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Bio1.J. Linn. Soc., 1, pp. 51-75.

With 2figures

April 1969

Speciation in tropical freshwater fishes
R. H. LOWE-McCONNELL, F.L.S.
Streatwick, Streat, Hassocks, Sussex
Among freshwater fishes greater diversity is shown, both within taxa and in communities,
at low than at high latitudes.
This paper discusses contributions that ecological studies on tropical fishes make towards
understanding this diversity, the mechanisms of speciation, and the ability of so many species
to coexist in these complex communities.
In Africa studies have centred around the explosive speciation of cichlids in the Great
Lakes ; in the rivers non-cichlids have speciated more abundantly. In South America catfishes
and characoids exhibit the most spectacular adaptive radiations, evolving mainly under
riverine conditions. Rates of evolution are indicated by the presence of five endemic Huplochromis cichlids in L. Nabugabo, a small lake cut off from L. Victoria 4000 years ago (Greenwood, 1965), and of eighteen cyprinids in the 10,000 year old L. Lanao in the Philippines
(Myers, 1960).
Tropical fish communities are diverse wherever no extreme environmental conditions
restrict fish distribution. No one explanation can account for this diversity in all cases. In rivers
it seems mainly due to allopatrically evolved species coming together in the course of time.
Species flocks are characteristically found in lakes, and here microgeographical isolation can
allow ‘allopatric’ speciation within the lake, as amongst the ‘Mbuna’ cichlids of Nyasa, but not
among Nyasa TiZupiu.
Isolating mechanisms include physical, chemical and biotic barriers causing spatial isolation; temporal isolation ; and behavioural barriers, many of these fishes having complicated
courtship displays, and some ‘homing’ to particular places to spawn.
The numbers of habitat niches are enhanced by the complexities of the tropical environment. Nevertheless there is a good deal of overlap-in Victoria thirty HupZochromis species
share a major habitat (Greenwood, 1964). Specializations to use a particular food have
developed several times so that even in a single habitat several species may tap the same food
source; ten species of Mbuna share the Aufwuchs of Nyasa’s rocky shores (Fryer, 1959). This
appears to contradict Gause’s exclusion hypothesis, but no two species are likely to overlap
in all their living requirements throughout life. Recent work on variations in oxygen tolerances of young cichlids (Welcomme, 1967) indicates how many as yet unstudied factors must
affect the dynamic balance of numbers between co-habiting species.
Devices that reduce competition, and the role of predators, are both discussed.
Predators abound, and predator pressure appears to have been a key factor in moulding
evolution in these communities. Many fish species have developed some form of parental care,
with associated decrease in fecundity. Biotic pressures are of far greater importance than
climatic factors in the evolution of these tropical fishes.
CONTENTS

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Introduction
Diversity of tropical freshwater fish faunas .
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Types ofwaters
Composition of the fish faunas .
Speciation in African Cichlidae
Conditions in Great Lakes compared with rivers .
Conditions in rivers
Niche number and overlap in ecological requirements
A note on isolating mechanisms
Conclusions
Summary
Acknowledgements
References

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R. H. LOW-MCCONNELL
INTRODUCTION

Among freshwater fishes greater diversity is shown, both within taxa and within
communities, at low than at high latitudes, closely related species living sympatrically
under tropical conditions, even sharing the same food.
This paper discusses the contribution that ecological and evolutionary studies on
tropical fishes make towards understanding how these complex communities have
originated, and how so many species manage to coexist. Tropical fishes have been
much studied, as they are economically very important, but results are published in
widely scattered journals and reports. I am here interpreting the work of many people
in the light of my own field experience in various parts of Africa and South America.
Understanding the diversity of tropical communities involves knowing (1) how the
communities become so complex, and (2) how they manage to persist. The first
question involves mechanisms of speciation and is principally genetical. We do not
know whether any fish mutation rates are higher in the tropics, but we do know that
where life cycles are shorter and breeding seasons protracted there will be more
generations per unit time, and it is possible that certain breeding habits may favour
rapid recombinations of genes or spread of mutations. Barriers between breeding
populations of fishes may be mainly geographical or ecological, temporal or ethological.
Species evolved in isolation, or semi-isolation, may then be brought together by
environmental changes. When this happens how do the species manage to coexist ?
This second question, primarily an ecological one, involves understanding the interrelationships of species with one another and the physical environment. How much
competition exists ? Are ‘niches’smaller in the tropics, or is there greater tolerance of
overlap in requirements between species under tropical conditions ? What controls
the relative numbers of different species; the exigencies of the physical environment,
seasonal changes, biotic factors such as food, space or predation ?
Concepts put forward which might explain latitudinal gradients in various animal
groups (reviewed by Pianka, 1966-and see his paper for bibliography-but with
little reference to fish) include those dealing with time and climatic stability (whether
older communities tend to have more species, provided that climatic catastrophies,
such as ice ages, do not intervene); spatial heterogeneity (whether more niches exist,
allowing greater specialization); competition (whether this is greater in complex
communities);predation (whether there are more predators, and their role in holding
down prey populations enough to lower the level of competition between themwhich implies reduced competition amongst prey species); productivity (whether
productivity or biomass are greater under tropical conditions, and if so, whether this
allows greater diversity).
What do ecological studies on tropical fishes contribute to these various concepts ?

DIVERSITY OF TROPICAL FRESHWATER FISH FAUNAS

The richest freshwater fish faunas are to be found in the Amazon, equatorial
Africa and tropical Asia (Fig. 1).The numbers of fish so far recorded from these areas
are only provisional, as exploration is far from complete and new species are being

SPECIATION IN TROPICAL FRESHWATER FISHES I I I I l I a 2 53 .

