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The inland water fishes of Africa

 | 
Didier Paugy
, 
Christian Levêque
, 
Olga Otero

General characteristics of ichthyological fauna

Christian Levêque et Didier Paugy

Résumé

Currently, among nearly 11,000 fresh and brackish water fish species described in the world (Nelson, 1994; Froese & Pauly, 2016), 3,360 species have been described from Africa, belonging to 95 families (Lévêque et al., 2008). Initially, these species were listed in the Check-List of the Freshwater Fishes of Africa or CLOFFA (Daget et al., 1984, 1986a and b, 1991), but since its publication numerous species have been described or placed in synonymy following taxonomic revisions. These revisions also concerned the higher entities (genera, families). Thus, many families have been broken up into several others. For example the old family Bagridae has been divided into three families: Bagridae, Claroteidae and Austroglanididae (Mo, 1991). Conversely, some families who still retained an uncertain status could now be grouped. Thus, genetic studies have confirmed the monophyly of the Cromeriidae and Grasseichthyidae with the Kneriidae within the Gonorynchiformes (Lavoué et al., 2005).
The discovery and description of new species of fish still constitute an important scientific activity in Africa. Rich faunas of cichlids endemic to the Great Lakes of East Africa have not yet delivered all their secrets. Similarly, still poorly-explored areas, such as Angola and some areas including parts of the Congo basin, should still reveal many other species.
In countries where the fish fauna is considered to be relatively well-known, the introduction of new fishing techniques (electro-fishing for example) can lead to the identification of species that had not been captured using traditional techniques alone. Similarly, using new taxonomic tools (i. e., molecular techniques, specific parasitic markers, electrical signals, behaviour, etc.) has facilitated the revision of some taxonomic groups previously based solely on morphology.
For example, we can mention the striking species flock of Brienomyrus (Mormyridae) in Ogowe basin in Gabon (Sullivan et al., 2002; Sullivan et al., 2004). At least 41 different forms could be identified, while only five species are described (figure 4.1). However, the specific status of some of these forms remains uncertain. Thus, it is not possible to distinguish the two groups that have different electric waves using only morphological or genetic characters (Arnegard et al., 2005) (figure 4.2).

Texte intégral

FIGURE 4.1. The Gabon-clade Brienomyrus (from Sullivan et al., 2002).

FIGURE 4.1. The Gabon-clade Brienomyrus (from Sullivan et al., 2002).

FIGURE 4.2. Biphasic and triphasic electric organ discharges recorded from different Brienomyrus signal forms that cannot be discriminated through genetic and morphologic characters alone.

FIGURE 4.2. Biphasic and triphasic electric organ discharges recorded from different Brienomyrus signal forms that cannot be discriminated through genetic and morphologic characters alone.

Composition of the ichthyofauna

1Cyprinidae, Alestidae, and a few Siluriformes families constitute the bulk of the riverine fish fauna with Cyprinodontiformes and Mormyridae (table 4.I). The Cichlidae are by far the most abundant with some 1,150 species (more than 2,000 estimated) and about 143 genera recorded, most of them endemic to East African lakes where many species still remain to be described.

2Ichthyologists used to distinguish three major groups of fish according to their tolerance of salt water and their ability to disperse across the sea (Myers, 1951):

  • the primary division comprises the “primary fish” which are strictly intolerant to salt water and are therefore limited exclusively to freshwaters. They are usually much longer established in continental waters than the other groups. For example, all Mormyrids belong to this group.
  • the secondary division includes “secondary fish” rather strictly confined to fresh water but evidently capable of occasionally crossing narrow sea barriers. Many of them are salt-tolerant for short periods. Some species of Cichlidae and Cyprinodontiformes belong to this division.
  • the “peripheral” division includes representatives of extant marine families which have colonized inland waters. An example is the genus Lates (Latidae), of marine origin, but now widespread in Africa where it has colonized most freshwater habitats. This group includes diadromous fish which are few in African freshwater contrary to what is observed in Europe.

3In Africa, 28 out of 94 indigenous families (six families were introduced), are “primary” division fishes and represent about 55% of the recorded species. Another 32% belong to the 3 families of “secondary” division fishes, of which the cichlids alone represent around 30%. The remaining 12% belong to 57 families of the “peripheral” division (table 4.I).

