By Jean Henri HUBER (1)
Following the identification of a new collection of Cyprinodontids (Pisces, Atheriniformes) from Ivory Coast by MM. Lévêque, Paugy et al., a systematic review is proposed: new keys for all species are presented, the distributions are studied according to vegetation and even extended for Epiplatys spilargyreius, Ep. chaperi s.l. and Micropanchax pfaffi; two new synonymies are forwarded: Ep. kassiapleuensis with Ep. olbrechtsi and Mic. monikae with Mic. schioetzi.
The Cyprinodontid fauna of the Ivory Coast is quite well known as a result of the collecting trips of Daget and Iltis (1965), Etzel (1974) and Roman and Schmitt (1978). There remained the need to study material from the untouched areas of the centre, northeast and north of the country. This has now been made possible by the collecting trips of Messrs. Lévêque and Paugy as well as other members of the Bouaké ORSTOM team. So we now have a reasonably satisfactory understanding of the Cyprinodontid fauna of the Ivory Coast. The fish collecting trips just mentioned were particularly fruitful thanks to the technique of electric fishing and the use of poison. And they were also of interest as they have shown that certain species have extended distribution in the south and west of the country.
Studying preserved Cyprinodontid material is particularly difficult owing to the small size of the fish, especially the Procatopodins, and this is not helped by their great variability. Therefore, in the new keys I have proposed, I have given greater importance to geographical location, general appearance of the body, maximum size, colouration type, even biology and behaviour rather than to meristic data and proportions which usually overlap. For each species there is an illustration by Mr. Wildekamp. The illustrations aim to show body shape and colouration patterns but are not intended to be used in any analysis.
The Ivory Coast is a small state in West Africa, covering an area a little over 300,000 sq. kms.
From a biogeographical point of view, several fundamental factors influence the presence, density and diversity of the species. The combination of these factors will enable us to define six distinct population zones. The climate factor is the most significant:
1. The 550km long coast has a particularly heavy rainfall - more than two metres a year - which gives rise to an area of dense forest and a large concentration of small rivers.
2. The interior as far as 8° North, that is up to the latitude of the town of Bouaké, gets less rain and its vegetation becomes reduced to wooded savannah.
3. North of Bouaké, the savannah becomes progressively drier: this is the "grassy savannah".
The hydrographical factor is important. The rivers - Cavally, Sassandra, Bandama & Comoé - drain the interior of the country following a fairly regular line north/south; but they are not of themselves sufficient to form real barriers. However, at each end two separate areas can be identified:
1. North of the parallel of Ferkessédougou, the flow of the rivers is slight especially in the dry season, which affects the fauna.
2. At the mouth of the rivers there are often brackish lagoons.
The altitude factor is important: the change of altitude from the coast inland is gradual, so that one cannot hope to find a clearly separated coastal fauna and plateau fauna as happens in other African countries. One area, however, is characterised by its relief: the triangle formed around Man, by the mountains of the Dans, Toura and Nimba that reach higher than 1.200m.
Lastly the surrounding countries must also be borne in mind. The absence of a natural frontier in the north and east would suggest continuity in distribution areas, whereas in the west - with the exception of the coastal strip - the mountains of Liberia and then the Guinea range form a major obstacle.
Fig.1. Map of Collecting Places.
Then, we can identify six distribution zones but their frontiers are not impassable for the Cyprinodont fauna.
I. The area of brackish water lagoons and mangroves - which continues into Liberia and Ghana.
II. The coastal forest strip as far as the vicinity of the latitude of Agboville - which continues into Ghana.
III. The bushy savannah as far as Bouaké - which also continues into Ghana.
IV. The grassy savannah as far as Ferkessédougou - a not very clear boundary.
V. The relatively dry savannah of the extreme North.
VI. Lastly a special area: the western forested plateaux around Man, which should be linked with the Guinea range.
COLLECTING PLACES AND IDENTIFICATION OF FISH CAUGHT.
You will find below the list of collecting places grouped in geographical areas. For each location the following information is given: the species collected, the date, the numbers of specimens and their sizes, and the number under which they are entered in the collections of the Museum National d'Histoire Naturelle de Paris (MNHN 1981-645 to 703).
1. Tributary of the Bandama at Bankanda
Micropanchax (Poropanchax) rancureli - 25 Feb. 1977 - 343 specimens 15- 30 mm - 1981-645.
2. Tributary of the Go, near the Tagba lagoon.
Micropanchax (Poropanchax) rancureli - 19 Feb. 1977 - 23 specimens 20mm- 1981-646.
Epiplatys dageti dageti - 19 Feb.1977 - 10 specimens 20-30mm - 1981-649.
3. River Boubo at Ekradon.
Micropanchax (Poropanchax) rancureli - 22 Feb 1977 - 24 specimens 15-25mm- 1981-650.
4. River Agnébi at Akoupé.
Micropanchax (Poropanchax) rancureli - 30 Jan 78 - 80 specimens 15-25mm 1981-648.
Epiplatys chaperi sheljuzhkoi - 30 Jan 78 - 12 specimens 25-40mm - 1981 652.
Micropanchax schioetzi - 30 Jan 78 - 23 specimens 20-30mm - 1981-654.
5. River M'pédo - tributary of the Agnébi.
Epiplatys chaperi sheljuzhkoi - 1 Feb 78 - 2 specimens 35-50mm-1981-653.
Micropanchax schioetzi - 1 Feb 1978 - 107 specimens 25-40mm - 1981-655.
6. River Agnébi at Dabou.
Epiplatys daqeti dageti - 2 Feb 75 - 2 specimens 20-35mm - 1981-651.
Epiplatys bifasciatus - 2 Feb 75 - 50 specimens 20-45mm - 1981-662.
7. Kan - 15km southeast of Bouaké - and Konkodekro.
Micropanchax schioetzi - 1 Sep.75 - 9 specimens 20-35mm - 1981-656.
