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14 Cichlid Fish Diversity and Speciation

Article · December 2006

DOI: 10.1201/9781420009538.ch14

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14 Cichlid Fish Diversity

and Speciation
J. R. Stauffer, Jr. and K. E. Black
School of Forest Resources, Penn State University,
University Park, Pennsylvania, USA

M. Geerts
Swalmen, The Netherlands

A. F. Konings
Cichlid Press, El Paso, Texas, USA

K. R. McKaye
Appalachian Laboratory, University of Maryland System,
Frostburg, Maryland, USA

CONTENTS

14.1 Introduction ....................................................................................................................... 214

14.2 Cichlid Phylogeny............................................................................................................. 214
14.2.1 Higher Level Taxonomic Relationships of Cichlids.......................................... 214
14.2.2 Intrafamilial Relationships of Cichlids .............................................................. 216
14.3 Cichlid Distribution........................................................................................................... 217
14.4 Cichlid Diversity and Speciation ...................................................................................... 217
14.5 Cichlid Adaptive Radiation............................................................................................... 218
14.5.1 Feeding Adaptations ........................................................................................... 218
14.5.2 Breeding Adaptations ......................................................................................... 219
14.6 Future Directions............................................................................................................... 220
References .................................................................................................................................... 220

ABSTRACT
Cichlids are one of the most species rich families of vertebrates, with conservative estimates citing
more than 2,000 extant species. Although native to tropical areas of the world, with the exception
of Australia, some 70–80% of cichlids are found in Africa, with the greatest diversity found in the
Great Lakes (lakes Victoria, Tanzania and Malawi). Their highly integrated pharyngeal jaw appa-
ratus permits cichlids to transport and process food, thus enabling the oral jaws to develop special-
isations for acquiring a variety of food items. This distinct feature has allowed cichlids to achieve
great trophic diversity, which in turn has lead to great species diversity. The high species diversity of
this vertebrate family is not accompanied by an appropriately high genetic diversity. The combination

213
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214 Reconstructing the Tree of Life

of rapid radiation of the group and relatively low genetic diversity has confounded attempts to
diagnose species and discern phylogenetic relationships. Behavioural traits appear to be important
characters for diagnosing many cichlid species.

14.1 INTRODUCTION
Cichlids (Cichlidae) are a species rich group of fish from the lowland tropics1 and are indigenous
primarily to the fresh waters of Africa, South America and Central America, with one species
extending its range to the Rio Grande River in southern North America. In addition, cichlids are
found in Madagascar, the Levant and India and have also been introduced into nearly all tropical
and subtropical regions of the world, either through escapes from aquaculture or ornamental fish
operations or intentionally to provide sport fishing opportunities or to control exotic plants2. They
have established breeding populations in warm waters of industrial effluents in temperate areas3
and have been introduced into some marine environments4. Numerous investigators5–14 have focused
on cichlid fishes for their ecological, evolutionary and behavioural research.
Without doubt, the cichlids’ explosive speciation, unique feeding specialisations, diverse mating
systems and great importance as a protein source in tropical countries have been factors stimulating
research interest in this group15–18. In fact, Greenwood19 referred to the cichlid species flocks as
“evolutionary microcosms repeating on a small and appreciable scale the patterns and mechanisms
of vertebrate evolution”. Many of these research efforts, however, have been slowed and results
often confused because of the uncertain systematic status of some of the cichlids being studied20,21.
The conservative bauplan of cichlids1 and relatively low genetic divergence is coupled with a great
morphological diversity that makes it difficult to diagnose species using morphological criteria
alone (Figure 14.1). Systematic confusion exists within Cichlidae and also within and between its
higher taxonomic ranks such as suborders. Such relationships are currently being debated. The
reasons why cichlids have managed to speciate so successfully, often within a restricted geographic
range such as the Great Lakes of Africa, have also been under investigation14,15,22.

