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 Nyasas rocky shores (Fryer, 1959). This
appears to contradict Gauses 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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52

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 nichessmaller 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

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R. H. LOW-MCCONNELL

found all the time. In South America the equatorial country of Brazil has 1383 species
belonging to some 46 families (Fowler, 1954), compared with 339 species in Argentina
(Ringuelet, Aramburu & Aramburu, 1967), and 456 species in Central America
(Miller, 1966). Myers (1947) pointed out that the Amazon system alone has well over
1000 species, a number which might be almost doubled when exploration is complete;
Eigenmann (1912:41) took 70 to 90 species in one haul of a 150-foot seine in the
Essequibo, and 60 species in two or three hours from a small brook in this area. In
equatorial Africa two areas of the Congo support 408 species (Poll & Gosse, 1963)
representing 24 families; the Upper Niger has at least 134 species (Daget, 1954), the
Zambezi 155 species, belonging to fewer families (Bell-Cross, 1965a), whereas the
number falls to 92 species south of the Zambezi (Jubb, 1967) and over half of these
belong to one family, the Cyprinidae. In tropical Asia, Thailand has 546 species
belonging to 49 families (Smith, 1945). For comparison the whole of Europe has
192 species representing 25 families (Ladiges & Vogt, 1965), and the Great Lakes
of North America have 172 species of 29 families (Hubbs & Lagler, 1947).
TYPES OF WATERS

Waters are predominently riverine in South America and Asia, though huge areas
alongside the rivers are flooded in wet seasons, giving temporarily lacustrine conditions,
and many of the South American rivers are very convoluted, forming ox-bow lakes
and pools where fishes get cut off from the main river until released by floods in
subsequent years, making a heterogeneous environment. Many of the South American
river systems interconnect (Orinoco to Amazon for example) and some fish species
are very widely distributed, from Central America to the Argentine.
In Africa there are four main river systems, the Niger and Nile, which have many
species in common, the Congo, which also has many of the same species, and the
Zambezi, as well as other eastward- or westward-flowing rivers and inland drainage
areas, such as Lake Chad. The equatorial Congo shows much less seasonal variation
than the Niger, Nile and Zambezi, which are at higher latitudes; the Congo, like
most of the Amazon, flows through high forest, many tributaries having a low pH
and forest debris causing de-oxygenated conditions.
Africa has also the Great Lakes of east and central Africa, formed only a few million
years ago and offering new opportunities to the fishes which could take advantage of
the lacustrine conditions. Conditions are much more stable in these lakes than in the
rivers, though Lake Malawi (= Nyasa) which is well south of the equator shows
greater seasonal changes than the equatorial L. Victoria, Large endemic fish faunas
(particularly of cichlid fishes) have evolved in these lakes, notably in Tanganyika,
Malawi and Victoria; differentiation (as measured by the numbers of endemic genera
present) is greatest in Tanganyika (which has 38 endemic genera), then Malawi (20
endemic genera), and least in Victoria (only four endemic genera) (Table 1); this
probably reflects the comparative ages of these three lakes. These lakes are very large,
Victoria, 26,000 square miles, the size of Northern Ireland, but only 90 m deep (the
depth of Windermere), lies like a saucer between the Rift Valleys, Tanganyika (over
a mile deep) and Malawi (over 400 miles long) lie in the rift valley.

SPECIATION
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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. 2. The South American freshwater fish fauna is mainly of ostariophysian fishes.
T h e Ostariophysi as a group dominate these tropical freshwater fish faunas. I n South
America these are represented by two main groups, the catfishes (Siluriformes) and
the Characoidei suborder of the Cypriniformes. T h e South American catfishes belong
to about 13 families, some of which are scaleless skinfish, ranging in size from small
parasitic species to some of the largest freshwater fishes known, others (belonging
to three distinct families) are heavily armoured. These catfishes make up 43% of
Table 1. The composition of tropical freshwater fish faunas
African rivers

S. America
Brazil

1383
c.46

No. spp.
No. families

Mormyriformes No.