Micropanchax schioetzi - June 75 - 2 specimens 25-40mm - 1981-658.
Micropanchax schioetzi - 2 Jul 76 - 55 specimens 30-40mm - 1981-659.
Epiplatys chaperi spillmanni - 23 Feb 76 - 19 specimens 40-60mm-1981-668.
Epiplatys chaperi spillmanni - 1 Sep.75 - 3 specimens 40-50mm - 1981-669.
8. River Béré - tributary of the River Maraoué.
Epiplatys bifasciatus - 26 Apr 76 - 3 specimens 30-35mm - 1981-663.
Micropanchax normani - 10 Aug 77 - 10 specimens 20-25mm - 1981-672.
Micropanchax normani - 20 Oct 77 - 5 specimens 15-25mm - 1981-673.
9. Kan near Bouaké.
Micropanchax schioetzi - 23.Feb 76 - 22 specimens 20-35mm - 1981-657.
10. River Maraoué at Niondjé.
Epiplatys bifasciatus - 2 Aug 77 - 2 specimens 30-35mm - 1981-664.
Micropanchax pfaffi - 16 Mar 77 - 15 specimens 15-20mm - 1981-676.
Micropanchax pfaffi - 2 Aug 77 - 29 specimens 20-25mm - 1981-677.
Micropanchax pfaffi - 9 Jan 75 - 2 specimens 20-25mm - 1981-678.
11. River Mbé - tributary of the Nzi.
Epiplatys chaperi spillmanni - 25 May 76 - 6 specimens - 35-50mm -1981-670.
12. River Comoé at Gansé - entrance to the National Park.
Epiplatys bifasciatus - 16.Jan 75 - 8 specimens 20-25mm - 1981-665.
Micropanchax normani - 16 Jan 75 - 5 specimens 20mm - 1981-674.
Epiplatys spilarqyreius - 16 Jan 75 - 2 specimens 25-35mm - 1981-693.
13. River Kovin - tributary of the Comoé.
Micropanchax pfaffi - 20 Feb 75 - 3 specimens 20-25mm - 1981-679.
14. River Léraba - tributary of the Comoé.
Micropanchax pfaffi - 20 Feb 74 - 2 specimens 15-20mm - 1981-680.
15. Tributaries of the Bandama near Koumballa and Ferkessédougou.
Micropanchax pfaffi - 7 Mar 75 - 19 specimens 15-20mm - 1981-681.
Epiplatys spilargyreius - 7 Mar 75 - 1 specimen 35mm - 1981-694.
16. River Solomougou - tributary of the Bandama near Guiembe.
Micropanchax pfaffi - 31 Mar 77 - 77 specimens 20-30mm - 1981-682.
17. River Badénou at M'bingué.
Epiplatys bifasciatus - 15 Jun 77 - 3 specimens 30-35mm - 1981-666.
Epiplatys bifasciatus - 30 Mar 77 - 1 specimen 35mm - 1981-667.
Micropanchax pfaffi - 15 Jun 77 - 68 specimens 15-25mm - 1981-683.
18. River Bagou, 10km east of Boundiala - Niger basin.
Micropanchax normani - 4 Apr 77 - 4 specimens 20mm - 1981-675.
Micropanchax pfaffi - 12 Jul 77 - 20 specimens 20mm - 1981-684.
Micropanchax pfaffi - 17 May 77 - 71 specimens 20-25mm - 1981-685.
Micropanchax pfaffi - 4 Apr 77 - 33 specimens 15-20mm - 1981-686.
Epiplatys spilargyreius - 12 Jul 77 - 2 specimens 35mm - 1981-695.
Epiplatys spilargyreius - 17 May 77 - 2 specimens 30-35nm - 1981-696.
19. River Bou - tributary of the Bandama - at Sirasso.
Micropanchax pfaffi - 15 Jun 77 - 14 specimens 20-30mm - 1981-687.
Epiplatys bifasciatus - 15 Jun 77 - 2 specimens 30mm - 1981-702.
20. River Yore-Lohro, 10km East of Tarato and 30km West of Korhogo.
Micropanchax pfaffi - 16 Jun 77 - 27 specimens 15-25mm - 1981-688.
21. Niaka on the River Bandama between Dikodougou and Niakaramandougou.
Epiplatys chaperi spillmanni - 5 Dec 74 - 1 specimen 25mm - 1981-671.
Micropanchax pfaffi - 13 Jun 77 - 2 specimens 15-20mm - 1981-689.
22. River Yani, at Madji, 19km south-west of Banandjé.
Micropanchax pfaffi - 3 Aug 77 - 31 specimens 20-30mm - 1981-690.
Micropanchax pfaffi - 15 Mar 77 - 51 specimens 15-30mm - 1981-691.
23. River Oyo, near Lokolo.
Epiplatys olbrechtsi - 17 Mar 77 - 2 specimens 20-40mm - 1981-698.
24. Tributary of the River Cavally near Danané.
Micropanchax schioetzi - 10 Apr 77 - 14 specimens 15-30mm - 1981-660.
Epiplatys olbrechtsi - 10 Apr 77 - 3 specimens 40-50mm - 1981-699.
25. River Cavally at Vatoua.
Epiplatys olbrechtsi - 10 Apr 77 - 3 specimens 35-40mm - 1981-700.
26. Tributary of the River Nipoué near Zouan (Houmien).
Micropanchax schioetzi - 10 Apr 77 - 104 specimens 15-40mm - 1981-661.
Epiplatys olbrechtsi - 10 Apr 77 - 8 specimens 40-55mm - 1981-701.
27. River Sassandra near the Sémien Ferry.
Micropanchax pfaffi - 21 Feb 75 - 19 specimens - 1981-692.