4The main feature of African fish fauna is the existence of a high proportion of endemic families (15 of the 28 families belong to the “primary” group) (figure 4.3). They are probably among the most ancient families of fish, given that the African continent has remained above sea level since the Precambrian, more than 600 million years ago. This long period of exundation may explain why Africa has a far more diverse fish fauna than South America, and an unparalleled assemblage of archaic families, such as Protopteridae, Arapaimidae, whose distribution extends far beyond Africa to Australia, Asia and South America, but that have evidently been there for a long time.

5The African inland waters house elements of marine origin which either evolved in freshwaters, or moved seasonally or sporadically between the sea and freshwaters: Perciformes, Pleuronectiformes and Tetraodontiformes provide examples.

6The number of inland waters fishes of Africa is a little lower than the number estimated for South America. Compared to African fish fauna, there are no Cyprinidae in the South American fauna which mostly consists of Siluriformes and Characiformes (table 4.II). Conversely, in Asia Cyprinidae are dominant, the Siluridae (or Siluriformes) abundant, and there are many species of marine origin (table 4.II).

TABLE 4.I. African fish families with representative in fresh and brackish waters (in orange-brown: introduced families).

TABLE 4.I. African fish families with representative in fresh and brackish waters (in orange-brown: introduced families).

Main sources of taxonomic information

7Scientific knowledge of African fish fauna is relatively recent and remains a poorly-explored area. The 10th edition of Linnaeus’s “Systema naturae”, published in 1758, cited only 13 African freshwater fish species – the ones collected in Egypt by Hasselquist. He died near Smyrna on his way home but his collections reached home in safety, and five years after his death his notes were published by Linnaeus and translated into French, German and English as “Voyages and Travels in the Levant in the Years 1749-1752”.

FIGURE 4.3. Endemic families of Africa.

FIGURE 4.3. Endemic families of Africa.

8The true father of modern African ichthyology, Michel Adanson (1727-1806), was a Frenchman who lived in Senegal from 1749 to 1753. Although he published nothing on fishes during his lifetime, he built a large collection in the form of a “herbarium”, part of which was given to the Cabinet du Roi in 1756. Another part was deposited at the Muséum de Paris, and the set was used by Cuvier and Valenciennes for their “Histoire naturelle des Poissons” whose 22 volumes were published between 1828 and 1849. About 130 to 140 species of African fresh and brackish water fishes were cited in the work. A third part of Adanson’s collection, kept in the family château, was only rediscovered in the early 20th century and was given to the Muséum de Paris in 1939. After Adanson’s death, part of his handwritten notes and field drawings were used for the “Cours d’Histoire naturelle” published in 1844-45 (Paugy, 2010a).

TABLE 4.II. Compared composition of the fish fauna in four representative large basins of Africa (Niger), South America (Mamoré), North America (Mississippi) and Asia (Mekong).

Mamoré South America

Mekong Asia

Mississippi North America

Niger Africa

Drainage area (km2)
Length (km)

159,000
1,050

783,000
,200

3,200,000
3,780

1,125,000
4,183

Agnathes

7

Elasmobranches

4

2

1

Dipnoi

1

1

Chondrostei

6

4

Semionotiformes

4

4

Amiiformes

1

Osteoglossiformes

3

2

32

Anguilliformes

1

Clupeiformes

4

2

4

6

Gonorynchiformes

1

Characiformes

138

35

Beloniformes

1

Cypriniformes

168

81

35

Siluriformes

146

65

65

Gymmnotiformes

20

Esociformes

2

Atheriniformes

4

2

Percopsiformes

1

Cyprinodontiformes

3

1

1

23

Perciformes of which Cichlidae

21
18

17

63

33
17

Gasterosteiformes

1

Synbranchiformes

1

7

3

Scorpeniformes

2

Pleuronectiformes

1

5

2

Tetraodontiformes

7

1

TOTAL

350

278

177

242

Continental level

9The only general fish fauna available on a pan-African scale is Boulenger’s Catalogue (1909-1916) which is only of historical value at present. A major step forward was the publication of the Check-List of the Freshwater Fishes of Africa (CLOFFA) which provides a current list of brackish and freshwater fish species with full synonymies as well as a full bibliography (Daget et al., 1984, 1986a and b, 1992). The CLOFFA is particularly useful for taxonomists as a major source of information for further taxonomic revisions.