594
43
553
40
0
0
81
6
0

Siluriformes
No.
% of fauna
No.
Characoidei

70
Cyprinidae

No.

Cichlidae

No.

YO

Niger

Congo

Zambezi

Asia
Thailand

134
26

408f
24

155
15

546
49

37

112

27

34
22
18
11
47
30
23
15
16

27

22

73

16

18

24

52
13
32

18
10

7
75

19
14

18

99

18
0

0
209
38
0

0
0

10

African Great Lakes

I

No. spp.
No. families
Cichlidae No. spp.
Yo of fauna
% Endemism
Cichlids
Non-cichlids
No. of genera
Cichlids
Non-cichlids
No. endemic genera
Cichlids
Non-cichlids

Albert

Rudolf

Victoria

Tanganyika

Malawi
(Nyasa)

46
14

37
14

177+
12

214
14

245f
7

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. 2 mainly from the following sources:- S.America: Brazil, Fowler, 1954. Asiu:
Thailand, Smith 1945. African t i w e n : Niger, Daget 1954; Congo, Poll & Gosse 1963; Zambezi, BellCross 1959. Africun Great Lakes: Albert, Greenwood 1966; Rudolf, Worthington & Ricardo 1936;
Victoria, Greenwood 1966; Tanganyika, Poll 1953, 1956; Malawi (Nyasa), Jackson et ul. 1963.

R. H. LOWE-MCCONNELL

56

the Brazilian fauna, and within the group it is the loricariid catfishes, particularly
the plecostomine group of rather small armoured fishes, primarily algal-grazing,
often found in rocky streambeds, that have speciated most (135 species in Brazil).
Another closely-related armoured group, the loricarines, 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 proportional composition of the freshwater fish faunas of S. Amm*cu (Brazil),
Asia (Thailand), Africun r i w m and Aficun Great Lakes. Numbers of species expressed as
percentage of known fauna (1% = 3.6"). Note relative importance of Cichlidae in lakes and
of Cypriniform fishes (Characoidei and Cyprinidae) in rivers.

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

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R. H. LOWE-MCCONNELL

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

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in number, mark returns also indicated prolonged association or re-encounter between

adult fish, a male and a female in one case, two females in another case. Albrecht
( 1963) studying Haplochromis (wingati and burtoni) in aquaria found that the females
became territorial when the young reached the free-swimming stage, and that the
young learnt the mothers territory and did not leave this home area until brooding had
ceased. Such behaviour would help populations to develop in semi-isolation.
The genera of cichlids which could adaptively radiate to use different foods have
speciated most; the outstanding example is the genus Haplochromis of which there
are 112 species in L. Malawi (Jackson, Iles, Harding & Fryer, 1963) and well over
120 species in Victoria (Greenwood, 1964; his more recent estimates put it at over
170 species), compared with Tilapia which are restricted to algae or plant food, and
of which the largest species flock is of four species in L. Malawi (Lowe, 1953).
The cichlidsforte is specialization to use different foods; Greenwood has pointed
out that they have the morphological and physiological apparatus to do this easily
(and so have the characoids, but the cyprinids, which are toothless, lack this ability
for trophic specialization). However, as Matthes (1964) has pointed out, fishes that
are too specialized in their food requirements could not survive in rivers, where
conditions change seasonally, which may explain the paucity of the cichlid fauna in
rivers compared with lakes.
CONDITIONS IN GREAT LAKES COMPARED WITH RIVERS