14.2 CICHLID PHYLOGENY

14.2.1 HIGHER LEVEL TAXONOMIC RELATIONSHIPS OF CICHLIDS
Morphological studies have provided insight into the phylogenetic position of Cichlidae. The
resultant relationships, however, differ considerably from that of molecular investigations. Kaufman
and Liem23 (see also Stiassny and Jensen24) grouped cichlid fishes (Acanthopterygii: Perciformes)
with Embiotocidae, Labridae and Pomacentridae in the suborder Labroidei on the basis of the
following three pharyngeal characters: fusion of the fifth ceratobranchial bones into one unit; contact
of the upper pharyngeal jaws and the basicranium; and lack of a dorsal subdivision of the sphincter
oesophagi muscle. As it is now recognised, this suborder includes some 1,800 species and represents
some 5–10% of all extant fishes24. Müller25 was the first to group the above families together, and
morphological data support the view that Labroidei has phyletic integrity24 based on the following
synapomorphies: bladelike keel on the lower pharyngeal bone and the change in insertion of at
least part of the transversus ventralis onto the keel; division of the transversus dorsalis anterior
muscle26; bony facets of the third pharyngobranchials of the upper pharyngeal jaws are exposed;
ventrally projecting rounded form of the neurocranial apophysis; and no subdivision of the sphincter
oesophagi muscle. The cichlids appear to be the sister group of all the other labroids1.
The monophyly of Labroidei is, however, confused, because many homoplasies exist in the
morphological data sets24,27. Moreover, studies of nuclear DNA suggest that Labroidei are not
monophyletic, and that embiotocids and pomacentrids are more closely related to more basal
perciforms28. This view is corroborated by Sparks and Smith29 who believe that the sister group to
the cichlids “might comprise a large assemblage of diverse perciform lineages, including but
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Cichlid Fish Diversity and Speciation 215

FIGURE 14.1 (A colour version of this figure follows page 240) Cichlid fishes. Cichlids have a conservative
bauplan, and specialised attributes, such as hypertrophied lips are the result of parallel evolution, thus
making species and higher level diagnoses difficult. (a) Amphilophus sp. ‘fatlip’ in Lake Xiloa, Nicaragua;
(b) Placidochromis milomo at Nkhomo Reef, Lake Malawi, Malawi; (c) Lobochilotes labiatus at Nkondwe
Island, Lake Tanganyika, Tanzania. (Photos reproduced with permission from A.F. Konings.)

presumably not limited to the other ‘labroid’ lineages, sparids, anabantids-nandids, haemulids,
percids, moronids, and kyphosids”. Westneat and Alfaro27 reported maximum RAG2 DNA sequence
divergence between wrasses and outgroups ranging as high as 23% between parrotfishes and some
of the cichlids they examined. Nevertheless, they supported the inclusion of the parrotfish as a
subgroup of Labridae; thus, it is doubtful if these families belong to the same suborder.
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216 Reconstructing the Tree of Life

Boulenger30 first speculated that the cichlids form a natural group within the perciform Acantho-
pterygians. Stiassny1 recognised Cichlidae as a monophyletic group based on five apomorphic characters:

• Loss of a structural association between parts A2 and A10 of the adductor mandibulae
muscle and the attachment of a large ventral section of A2 onto the posterior border of
the ascending process of the anguloarticular
• An extensive cartilaginous cap on the front margin of the second epibranchial bones
• An expanded head of the fourth epibranchial bones
• Presence of characteristically shaped and distributed microbranchiospines on the gill
arches
• The subdivision of the traversus dorsalis anterior muscle into three distinct parts as
described by Liem and Greenwood26

The monophyly of Cichlidae is further supported by the morphology of otoliths and configu-
ration of the digestive tracts. Gaemers31, based on the structural configuration of the sagitta, also
hypothesised that the cichlids are monophyletic. The sagitta is usually the largest otolith in most
teleosts, including cichlids. The sagitta of cichlids is strong, thick, with a more or less oval, short,
elliptical to rounded pentagonal shape31. If the pseudocolliculum in the sagitta of cichlids is a
synapomorphic character, it supports other evidence of monophyly of the family26,32. Finally, three
structural attributes of a cichlid’s digestive tract support cichlid monophyly: the stomach’s extend-
ible blind pouch; the left hand exit to the anterior intestine; and the position of the first intestinal
loop on the left side33.