Regional level

10For several decades, ichthyologists devoted their research to the inventory of the fish fauna of African continental waters, which allowed the description of many new species. In some cases, their work led to the publication of local or regional inventories and fauna lists. In other cases, regional taxonomic summaries were not yet made. A detailed bibliography of taxonomic publications on African fishes can be found in volumes 3 and 4 of CLOFFA (Daget et al., 1986b, 1991). The websites FishBase (www.fishbase.org) and Faunafri (http://www.poissons-afrique.ird.fr/​faunafri/​) are, of course, essential tools.

Maghreb

11Pellegrin (1921) remains an important reference for the Maghreb. Other works on the systematics and phylogeny of various groups, notably the Cyprinidae, have been published since, but a summary of these works would be welcome (Doadrio, 1994).

Western Africa

12Ichthyological inventories for West Africa began in the late 19th century. F. Steindachner from Austria (1870, 1895) as well as A. Günther from Britain, then H.E. Sauvage and J. Pellegrin from France and the Belgian-born British scientist G.A. Boulenger, described numerous species based on the collections that were brought back to them by travellers. A fauna of the freshwater fishes of West Africa written by Pellegrin (1923) summarized the state of knowledge at that time. Later, especially after the Second World War, inventories covered whole basins or regions. These included the following monographies by country or basin: Gambia (Svensson, 1933; Johnels, 1954), Ghana (Irvine, 1947), Upper Niger (Daget, 1954), Chad basin (Blache et al., 1964), Côte d’Ivoire (Daget & Iltis, 1965), Volta (Roman, 1966), coastal rivers of Côte d’Ivoire, Ghana, Guinea, Sierra Leone, Liberia, and Togo (Schültz, 1942; Daget, 1962b; Teugels et al., 1988; Lévêque et al., 1989, 1991; Paugy et al., 1989, 1994; Paugy & Bénech, 1989). The many inventories established later, most notably by teams from Orstom (later IRD) and the Tervuren Museum (Belgium) made it possible to publish an updated fauna of fresh and brackish water fishes of West Africa (Lévêque et al., 1990, 1992; Paugy et al., 2003a, 2003b) which provide the state of knowledge on the taxonomy and distribution of species.

Nile Valley

13By the late 18th century, European ichthyologists had begun studying Nile fishes. In 1789, Gmelin cited 17 Nile species based on the collections of Hasselquist (1722-1752) and Forskål (1736-1765). In 1799, Geoffroy Saint-Hilaire accompanied General Bonaparte during his Egyptian expedition and assembled large collections deposited with the Muséum d’Histoire naturelle de Paris. He published the description of Polypterus in 1802, and that of other species in 1809. The first overview of Nile fishes was published in 1907 by Boulenger and remains a useful reference.

14Lake Tana was the focus of recent work that revealed the existence of a large Barbus species flock (Nagelkerke & Sibbing, 2000).

Lower Guinea

15The first collections from Cameroon were collected by the German scientist R. Buchholz in the region of Douala and deposited at the Zoologissches Museum der Humboldt Universität in Berlin where they were studied by W.C.H. Peters. Later, Lönnberg of Sweden published the results of collections carried out in the area of Mount Cameroon. Between 1903 and 1913, G.A. Boulenger wrote a summary of the collections made by the American scientist G.L. Bates deposited at the British Museum in London. The Austrian professor K. Haberer also carried out important collections from the Sanaga River between 1907 and 1909, deposited at the Naturhistorisches Museum in Vienna but described only twenty years later by Holly. In 1930, he published a synopsis of the freshwater fishes of Cameroon. Later, E. Trewavas (1962) examined the crater lake fishes deposited in the British Museum’s collections. They were collected in 1948 by the British scientist P.I.R. MacLaren from the southwestern part of the country.

16The first specimens from the Ogowe River collected by R.B.N. Walker in 1866 were investigated by British scientist A. Günther of the British Museum. Later, the river was explored in 1874 by Savorgnan de Brazza, who deposited large collections at the Muséum d’histoire naturelle de Paris. Sauvage studied them and later published the first ichthyological inventory of the Ogowe River (Sauvage, 1880). Finally, Mrs Kingsley arrived on the banks of the Ogowe River in 1895 to study the civilizations of Gabon, but she also collected 65 fish species (of which 46 were then unknown). J. Pellegrin continued this work later.