Lakes are younger than rivers and are colonized from them. 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.
Among the freshwater fishes the cichlids were pre-eminently able to take these
opportunities. This is particularly clear in the case of Lake Tanganyika, where the
formation of the lake cut across the Malagarazi river drainage flowing westward to
the Congo basin. This left a distributional gap for most of the non-cichlids, Congo
species (such as Polypterus congim and P . ornat+innis, Mormyrus longirostris,
Distichodus maculatus, Tetraodon mbu and Labeo weeksii) being found in the Malagarazi but not in the lake, whereas the cichlids have speciated most abundantly in the
intervening lake (Poll, 1951). Some non-cichlids have also colonized these Great Lakes,
but seem to have taken longer to become adapted to lake conditions, many still having
to migrate up inflowing streams to spawn. Tanganyika lake has eight endemic genera
of non-cichlids (compared with 30 endemic cichlid genera), and species flocks of zooplankton-feeding clupeids (two species), predatory Lutes and Luciolates (five species),
Chrysichthys catfish (six species) and Mastacembelus (12 species) (Poll, 1953; Matthes,
1962). Lake Malawi has no endemic genera of non-cichlids, but there is a species
flock of offshore-living large Dinotopterus catfish (Jackson, 1959; Greenwood, 1961).
In the Philippines Lake Lanao has a cyprinid species flock studied by Myers (1960).
Myers commenting on this Lanao flock says that lack of competition allows this
explosive adaptive radiation. But what factors push the fishes into these adaptive
radiations 1 Presumably intraspecifx competition at first. Cichlid numbers build up
very rapidly after a new lake is stocked with them, as the changes in fish fauna after

60

R. H. LOWE-MCCONNELL

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

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Heterogeneity of the environment is not so easy to detect in the shallow lakes, and
in small lakes differences will be less permanent. In Victoria Greenwood (1964)
stressed that despite its vast size, the number of major habitats it offers is smallpapyrus-filled bays, bottomed with thick organic mud, exposed sandy shorelines, and,
less commonly, rocky cliff shores; 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).
The distribution of young cichlids (Tilapia and Haplochromis) along L. Victorias
shoreline was studied by Welcomme (1964) who showed that the various species
have different temperature and oxygen concentration tolerances, governing the depth
and the distance from the shore at which they live at various times of day; this
work indicates the interaction of many factors we have as yet only just started to
measure. It is clear, too, that predator pressures will also vary according to where the
fishes are living; the TiZapia are likely to escape some fish predation by being able to
move into the warmer shallower water than the Haplochromis, though here they may
be more subject to bird predation.
The effect of the recent dramatic rise in lake level in L. Victoria on the relative
numbers of different TiZapia species (Welcome, 1966) also stresses the interplay of
environmental and biotic changes in the lakes. 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.
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. Petr (1967)
compares changes in the Volta with those in L. Kariba studied by Jackson and J.F.R.O.
staff;in both lakes the proportion of cichlids, plankton-feeding Tilapia, rose sharply,
and some more riverine species (notably the mormyrids from Volta) decreased in
numbers rapidly (Mormyridae are absent from L. Tanganyika and L. Rudolf).
The ability of non-cichlids of fluviatile origin to colonize lakes was discussed by
Corbet (1960), who pointed out that rocky patches were of particular significance for
this, providing the most riverine conditions within the lakes; many of these noncichlids retain their old habit of running up streams from the lakes to spawn, and of
being more facultative in feeding habits than the more specialized lacustrine fishes.
As Whitehead (1959) has remarked, migrating fishes must have the ability to tolerate
a wide range of environmental conditions, such as the changes between lake and river.
CONDITIONS IN RIVERS

Rivers are older than lakes and support older communities of fishes, 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. However, extensive speciation has occurred in the rivers within certain
subfamilies, for example, among the tetragonopterid characoids and the plecostomine
and pimelodid catfishes of South America, and the mormyrids, characoids and
mochocids of the Congo ; and the cyprinids have undergone geographic speciation in
the numerous rivers of east, southern and west Africa. Myers (19604 stated that in