28. River Daka, Tributary from Lake Volta towards Yendi (Ghana).
Epiplatys spilargyreius - 4 Mar 78 - 5 specimens 25-35nm - 1981-697.
Epiplatys bifasciatus - 4 Mar 78 - 12 specimens 25-40mm - 1981-703.
Fig. 2a Distribution map of the Aphyosemion species as in the areas marked on Fig. 1. M = A. maeseni, P = A. petersi, W = A. walkeri.
Review of the Cyprinodontid Fauna.
The collecting trips under discussion would indicate the value of a review of the systematics of the Ivory Coast Cyprinodontids after a re-examination of the types of the species concerned.
The Ivory Coast is amply supplied with Cyprinodontids, but as in other countries west of Benin, the biotopes - and consequently the number of species - are less diversified than in the east, where the forest covering and the mountains play an important role.
Rivulins are represented by two genera in the Ivory Coast: Aphyosemion and Epiplatys. The first differs from the latter in a more cylindrical body, a less broad head, a very marked sexual dichromism; it also differs in its behaviour and its preference for shaded biotopes.
Aphyosemion is restricted to the forest biotopes: area II on the coastal plain and area VI around Man.
Epiplatys is present in these areas but it is also found in the open savannah biotopes: areas III, IV and V, that is all the interior of the country. It tends to become less common further north.
The Genus Aphyosemion.
Only three species live in the forest of the Ivory Coast; they represent two distinct superspecies - none of them were caught by Lévêque and colleagues.
1. Long morphology, fairly well marked sexual dichromism, D = 10-12, A = 14-15, D/A = +6 to +7; not annual.
1-1 - restricted to the coastal strip; yellow phase, rounded caudal edged in black: petersi
1-2 - from the western forest plateau around Man, blue phase, fasciation
only on the body of the female: maeseni
2. More massive morphology, very marked sexual dichromism, Dm = 14, Am = 16
D/A = +2, semi-annual, very variable colouration with red fasciation or red spots on the body and unpaired fins: walkeri
1. Aphyosemion petersi (Sauvage,1882)
Haplochilus petersi Sauvage,1882 - Bull. Soc. Zool. France: 324.
Types come from Kouakoukro in south-east Ivory Coast, as do those of Epiplatys chaperi.
Roloffia petersi Clausen,1966.
Relict species, individual in behaviour. This fish should be linked with the species from the plateaux in a special group comprising: A. maeseni Poll,1941; A. guineense Daget,1954; A. guignardi (Romand,1981) from Guinea, A. viride (Ladiges & Roloff, 1973) from Liberia and the fish from Bobo Dioulasso in Upper Volta, now named A. banforense (Seegers, 1982).
A. petersi lives in the coastal springs and streams containing exclusively soft water, in the centre and the east of the south of the Ivory Coast and in the west of southern Ghana; it prefers clear, very shallow water - less than 5cm. Meristics: Dm = 10, Am = 15, D/A = +6. Karyotype: n = 20, A = 30 (Scheel, 1968a). Characteristic colouration: temporary black fasciation in both sexes.
2. Aphyosemion maeseni Poll, 1941.
Aphyosemion maeseni Poll, 1941 - Rev. Zool. Bot. afr.,34: 141-142.
Roloffia maeseni : Clausen, 1966.
Aphyosemion liberiense (not Boul.); Daget & Iltis 1965 (MNHN 32-304, 63-420 and 421: 59-127). Described from the River Masei, a Tributary of the River Nuon (or Noun) in the western forest of the Ivory Coast, this species has been found on several occasions in this area, that is to say in the triangle formed by the towns of Touba, Man and Danané, and then, in the west, between Salayio - northeastern Liberia - and N'zérékoré - southeastern Guinea - where it is sympatric with viride (Scheel, 1968a; Ladiges & Roloff, 1973). The two forms are usually sympatric and the most easterly point for A. viride is only about 60 km west of the type location of A. maeseni . Consequently it is reasonable to consider, in the absence of geographical barriers, A. viride as a possible element of the Ivorian fauna. A. maeseni differs from A. viride in minor meristic differences - Dm = 11 and Am = 15, the dorsal joining the body further forward. Its body is slightly more slender and, there is considerable difference in the colouration: marbled dorsal, bluish white marginal bands; black fasciation on the unpaired fins of females; black spot behind the operculum in the female and young male. Karyotype: n = 21 for both species (Scheel,1974).
3. Aphyosemion walkeri (Boulenger,1911.)
Fundulus walkeri Boulenger,1911 - Ann. Mag. Nat. Hist.,B(8): 262.
Fundulus spurrelli (Boulenger,1913).
Aphyosemion litoriseboris Radda,1976.
For a long time the status of this species has been the subject of contro versy. A. walkeri was described from a female in bad condition from the Bokitsa mine in southern Ghana; A. spurrelli was described from Dunkwa, a few tens of kilometres south-west of the type location of A. walkeri .
An important point is that Boulenger published a drawing with vertical fasciation. Up to 1976 two colouration types were reported:
the striated or spotted type from the Ivory Coast around Abidjan - imported by Sheljuzhko, Broche
the fasciated type from Ghana - imported by Schreiber,
and two different interpretations have been put forward. Radda made walkeri and spurrelli synonymous and suggested the new name litoriseboris for the spotted populations from the Ivory Coast.
Berkenkamp & Etzel proposed a single species with two sub-species: A. walkeri in the west (spotted- striated) and A. spurrelli in the east (fasciated).
I have shown (1981b) that neither of these interpretations was correct: existence of two species not established in the first case; geographical inversion of the type locations and a spotted eastern population at Kumasi in the second.
Apart from a comparison of the types and the absence of a geographical barrier, two western populations have been crossed to produce completely fertile progeny, giving fasciated individuals in the F3 generation. In addition, there is the general variability of annual Cyprinodontids, and all these factors suggest that all these fish are polymorphs of one species, and: so one taxon is retained here.