Africa has also the Great Lakes of east and central Africa. LOW-MCCONNELL found all the time. In equatorial Africa two areas of the Congo support 408 species (Poll & Gosse. though Lake Malawi (= Nyasa) which is well south of the equator shows greater seasonal changes than the equatorial L. Malawi and Victoria. such as Lake Chad. flows through high forest. Victoria. which have many species in common. 1947). but only 90 m deep (the depth of Windermere). Many of the South American river systems interconnect (Orinoco to Amazon for example) and some fish species are very widely distributed. lies like a saucer between the Rift Valleys. For comparison the whole of Europe has 192 species representing 25 families (Ladiges & Vogt. 1965). Eigenmann (1912:41) took 70 to 90 species in one haul of a 150-foot seine in the Essequibo. In Africa there are four main river systems. which are at higher latitudes. Myers (1947) pointed out that the Amazon system alone has well over 1000 species.000 square miles. and the Great Lakes of North America have 172 species of 29 families (Hubbs & Lagler. and many of the South American rivers are very convoluted.R. notably in Tanganyika. the Zambezi 155 species. and 60 species in two or three hours from a small brook in this area. belonging to fewer families (Bell-Cross. TYPES OF WATERS Waters are predominently riverine in South America and Asia. as well as other eastward. which also has many of the same species. the Upper Niger has at least 134 species (Daget. In South America the equatorial country of Brazil has 1383 species belonging to some 46 families (Fowler. Large endemic fish faunas (particularly of cichlid fishes) have evolved in these lakes. the Cyprinidae. from Central America to the Argentine. formed only a few million years ago and offering new opportunities to the fishes which could take advantage of the lacustrine conditions. In tropical Asia. 1945). 1966). 1967). Aramburu & Aramburu. whereas the number falls to 92 species south of the Zambezi (Jubb. and 456 species in Central America (Miller. the Congo. then Malawi (20 endemic genera). 1967) and over half of these belong to one family. many tributaries having a low pH and forest debris causing de-oxygenated conditions. 1965a). Tanganyika (over a mile deep) and Malawi (over 400 miles long) lie in the rift valley. Thailand has 546 species belonging to 49 families (Smith. a number which might be almost doubled when exploration is complete. 1954). the Congo. forming ox-bow lakes and pools where fishes get cut off from the main river until released by floods in subsequent years. Victoria.or westward-flowing rivers and inland drainage areas. though huge areas alongside the rivers are flooded in wet seasons. this probably reflects the comparative ages of these three lakes. the Niger and Nile. . differentiation (as measured by the numbers of endemic genera present) is greatest in Tanganyika (which has 38 endemic genera). and the Zambezi. compared with 339 species in Argentina (Ringuelet. 1954). making a heterogeneous environment. the size of Northern Ireland. giving temporarily lacustrine conditions. These lakes are very large. like most of the Amazon. and least in Victoria (only four endemic genera) (Table 1). H. The equatorial Congo shows much less seasonal variation than the Niger. 26. 1963) representing 24 families. Nile and Zambezi. Conditions are much more stable in these lakes than in the rivers.

S. others (belonging to three distinct families) are heavily armoured. Smith 1945. Greenwood 1966. Tanganyika. Fowler. Rudolf. % of fauna No. Malawi (Nyasa). T h e Ostariophysi as a group dominate these tropical freshwater fish faunas. the catfishes (Siluriformes) and the Characoidei suborder of the Cypriniformes. Cichlidae No. African t i w e n : Niger. Congo. families Mormyriformes No. Characoidei 70 Cyprinidae No. 594 43 553 40 0 0 81 6 0 % 0 Siluriformes No. T h e South American catfishes belong to about 13 families. Worthington & Ricardo 1936. 2 mainly from the following sources:. some of which are scaleless ‘skinfish’. 2.America: Brazil. Poll 1953. The composition of tropical freshwater fish faunas African rivers S. of genera Cichlids Non-cichlids No. BellCross 1959. 1963. No. ranging in size from small parasitic species to some of the largest freshwater fishes known. Zambezi. 1954.46 No. These catfishes make up 43% of Table 1. families Cichlidae No. The South American freshwater fish fauna is mainly of ostariophysian fishes. Africun Great Lakes: Albert. Poll & Gosse 1963. Victoria.SPECIATION IN TROPICAL FRESHWATER FISHES 55 COMPOSITION O F T H E FISH FAUNAS T h e fish groups which show the greatest diversity are indicated in Table 1 and Fig. No. Jackson et ul. spp. spp. spp. Asiu: Thailand. Yo of fauna % Endemism Cichlids Non-cichlids No. 1956. . Daget 1954. America Brazil 1383 c. I n South America these are represented by two main groups. Greenwood 1966. YO % Niger Congo Zambezi Asia Thailand 134 26 408f 24 155 15 546 49 37 112 x t 27 34 22 18 11 47 30 23 15 16 27 22 73 16 18 24 52 13 32 18 10 7 7 75 19 14 18 99 18 0 0 209 38 0 0 0 10 0 African Great Lakes I No. endemic genera Cichlids Non-cichlids Albert Rudolf Victoria Tanganyika Malawi (Nyasa) 46 14 37 14 177+ 12 214 14 245f 7 9 5 127f 134 62 193+ 19 13 71 44 5 40 12 98 58 78 98 97 57 65 2 26 3 22 6 20 34 42 23 22 0 0 0 0 4 0 30 0 20 0 Data here and Fig.

particularly the plecostomine group of rather small armoured fishes. primarily algal-grazing. The proportional composition of the freshwater fish faunas of S. LOWE-MCCONNELL 56 the Brazilian fauna. Note relative importance of Cichlidae in lakes and of Cypriniform fishes (Characoidei and Cyprinidae) in rivers. that have speciated most (135 species in Brazil). H. Numbers of species expressed as percentage of known fauna (1% = 3. Another closely-related armoured group. . Amm*cu (Brazil).6"). often found in rocky streambeds. Africun r i w m and Aficun Great Lakes. living mainly in submerged SOUTH AMERICA ASIA AFRICA RIVERS Niger Congo Albert Victorio Key: Cichlids Cotfishes Zombez i Rudolf Tongonyiko 6) Chorocoids Cyprinids Molowi Morrnyroids Others FIGURE 2. the loricarines.R. Asia (Thailand). and within the group it is the loricariid catfishes.

Most of these characoids move upstream and out into the flooded country to spawn during the rains. and cyprinids which have evidently come in from Asia and spread down eastern rivers to southern Africa. taken by Eigenmann (1912) as one family. Characoid fishes present one of the most extreme cases of evolutionary adaptive radiation among living groups of vertebrates (Weitzman. Africa also has many catfishes (some of these most closely related to Asian species). but the pimelodid skmfishes. but as yet ecological studies are only in their infancy. most species are widely distributed. a rather omnivorous group. but the richest complex of species is among the small rather generalized tetragonopterids studied by Eigenmann (1917). The Cichlidae make up only 6 % of the Brazilian freshwater fish fauna. however. Cichlids are. of which Brazil has 179 species. make up 40 % of the South American fish fauna. however. for the complexity of the fauna makes these difficult until the species are better known. has 64 species. enough to enable us to compare conditions here with those in the better-studied African waters. again perhaps because it was a less saturated river. though here they are somewhat less well-represented. T h e rivers of Africa have far fewer cichlids than have the Great Lakes. 1962). many hundreds of kilometres (see page 65). various characoids fill niches in South American waters comparable to those occupied by cyprinids in Africa. and have undergone extensive adaptive radiations to use all kinds of food. We do. and unlike the African cichlids they are not true mouthbrooders. though these two groups are not at all closely related. they vary very much in body size and form. but now considered to represent 6 to 13 families. many of them move very long distances. where the genus CichZusomu is represented by 14 species (Miller. Africa shares the characoids with South America. the gymnotoids endemic to South America (four families and including the electric eel Electrophorus electrim) and the mormyroids (two families) endemic to Africa. 2). but in some of the Great Lakes (Victoria. the Zambezi has a higher percentage (15%). now know something about the general natural history of some of the fishes and fish groups. South America completely lacks the Cyprinidae. an important family in Africa and the predominant family in Asia. 1962). The characoids. Cyprinids are also well-represented in West Africa. also have high representation (112 species). very important in Central America. These South American fish faunas have attracted a good deal of systematic study. and occur in the Congo. 1966). probably because environmental saturation with better adapted fishes had occurred there before they arrived. and these are most abundant in the Congo and West Africa (Table 1 . Ecological counterparts are also provided by two groups of electric fishes. in the Congo and Niger cichlids make up only 7% of the fauna (compare 6 yoin South American rivers). 1966).SPECIATION IN TROPICAL FRESHWATER FISHES 57 forest debris. and their whole biological regime is governed by seasonal flooding. the African genus Alestes greatly resembles the South American genus Brycon (Weitzman. Tanganyika and Malawi) . Fig. their success here has been attributed to their salt tolerance and ability to move into an area not yet saturated with primary freshwater fishes when South America became connected with Central America at the end of the Pliocene after its long period of isolation (Myers.