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tropical continental streams the number of species is often much larger than in any
known lake fauna, and narrow trophic specializations are the rule. As already mentioned
the cichlids form only a relatively small element of the riverine faunas.
Speciation in the rivers generally appears to be geographical in the first instance,
but the many changes of drainage and river capture have led to changes in fish
distribution, with consequent overlapping of species, many of which have similar
ecological requirements. The sheer size of these river systems is important in allowing
many species to evolve within a river system; species may develop in geographical
isolation in tributary headwaters, and barriers between populations may be physical
(such as speed of water flow or type of river bed), chemical (such as the black and
white water types of Amazonia), 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, but alternate with lake-like stretches.
Such alterations in ecological conditions allow microgeographical isolation within a
river system, the various species being restricted to different stretches of the river
in some groups of fish. Other fish species, the more facultative ones and those able to
withstand conditions in headwaters (which may be in rocky streams, or large swampy
areas on drainage divides as is often the case on these old land surfaces), cover extremely wide ranges from river system to river system (Central America to Argentine
for example) ; the present-day movement across the Congo/Zambezi watershed has
been studied by Bell-Cross (196%). Other species, such as many of the rock-dwelling
plecostomines and other small catfishes appear to have very restricted ranges.
Conditions in rivers fluctuate more than in the Great Lakes, both the actual size
of the environment and the food supplies varying seasonally with the rains. This
variation is on a far greater scale in the tropics than in temperate regions, thousands
of square kilometres being flooded each year. This seasonal variation is least marked
in rivers in the great equatorial forest region, where it rains throughout the year.
Even here, however, there are seasonal changes in level, generally with two periods
of high and two of low water each year, due to the two peaks in rainfall as the sun
passes overhead, or one mainly due to rains in the tributary headwaters many hundreds of kilometres upstream, the other to more local rains, as in the Amazon, and
also in the Niger (the blackand white floodsof the Niger) and Zambezi. Seasonality
in the rivers increases with the distance from the equator, until the rivers dry up to
such an extent that the fish fauna is limited by the physiological requirements of
dry season conditions. Between these two extremes, the large rivers running through
forest and savanna which flood annually, inundating hundreds of square kilometres
along their banks, carry some of the most productive, though seasonal, fisheries of
Africa, South America and Asia.
Myers (1960~)pointed out that no attempt had yet been made to define the niches
in tropical American rivers, and that such definitions are necessary before overlap
between fish species can be understood. This is still the case, though the limnology
of Amazonia is now much better known through the studies of Sioli (1964, 1967),
Fittkau (1967), Marlier (1967) and others. The Amazonian environment is in many
ways comparable with that of the Congo. I n the Congo, fish ecology has been studied
at three widely separated points, the Stanley Pool (near Kinshasa, Leopoldville) by