A. walkeri is characterised by its southern forest habitat, its annual character and its deep morphology. Meristics: see key; Karyotype: n = 18, A = 24 (Scheel, 1968). It is the only annual living in the Ivory Coast. In the west it is replaced by the Callopanchax group ( A. occidentale, etc.), and in the east by the Pronothobranchius. These two groups do not seem to be related to walkeri, which is probably a relict form.
The Genus Epiplatys.
The representatives of the genus Epiplatys predominate numerically in the open habitats (stagnant or sluggish) of the south-east and the centre of the Ivory Coast.
1. Rather massive morphology, large size, temporary bars, numerous spots over a wide body area, submarginal black band in the anal (Superspecies fasciolatus ) D = 10-14, A 1 = 15-18: E. olbrechtsi
2. Very slender morphology D = 6-10. A 1= 14-18.
2-1 Oblique fasciation on the sides, particularly pronounced in the rear part of the body: E. spilargyreius
2-2 Two longitudinal bands on the sides, morphology less deep:1 E. bifasciatus
3. Intermediate morphology 0 1= 9-11, A = 14-16.
3-1 Small fish - less than 5 cm.- black vertical lines, one of which is on the caudal peduncle: E.1 dageti1
3-2 Medium sized fish - up to 7 cm.- slightly oblique black fasciation, absent on the caudal peduncle, temporary or variable according to the populations (sub-species): Superspecies E. chaperi
3-2-1 Well marked fasciation.
3-2-1a Around Kouakoukro (Southeast Ivory Coast): E. chaperi chaperi
3-2-1b Around Akroaba (Southeast Ivory Coast) filaments on male's ventral fins: E.1 etzeli
3-2-2 Temporary fasciation, heavily spotted and with variably coloured reflections.
3-2-2a Around Abidjan (South Ivory Coast): E. chaperi sheljuzhkoi
3-2-2b Around Bouaké, in savannah in the middle of the Ivory Coast: E.1 chaperi spillmanni
4. Epiplatys olbrechtsi Poll,1941.
Epiplatys olbrechtsi Poll, 1941 - Rev. Zool. Bot. Afr.,34: 139.
Epiplatys fasciolatus olbrechtsi: Daget & Iltis,1965.
Epiplatys kassiapleuensis Berkenkamp & Etzel,1977.
Described from the same place as maeseni , the more recent collecting places extend the distribution further west. Unfortunately, until recently, fish caught by Roloff at Salayio - northern Liberia - in 1977 were attached to olbrechtsi. Berkenkamp & Etzel (1977) considered Roloff's fish to be E. olbrechtsi and they re-described the fish from Man-Kassiapleu in the western part of the Ivory Coast under the new name E. kassiapleuensis . In the opinion of Romand (1978), Roloff's fish represent a new species E. roloffi distinct from olbrechtsi. I have studied the types of the two species and also the material collected by Lévêque (coll. loc. No.23, 24, 25 & 26) and also Daget & Iltis's material (MNHN 1963-423). I am convinced that E. roloffi is quite distinct from E. olbrechtsi , but that E. olbrechtsi and E. kassiapleuensis cannot be separated by either morphology or colouration.
E. roloffi, like E. lamottei Daget, 1954, differs appreciably from E. fasciolatus Günther, 1866 and the closely related fish E. f.1 tototaensis Romand,1978 and E. olbrechtsi Poll, 1941 (and other taxa of doubtful status) in the following respects: the vertical bars, the single form of the unpaired fins, and some details in the frontal scalation pointed out by Romand (1978). Besides, E. olbrechtsi and E. kassiapleuensis inhabit the same biogeographical region, the same river basin, the same inland plateau forest, and the two type locations are less than 100km apart and without a barrier of any sort. The colouration types are comparable and the differences are surely only due to polymorphism between populations, and so I have no hesitation in saying that 1 E. kassiapleuensis and E. olbrechtsi are synonyms.
Fig 2b Distribution map of the Epiplatys species, as in the areas marked in Fig.1. B = E. bifasciatus , C 1 = E. chaperi chaperi , D 1= E. dageti , E 1= E. etzeli , H 1 = E. chaperi sheljuzhkoi , O 1= E. olbrechtsi , P = 1E. chaperi spillmanni and S = E. spilargyreius.
Since Scheel (1968) it has been accepted that E. olbrechtsi represents a distinct species, whereas Daget & Iltis (1965) maintained it was only a subspecies of E. fasciolatus . Crossing experiments and more detailed study seem necessary to determine the degree of separation between these fish. This is now all the more important since 'new species' of doubtful status have been described recently, and there is a possibility of a con tinuum of variability.
Within the Ivorian fauna, E. olbrechtsi owes its clearly defined position to its massive appearance, its being a member of the E. fasciolatus superspecies and the bars on the rear part of the body.
Meristics: see key.
5. Epiplatys spilargyreius (Duméril, 1861)
Poecilia spilargyreia Duméril,1861 - Arch. Mus. Nat. 10: 258 - types from the Mandingues coast (Senegambia).
Haplochilus senegalensis Steindachner, 1870.
Haplochilus marnoi Steindachner, 1881.
Haplochilus senegalensis var. acuticaudata Pellegrin, 1913.
Panchax grahami var. decemfasciata Pellegrin, 1934.
Epiplatys spilargyreius has the widest distribution of all African Cyprinodontids, from Senegambia and the Nile to as far as Kinshasa in Zaïre. Considering its distribution, there is very little variability in morphology or colouration. It prefers the areas of dry savannah on inland plateaux, but it is occasionally found near the coast, especially when the trans ition in relief is gradual. To a great extent it shares the distribution area of E. bifasciatus , but curiously enough they have only rarely been caught together in the Ivory Coast. They have probably chosen different biotopes.