14.2.2 INTRAFAMILIAL RELATIONSHIPS OF CICHLIDS

Many subfamilial names have been used to indicate groupings within Cichlidae, but do not neces-
sarily represent hierarchical patterns34. Etroplinae (that is Etroplus and Paretroplus) has been
proposed to be the sister group of all other cichlids29. Following this separation, Ptychochrominae
were considered to be the sister group to the remaining cichlids. What then is left has been divided
further into two purportedly monophyletic groups, the African Pseudocrenilabrinae and the New World
Cichlinae. It should be noted that Schliewen and Stiassny35 used the term Haplotilapiines to refer
to non-Heterochromis cichlids from Africa and the Levant.
Within the family, a Congolese genus, Heterochromis, was proposed to be the most primitive
of the African cichlids36. Stiassny37 diagnosed Heterochromis to be the sister group of all the other
cichlids, based on the presence of a single ligament attaching the lower pharyngeal bone to the
cartilaginous fourth basibranchial. The monophyly of Neotropical cichlids was questioned by
Cichocki38, who postulated that Cichla, a Neotropical genus, was grouped with the cichlids of
Africa. This was debated by Stiassny1, who favoured a Cichla-crenicichline clade within the
Neotropics. Stiassny1 postulated a close etropline relationship with Heterochromis, while
Kullander34 regarded Heterochromis as the sister group of the majority of Neotropical cichlids.
Despite this relationship, Kullander34 emphasised that the dichotomy of Old and New World cichlids
was well supported by the following character states:

• Short anterior arm of epibranchial, which is a reduction that occurs in several lineages,
including the Etroplinae and Astatotilapia, and which is reversed in groups such as
Cichlasomatinae
• Interdigitating suture connecting the vomerine shaft and the parasphenoid bar (requires
independent development in Ptychochromis and reversal in Biotoecus, Dicrossus and
Nannacara)
• Presence of an anterior palatoethmoid ligament, which occurs in all Neotropical cichlids
and Heterochromis, but in no other Old World cichlids
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Cichlid Fish Diversity and Speciation 217

Retroculus is regarded as the earliest diverging lineage of Neotropical cichlids and the sister group
of Cichla-Crenicichla, which is placed in a new subfamily, Cichlinae34. Kullander34 recognises the
following Neotropical subfamilies: Astronotinae (Astronotus, Chaetobranchus), Geophaginae (for
example, Geophagus, Apistogramma) and Cichlasomatinae (Cichlasomines, Heroines Acaronia).
Mitochondrial DNA supports the monophyly of Cichlidae, but differs in its interpretation of
intrafamilial relationships39. In particular, Heterochromis is sister to the remaining African cichlids,
and Retroculus is the most basal taxon of the Neotropical cichlids. Farias et al.40 believed that
Astronotus, Cichla and Retroculus formed three independent basal lineages, even though one of
their trees favoured Astronotus as the sister group to Cichla. Therefore, Astronotinae, sensu
Kullander, is not accepted by molecular biologists, whereas Chaetobranchus and Chaetobranchopsis
are grouped together as Chaetobranchines. Farias et al.40 remove Crenicichla and Teleocichla from
Cichlinae (sensu Kullander) and transfer that group to Geophaginae. In the molecular phylogeny,
Acaronia is no longer regarded as the sister group of the Heroini/Cichlasomatini group40.