also Fryer (1959a. and the food of the non-cichlids by Corbet (1961). 1962). particularly their ability to spawn in still water and freedom from the need to run up streams to spawn. in Central America they exploited many types of environment.b. which makes them independent of the rains for spawning. and with the parent. and often in marginal habitats such as crater lakes (e. The prolonged breeding period also allows genetic recombinations to be tried out in a relatively short time. The breeding males develop specific breeding colours and establish territories where they are visited by the ripe females. Welman (1948) and Holden (1963) in Nigeria. More recent evidence suggests that the females may home to certain restricted brooding grounds and become territorial when the young first reach the free-swimming stage: Fryer (1961) found marked Tilapia wuriubilis returning to brooding areas that they had previously used. for references). This production of young throughout the year keeps populations very stable. Studies on the ecology of riverine fishes in Africa include those by Poll (1959) Poll & Gosse (1963) Gosse (1963). 1966). LOWE-MCCONNELL cichlids make up more than 60 to 70%of the fauna. more or less throughout the year in some cases. producing families at frequent intervals over a protracted breeding period. the Malawi fauna by Trewavas (1935). which Poll (1956) considered to be of great evolutionary significance. Jipe: Lowe. L.58 R. Johnels (1954) on the Gambia. 1957) on the Niger and other west African rivers. Poll (1966. (1964) summarize work on L. 1967) on the rivers of Angola. Chad fishes. life cycles are short (many species mature in less than a year) so the number of generations per unit time is high. This mouth-brooding habit keeps the young together. and Matthes (1964). Many other members of the staffs of the East African Freshwater Fisheries Research Organization and the Joint Fisheries Research Organization for North Rhodesia and Nyasaland have also contributed to these studies. in the Cameroons: Trewavas. 1952. and though figures as yet are small .b) on the ‘Mbuna’ rockfish.g. and (ii) their ability to undergo morphological and physiological changes allowing them to exploit all kinds of food. and the soda lakes and hot springs of east Africa (Coe. 1966). 1953) on the Tilapiu. Daget (1954. and Lowe (1952. new young entering stocks at all times of year. 1956) and Coulter (1962). on the Congo. The African cichlids are nearly all mouth-brooders. H. Many studies have centred around the explosive speciation of cichlids in the Great Lakes of Africa.g. 1955). speciating in many lakes (Miller. Mating appears to be polygamous. SPECIATION IN AFRICAN CICHLIDAE What is especial about the cichlids which has enabled them to colonize these Great Lakes of Africa ? The Cichlidae were preadapted for lacustrine life by (i) their breeding habits. and fishes at all stages of growth being present in the populations. the Tanganyika cichlid fauna (134 species) has been studied by Poll (1951. Blache et al. Jackson (1961a. which pick up the eggs and move off to brooding grounds. the Victoria cichlids by Greenwood 1956-67.b) and BellCross (1965~)on the Zambezi. 1960a. Elsewhere in Africa they have colonized and speciated in small lakes (e. and 97 to 98 % of these cichlids are endemic to a particular lake. 1964. Cichlids are genetically plastic and good at moving into new environments.

whereas the cichlids have speciated most abundantly in the intervening lake (Poll. Mormyrus longirostris. Malawi (Lowe. Chrysichthys catfish (six species) and Mastacembelus (12 species) (Poll. The formation of a large lake presents a new unpopulated environment (comparable with an island) offeringopportunities for creatures which can take advantage of lacustrine conditions. ornat+innis. The genera of cichlids which could adaptively radiate to use different foods have speciated most. Greenwood. Congo species (such as Polypterus congim and P . 1962). where conditions change seasonally. as the changes in fish fauna after . Myers commenting on this Lanao flock says that ‘lack of competition’ allows this explosive adaptive radiation. lack this ability for trophic specialization). the outstanding example is the genus Haplochromis of which there are 112 species in L. Lake Malawi has no endemic genera of non-cichlids. as Matthes (1964) has pointed out. Iles. Malawi (Jackson. 1963) and well over 120 species in Victoria (Greenwood. In the Philippines Lake Lanao has a cyprinid species flock studied by Myers (1960). compared with Tilapia which are restricted to algae or plant food. Among the freshwater fishes the cichlids were pre-eminently able to take these opportunities. a male and a female in one case. Albrecht ( 1963) studying Haplochromis (wingati and burtoni) in aquaria found that the females became territorial when the young reached the free-swimming stage. two females in another case. CONDITIONS IN GREAT LAKES COMPARED WITH RIVERS Lakes are younger than rivers and are colonized from them. 1959. This is particularly clear in the case of Lake Tanganyika. But what factors push the fishes into these adaptive radiations 1 Presumably intraspecifx competition at first. where the formation of the lake cut across the Malagarazi river drainage flowing westward to the Congo basin. 1953). Distichodus maculatus. and of which the largest species flock is of four species in L. However. and that the young learnt the mother’s territory and did not leave this home area until brooding had ceased. his more recent estimates put it at over 170 species). many still having to migrate up inflowing streams to spawn. The cichlids’forte is specialization to use different foods. but the cyprinids. 1961). Such behaviour would help populations to develop in semi-isolation. mark returns also indicated prolonged association or re-encounter between adult fish. which are toothless. 1964. Tanganyika lake has eight endemic genera of non-cichlids (compared with 30 endemic cichlid genera). Some non-cichlids have also colonized these Great Lakes. which may explain the paucity of the cichlid fauna in rivers compared with lakes. 1953.SPECIATION IN TROPICAL FRESHWATER FISHES 59 in number. Matthes. and species flocks of zooplankton-feeding clupeids (two species). Cichlid numbers build up very rapidly after a new lake is stocked with them. Tetraodon mbu and Labeo weeksii) being found in the Malagarazi but not in the lake. This left a distributional gap for most of the non-cichlids. Harding & Fryer. but seem to have taken longer to become adapted to lake conditions. Greenwood has pointed out that they have the morphological and physiological apparatus to do this easily (and so have the characoids. 1951). fishes that are too specialized in their food requirements could not survive in rivers. predatory Lutes and Luciolates (five species). but there is a species flock of offshore-living large Dinotopterus catfish (Jackson.

