig. 1A) and
Barbus macrops
Boulenger,
1911
(Fig.
1B)
For a long time, there has been considerable
confusion between both species. Re-examination
of the type specimens of
B. deserti
allowed
Hopson
&
Hopson (1965) and Lévêque (1989) to
conclude that both are valid species. They share
similar scale counts, and show a black spot
on
the
dorsal
fin.
But the black spot covers the tip of the
fin in
B. macrops,
whereas
B. deserti
bears a spot
on
the distal half of the first branched rays,
distantly set from the distal margin.
Barbus deserti
was described
on
the basis of four specimens col-
lected in Tassili n’Ajjer at Redir de Sfedil (Pelle-
grin, 1909) which is apparently a pool of the river
Edefil (or Idefil, or Efedil), a seasonal tributary of
the river Imirhou which carries the periodical
rainfall of the northern part of the Tassili. Sko-
repa (1977) examined 36 specimens of
B. deserti
collected in two “gueltas” of the Imirhou river,
and a series in MNHN (no 1959-120) originates
from Oued Djerat, few kilometers north to the
Imirhou river. Specimens of
B. deserti
were also
collected at EI Barcat (Barkat) near Ghat, and are
housed in
MNHN
(no
1935-59) and BMNH
(no
1937.7.8: 4-6) (see Hopson
&
Hopson, 1965;
Lévêque, 1989).
Barbus deserti
is apparently re-
stricted to northern slopes of Tassili n’Ajjer. As
a result, most of the other series previously
identified as
B. deserti
in literature, are in fact
B.
inncrops
which is a widely distributed species
known from most West African river basins.
Barbus anema
Boulenger, 1903 was reported
from Totous (Tibesti) by Pellegrin (1919a). Ac-
cording to Daget (1959a) this identification is
doubtful. The specimens are badly preserved
and Daget (op. cit.) observed posterior barbels in
one specimen. I was able to observe anterior and
posterior barbels in two of the Totous specimens
(MNHN
no
1919-65). The tip of the dorsal fin
is
blackish and there are some remains of a black
longitudinal band. I consider these specimens as
juveniles of
B. macrops
which has already been
collected at Totous.
Barbus pobegtrini
Pellegrin, 1911 (Fig. 1A)
Numerous specimens were collected in diffe-
rent sites of the Adrar area (Mauritania). The
species was identified
Barbus (Capoeta) pobeguini
~zauvitanica
Pellegrin, 1937, by Estève (1952), and
Barbzis (Hemigrammocnpoeta) mirei
Estève, 1952, a
junior synonym of
B. pobeguini
(see
Monod,
1954). This species is known from the Niger,
Senegal and Volta basins but has never been
collected in the Chad and Nile basins.
Labeo niloticzis
(Forsskål, 1775)
/
Labeo senega-
lensis
Valenciennes, 1842 (Fig.
1B)
The specimens from Totous (Tibesti) identi-
fied
as
Labeo korie
by Pellegrin (1919a) were
juveniles. Daget (1959a) mentioned that they
were in poor condition and reidentified the se-
ries, as well as another sample collected
in
the
same water body, as
L. niloticus.
The three speci-
mens from Totous examined by Daget have 15
branched dorsal rays, 42 scales in longitudinal
series, 8.5 scales between lateral line and dorsal
fin,
5 to 5.5 between lateral line and base of
pectoral fin, 20 scales around caudal peduncle.
The number of scales, as well as the number of
dorsal rays, are characteristics of
L.
nilotbis,
compared to
L. korie
and
L. senegalensis,
accor-
ding to Reid (1985).
Daget (1968) also reported specimens identified
as
Labeo
sp. from Ennedi, and mentioned that
they were close to
L.
korie
/
L. nilotictis
from
Tibesti. The specimens
(MNHN
no 1967-658)
have meristic counts slightly different from the
Tibesti ones: 13-15 (mode 14) branched dorsal fin
rays; 38-40 scales inlongitudinal series; 6.5 scales
between lateral line and dorsal fin origin; 4.5
scales between lateral line and base of pectoral
Lévêque:
Relict
fishes
in
Sahara
43
fin;
16 scales around caudal peduncle. The Enne-
di specimens are therefore very close to
L.
senega-
lensis
occuring in Chad basin and distinguished
from the Nile species by lower scale counts (see
Reid, 1985). In fact, it may well be that
L. senega-'
lensis
present in the Chad and western basins,
and
L. lzorie
and/or
L. niloticus
are local popu-
lations of the same species. Nevertheless, the En-
nedi specimens appear to be related to the Chad
fauna whereas the Tibesti
Labeo
are closer to the
Nile ones.