Identification is no problem (diagonal bands); its presence in the Ivory Coast is confirmed by locs. No.12, 15, 18, & 28 (sympatric with E. bifasciatus )
The synonymies mentioned above have been included as a formality, as there is total agreement on the subject.
Meristics: Dm=8-10, Am=15-18; Karyotype: n = 17, population from the Niger basin (Scheel,1968).
6. Epiplatys bifasciatus (Steindachner, 1881)
Haplochilus bifasciatus Steindachner, 1881 - Sitzber. Akad. Wiss. 83: 199.
Types from Bahr-el-Seraf and Bahr-el-Gebel (Nile basin)
Panchax taeniatus Pfaff, 1933.
Epiplatys ndelensis Fowler, 1949.
Epiplatys bifasciatus shares many common features with E. spilargyreius :
a wide distribution, but less extensive towards the south, since it probably stops at Oubangui in Central Africa. It too shows a preference for the dry savannah biotopes of the inland plateaux, but it is frequently found in the coastal forest biotopes (loc.6) where E. spilargyreius is replaced by the E. fasciolatus superspecies. In view of its wide distribution, surprisingly little variability in morphology and colouration. There is no problem identifying fish like E. bifasciatus : very slender, shorter dorsal, double longitudinal band and intense reticulation on the sides; its presence in the Ivory Coast is confirmed here by locs. No. 6, 8, 10, 17, 19 & 28 (sympatric with E. spilargyreius ) and the synonymies above are accepted.
E. bifasciatus can be said to be related to E. barmoiensis Scheel,1968, a relict species restricted to the coastal areas of Liberia and Sierra Leone, and also to E. biafranus Radda,1970, another relict species, this one coming from the coastal areas of South-East Nigeria.
Meristics: Dm = 6-9, Am = 14-17. Karyotype n.= 20 (Scheel,1968a).
7. Epiplatys dageti dageti Poll, 1953.
Epiplatys dageti Poll, 1953 - Rev. Zool. Bot. Afr.,48: 265. Types from Port Bouet, near Abidjan.
Poecilia (Lycocyprinus) sexfasciata : Peters,1863 ( not Gill, 1862 ) preoccupied name.
A small species, Epiplatys dageti occupies the coastal area of Liberia, the Ivory Coast and Ghana. It replaces E. annulatus east of Monrovia (Huber, 1980). The population with a red throat from Monrovia (Liberia) has been described by Daget and Arnoult (1964) as the sub-species E. d. monroviae .
A very detailed study of 1E. d. dageti and E. d. monroviae was published by Clausen and Scheel in 1966. At the present time we do not know their geographical separation, if it does exist. It would be very interesting to know the population from the mouth of the River Cavally.
Epiplatys dageti has about six vertical bars on the sides, a characteristic also found. in the superspecies E. sexfasciatus , E. multifasciatus and E. chaperi . It differs from the first two in its smaller size and in the presence of one of the bars on the caudal peduncle.
Meristics: Dm = 9- 10, Am = 14-16. Karyotype: n = 25 (Scheel,1968).
8. Epiplatys chaperi (Sauvage, 1882) and related fish.
Preliminary note: recently this species has been divided into 4 sub-species - three of which live in the Ivory Coast and one, E. c. schreiberi , is found in Ghana. In addition there is a distinct species, E. etzeli . I do not know enough about them to suggest any change in their status, and so I am giving below the systematic data of Berkenkamp & Romand, together with comments.
Collecting places: Loc. Nos. 4, 7, 11 and 21.
8a. Epiplatys chaperi chaperi (Sauvage,1882)
Haplochilus Chaperi Sauvage,1882 - Bull. Soc. lool. Fr.,7: 323. Types from Kouakoukro, 90km east of Abidjan, sympatric with Aphyosemion petersi.
8b. Epiplatys chaperi sheliuzkhoi Poll,1953.
Epiplatys sheljuzhkoi Poll - Rev. Ecol. Bot. Afr., 48: 262-265 Types from the Abidjan area.
8c. Epiplatys chaperi spillmanni Arnoult, 1960.
Epiplatys spillmanni Arnoult - Bull. Mus. Nat. Hist. Nat.,2: 219. Types from Bouaké - Central Ivory Coast - 200km. east of Abidjan.
8d. Epiplatys etzeli Berkenkamp, 1975.
Epiplatys etzeli Berkenkamp - Aquarienfreund (10): 183-198. Types from Akroaba 2:, Ono region, 60km. east of Abidjan.
All these fish are isomorphs. Meristics: D = 8-12, A = 14-19, L.L. = 28-32.
Karyotype for chaperi and sheljuzhkoi: n = 25 (Scheel,1968a). Berkenkamp (1975) states that etzeli has a more slender morphology and a filamentous extension in the male's ventral fins. The first difference I have not been able to find, and the second surely cannot be a characteristic warranting species status.
Berkenkamp and Romand's nomenclature is based on geographical separation and colouration.
Geographical separation: chaperi and etzeli in the southeast, sheljuzhkoi around Abidjan - three forest forms - and spillmanni in the north around Bouaké - a form of savannah. However, several comments must be made. Epiplatys are good swimmers, which allows them to encroach on considerably large areas (cf. E. spilargyreius) , and no geographical barrier appears between these regions. The type locations of the first three species are situated inside a 30km. segment, and Berkenkamp (1975: 193) reports that spillmanni is sympatric with etzeli near the type location of the latter - Akroaba2. As for colouration, sheljuzhkoi is the only one not to have fasciation in live fish behind the ventrals. The other three have four or five cross bars. And then the body of sheljuzhkoi and spillmanni is densely coloured with large red spots in regular lines, whereas in E. chaperi and E. etzeli these are irregular. The first two fish often have a black crossbar immediately behind the pectorals; this is absent in the others.