1966. Myers (19604 pointed out that the explosive evolution of endemic fish faunas in older lakes produces specializations far beyond those exhibited outside the lake. separated off from Lake Victoria by a sandbar dated 4000 years old. that is. but a small lake. Myers. 1966). Petr. LOWE-MCCONNELL the formation of new man-made lakes show (Jackson. formed by a larva flow 10. 1965). so the fishes are concentrated around the shore or in surface waters. 1956).60 R. for this reason these lakes may be regarded as ‘evolutionary traps’ (Briggs. These deep tropical lakes remain permanently stratified. and he suggested that such evolutionary ‘spurts’ may be important for the origin of higher taxonomic groups. as do most of the members of the Mastucembelus species flock in this lake (Matthes. Evolution may be very rapid in these new environments. The availability of foods throughout the year allows extreme specializations to use a particular food source (so food niches may become very small). The normal fate of a lake is to fill up and become a swamp. Malawi (Nyasa) it supports the complex of cichlid species known locally as ‘Mbuna’ (27 species belonging to nine genera) whose ecological and evolutionary relationships were studied in detail by Fryer (1959). . or when. 1962). These rocky stretches are like islands. H. from other such rock ‘islands’. Fishes dwelling among rocks are generally rather small. shelter from predators. in L. 1960a). cichlids multiply so fast that they very soon ‘runt’. now hasl8endemic cyprinids (a species flock of 13 and five species belonging to four endemic genera. in this case in the absence of predators. In the Victoria basin Corbet (1961) attributed the stenophagy of Mastacembelus for an insect diet to stem from its restriction to a rocky habitat (in contradistinction to most other non-cichlids in Victoria which are Inore facultative feeders). Should. seasonal fluctuations in level are small in relation to the volume. fishes that have specialized for conditions in lakes will be at a disadvantage and most will die out. have probably had an important role in allowing fishes from one rock ‘island’ to gain access to other rock areas. In L. where the shoreline is broken into alternating rocky or sandy or swampy stretches (the latter generally where a stream enters the lake). However. and produce small families. or in types of food available. has five endemic Haplochromis species (Greenwood. and when stocked in dams. conditions change. The well-lit rocky environment seems to be a potent one for fish speciation. Rocks not only provide a rich food supply where there is enough light. or semi-isolation. due to long-term wet or dry meteorological cycles. 1967). Heterogeneity of environment within the lake is much more obvious in the deep lakes such as Malawi and Tanganyika.000 years ago. Changing lake levels. Actual rates of speciation are difficult to determine. Tanganyika most of the endemic cichlids live round the rocks (Poll. 1960. but these lakes are so large that they have persisted long enough for speciation to occur in them. capable of developing their own species of fish in isolation. so there is little seasonal change in size of the environment. Harding. Nabugabo. unless earth movements let the water out. among fishes small enough and specialized enough to be able to fulfil all their needs for the whole life cycle (including breeding) among the rocks. Within the Great Lakes conditions are much more stable than in rivers.but also provide cover. of algae with associated microfauna (Aufwuchs). Lake Lanao. the deep waters devoid of oxygen.

rocky cliff shores. The distribution of young cichlids (Tilapia and Haplochromis) along L. In Victoria Greenwood (1964) stressed that despite its vast size. among the tetragonopterid characoids and the plecostomine and pimelodid catfishes of South America. Rudolf). which have been subject to environmental selection for much longer. Among these riverine fishes adaptive radiations are shown between families or subfamilies rather than within genera. Victoria on the relative numbers of different TiZapia species (Welcome. The ability of non-cichlids of fluviatile origin to colonize lakes was discussed by Corbet (1960). The creation of large man-made lakes in recent years has provided unique opportunities to study changes in fish populations as a river becomes a lake.in both lakes the proportion of cichlids. It is clear. exposed sandy shorelines. this work indicates the interaction of many factors we have as yet only just started to measure. extensive speciation has occurred in the rivers within certain subfamilies. though here they may be more subject to bird predation. However. too. and the mormyrids. many of these noncichlids retain their old habit of running up streams from the lakes to spawn. governing the depth and the distance from the shore at which they live at various times of day. As Whitehead (1959) has remarked. providing the most riverine conditions within the lakes.O. southern and west Africa. the TiZapia are likely to escape some fish predation by being able to move into the warmer shallower water than the Haplochromis. and. staff. and of being more facultative in feeding habits than the more specialized lacustrine fishes. and the cyprinids have undergone geographic speciation in the numerous rivers of east. The effect of the recent dramatic rise in lake level in L. and in small lakes differences will be less permanent. for example.SPECIATION IN TROPICAL FRESHWATER FISHES 61 Heterogeneity of the environment is not so easy to detect in the shallow lakes. such as the changes between lake and river.R. Kariba studied by Jackson and J. The introduction of five exogenous TiZapia species into Victoria has provided a large-scale experiment on how the introduced species interact with the two indigenous endemic species of Tilapia and other fishes.F. who pointed out that rocky patches were of particular significance for this. the number of major habitats it offers is smallpapyrus-filled bays. 1966) also stresses the interplay of environmental and biotic changes in the lakes. characoids and mochocids of the Congo . migrating fishes must have the ability to tolerate a wide range of environmental conditions. plankton-feeding Tilapia. bottomed with thick organic mud. less commonly. CONDITIONS IN RIVERS Rivers are older than lakes and support older communities of fishes. and some more riverine species (notably the mormyrids from Volta) decreased in numbers rapidly (Mormyridae are absent from L. he found as many as 30 Haplochromis species in one major habitat-as well as various non-cichlid species (only one or two per genus). Petr (1967) compares changes in the Volta with those in L. Tanganyika and L. Myers (19604 stated that in . Victoria’s shoreline was studied by Welcomme (1964) who showed that the various species have different temperature and oxygen concentration tolerances. that predator pressures will also vary according to where the fishes are living. rose sharply.