Labeo parvus
Boulenger, 1902 (Fig. 1A)
For Reid (1985)
L. tibestii
Pellegrin, 1919 is a
junior synonym of
Labeo oguizensis.
According to
Jégu
&
Lévêque (1984)
L. ogziizensis
as well as
L.
chariensis
are synonyms of
L.
parvus,
a wide-
spread species known from the West African
river basins, including Chad, but absent from the
Nile. Daget (1959a) already considered
L. tibestii
to be close to
L. annecteizs
which could be a
synonym of
L.
parvus
according to Reid (1985).
Raiamas senegalensis
(Steindachner, 1870)
(Fig. 1A)
A single specimen was collected in the Yezei
(Tibesti) and identified
Barbus
loati
by Pellegrin
(1919a). Daget (1959a) reidentified the specimens
as
B.
senegalensis.
For Lévêque
&
Bigorne (19831,
R.
loati
is a junior synonym of
R.
sei?egalensis.
Clariidae
Clarias aizguillaris
(Linnaeus, 1758) (Fig. 1C)
C.
senegalensis
Valenciennes, 1840 reported
from Adrar (Mauritania) and central Sahara, is a
junior synonym of
C. arzguillaris
(see Teugels,
1986).
Clarias gariepinus
(Burchell, 1822) (Fig. 1C)
Clarias lazera
Valenciennes, 1840 reported
from Ennedi and Tibestiis a junior synonym of
C.
gariepinus
(see Teugels, 1986).
Cvprinodontidae
Epiplatys spilargyreius
(Duméril, 1861)
(Fig.
1D)
Specimens collected at Tigui (Borkou) were
identified as
Aplocheilus marni
(incorrect spelling
for
Haplochilus marnoi
Steindachner, 1881) by
Estève (1952) and reidentified as
Epiplatys senega-
lensis
(Steindachner, 1870) by. Daget (1959a).
Scheel (1968) introduced the synonymy with
E.
spilargyreius.
Cichlidae
Astatotilapia desfontainesi
(Lacépède, 1803)
(Fig. 1D)
The species previously identified
Haplockro-
inis desfontainesi
is restricted to Tunisia and Alge-
ria (Regan, 1922), despite the wide distribution
range of the genus in Africa (Greenwood, 1979).
The mention of
A.
desfontainesi
from Chad basin
by Dumont (1982: Tab.
1)
is erroneous. It is
A.
bloyeti
Sauvage, 1883 which is known from the
Chad area and Upper Niger (Greenwood, 1979).
Hemiclzvomis bintaciilatus
Gill, 1862 (Fig. 1D)
Atlas.
Reported from different localities up to the
Sarotherodon galilaeus
(Linnaeus, 1758)
(Fig. 1E)
According to Trewavas (1983),
Tilapia borkua-
na
Pellegrin,
1919
which was reported from
many localities in the Borkou-Ennedi-Tibesti is a
subspecies of
Sarotherodon galilaeus.
The most
striking character is the pigmentation. Another
subspecies,
Sarotherodon galilaeus galilaeus
is
known from the Adrar (Mauritania) (Monod,
1951; Estève, 1952).
Tilapia
zillii
(Gervais, 1848) (Fig. 1E)
The species is reported from numerous lo-
calities up to the Atlas. It is widespread and ap-
parently resistant to salinity.
Discussion
At the present time, as throughout the Pleisto-
cene, the Atlas watershed is a geographical bar-
rier between Sudanian fish in the desert and
those of Eurasian origin in the Maghreb. But
Greenwood (1973) found in fish remains from
late Miocene deposits of Tunisia (Bled el Doua-
rah), a diversified freshwater ichthyofauna
showing strong affinities with the contemporary
fauna of Egypt and tropical Africa. Among iden-
tified fossils, representatives of the genera
Polypterus, Lates, Heterobranchus
and
Clarias,
?
Clarotes
and
?
Synodoizfis
are present. Thus the
Ichthyol. Explor. Freshwaters,
Vol.
1,
No.
1
44
Table
1.
Occurence of afrotropical fish species in different areas of the Sahara. Modified from Dumont
(1982).
1,
Adrar (Mauritania);
2,
Tunisia and south Algeria;
3,
Air;
4,
Ahaggar;
5,
Tassili N'Ajjer;
6,
Ghat;
7,
Tibesti;
8,
Ennedi;
9,
Borkou.