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Poll (1959), Lac Tumba and a tributary river at Ikela in the heavily forested centra
Congo basin by Matthes (1964), and higher up the main river at Yangambi (near
Kisangani, Stanleyville) by Gosse (1963; Poll & Gosse, 1963). Matthes has attempted
to define the niches in the heavily forested river at Ikela and the more open waters of
Lac Tumba.
The picture emerging from these Congo studies is that there are considerable
seasonal variations in physical and chemical conditions, with two periods of high and
two of low water each year, 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, their main spawning
and feeding grounds, remain accessible to them. These forest waters are very acid
(pH 4.0 in places but rising in the rains). The forest debris makes the water very
&-oxygenated ; many of the fishes have accessory respiratory organs, generally
associated with the respiratory system, such as the respiratory trees of clariids and
labyrinthine organs of anabantids, not modifications of the alimentary canal as in
many of the South American fishes which cease to feed in the dry season. The heavy
shading and shortage of mineral salts makes the waters poor in plankton, except
locally and seasonally; forest detritus is an important source of food. Rocky environments here are poor in number of species as they are too shaded to support an algal
flora. Forest debris more commonly fills the role of providing refuges for the lucifuge
fishes. The peripheral marginal shallows, with calm water, dense vegetation and rich
in invertebrates as food, carry the richest ichthyofauna, many species spending some
stage of their life history here. Differences in water temperatures between the various
biotopes present effective barriers, but fishes have to be more eurythermic in the
riverine than in the lacustrine environment. Matthes concludes that niches are very
diverse, also changing in time, particularly seasonally, so it is no surprise that so
many species can co-exist, but that great trophic specialization is not possible where
seasonal changes are great : the Niger fish fauna, for instance, much poorer than that
of the Congo, is particularly lacking in more specialized feeders.
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, von Ihering, 1930), and (2) the small-brood fishes showing parental
care and producing few young at a time. There is a close relationship between batch
size, amount of movement and food resources; where conditions are seasonal there
are great variations in numbers, which allow the seasonally fluctuating resources of
the environment to be fully exploited, and fishes are also more facultative feeders.
There is strong selection pressure for rapid growth and early maturity, life cycles are
short, and fluctuations in numbers from year to year, due to variations in spawning
success, are very quickly reflected throughout the population ; for comparable reasons
populations are able to recover rapidly after severe depletion in bad years. This was
noticable in the Rupununi savanna area of Guyana, where the predatory Seiymalmus
were extremely abundant some years, scarce in others (Lowe-McConnell, 1964)
which could be related to local spawning conditions. Such fluctuations in abundance
of piscivorous fishes must greatly affect the co-existence of prey species.
The piracema fishes run up the rivers and streams, either spawning in the rivers

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

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

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R. H. LOWE-MCCONNELL

Serrmulmus for example may contain boluses of vegetable matter. There is much
overlap, particularly in the dry season, when few food resources are available except
bottom debris or exogenous forest products (fruits, flowers or aerial insects) dropping
in from overhanging trees.
NICHE NUMBER AND OVERLAP IN ECOLOGICAL REQUIREMENTS

The number of habitat niches are enchanced by the complexities of the environment, and each species present contributes to this complexity. Nevertheless there is
a good deal of overlap in the requirements of different species. Myers (1960~)questions
whether there really are more niches than in temperate zone rivers, which he doubts;
Fryer (1959b) discusses the non-diversity of niches in L. Nyasa, and Greenwood
(1964) the scarcity of major habitats in L. Victoria. Matthes (1964) on the other hand,
considers that there are abundant niches in the riverine environment of the Congo,
especially when changes of niche with time, particularly with seasonal changes, are
considered.
In Victoria, where Greenwood found 30 Haplochromis species in a major habitat,
specializations to use a particular food have developed several times; 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, about 12 species feed on molluscs (extracting them from their shells
in two quite different ways), four species on algae, one on plants, and that a surprisingly
high number are piscivorous-at least 50 species, including a group of at least eight
species that specialize in feeding on the eggs and larvae of other Haplochromis,
which are presumably sucked from the mouth of the parent. The non-cichlids of
L. Victoria studied by Corbet (1961) tend to be more facultative feeders (a riverine
characteristic retained in the lacustrine environment), changing the food eaten with
the size of fish, the habitat, and availability of food at different times; thus for example
Corbet found five mormyrid species feeding mainly on chironomid larvae (which
show lunar emergence cycles), and he comments (p. 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. 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. 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, sand or from amongst rocks or vegetable debris.
Amongst the more specialized cichlids, several species may share the same food
source; Fryer (1959~)found ten species of Mbuna in L. 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, a definition which does not specify the factors
or relations which compose it, and no two species are likely to overlap in all their living requirements
throughout life. Weatherley (1963) working on freshwater fishes, proposed restricting the definition of
niche to the nutritional role of an animal in its ecosystem, that is in its relation to all the foods available
to it, but such a restriction is not acceptable to all biologists (including myself), though Weatherley
did broaden his definition by pointing out that though competitive interaction may be of the direct
kind for food, it probably more frequently takes the form of a behavioural inhibition that limits feeding
territory.