inundating hundreds of square kilometres along their banks. and barriers between populations may be physical (such as speed of water flow or type of river bed). thousands of square kilometres being flooded each year. the various species being restricted to different stretches of the river in some groups of fish. due to the two peaks in rainfall as the sun passes overhead. and that such definitions are necessary before overlap between fish species can be understood. or one mainly due to rains in the tributary headwaters many hundreds of kilometres upstream. where it rains throughout the year. the more facultative ones and those able to withstand conditions in headwaters (which may be in rocky streams. Speciation in the rivers generally appears to be geographical in the first instance. both the actual size of the environment and the food supplies varying seasonally with the rains. South America and Asia. fisheries of Africa. This is still the case. Leopoldville) by . fish ecology has been studied at three widely separated points. with consequent overlapping of species. as in the Amazon. Marlier (1967) and others. such as many of the rock-dwelling plecostomines and other small catfishes appear to have very restricted ranges.62 R. Such alterations in ecological conditions allow microgeographical isolation within a river system. LOWE-MCCONNELL tropical continental streams the number of species is often much larger than in any known lake fauna. but alternate with lake-like stretches. Other species. generally with two periods of high and two of low water each year. Conditions in rivers fluctuate more than in the Great Lakes. Seasonality in the rivers increases with the distance from the equator. or biotic. land surface movements have led to the rejuvenation of rivers along their courses-fast flowing stretches are not restricted to headwaters as in most smaller temperate rivers. Between these two extremes. the Stanley Pool (near Kinshasa. As already mentioned the cichlids form only a relatively small element of the riverine faunas. cover extremely wide ranges from river system to river system (Central America to Argentine for example) . H. however. and narrow trophic specializations are the rule. Myers (1960~)pointed out that no attempt had yet been made to define the niches in tropical American rivers. though seasonal. The sheer size of these river systems is important in allowing many species to evolve within a river system. many of which have similar ecological requirements. 1967). chemical (such as the black and white water types of Amazonia). This seasonal variation is least marked in rivers in the great equatorial forest region. the large rivers running through forest and savanna which flood annually. species may develop in geographical isolation in tributary headwaters. Other fish species. Even here. This variation is on a far greater scale in the tropics than in temperate regions. I n the Congo. though the limnology of Amazonia is now much better known through the studies of Sioli (1964. the present-day movement across the Congo/Zambezi watershed has been studied by Bell-Cross (196%). and also in the Niger (the ‘black’and ‘white’ floodsof the Niger) and Zambezi. Fittkau (1967). the other to more local rains. there are seasonal changes in level. or large swampy areas on drainage divides as is often the case on these old land surfaces). The Amazonian environment is in many ways comparable with that of the Congo. until the rivers dry up to such an extent that the fish fauna is limited by the physiological requirements of dry season conditions. but the many changes of drainage and river capture have led to changes in fish distribution. carry some of the most productive.

many species spending some stage of their life history here. their main spawning and feeding grounds. remain accessible to them. though the extent and duration of flooding at these times varies very much from year to year-a factor of vital consequence for the fishes as this determines how long the flooded forest and marshy swamps. except locally and seasonally. either spawning in the rivers . Matthes has attempted to define the niches in the heavily forested river at Ikela and the more open waters of Lac Tumba. life cycles are short. where the predatory Seiymalmus were extremely abundant some years. but fishes have to be more eurythermic in the riverine than in the lacustrine environment.SPECIATION IN TROPICAL FRESHWATER FISHIS 63 Poll (1959). are very quickly reflected throughout the population . The breeding habits of the riverine fishes of South America fall into two groups: (1) the fecund ‘piracema’ fishes (the Amerindian name for the great fish runs up Brazilian rivers. 1963). Lac Tumba and a tributary river at Ikela in the heavily forested centra Congo basin by Matthes (1964). There is strong selection pressure for rapid growth and early maturity. and (2) the ‘small-brood’ fishes showing parental care and producing few young at a time. There is a close relationship between batch size. The picture emerging from these Congo studies is that there are considerable seasonal variations in physical and chemical conditions. also changing in time. 1964) which could be related to local spawning conditions. for comparable reasons populations are able to recover rapidly after severe depletion in bad years. and fishes are also more facultative feeders. with calm water. von Ihering. but that great trophic specialization is not possible where seasonal changes are great : the Niger fish fauna. for instance. amount of movement and food resources. Differences in water temperatures between the various biotopes present effective barriers. due to variations in spawning success. forest detritus is an important source of food.0 in places but rising in the rains). Matthes concludes that niches are very diverse. dense vegetation and rich in invertebrates as food. These forest waters are very acid (pH 4. which allow the seasonally fluctuating resources of the environment to be fully exploited. The ‘piracema’ fishes run up the rivers and streams. The peripheral marginal shallows. Stanleyville) by Gosse (1963. The heavy shading and shortage of mineral salts makes the waters poor in plankton. Rocky environments here are poor in number of species as they are too shaded to support an algal flora. many of the fishes have accessory respiratory organs. carry the richest ichthyofauna. with two periods of high and two of low water each year. not modifications of the alimentary canal as in many of the South American fishes which cease to feed in the dry season. much poorer than that of the Congo. such as the respiratory trees of clariids and labyrinthine organs of anabantids. where conditions are seasonal there are great variations in numbers. generally associated with the respiratory system. Forest debris more commonly fills the role of providing refuges for the lucifuge fishes. scarce in others (Lowe-McConnell. so it is no surprise that so many species can co-exist. particularly seasonally. The forest debris makes the water very &-oxygenated . Poll & Gosse. is particularly lacking in more specialized feeders. 1930). Such fluctuations in abundance of piscivorous fishes must greatly affect the co-existence of prey species. This was noticable in the Rupununi savanna area of Guyana. and fluctuations in numbers from year to year. and higher up the main river at Yangambi (near Kisangani.

1928). von Ihering. and since the rising water level often falls temporarily. or drift there.). The small characoids are rather generalized feeders (von Ihering. 1930a. to the bottom. the rains. showing parental care and producing few young at a time. there is strong selection pressure €or increased fecundity and for rapid development of the eggs and young. but other species appear to be territorial and they or their replacements were observed in the same places day after day as long as conditions stayed more or less the same. or most of. Miyadi. often on the caudal fin. LOWE-MCCONNELL or running out into the flooded savanna or forest to spawn (as in the ‘lateral migrations’ of West African fishes. This group includes most of the characoids and many large catfishes inSouth America (von Ihering. The ‘small-brood’ fishes. such as the predatory characoids Sewasalmus and Hoplias. Fryer & Whitehead. which are labelled by readily recognizable specific colour marks. and develop in the warm shallows. all sizes of fish being found together in one area. A few other species also show some parental care. as the water falls. which results from their reactions to the current. solitarily or in shoals etc. they appear to take whatever food is available at their particular position in the water. The young fishes grow fast in the streams and flood pools during the rains. living nearer the bottom or bank. and the young are soon free-swimming. laying all. all swimming against the current) line the edges of the rivers and around the numerous islands. b. and to the presence of numerous other species (cf.g. 1959). or rises so fast that they are swept away. itself determined by relationships with current. 1930. This may account for the females being larger than the males in most of this group of fishes. Many of these South American fishes have ingenious ways of guarding the eggs. Male Aspredo carry them attached to the belly. bottom and other species. Barros & Planet. 1936) . producing relatively large eggs. and some male loricarines have them attached to a special prolongation of the lower lip. These are very fecund fishes. in pools or amongst rocks where some food is available throughout the year. Greenwood. In these fishes the eggs generally hatch within two or three days (von Ihering. others. There is here selection for small families. as do cichlids and plecostomine algal-grazing catfishes. and the females are generally smaller than the males in these fishes. or drop downstream. Daget. H. deposit them in small holes in stream banks or below stones. or early in. obs. Conditions change rapidly. often start to spawn in the dry season before the rains. their eggs at one time. such as Hypostomus. Towards the end of the rains bands of small tetragonopterids of mixed species (a dozen or so species together. swiftly-flowing stretches becoming calm pools behind obstructions such as fallen trees or sandbanks emerging at the lower water levels. keeping out of the main current. the various species. e. the latter a predator living mainly in ponds. 1957). and some of the fishes move elsewhere. the number of eggs produced increasing with size of female. however.64 R. 1955. stranding eggs and young. Thus in . at the start of. and in Africa the cyprinids (such as Barbus and Labeo) as well as characoids such as Alestes and catfishes such as Clanks and Schilbe. so the communities are continually changing (pers. producing large numbers of eggs per female. These are mostly fishes which remain in one habitat. Their position in the water is determined by their behaviour. The community is in a continual state of flux as some species move upstream. Eggs are deposited in the flooded vegetation at the sides of the rivers. show characteristic differences in behaviour. 1960).