14.3 CICHLID DISTRIBUTION

Apart from Australia and Antarctica, cichlids occur on all the major fragments of the former
supercontinent Gondwana. The extant distribution of cichlids and phylogenetic relationships suggest
that the family was well established before the separation of Gondwanaland. Thus, the family
should be at least 160 million years old41,42. This view, however, is not supported by studies based
on the fossil record, and the time of origin of this taxon is still hotly debated. For example, many
authors believe that the cichlids are not old enough to have been present on that supercontinent43.
India and Madagascar drifted away during the Late Jurassic (160 million years before present,
mybp) and not during the Middle Cretaceous, as Van Couvering44 apparently believed. Molecular
evidence presented by Vences et al.45 suggests that the extant distribution pattern of the cichlids
came into being well after Gondwana fell apart. In that respect, it should be mentioned that Murray46
believed that a Gondwanan origin is not needed to explain the present distribution of cichlids.
The oldest known fossil cichlids, the species of the genus Mahengochromis, have an age of
46.3 mybp, which means that they date from the Middle Eocene46. Fossil cichlids from the Oligocene
of Africa have been described by Van Couvering44, and because of the highly specialised dental
features of these fishes, Stiassny1 concluded that the origin of the family predates the earliest fossils
considerably and that the cichlids probably arose in the Early Cretaceous, some 135 mybp.
This view is incompatible with the fossil record. In a recent paper, Arratia et al.47 have
emphasised, once more, that thus far no perciform fishes have been found that are older than the
latest Cretaceous (85–65 mybp). That age differs dramatically from that postulated by molecular
studies (Salmoniformes and Gadiformes versus Perciformes divergence of some 285 mybp). In this
respect, it should be remembered that the cichlids are regarded as advanced perciforms, which
means that they should be much younger than the primitive forms that have been found in the Late
Cretaceous sediments. Cichocki38 postulated that the cichlids with the most plesiomorphic charac-
ters are found in Madagascar and India, which was endorsed by Kullander34, discussed by Stiassny1,
and supported by mtDNA analysis by Farias et al.39.

14.4 CICHLID DIVERSITY AND SPECIATION

Between 70 and 80% of all cichlid fishes are native to African freshwaters, of which the majority
are part of the major lacustrine radiations; for example, Lake Victoria (c. 500 species), Lake
Tanganyika (c. 200 species) Lake Malawi (c. 850 species)15,48. The remainder occur in the fresh-
waters of the New World, with the exception of a single genus, Etroplus, which is found in the
coastal waters of Southern India and Sri Lanka; five genera, which are found on Madagascar; the
genus Tristramella, which is endemic to the Levant; and one genus, Iranocichla, which is endemic
to Southern Iran.
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218 Reconstructing the Tree of Life

By far the greatest radiation of cichlids is found in the Great Lakes of Africa, with Lake Malawi
alone having as many as 850 species49. The phylogenetic diversity ranges from the single invasion
of Lake Malawi, which resulted in the endemism of all but a few species, to multiple invasions in
Lake Tanganyika, which resulted in the presence of 12 different tribes50. The rich fauna of these
lakes is primarily attributable to the explosive adaptive radiation and speciation51–53 of the haplo-
chromines sensu lato (see Schliewen and Stiassny35). The driving mechanism for these speciation
events is unknown. The two most widely proposed methods are allopatric speciation5,15,54–57 and
intrinsic isolating mechanisms14,22,58–64. Furthermore, biologists generally agree that female mate
choice can act as a strong driving force in runaway speciation where the average female preference for
a specific male trait differs between two allopatric populations65–69. Thus, behavioural traits are
important tools for the diagnosis of these African cichlids, primarily because behavioural traits
played a very important role in and facilitated the rapid radiation of these fishes, which may not
always be accompanied by discernable morphological changes70.
Certainly, there are fewer species of cichlids in South and Central America than in the Old
World. Greenwood19 noted that the focus on African cichlids has distracted attention from the South
American Cichlidae, a fact which he regarded as ‘sad but understandable’. Nevertheless, the
neotropical cichlid fauna is varied and diverse1, comprising some 50 genera and 450 species34, with
new species still being discovered71. In the last decade, on average more than 20 species of cichlids
have been formally described each year. The existence of literally hundreds of species awaiting
description is likely to continue this trend.

14.5 CICHLID ADAPTIVE RADIATION

The cichlids of the Great Lakes of Africa have undergone one of the most rapid radiations of any
known vertebrate group47,72–74. This rapid speciation rate, however, is not correlated with a high
genetic diversity74, as discussed earlier. Conversely, the neotropical cichlids, especially the geoph-
agines, have a significantly higher rate of genetic divergence than their African counterparts39,75,
but this does not seem to be expressed in the more diversified group. This makes the taxonomy
and systematics of the group challenging. The high genetic diversity of the Neotropical cichlids,
when compared to the ones from Africa, is somewhat surprising considering the greater species
diversity of Old World cichlids. A number of the following adaptations may be associated with
cichlid speciation.