1
2
3 4
5
6 7 8 9
B. apleurogramma
B. occidentalis
B. deserti
B. macrops
*
B. pobeguini
t
L.
niloticus
R.
semgalensis
C.
anguillaris
*
C.
gariepinirs
E.
spilarpjreius
H.
bimaciilatzrs
*
S.
g.
galilaeiis
*
S.
g.
borkiianus
T.
Alii
.*
L.
parvus
t
.%
.%
.%
*
*
*
*
t
*
*
t
*
*
*
*
t
* *
*
*
*
fish fauna
of
the late Miocene in Tunisia, con-
trasts strongly with that of the present, which is
extremely depauperate and show affinities with
species in Europe and Asia Minor
(Barbus capito,
Aphaniirs
and
Phoxinella).
The cichlid
Astato-
tilapia desfontainesi
is the only species of
un-
questionable African affinities. The Miocene
fauna from Tunisia
is
fairly similar to contempo-
rary records from Egypt (Priem, 1914, 1920),
Syrte Major, Cyrenaica (Arambourg, 1963) and
East Africa (Greenwood, 1951; Greenwood
&
Howes, 1975). Van Cowering (1982) also repor-
ted that
Palaeochromis,
a late Miocene cichlid
discovered in the Seybouse Valley near Guelma
(Algeria), is closely related with Central and
West African
Pelmatochromis.
According to
Greenwood (1973), "the faunal similarity bet-
ween different Miocene deposits, gives the im-
pression that during the Miocene, there was a
fairly uniform freshwater fauna (at least at the
generic level) widely distributed in Africa north
of the equator. Due to the relative paucity of
materials, it is difficult to determine just how
widespread this fauna was or when it made its
first appearance. But Eocene deposits in Libya
(Arambourg
&
Magnier, 1961) have a very simi-
lar fish fauna
(Polypterus, Lates
and unidentified
Siluriformes)".
Protopterus
sp. were also indenti-
fied in Lower Eocene fossils from EI Kohol, near
Brezina, on the southern flank of the Saharan
Atlas, Algeria (Mahboudi et al., 1984), The fossil
records from Lake Edward (Greenwood, 1959;
Greenwood
&
Howes, 1975) revealed that during
Pliocene to late Pleistocene, the fish fauna was
more similar to the Nile one than
it
is today and
included several of the genera now absent. For
Lake Victoria, only few data indicate that some
Nile type elements (now absent) were present
before the modern lake was formed (Green-
wood, 1951,1974).
As
a result of taxonomic revisions, it appears that
the relict fauna of the Borkou-Ennedi-Tibesti area
which exhibits the highest diversity (Table
l),
is
clearly related to that fauna existing in the Chari
and Nile basins. All species reported from this
area, except for
B.
apleirrogramma,
are found in
those basins. There
is
good evidence that during
the last Holocene humid period, the Chad basin
extended to the foot of the Ennedi-Tibesti whose
drainages had flowed into the so-called mega-
Chad (see Talbot, 1980; Servant, 1973). The iso-
Lévéque:
Relict fishes
in
Sahara
45
Fig.
2.
Estimated limits
of
potential catchments
of
the Senegal, Niger and Chad systems during the earlier Holocene
when the whole catchment areas were probably active. Modified from Talbot (1980) and using informations from
Riser
&
Petit-Maire (1986).
lation of these populations happened 5-6,000 yrs
ago and this apparently explains the relative rich-
ness of the ichthyofauna. It is also clear that in
spite of the present extreme isolation of the area,
this time period is apparently insufficient to al-
low speciation and appearance of endemics,
even if some morphological variability can be ob-
served.
The record of
B.
apleurogramma
in the Ennedi,
is nevertheless more surprising as this species is
nowadays only known from
L.
Victoria and the
connected river systems. Until now the impor-
tance of this record and its significance have
rather been neglected. Knowing that the species
no longer occurs in the Chad and the lower Nile,
it could well be the relict of a more ancient fish
fauna extending northwards, and whose repre-
sentatives later disappeared from Sudanese river
basins. There are many other evidences of faunal
exchanges during the geological times.