17The first collections from the coastal basins of the Congo were carried out in 1929 by French scientist Baudon and deposited at the Muséum de Paris, where they were also examined by J. Pellegrin.

18For Lower Guinea (Cameroon, Gabon, Congo), partial contributions had to suffice for a long time: Amiet et al., 1987; Daget, 1978, 1979; Daget & Depierre, 1980; Roman, 1971; Mbega & Teugels, 2003; Thys van den Audenaerde, 1966, 1967; Teugels et al.., 1992; Teugels & Guégan, 1994; Mamoneke & Teugels, 1993. But an inventory of N’Tem fauna was nonetheless published (Kandem Toham, 1998) and more recently, an overview of the fresh and brackish water fauna of Lower Guinea was also published (Stiassny et al., 2007a and b).

Congo

19The situation is much less favourable for the Congo, for which Boulenger (1901) proposed an initial inventory. Major contributions from the first half of the 20th century came from Nichols & Griscom (1917), Fowler (1936) and Poll (1938). These were followed after the Second World War by major regional contributions such as those by Banister & Bailey (1979), De Kimpe (1964), Gosse (1963, 1966, 1968), Matthes (1964), Poll (1967, 1976), and Poll & Gosse (1963). There are numerous collections from the Congo basin, and it is urgent to summarize and update the available information (Teugels & Guégan, 1994) because this work was not yet realized for the largest African basin. Nevertheless, several recent local contributions (Wamuini Lunkayilakio et al., 2010; Ibala Zamba, 2010) have improved our knowledge of the ichthyological fauna of the Congo basin, which most certainly contains the greatest species diversity.

Angola

20Despite Poll’s work (1967) over a very limited area, Angola remains a poorly known region in which there remains much to be discovered.

Eastern Africa

  • 1 The site http://www.destin-tanganyika.com/ also provides a fully updated overview of the ichthyolo (...)

21The fauna of the East African great lakes is composed in large part of endemic species (Lowe-McConnell et al., 1994). For Lake Victoria, taxonomic data can be found in Barel et al. (1977), Greenwood (1980, 1981), Oijen (1991, 1996), Oijen & Witte (1996), Witte & Oijen (1990), Fermon (1997). For Lake Tanganyika, the main sources are Brichard (1989), Coulter (1991), De Vos & Snoeks (1994) and Poll (1986).1 References for Lake Malawi include Konings (1990), Eccles & Trewavas (1989) and Duponchelle & Ribbink, 2000); for Lake Kivu, Snoeks (1994); and for Lake Rukwa, Seegers (1996).

Zambezi

22The fauna of the Zambezi is relatively scarce and is rather well known with the work of Jubb, 1965, 1967; Bell-Cross, 1972, 1976; Gaigher & Pott, 1972; Bruton & Kok, 1980; Skelton et al., 1985; Jackson, 1986; Skelton, 1994; and Marshall, 2011.

East Coast

23An exhaustive list of the fish species of the Ethiopian Rift Valley was recently published by Golubtsov et al., 2002. For the East Coast of Africa, Eccles (1992) published a summary of the current state of knowledge, and a detailed inventory was proposed by Skelton (1994).

Southern Africa

24In South Africa, the inventory of freshwater fishes began with Burchell’s expedition (1811-1812), whose results were published in 1822. A. Smith, who was in charge of the Cape Town museum built in the 1820s, carried out missions inside the country between 1834 and 1836, and published the results of his collections between 1840 and 1845; they included several fish species. During this period, German scientist Wilhelm Peters went on an expedition in what is now Mozambique and described many species. In 1861, France’s consul at Cape Town, F. Castelnau, described 21 fish species including some from the Okavango. Several other European scientists, including F. Steindachner of Austria, M. Weber of the Netherlands, and A. Günther and G. Boulenger of Britain, also made significant contributions in the late 19th and early 20th centuries. G.D.F Gilchrist and W.W. Thompson of the South African Museum were the first resident scientists to work on freshwater fishes. They published an initial catalogue between 1913 and 1917. Their successor, K. Barnard (1943), then R. Jubb after the Second World War, pursued the inventory of species with Jubb’s 1967 release of “Freshwater Fishes of Southern Africa”. Skelton (2001) provides an updated summary of knowledge of this relatively poor fauna dominated by the Cyprinidae.