14.5.1 FEEDING ADAPTATIONS

High trophic diversity of the Old World cichlids results in the consumption of virtually every food
type available in the environment76,77. The successful radiation of cichlids in the rift valley (for
example, in Lake Malawi and Lake Tanganyika) is purported to be a result of cichlids’ differential
ability to acquire food78. In cichlids, mouth structure, dentition, gill raker number79 and jaw structure
vary tremendously, and this variation in structure seems to be tied to a variety of feeding
techniques15,80. Documented feeding strategies of Lake Malawi fishes, for example, illustrate numer-
ous feeding specialisations including death feigning81, paedophagy82,83, lepidiophagy84, cleaning85
and scraping and raking of an algal, diatomaceous and detrital biolayer (aufwuchs) from the rock
surfaces5. Phenotypes, such as reverse counter shading, are associated with such bizarre strategies
as hunting upside down86.
Primarily through the work of Liem and his coworkers26,76,77,87 it became evident that the trophic
diversification of cichlids does not require major structural adaptations. Liem72 attributes the great
colonising success of cichlids to their possession of a highly integrated pharyngeal jaw apparatus.
Thus, the pharynx has been a particular focus of modification in Labroidei, including the cichlids1.
Liem72 further states that this specialised innovation allows cichlids to transport and prepare food,
enabling the premaxillary and mandibular jaws to develop specialisations for collecting diverse
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Cichlid Fish Diversity and Speciation 219

food items. It is this distinct feature of cichlids that has permitted them to dominate colonisation
of many habitats and adopt feeding opportunities available in lacustrine environments72,76.
Phenotypic plasticity is defined as the possible environmental modification of the phenotype88.
The degree of phenotypic plasticity that cichlids exhibit is congruent with the ability of cichlids
to take advantage of many habitats and feeding opportunities. Despite the morphological plasticity
observed in other fishes89–91, the morphology of African cichlids initially was thought to be rigid92;
however, much morphological variability has been observed93–98. In particular, the observed
phenotypic plasticity in some instances has involved the pharyngeal jaw apparatus. For example,
the phenotypic diversity of the pharyngeal jaws of the New World Herichthys minckleyi99,100 has
been documented extensively. Furthermore, differences in bone structure of the lower pharyngeal
jaw of Astatoreochromis alluaudi resulted from different diets101–103. In addition to these obser-
vations, Meyer94 and Wimberger97,98 have experimentally demonstrated the effects of diet on
plasticity of head morphology in New World cichlids, but Meyer94 hypothesised that the plasticity
of mouth brooding Old World cichlids may not be as pronounced due to constraints on jaw
morphology for mouth brooding. Stauffer and Van Snik Gray104 effected significant differences
in head morphology of Lake Malawi rock-dwelling cichlids by experimentally manipulating diets.
The magnitude of plasticity in these mouth brooding Lake Malawi cichlids, however, was not
as pronounced as that observed for the New World substrate brooder Herichthys cyanoguttatum.
They did postulate, however, that phenotypic plasticity might have contributed to the extensive
trophic radiation and subsequent explosive speciation observed in Old World haplochromine
cichlids. Lewontin105 postulated that colonising species, possessing a high degree of phenotypic
plasticity, may have a selective advantage because of their ability to exploit additional resources
in differing environments.