Some fishes make long migrations between feeding and spawning grounds . and down. of many species are well known in Africa. (representing at least three different evolutionary lines). Competition for food in these pools is reduced by reduction in appetites during the dry season. that there appears to be little room for food in the body cavity. Such migrations will take the fishes through many types of water. in contrast with stenophagous species. 1959). huge numbers of small fishes living together in the rapidly shrinking waters. Competition is also decreased by the cichlids and characoids being active by day. in which where they live is governed by the availability of a particular food supply.pugnacious fishes which appear to be territorial. 1964). Specializations to use various foods probably come into maximum effect at this time. fruit eaters. but the fish often take other foods-the predatory 5 . 1935). the catfishes and gymnotoids (in Africa the mormyrids) by night (Lowe-McConnell. when fat stores are laid down which help the fish to live through the dry season. with associated temperature and other changes. as the fishes are very difficult to catch when the water is high. unless eaten before then. e. The rains are the main feeding and growing time for all these fishes. among the cypriniformes this is much more marked among the characoids. 1952 showed for Alestes in West Africa). the different families of the larger characoids are basically predators. with their widely varying dental morphology. insectivores. Under riverine conditions the fishes have to be more facultative feeders. others (such as the characoid Prochilodm) have the ovaries so enlarged at this time. In the flood pools the growth of aquatic plants is most abundant during the rains. The biomass of prey species is at its greatest at the end of the rains. and longitudinal migrations. so these fishes must be eurythermic and able to withstand very different physical and chemical conditions. Many of the fishes in these South American waters use parts of the alimentary canal as accessory respiratory organs for aerial respiration in these stagnant pools (Carter. The small generalized and more basic tetragonopterids seem to be not very specialized in food habits. Elsewhere crevices in rocks are used. 1931. Riverine fishes do show adaptations to use different types of food. these crevices are of particular significance in providing hiding places above the de-oxygenated bottom waters. but some fishes become locked in the pools if the water falls too suddenly. changing their food with what is available at different times of year (as Daget. marking experiments have shown that Prochilodm for example move 1000 km up and down the Mio-Guassu river in Brazil at comparable times each year (Godoy. Thus in these waters the fishes have to be able to withstand a wide range in conditions from a rushing torrent at one time of year to de-oxygenated stagnant pools in the dry season. as well as lateral migrations. preyed on by large numbers of piscivorous fishes and birds. the main rivers. the holes and crevices in submerged tree debris criss-crossing the forested creeks become packed with fish. or even toothless mudsuckers.g. by gymnotoids such as Sterfiurchm.SPECIATION IN TROPICAL FRESHWATER FISHES 65 these fishes the food eaten depends on where they are living. but little is known about this. than amongst the toothless cyprinids. providing cover and shelter for diverse food organisms. where they have to stay until released by the next rains. itself governed by other factors. The development of this nocturnal fauna (the ‘lucifuge’ fishes of Matthes) is dependent on the fishes having places where they can hide by day. Many species move back to.

Fryer (1959b) discusses the non-diversity of niches in L. sand or from amongst rocks or vegetable debris. including a group of at least eight species that specialize in feeding on the eggs and larvae of other Haplochromis. . Myers (1960~)questions whether there really are more niches than in temperate zone rivers. it probably more frequently takes the form of a behavioural inhibition that limits feeding territory. the habitat. specializations to use a particular food have developed several times. proposed restricting the definition of niche to the ‘nutritional role of an animal in its ecosystem. NICHE NUMBER AND OVERLAP IN ECOLOGICAL REQUIREMENTS The number of habitat niches are enchanced by the complexities of the environment. flowers or aerial insects) dropping in from overhanging trees. that is in its relation to all the foods available to it’. and Greenwood (1964) the scarcity of major habitats in L. and each species present contributes to this complexity. In Victoria. and that a surprisingly high number are piscivorous-at least 50 species. kunnume than from certain other species-a warning of how misleading assessments of food preferences can be if based on a few specimens localized in time and space. Amongst the more specialized cichlids. are considered. particularly with seasonal changes. several species may share the same food source. which he doubts. Weatherley (1963) working on freshwater fishes. Victoria studied by Corbet (1961) tend to be more facultative feeders (a riverine characteristic retained in the lacustrine environment). Matthes (1964) pointed out that the long snout of the long-snouted mormyrid Gnuthonemus species found in the Congo is equally useful in probing insects out of mud. among the 120 species known in 1964 Greenwood estimated that the least specialized (most like the riverine ancestors) feeding on insect larvae and detritus make up 30% of the Haplochromis flock. where Greenwood found 30 Haplochromis species in a major habitat. especially when changes of niche with time. There is much overlap. Matthes (1964) on the other hand. and no two species are likely to overlap in all their living requirements throughout life. The non-cichlids of L. H. Fryer (1959~)found ten species of Mbuna in L. and he comments (p.66 R. changing the food eaten with the size of fish. and availability of food at different times. Nyasa sharing the Aufwuchs The niche in the Eltonian sense as defined by Uvardy (1959) is a functional entity within the biotic community which can be studied in a comparative way. considers that there are abundant niches in the riverine environment of the Congo. one on plants. particularly in the dry season. a definition which does not specify the factors or relations which compose it. though Weatherley did broaden his definition by pointing out that though competitive interaction may be of the direct kind for food. Nyasa. four species on algae. 41) that it can happen that large individuals of Mormym kannume feeding over a hard bottom at full moon can differ more in their food from other M. Victoria. Nevertheless there is a good deal of overlap in the requirements of different species. about 12 species feed on molluscs (extracting them from their shells in two quite different ways). when few food resources are available except bottom debris or exogenous forest products (fruits. which are presumably sucked from the mouth of the parent. thus for example Corbet found five mormyrid species feeding mainly on chironomid larvae (which show lunar emergence cycles). but such a restriction is not acceptable to all biologists (including myself). LOWE-MCCONNELL Serrmulmus for example may contain boluses of vegetable matter.