Madagascar

25Malagasy fauna mainly comprises species of marine origin (Pellegrin, 1933; Arnoult, 1959; Kiener, 1963; Stiassny & Raminosoa, 1994). If species that only penetrate fresh water occasionally are excluded, known species would number around sixty (de Rahm, 1996; Sparks & Stiassny, 2003).

Taxonomic revisions

26Many taxonomic revisions of African fishes at the continent level have been published during past 30 years: Among the main works, we can list:

  • Mormyridae: phylogeny of the family (Lavoué, 2001); phylogeny of the Gabon-clade Brienomyrus (from Sullivan et al., 2002)
  • Alestidae: Alestes and Brycinus (Paugy, 1986); Hydrocynus (Paugy & Guégan, 1989); Hubert et al., 2005
  • Cyprinidae: Neobolines (Howes, 1984), Labeo (Reid, 1985)
  • Bagridae: in Africa, the Bagridae are now separated in three: Bagridae, Claroteidae and Austroglanididae (Mo, 1991)
  • Claroteidae: Chrysichthys (Risch, 1986); Auchenoglanis and Parauchenoglanis (Teugels et al., 1991)
  • Schilbeidae: Schilbe (De Vos, 1984; 1995)
  • Mochokidae: Synodontis (Poll, 1971); Microsynodontis (Ng, 2004); phylogeny of Synodontis (Pinton et al., 2012)
  • Clariidae: Clarias (Teugels, 1986; Hanssens, 2009) and Heterobranchus (Teugels et al., 1990)
  • Malapteruridae: Malapterurus description of 11 new species; a new genus Paradoxoglanis with 3 new species (Norris, 2002),
  • Cyprinodontiformes: Poeciliidae (Huber, 1999; Ghedotti, 2000), Aplocheilidae (Murphy & Collier, 1997; Murphy et al., 1999).
  • Cichlidae: Oreochromis and Sarotherodon (Trewavas, 1983), Tylochromis (Stiassny, 1989)
  • Mastacembelidae: Aethiomastacembelus and Caecomastacembelus grouped into a single genus Mastacembelus (Vreven, 2001; 2005)
  • Channidae: Parachanna (Bonou & Teugels, 1985).

27It is now possible to keep track of developments in the identification of fauna in real time by consulting reliable specialist sites.

Notes

1 The site http://www.destin-tanganyika.com/ also provides a fully updated overview of the ichthyological fauna of Lake Tanganyika.

Table des illustrations

Titre FIGURE 4.1. The Gabon-clade Brienomyrus (from Sullivan et al., 2002).
URL http://books.openedition.org/irdeditions/docannexe/image/25187/img-1.jpg
Fichier image/jpeg, 534k
Titre FIGURE 4.2. Biphasic and triphasic electric organ discharges recorded from different Brienomyrus signal forms that cannot be discriminated through genetic and morphologic characters alone.
URL http://books.openedition.org/irdeditions/docannexe/image/25187/img-2.jpg
Fichier image/jpeg, 70k
Titre TABLE 4.I. African fish families with representative in fresh and brackish waters (in orange-brown: introduced families).
URL http://books.openedition.org/irdeditions/docannexe/image/25187/img-3.jpg
Fichier image/jpeg, 318k
URL http://books.openedition.org/irdeditions/docannexe/image/25187/img-4.jpg
Fichier image/jpeg, 390k
URL http://books.openedition.org/irdeditions/docannexe/image/25187/img-5.jpg
Fichier image/jpeg, 257k
Titre FIGURE 4.3. Endemic families of Africa.
URL http://books.openedition.org/irdeditions/docannexe/image/25187/img-6.jpg
Fichier image/jpeg, 373k

Auteurs

Research director at IRD. He has published numerous works on the systematics, distribution, and ecology of West African fresh water fishes, and expanded the collections of the MNHN in Paris.

Research director at IRD. He has published numerous works on the systematics, distribution, and ecology of West African fresh water fishes, and expanded the collections of the MNHN in Paris.

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