14.5.2 BREEDING ADAPTATIONS

Cichlids have also a number of reproductive strategies associated with their speciation. Barlow65,66
and Keenleyside16,17 give an excellent review of reproductive strategies and parental care in cichlid
fishes. The breeding tactics of cichlids can be separated into two major groups: substrate brooders
and mouth brooders65,106–109. Substrate brooding cichlids are generally monogamous and exhibit
biparental care of the eggs and brood, although some species are polygynous110. Substrate brooding
appears to be the plesiomorphic character state. The primitive cichlids of Asia and Madagascar
(that is Etroplinae and Ptychochrominae), most of the New World species, Old World Tilapia spp.,
and selected Old World pseudocrenilabrines are substrate brooders109,111–113.
Most mouth brooders are polygamous, although monogamy is found in a few species107,114.
Some New World cichlid mouth brooders begin as substrate brooders, but then gather either the
eggs (ovophilous mouth brooders) or the hatched fry (larvophilous mouth brooders) into their
mouths, including some Aequidens, Geophagus, Gymnogeophagus and Satanoperca115,116. Some
Chromidotilapia from Africa employ a similar delayed mouth brooding strategy117,118, while male Chro-
midotilapia guentheri brood the eggs and larvae. Saratherodon galilaeus exhibits an intermediate form
of biparental care, where both the male and female gather the eggs into their mouths after fertilisation
and then separate65.
The vast majority of African mouth brooders are polygamous. The rock dwelling cichlids of
lakes Malawi and Tanganyika defend territories and attract females to spawn in either rock crevices,
small caves, on the rock surface, or in the water column above rocks. Males of these fishes
presumably rely on their brilliant and diverse colour patterns to attract females70. In addition, many
of the Lake Malawi sand dwelling cichlids construct a wide variety of bower forms in leks with a
hundred to many thousand individuals119–123. These bowers can be broadly divided into 10 types121
and range in size from giant craters three metres in diameter124 to small depressions in the sand.
The females move over the arena and lay eggs with several males. The entire egg laying process
takes 25–60 minutes from the time a female lays her first egg125 until she leaves the arena with
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220 Reconstructing the Tree of Life

eggs in her mouth. Genetic studies of paternity for several Lake Malawi species show that females
mate with as many as six males per brood126.
The process of female choice is complex. Female Otopharynx c.f. argyrosoma selectively chose
males that occupy bowers in the centre of the arena121, while female Mehenga conophoros and
Lethrinops c.f. parvidens chose males that build the largest bowers122,125,127. If the rapid radiation
of the Lake Malawi cichlid flock was accelerated by sexual selection, the observed differences
in behaviour might be the best way to distinguish between sibling species that differ little in
morphology71.

14.6 FUTURE DIRECTIONS

Barlow66 dedicated an entire chapter in his excellent book, The Cichlid Fishes: Nature’s Grand
Experiment in Evolution, to the problems associated with the Earth’s growing population. Whilst
he discussed the risks of burgeoning human populations and the associated pattern of diminishing
resources on the Earth’s biota, he focused on cichlids in particular. Certainly, one of the detrimental
aspects of human populations is the decrease in genetic diversity connected to the increase in the
rate of extinction of natural populations. Probably the most drastic incident of mass extinction in
our lifetimes occurred with the destruction of the cichlid species flock in Lake Victoria that was
associated with the introduction of the Nile Perch128,129. The relatively small and colourful cichlids
endemic to the lake were suddenly confronted with a huge predator, which, a few decades after its
introduction, had severely decimated the cichlids and established itself as the new food source for
the local populace. In addition to the loss of cichlids due to introductions, the introduction of
cichlids has had detrimental impacts on native fish faunas throughout the world2,3,130.
The high morphological diversity of the cichlids coupled with their conservative bauplan and
the relative low genetic divergence of the Old World cichlids makes it difficult to diagnose species
using morphological criteria alone. The extremely recent diversification of several cichlid lineages
presents serious challenges for the use of genetic techniques for alpha and beta taxonomy. Initially,
the attempts to use allozymes to resolve phylogenetic relationships were limited131. Kocher et al.132
have discovered a large number of polymorphisms in the tilapia genome, and Arnegard et al.133
and Markert et al.134 have used microsatellites to study population structure of several Lake Malawi
rock dwelling cichlids. The use of amplified fragment length polymorphisms by Albertson et al.135
was promising for the determination of supraspecific relationships and species diagnoses. As stated
previously, behavioural traits are important characters for diagnosing many of these cichlids and
determining phylogenetic relationships70. Thus, Stauffer and McKaye71 recommended that a com-
bination of morphological, genetic and behavioural data be used to diagnose these species and
determine phylogenetic relationships.

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