It should also be noted that
B.
apleurogramnza
belongs to a group of
Barbus
showing a serrated
last simple dorsal fin ray. Without giving to
much emphasis to this morphological feature as
its phylogenetic significance is not yet known, I
can nevertheless ascertain that nowadays this
group is represented by only few species, with a
limited distribution, in Africa north of the equa-
tor:
B.
cadenati,
B.
dialonensis
and
B.
guineensis,
restricted to the upper courses of the rivers
Konkoure, Gambia and Senegal in the Fouta
Djalon (Guinea) (Daget, 1962);
B. tniolepis
known
from the upper reaches of the Chari in Central
African Republic but widespread in Zaire and
coastal rivers in Cameroon;
B.
kerstenii, B. netr-
mayeri
and
B.
pellegrini
occuring in the Nile drai-
nage only in the Lake Albert-Edward region, but
widespread in East Africa (Banister, 1987). It
could then well be that
B.
aplezirogranznza
from
Ennedi, was part of an ancient fauna which was
more widely distributed than nowadays, and
whose representatives disappeared probably
during an arid climatic phase, relict populations
subsisting in particular zones. That could imply
that the isolation of
B.
apleurogranztna
would be
older than the last humid period.
The phylogenetic affinities of
B.
deserti
known from the northern slopes of the Tassili
n’Ajjer are not clear, but morphologically it
appears t9 be related to the small tropical
Barbus
with which it was confused. There are super-
ficially close similarities with Sudanese species
and comparing morphometric characteristics
and colour pattern,
B.
deserti
resembles
B.
callip-
terus,
a presently widespread species in Niger
and Chad basins. Some other tropical species
were also collected in the Tassili:
Clarias
gariepiizus, Tilapia zillii
and
Hemichrotnis bimacu-
Ichthyol.
Explor.
Freshwaters, Vol.
1,
No.
1
46
lattis.
It
is
possible that the above species are relict
fauna of the Chad basin fauna which were able to
survive severe droughts and to colonize the nor-
thern slopes of the Tassili. To support this idea, it
should be recalled that the Tassili represents the
northern limit of the maximum potential catch-
ment of Chad (see Talbot, 1980 and Fig.
2).
Du-
ring periods of maximum humidity, including
the earlier Holocene, the whole catchment was
probably active and relict drainages from Tassili
to Chad are still apparent.
Barbus
biscnrensis,
a
northern colonizer, is also present
on
the nor-
thern slopes of Tassili.
The relict fish fauna of the Ahaggar is poor
compared to the Tibesti-Ennedi one and there is
no
record from
Aïr
or Adrar Ifora. That is surpri-
sing
if
we consider the dense relict drainage of the
central Sahara including prominent watercour-
ses such as the Tilemsi, Dallol Bosso, Dallol
Maouri and Tarka which were active during late
Pleistocene and upper Holocene (Talbot, 1980).
During the early Holocene, there were large tro-
pical lakes (Riser
&
Petit-Maire, 1986) and a
diversified fossil fish fauna was collected all over
this area (Daget, 195913,1961; Gayet, 1983).
Lates
inaliensis
Gayet, 1983 was described from the
Taoudeni depression of Malian Sahara, but is
now considered as a synonym of
L. niloticus
(Van
Neer, 1987). The paucity of the present fish fauna
could be explained by the scarcity
of
suitable
perennial habitats of the area, but this hypothesis
can hardly be tested.
Bailey-Watts
&
Rogers (1970) provided some
information on the fish fauna of the Jebel Marra.
The samplings were restricted to a stream (Wadi
Golol) tributary of the Wadi Azum, and an ad-
jacent pool. Such habitats are connected to the
main river during the flood and the fish fauna
collected is therefore representative of the Chari
basin. Green et al. (1984) also provided fragmen-
tary information
on
fish fauna inhabiting Lakes
Keilak and Kundi, respectively in south Kordo-
fan Province and southern part of Darfur in
Sudan.
Clarias lazera
(=
C.
gariepintis), Tilapia
zillii
and
Schilbe
mystus
were caught in those
isolated water bodies from the Upper White Nile
basin, which nevertheless probably contribute
water to the Nile system in exceptional wet sea-
sons.
The Saharan relict tropical fish fauna sur-
vived for thousands of years in isolated small
water bodies. But
one
could expect that some of
the records will become of historical interest, as
long as man impact (mainly water pollution) will
progressively modify the present distribution, in
eradicating fish species in some
of
the water
bodies.
Acknowledgements
This work was part of PEDALO programme
(Poissons d'eau douce <Afrique de l'Ouest)
sponsored by ORSTOM and PIREN (CNRS). The
author is grateful to D. Paugy and G. Teugels for
their valuable comments.
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I
-'
Arambourg,
C.
&
P.
Magnier. 1961. Gisements de verté-
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C.
R.
Hebd. Séanc. Acad. Sci. Paris, 252: 1181-1183.
Bailey-Watts, A.
E.
&
J.
F. Rogers. 1970. The fish collec-
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12:
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Banister, K.
E.
1987. The
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Received
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Revised
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Accepted
17
November
1989
Lévêque: Relict fishes
in
Sahara