and distinctively marked fishes of different genera mimicing one another and living together. not just feeding territory. 1959). may limit numbers . and in South American waters three groups of armoured catfishes have evolved . Actual living space. leading to a fall in recruitment (Weatherley. 1960. also come to the conclusion that predation has had a greater role in South America. This sharing of food appears to contradict Gause’s principle (better called the exclusion principle. and that predator pressures have here a greater role in the evolution and control of the communities. except for the partially armoured Synodontis and a few small Congo species. and also if spawning is impaired through overcrowding.SPECIATION IN TROPICAL FRESHWATER FISHES 67 in one rocky shore locality. nocturnal fishes which hide in crevices by day may be territorial about their crevices. the ‘Utaka’ group. 1963:68) and suggests that food is not limiting numbers in these cases. may be the only ones to do so throughout the year). Fryer (1959) suggested that overlap was possible among many of the Mbuna because Aufwuchs was so abundant. Offence is matched with defence. amongst these fishes it is not only herbivores which share the same food (though they may do so more often. parasites. the counterplay of offence and defence producing the amazingly well-concealed and mimetic forms in such abundance in South America. Field work in South America after Africa gave me the impression that there were many more types of predators in the South American waters. other species may have chemical or electrical effects on their co-habiting species as yet unstudied. chemical and biotic factors governing Tilapia distribution in the lake studied by Welcomme (1966) under high lake conditions have already been mentioned. It is interesting that Kettlewell (1959) looking at South American insects after African ones. Nyasa Mbuna and Greenwood for Victoria Haplochromis. diseases or other density-dependent factors. Lowe (1953) found phytoplankton-feeding Tilapia in L. and in this lake 16 species of Haplochromis. Mayr (1963:80) noted that generalized herbivores are often exceptions to the exclusion principle. The South American fish fauna is also rich in mimics. Victoria when the lake was low (Beauchamp. Among fishes the effects of competition for food can be mitigated as fishes have such flexible growth rates. However. Lowe-McConnell. and the interacting physical. Slobodkin (1964) has shown experimentally with hydrids that the addition of a predator can help two species to co-exist. this may. The presence of so many predators in these tropical haters is likely to help the co-existence of prey species by keeping their numbers below the levels at which they would compete for food. 1963). such armoured fishes are noticably lacking in African waters. cryptic fishes which resemble leaves and twigs. Nyasa living in mixed shoals for much of the year (and two algal-eating Anubleps species were sometimes seen shoaling together in Guyana) . and (b) the huge seasonal fluctuations in abundance of prey species in rivers may allow predators more scope. (a) because the riverine communities have had longer in which to evolve. 1958. are predominently zooplankton-feeders (Iles. as they are seldom food-limited. Mayr. numbers being controlled by predators. reflect conditions in the older communities of the rivers compared with the lakes. depending on population density. however. Kettlewell . and apart from zooplankton-feeders. cichlid breeding males or couples become territorial. as was suggested by both Fryer for L. The availability of suitable spawning sites appeared to limit the numbers of indigenous Tilapia in L. 1963).

Annual Report for 1952). carpeting the swamp. Malawi have well-defined breeding seasons. buds and fruits of the waterlilies. saka breeds in deeper water during the rains (December to February). rich in micro-organisms. by slowing down production cycles. World-wide climatic changes associated with the Ice Ages are likely to have had more profound effects on the waters on Africa than of the interior of South America. sparnipinnis a sky-blue male. Differences in time and place of spawning are also reinforced by differences in male breeding colours. The role of predators in restricting or promoting speciation has been discussed since Worthington (1937) correlated the absence of species flocks from the Great . whereas at the lower elevations prevailing in much of the interior of South America (the Orinoco. leaving a semi-digested flock of plant material. Thus the exceptional productiveness of the Malagarazi waterlily swamps (Tanzania).68 R.H. here separation depended on specificity of displays and the unlearnt selective responsiveness of the female to them-habitat. Beauchamp (1958. but T . the nature of some of the rivers would be changed in wetter and drier cycles (as Jackson (1963) has suggested to explain the puzzling discontinuous distributions of some African fishes). 1953). T. the leaves. wet or dry cycles would merely mean more or less extensive flooding and increased or decreased interconnections between long-existing river systems. Ethological isolating mechanisms were found to be of major importance between four intimately sympatric poeciliid fishes from Guyana trenches studied in aquaria by Liley (1966) . a hypothesis discussed by Hickling (1961). but T . A NOTE ON ISOLATING MECHANISMS In addition to microgeographical and ecological barriers between fish populations in lakes and rivers already mentioned. barriers may be principally temporal or ethological. T . the small fish then forming the food for piscivorous fishes (E. because the general land surface of the African continent is higher. Some fishes may actually ameliorate the habitat for others. which is then used as food by two species of Tilapiu and two Haplochromis. 1964) has pointed out that since breakdown of plant matter in the bottom mud of tropical lakes is often much slower than through herbivores. The Tilapia of L. compared with less productive ones such as L. the removal of large numbers of herbivores (such as Tilapia and hippopotami) from a lake will itself decrease the productivity. The principle of using ecologically complementary species to ameliorate the environment for one another.F. Amazon. These feed on. saka breeds in shallow water before the rains (August to November). in the Malagarazi. Kyoga (Uganda). and partially digest. seasonal and mechanical isolating mechanisms were of minor importance.O.R. Paraguay systems). appeared to be mainly due to the presence of the characoids Alestes macrophthalrnus and Distichodussp. lidole at about the same time but in much deeper water or off open sandy beaches. where this is so. Zidole having black males. T .A. saka and T . LOW-MCCONNELL attributed this to a longer period of uninterrupted evolution in isolation in South America. sparnipinnis which most nearly resembles T . often with a white head (Lowe. and so increase production has long been used in ponds by Chinese fish culturists.