It would be nevertheless a very good thing if these features were the subject of more detailed study, especially with the specimens caught in the intermediate areas. It should be noted that the specimens from location 11 are here called spillmanni for geographical reasons (cf. map) but they differ considerably from the specimens from location 7, Kan-Bouaké, which are typical spillmanni. The fish from location 11 are closer to chaperi s.s. In the same way Berkenkamp & Etzel (1977: 151) publish the photograph of a male from Kouakoukro which, for geographical reasons, should be placed with E. chaperi but has no bar on the body.
In short it seems, as Scheel thinks (1968a), that there are a priori only two live phenotypes in these groups: 1 - fasciated, irregularly spotted, in the east E. chaperi ; 2 - with few or no bars, heavily and irregularly spotted, in the west and north, E. sheljuzhkoi . A sub-division into two further sub-units would not seem justified until after new collecting trips, the examination of new cases of sympatric fish and crossing experiments carried out to F3 and checked by the karyotype of the hybrids.
These are also represented by two genera in the Ivory Coast. Aplocheilichthys and Micropanchax. They have only been very recently separated, for I have shown (1981a) that only A. spilauchen should be considered to be an Aplocheilichthys for reasons of morphology, biology and behaviour. Most of the other Procatopodins come into the genus Micropanchax until a thorough revision has been made.
Aplocheilichthys lives in the marshy lagoons of area I and it is occasionally caught in the coastal streams of area 2 up to which it has been able to swim. Micropanchax lives in all the territory, but prefers such or such an area depending on the species; for example, M. rancureli , the forest plain of area 2; M. schioetzi prefers the bushy savannahs of the centre of area 3, but being adventurous, it is also found in forest areas 2 and 6.
Lastly M. pfaffi and M. normani are Sudanese species from area 5 which occasionally enter the southern areas, where they become progressively rarer.
Fig.2c Distribution map of the Aplocheilichthys and Micropanchax species as according to the areas marked on fig.1. N = 1 M. normani, P = M. p faffi, R 1 = M. rancureli , S 1= M. schioetzi and * 1 = A. spilauchen .
The Genus Aplocheilichthys.
A. spilauchen , the only species of the genus, is well represented in the Ivory Coast, probably more than in the other regions of its distribution, owing to the large coastal lagoons - the Abry, Ebrié and Lahou lagoons. Daget & Iltis (1965) even report it from nearly 100km. inland.
9. Aplocheilichthys spilauchen (Duméril,1859).
Poecilia spilauchena Duméril - Arch. Mus.,10: 258 Types from Sénégal.
Aplocheilichthys typus Bleeker, 1863.
Aplocheilichthys tschiloangensis Ahl,1928 (cf Huber, 1981a)
Aplocheilichthys spilauchen is characterised by a very extensive distribution from Sénégal to the Congo.
It is usually restricted to a coastal strip of less than 10 km., like the other Procatopodin Pantanodon, in the coastal regions of the African east; it lives in the lagoon mangroves or the sluggish streams at their mouth, sympatric with Periophthalms. More rarely it is caught further inland in company with Aphyosemion and Epiplatys. It is easy to identify: average slender body, translucent with slight reticulation of the scales and black bars on the body and fins of the male.
Meristics: D = 6-8, A 1= 11-14, L.L. = 25-28. Karyotype: n = 24 (Scheel, 1968). Maximum size: 6-7cm.
A. spilauchen is not closely related to any other Cyprinodontid. It is a relict species. It differs from the other Procatopodins in its larger size, its brackish water habitat, its less gregarious behaviour and the position of the pectoral fins. Only three groups have any similarity in morphology: Procatopus, Hypsopanchax, which are not found in the Ivory Coast, and the M. nimbaensis group of the West African plateaux. This last group is represented by M. schioetzi in the five ways described above, and also in the slight reticulation, the black bars on the male, the shorter anal fin (A 1 = 11-14 as against 14-17) and the usual micromorpho logical details of the head.
The Genus Micropanchax.
The representatives of Micropanchax are the most common Cyprinodontids in the Ivory Coast. The many rivers, the large areas of savannah shrubby or otherwise, combined with the drier climate favour their reproduction at the expense of the Rivulins. These aspects are reflected by the recent collecting trips under discussion. Their density increases towards the regions of Sudan, whilst that of the Rivulins decreases, the transition taking place around Bouaké. (area 3-4).
Four species belonging to distinct evolutionary lines have been recognised:
1- Slender morphology, short dorsal fin (D =7-8), small in size, less than 30mm.
1-1 Only coastal (forest), very pronounced sexual dimorphism, very slender morphology, cephalic pores: M. rancureli
1-2 Only inland (in the Ivory Coast) less pronounced sexual dimorphism, morphology not so slender, considerable reticulation in preserved material, cephalic pores: M. normani
1-3 Only inland (tendency to be further north than the previous fish, occasionally sympatric) very slender, no sexual dimorphism, slight dichromism, cephalic grooves: M. pfaffi
2- Deep morphology, wide dorsal (long?) (D = 8-11), reaching 50mm in size, coastal and inland, strong reticulation on the sides, spotted fins on the male, cephalic grooves: M. schioetzi
10. Micropanchax (Poropanchax) rancureli (Daget, 1964)
Aplocheilichthys rancureli Daget - Bull. Mus. Nat. Hist., 36 (4): 592. Types from the Bance near Abidjan.