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

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R.H. LOW-MCCONNELL

attributed this to a longer period of uninterrupted evolution in isolation in South

America. 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, because the general land surface of the African continent is higher; where
this is so, 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), whereas at the lower elevations prevailing in much of the
interior of South America (the Orinoco, Amazon, Paraguay systems), wet or dry
cycles would merely mean more or less extensive flooding and increased or decreased
interconnections between long-existing river systems.
Some fishes may actually ameliorate the habitat for others. Thus the exceptional
productiveness of the Malagarazi waterlily swamps (Tanzania), compared with
less productive ones such as L. Kyoga (Uganda), appeared to be mainly due to the
presence of the characoids Alestes macrophthalrnus and Distichodussp. in the Malagarazi.
These feed on, and partially digest, the leaves, buds and fruits of the waterlilies,
leaving a semi-digested flock of plant material, rich in micro-organisms, carpeting the
swamp, which is then used as food by two species of Tilapiu and two Haplochromis,
the small fish then forming the food for piscivorous fishes (E.A.F.R.O. Annual
Report for 1952). The principle of using ecologically complementary species to
ameliorate the environment for one another, and so increase production has long
been used in ponds by Chinese fish culturists.
Beauchamp (1958, 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
removal of large numbers of herbivores (such as Tilapia and hippopotami) from a lake
will itself decrease the productivity, by slowing down production cycles, a hypothesis
discussed by Hickling (1961).
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. The Tilapia of L. Malawi have well-defined breeding seasons; T . saka
breeds in shallow water before the rains (August to November), T . lidole at about the
same time but in much deeper water or off open sandy beaches, but T . sparnipinnis
which most nearly resembles T . saka breeds in deeper water during the rains (December to February). Differences in time and place of spawning are also reinforced by
differences in male breeding colours, T. saka and T . Zidole having black males, but
T . sparnipinnis a sky-blue male, often with a white head (Lowe, 1953).
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) ; here separation depended on specificity of displays and the unlearnt
selective responsiveness of the female to them-habitat, seasonal and mechanical
isolating mechanisms were of minor importance.
The role of predators in restricting or promoting speciation has been discussed
since Worthington (1937) correlated the absence of species flocks from the Great

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Lakes Albert and Rudolf with the presence there of the large predatory fishes Lutes
and Hydrocynus, and their absence from Victoria, Edward and Nyasa. However it is
now known from fossil evidence that both these genera occurred in L. Edward,
Lutes as recently as 12,000 years ago (Greenwood, 1959, 1966), and these genera are
still living in L. Tanganyika which has numerous endemic cichlid species. Fryer &
Iles (1955) pointed out that there are many predators amongst the cichlids of L.
Nyasa, and it seems probable that the aquisition of a fully differentiated riverine
fauna in Albert and Rudolf (see Fig. 2), and also possibly the absence of a rocky
shoreline providing cover in L. Albert, may best explain the absence of cichlid
species flocks from these lakes. Jackson (1961~)suggested that spawning migrations
up streams were devices to get the young away from predators, a paper commented
on fully by Fryer & Greenwood (1965). But spawning migrations are old features,
retained from riverine ancestry, though Jackson was right to stress the importance
of cover for small fishes. In L. Tanganyika cichlid speciation appears to occur mainly
in rocky areas; Coulter (1962:26) thought that the reason that speciation had proceeded in L. Tanganyika in the presence of Lutes, Hydrocynus and the predatory cichlid
Boulengerochromis was due to a number of special evasive mechanisms, most marked
of which are the regular daily migrations into deeper water, shelter in dense growths
of weed or shelter in the rocky bottom (and, in the case of potadromous species, in the
rivers and creeks). Coulter envisaged speciation occurring here despite the presence
of predators, but these appear to patrol the openwater areas, and the data could, I
think, be interpreted that their presence here may be both restricting speciation by
preventing the colonization by cichlids of areas without shelter, and promoting
speciation by enhancing the isolation of shelter areas, such as rock patches, predators
thus exerting their influence in both ways simultaneously.
CONCLUSIONS