in the rivers and creeks). the riverine species being more facultative in their feeding and other ecological requirements. Fryer & Iles (1955) pointed out that there are many predators amongst the cichlids of L. but by encouraging specialization lakes may become ‘evolutionary traps’. Coulter envisaged speciation occurring here despite the presence of predators. CONCLUSIONS It seems clear. Speciation in lakes may be rapid. Jackson (1961~)suggested that spawning migrations up streams were devices to get the young away from predators. Tanganyika cichlid speciation appears to occur mainly in rocky areas. In L. such as rock patches. But spawning migrations are old features. a paper commented on fully by Fryer & Greenwood (1965). in the case of potadromous species. The riverine communities are older. Hydrocynus and the predatory cichlid Boulengerochromis was due to a number of special evasive mechanisms. though Jackson was right to stress the importance of cover for small fishes. I think. and the data could. 1966). In both lakes and rivers there is often much overlap in food eaten by co-habiting species. that diversity in tropical freshwater fishes is very marked both in the stable conditions of the lakes. Specializations to use different foods are more marked under lake conditions (though several species in a lake may develop the same specializations and be found co-habiting). Albert. Nyasa. such as new lakes or around rocks if light penetration allows algal growth. However it is now known from fossil evidence that both these genera occurred in L. and promoting speciation by enhancing the isolation of shelter areas. Some genetically plastic families. and the fishes living there will have had to pass through a series of environmental filters during their evolution. may best explain the absence of cichlid species flocks from these lakes. Edward and Nyasa. Coulter (1962:26) thought that the reason that speciation had proceeded in L. shelter in dense growths of weed or shelter in the rocky bottom (and. are particularly prone to speciate. Also. and also possibly the absence of a rocky shoreline providing cover in L. most marked of which are the regular daily migrations into deeper water. Lutes as recently as 12. predators thus exerting their influence in both ways simultaneously. and in the seasonally fluctuating unstable conditions in the rivers. be interpreted that their presence here may be both restricting speciation by preventing the colonization by cichlids of areas without shelter.000 years ago (Greenwood. some places.SPECIATION IN TROPICAL FRE~HWATER FISHES 69 Lakes Albert and Rudolf with the presence there of the large predatory fishes Lutes and Hydrocynus. Edward. two species are unlikely to share the niche all the time (at all times of year or . 1959. It seems clear that two or more species can share the same niche if factors other than food keep the numbers below the level of competition for food (competition for living space and the abundant predators being amongst such controlling factors). and it seems probable that the aquisition of a fully differentiated riverine fauna in Albert and Rudolf (see Fig. Tanganyika which has numerous endemic cichlid species. therefore. are particularly conducive to speciation. Tanganyika in the presence of Lutes. 2). such as the Cichlidae. but these appear to patrol the openwater areas. and their absence from Victoria. retained from riverine ancestry. and these genera are still living in L.

though resources in rivers may be very seasonal and among the most riverine species reduced family size is not shown. The ability of the environment to receive and make room for introduced species (as in the case of the L. herbivores in lakes and swamps (and possibly detritus feeders in forest rivers) have a particularly important r81e in ameliorating the environment for other species. which may also vary from place to place. LOW-MCCONNELL throughout life). In rivers the extent and duration of the rains. there is no shortage of potential causes for a tropical increase in species diversity. changing all the time. and coexistence. operating in different ways on the various stages of the life cycles of co-habiting species. which vary from year to year. These fish studies make it very clear that the relative numbers of different species. studied by Welcomme has provided clearest evidence of changing lake level on the relative numbers of the various Tilapia species . as MacArthur adds. the diversity of tropical fish faunas fulfils many of these expectations. However. populations recovering rapidly after depletion in bad years (e. high-lighted predator pressures in these South American waters. elsewhere the failure of our attemps to introduce Tilapia for fish culture in the flood-fallow sugarcane fields of Guyana. In the complicated interaction of environmental and biotic factors. and the abundance of plant food throughout the year allows the presence. In lakes the situation amongst the L. degree of specialization and reduced family size . lack of seasonality in resources. the biotic factors are . and their spawning and survival success. but the dynamic equilibrium is such that the system is elastic enough to take in new species.g. favouring one species in one place. These tropical fish communities are extremely dynamic. and no reason to expect this increase to be controlled by any single potential cause. Rukwa). However. another in another part of their range. depends on small environmental changes interacting with predation and other biotic pressures.and more openwaterliving species depended on the varying lake levels. Victoria Tilapia. MacArthur (1965) maintained that the number of species within a habitat can be expected to increase with productivity (sometimes). 1953)had suggested that the relative numbers of inshore.70 R. The resilience of the environment may also be increased by the numbers of species present.this was no surprise as studies on Nyasa Tilapia (Lowe. Small oscillations in selection pressures probably have a vital importance in permitting coexistence. with their interconnecting food webs and other interrelationships. as demonstrated in the Malagarazi swamps and as put forward by Beauchamp . The relatively short life cycles in these tropical fishes also aid the resilience. with structural complexity of the habitat. So it seems clear that it is the small oscillations in conditions interacting with predation and other biotic pressures that give variations in relative numbers of species from year to year. as after drought in L. or in all sections of the habitat. determine how many fishes survive in the dry season retreats. The actual currying capacity of the environment may also be increased by the numbers of species present. aided by differential predation by man. Victoria Tilapia) is a product of this resilience-there is really no problem of ‘empty niches’ waiting to be filled in the well-established communities. H. their coexistence. The success of such introductions also points to the power of geographical barriers in keeping species apart until moved by man. are released from them. of many herbivorous species.

(6) In lakes the production of numbers of young at frequent intervals keeps numbers of fishes more or less constant throughout the year. amongst rocks) produce small batches of young. There is a close relationship between seasonality of spawning. restricted to small habitats (e. whereas in the tropics the creatures play a much more active part in creating the environment. particularly herbivores. moulding. and indiscriminate (density-independent) catastrophic mortalities are likely to occur. several species living in one locality having specializations to use one particular food resource. amount of movement and food resources. helps to control numbers and to permit the coexistence of prey species. so selection is here a more creative. particularly in South America. process. and the presence of numerous piscivorous species. selection will be directed to physiological adaptations to the controlling factor (such as cold or drought) and for increased fecundity. SUMMARY (1) Among freshwater fishes greater diversity is shown. this stability has allowed great trophic specialization amongst the fishes. (8) The actual carrying capacity of the environment may be increased by the presence of certain species. These fish studies bear out Dobzansky’s (1950) suggestion that the nature of selection differs in tropical regions from high latitudes. and there is much overlap in food eaten by different species. and it is perhaps not too much of an exaggeration to say that at high latitudes the environment creates or selects the creatures.SPECIATION IN TROPICAL FRESHWATER FISHES 71 relatively more important than they are in temperate regions. But under most tropical conditions the individuals that survive and reproduce will be those that are most attuned to the complex interrelationships of the organic community. or even beget. in both lakes and rivers. (7) Competition for living space. many fine examples of which are known amongst these tropical fishes. at low than at high latitudes. (9) The relative numbers of individuals of different species depends largely on small oscillations in environmental conditions interacting with predation and other biotic pressures. however. (4)The riverine fishes have to be more facultative feeders than lake fishes since conditions change seasonally. ( 5 ) The Cichlidae were the fishes best able to take advantage of lacustrine conditions when the Great Lakes of Africa were formed. . (2) This high tropical diversity is shown both in the very stable environment of the Great Lakes of Africa. but there is also considerable overlap. seasonally fluctuating conditions. increasing in seasonality with distance from the equator. both within taxa and within communities. complexity. (3) In lakes.g. Where the physical environment is harsh. Complexity appears to allow. some riverine fishes. capable of producing subtleties such as concealment behaviour and mimicry. being preadapted by their ability to breed in still water and to make adaptive radiations to use different foods. In rivers seasonal spawning and high fecundity leads to great fluctuations in numbers of fish. in the rivers. and in the very unstable. batch size.

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