Described from the south of the Ivory Coast, M. rancureli is restricted to the forest area - Ghana included - where it lives in large soft water streams. Its coastal distribution, its slender morphology, its marked sexual dimorphism (extended dorsal, anal and ventral fins in the male) make it belong to the best defined evolutionary group of Micropanchax, called Poropanchax by Clausen because of the cephalic pores. It is replaced in the east starting from Benin, by a very close form, M. macrophthalmus , then in Biafra by M. hannerzi and in the south of the Cameroons and in Equatorial Guinea by M. scheeli. Some authors say these forms can be separated at only the level of sub-species. It is not easy to separate it from another Ivorian form, M. normani , which extends to the coastal areas in the west of Sierra Leone. In fixed material M. normani differs from M. rancureli in the strong reticulation on the sides, the less slender body, the two post dorsal and post anal black bands and the strong cephalic pigmentation. However they are similar in having cephalic pores and sexual dimorphism, with the result that Clausen placed both of them in Poropanchax. It seems to me that this is not the case and that M. normani is nearer the inland forms from Sudan, the Cameroons and Congo-Gabon, namely M. camerunensis and M. stictopleuron . My reasons for thinking this are the fact that the cephalic structures in M. camerunensis are intermediate between the other two (Huber, 1981a).
M. rancureli has been caught in locs. 1, 2, 3 and 4 near the type location.
It is often sympatric with E. dageti that inhabits the same geographical area. The live fish display on the sides beautiful metallic blue colouration.
Meristics: D = 6-8, A 1= 11-15, L.L. = 27-29.
11. Micropanchax normani (Ahl, 1928)
Aplocheilichthys normani Ahl - Ann. Mag. Nat. Hist.,10 (2): 600. Types from the River Kiyawa, near Katagum, North Nigeria.
Aplocheilichthys gambiensis Svensson, 1933.
Micropanchax macrurus manni Schultz, 1942.
Described from north of Nigeria, M. normani covers the major part of the savannahs of West Africa from Senegambia (gambiensis) to Central Africa and the Upper Nile. It also extends to the coastal forest areas of Liberia (manni), Sierra Leone and Guinea and the savannah regions of Ghana and Togo. It has been accepted for a long time that the names manni and gambiensis are synonyms.
In the paragraph on M. rancureli , M. normani has been described and the references given. M. normani should probably be placed closely with the other more slender inland forms, which show considerable variability in their neuromasts: 1 M. camerunensis and M. stictopleuron .
It seems to be rare in the Ivory Coast. It has been caught at locs. 8, 12 and 18 - sympatric with M. pfaffi . Live fish show little colour: the sides of the male and the eye reflect a metallic green and a few irregular bars appear on the Caudal and the rear of the dorsal and anal.
Meristics: D =6-8, A=9-11, L.L.=24-27. Karyotype: n 1 = 24 (Scheel,1972).
12. Micropanchax pfaffi (Daget, 1954)
Aplocheilichthys pfaffi Daget - Mémoire IFAN No.36: 324, fig.124.Types from upper Niger.
Aplocheilichthys longicauda Blache & Miton,1960.
Described from Mali, M. pfaffi has been found in all the Sudanese savannahs1 , from Senegambia to as far as Central Africa and passing through Chad (longicauda). Its distribution area is largely shared by M. normani , but they are only rarely found sympatrically. M. pfaffi tends to be further north and it never reaches the coastal areas. M. pfaffi has a wide distri bution, but there is the possibility that it is even wider if one includes it in M. loati (Boulenger, 1901) which is very close in morphology and inhabits the same biotopes in East Africa. We are not yet in a position to say that they are synonyms as we have not been able to examine live fish. Seegers (1980) also links M. pfaffi and M. loati with M. antinorii (Vinciguerra, 1883) from Ethiopia. Scheel (1968b) was the first to put forward the hypothesis that M. lonqicauda was a synonym, and I was able to confirm this when I examined the types in Paris (1959-225 and 410).
Of the four Ivorian species, M. pfaffi is the most slender; the long shape is made even more pronounced by the vertical fins being short in comparison with M. rancureli and M. normani . The fish has very little colouration, and there are no spots on the fins. It has been caught in locs. 10, 13, 14, 15, 16, 17, 18 (sympatric with M. normani), 19, 20, 21, 22 and 27. This means that its Ivorian distribution area is larger than stated by Daget & Iltis's publication (1965). It now extends to the centre of the country and to the Sassandra.
Meristics: D = 6-9, A = 12-15, L.L. = 27-30.
13. Micropanchax schioetzi (Scheel, 1968)
Aplocheilichthys schioetzi Scheel - Rev, Zool. Bot. Afr., 78 (3-4): 277-283.
Aplocheilichthys macrurus (not Boulenger): Daget & Iltis,1965.
Aplocheilichthys monikae Berkenkamp & Etzel,1976.
Scheel (1968) described schioetzi after a well known fish from Ghana, which had been wrongly called M. macrurus , which is in fact a species found in Angola. The types of M. schioetzi were caught in the Bobir forest re serve, near Kumasi in Ghana. Subsequently other locations were recognised in central and eastern Ghana (Loiselle, 1974) and in the Ivory Coast (Daget and De Rham, 1970). In his publication, Loiselle points out an example of polymorphism in colouration in M. schioetzi . He takes the example of the population from the River Adansu which has black edges on the caudal and anal fins; he studies the biology and distribution of the fish compared with other Cyprinodontids of Ghana.
In 1976, Berkenkamp & Etzel described 1M. monikae from material collected by Etzel near Yo and the surrounding area on the west central savannah plateau of the Ivory Coast - Gouan or Bafing basin. They differentiated M. monikae from M. schioetzi by the structure of the frontal scalation (fusion of the 2 scales E in the second, separation in the first) and by the colouration. After re-examining the types of M. schioetzi at Tervuren and the Ivory Coast fish in Paris, I cannot agree with the retention of M. monikae as a valid name, and there are three reasons for this:
1. The two type locations are inland savannahs, which are not separated by any geographical or river barrier. They are only 650km apart, a short distance for Procatopodins.
2. The two fish are definitely isomorphs and the colouration types of the specimens are identical. The differences in fin colouration are due to variations in the populations, as was shown by Loiselle (1974), or as is also the case with the Procatopus from Nigeria to the Congo (Huber, 1981a).