It seems clear, therefore, that diversity in tropical freshwater fishes is very marked
both in the stable conditions of the lakes, and in the seasonally fluctuating unstable
conditions in the rivers. 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), the riverine species being more facultative in their feeding
and other ecological requirements.
Some genetically plastic families, such as the Cichlidae, are particularly prone to
speciate; some places, such as new lakes or around rocks if light penetration allows
algal growth, are particularly conducive to speciation. Speciation in lakes may be
rapid, but by encouraging specialization lakes may become evolutionary traps.
The riverine communities are older, and the fishes living there will have had to pass
through a series of environmental filters during their evolution.
In both lakes and rivers there is often much overlap in food eaten by co-habiting
species. 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).
Also, two species are unlikely to share the niche all the time (at all times of year or

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R. H. LOW-MCCONNELL

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

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relatively more important than they are in temperate regions. These fish studies bear
out Dobzanskys (1950) suggestion that the nature of selection differs in tropical regions
from high latitudes. Where the physical environment is harsh, and indiscriminate
(density-independent) catastrophic mortalities are likely to occur, selection will be
directed to physiological adaptations to the controlling factor (such as cold or
drought) and for increased fecundity. 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, so selection is here a more
creative, moulding, process, capable of producing subtleties such as concealment
behaviour and mimicry, many fine examples of which are known amongst these tropical
fishes, particularly in South America.
Complexity appears to allow, or even beget, complexity, and it is perhaps not too
much of an exaggeration to say that at high latitudes the environment creates or selects
the creatures, whereas in the tropics the creatures play a much more active part in
creating the environment.
SUMMARY

(1) Among freshwater fishes greater diversity is shown, both within taxa and within
communities, at low than at high latitudes.
(2) This high tropical diversity is shown both in the very stable environment of the
Great Lakes of Africa, and in the very unstable, seasonally fluctuating conditions, in
the rivers.
(3) In lakes, this stability has allowed great trophic specialization amongst the fishes,
but there is also considerable overlap, several species living in one locality having
specializations to use one particular food resource.
(4)The riverine fishes have to be more facultative feeders than lake fishes since
conditions change seasonally, increasing in seasonality with distance from the equator,
and there is much overlap in food eaten by different species.
( 5 ) The Cichlidae were the fishes best able to take advantage of lacustrine conditions
when the Great Lakes of Africa were formed, being preadapted by their ability to
breed in still water and to make adaptive radiations to use different foods.
(6) In lakes the production of numbers of young at frequent intervals keeps numbers
of fishes more or less constant throughout the year. In rivers seasonal spawning and
high fecundity leads to great fluctuations in numbers of fish; some riverine fishes,
however, restricted to small habitats (e.g. amongst rocks) produce small batches of
young. There is a close relationship between seasonality of spawning, batch size,
amount of movement and food resources.
(7) Competition for living space, and the presence of numerous piscivorous species,
in both lakes and rivers, helps to control numbers and to permit the coexistence of
prey species.
(8) The actual carrying capacity of the environment may be increased by the presence of certain species, particularly herbivores.
(9) The relative numbers of individuals of different species depends largely on small
oscillations in environmental conditions interacting with predation and other biotic
pressures.

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R. H. LOWE-MCCONNELL

(10) The relatively short life cycles of the fishes aid resilience, populations increasing
rapidly again after depletion in bad years.
(11) Tropical fish communities are extremely dynamic, changing all the time; the
nature of selection differs in tropical regions from high latitudes, biotic factors being
more important than they are in temperate regions.
ACKNOWLEDGEMENTS

I am most grateful to my colleagues Dr P. H. Greenwood and Dr E. Trewavas

for reading and commenting on the manuscript and draft of this paper, and to them
and Dr M. E. Brown (Varley) for many stimulating discussions on these problems.
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