3. The fish inhabits all the central region and even some coastal locations (the fish collections here extend the distribution: locs. 4, 5, 7, 9, 24, 26). The study of these specimens shows that there is a greater variation within one population than between populations when it comes to meristics and micromorphological characteristics (frontal neuromasts, frontal scalation). Finally, the two types of scalation reported by Berkenkamp & Etzel are to be found, more often it is true according to the separated type.
In conclusion, there seems no reason why the case of M. schioetzi should be different from the other Procatopodins which are sympatric. It seems inevitable that we should be thinking of the existence of only one species with deep-bodied morphology in the Ivory Coast and Ghana. We can now say that it is not improbable that eventually M. schioetzi and M. lamberti (Daget, 1962) will be declared synonyms. A study of the types of lamberti (MNHN 1959-118,1959-96 to 98) reveals a comparable morphology and an analo gous fixed colouration. Moreover, Scheel, in a description of schioetzi indicates that they belong to the same superspecies but that he has not been able to study specimens of lamberti. It seems too early to state that the two names are synonyms owing the absence of live material, and especially because the distribution areas are separated by the high plateau of the Guinea ridge, which forms an important barrier and a dividing line for water between the Niger basin and the coastal rivers.
Finally M. schioetzi shows a growth morphological allometry like the other groups of deep-bodied Procatopodins: the Procatopus sp. and the Hypsopanchax sp.
M. schioetzi and the closely related fish, M. lamberti (Daget), M. kabae (Daget) and probably M. nimbaensis (Daget) share too with these two genera the choice of a habitat ranging from the plain to the high plateaux. However, with insufficient information and live material, it is not possible to make further comparisons. The differences between M. schioetzi and A. spilauchen have been given in the paragraph on the second species. Fish of deep body (up to 33% of standard length), with a metallic blue body, the scales edged in brown forming reticulation, and with an orange-yellow edge to the dorsal, caudal and anal fins, sometimes emphasised with black.
Meristics: D = 7-11, A = 11-15, L.L. = 25-27.
With these recent collecting trips, considerable progress has been made in our knowledge of the Ivorian Cyprinodontids. It has been possible to state the distribution of the different species in terms of biogeographical constraints, and also to put forward a fairly satisfactory nomenclature. There remains the pressing need for living material for the study of the karyotypes. Crossing experiments are needed to F3, with checks on the karyotypes of the hybrids and on the variability of the colouration types. All this will give us a better picture of the relationships between the species and particularly the relationships between the Ivorian species and the other evolutionary groups, especially Procatopodins.
BERKENKAMP O., 1975. - Epiplatys etzeli spec. nov., ein neuer Hechtling aus der Südlichen Elfenbeinkuste. Aquarienfreund, 4(10): 183-198.
BERKENKAMP O. & ETZEL V., 1976. - Aquarienfische aus der Elfenbeinküste. 2- Aplocheilichthys monikae spec. nov., aus dem Westlichen Hochland. Aquarienfreund, 5(12): 223-238.
BERKENKAMP O. & ETZEL V., 1977. - Aquarienfische aus der Elfenbeinküste. 3- Roloffia maeseni (Poll,1941) aus dem Westlichen Hochland. - Aquarienfreund, 6(4): 63-78.
BERKENKAMP O.& ETZEL V., 1977. - Aquarienfische aus der Elfenbeinküste. 5- Epiplatys chaperi chaperi (Sauvage,1882) aus dem Sudosten. Aquarienfreund, 6(8): 145-158.
BERKENKAMP O.& ETZEL V., 1977. - Aquarienfische aus der Elfenbeinküste. 6- Epiplatys kassiapleuensis spec. nov. aus dem Westlichen Hochland. Aquarienfreund, 6(10): 187-198.
DAGET J., 1962.- Les poissons du Fouta Djalon et de la Basse Guinée. Mém. Inst. Fr. Afr. Noire,(65): 127-144.
DAGET J. & ILTIS A., 1965.- Poissons de Côte d'Ivoire. Mém. Inst. Fr. Afr. Noire, (74): 186-209.
HUBER J.H., 1978.- Caractères taxinomiques et tentative de regroupement des espèces du genre Aphyosemion. Rev. fr. Aquariol., 5(1): 1- 32.
HUBER J.H., 1981a.- A review of the Cyprinodont fauna of the coastal plain in Rio Muni, Gabon, Congo, Cabinda and Zaire. British Killifish Association Separatum, November issue: 1-48.
HUBER J.H., 1981b.- Aphyosemion walkeri Boulenger. Supplément à la Rev. fr. Aquariol. 8(1).
LOISELLE P.V., 1974.- Colour polymorphism in a Ghanian population of Aplocheilichthys schioetzi Scheel,1968. J.A.K.A. (J. Am.Killifish Association) 7(6): 201-208.
POLL M., 1941.- Poissons nouveaux de la Côte d'Ivoire. Rev. Zool. Bot. Afr.,34 (2): 133-143.
ROMAND R., 1978.- Cyprinodontidae around Abidjan with special reference to their biotypes. J.A.K.A. KN, 11(1): 3-12.
SCHEEL J.J., 1968a- Rivulins of the Old World. T.F.H. Pub.:1-480.
SCHEEL J.J., 1968b.- Description of a new species of Procatopodinae from Ghana, with remarks on the frontal patterns of scales and of neuromasts in West African procatopodin species. Rev. Zool. Bot. Afr. 78 (3-4): 277-283.
Translated for the B.K.A by Peter Watkins and prepared for publication by Tim Addis and Richard Cox. Illustrated by Ruud Wildekamp. A British Killifish Association publication © B.K.A. 1984. (2)
(1) Ichtyologie générale et appliquée, 43,rue Cuvier, 75231 Paris Cedex 05, France
(2) First publication (in French): Cybium.3rd series, 1982, 6 (2): 49-74.
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