Local adaptations in populations of rock-dwelling haplochromines (Pisces:Cichlidae) from southern Lake Victoria

The radiation of cichlid fishes in the African great lakes is often described as adaptive, because, at a superficial level, cichlid fishes seem adapted to the ecological niches they occupy. However, adaptedness has rarely been studied. We''ve investigated to what extent island populations of three species of the rock-dwelling genus Neochromis, endemic to Lake Victoria, are adapted anatomically to exploit locally abundant resources. Specifically, we asked whether different resource environments were reflected in differences in the feeding apparatus, both within species and between species. In populations of two specialized biters, the algae scrapers N. rufocaudalis and N. omnicaeruleus, the biting force of the lower jaw increased with increasing amount of items that require biting in the diet. N. greenwoodi is a less specialized biter; we found differences between two populations in the hyoid position and in the premaxilla that enhance suction feeding. These adaptations were related to the amount of items requiring suction. Comparing across three sympatric pairs of species, in each case different diets were reflected in differences in anatomy.     

Experimental evidence for adaptive phenotypic plasticity in a rock-dwelling cichlid fish from Lake Victoria

We demonstrate adaptive phenotypic plasticity (PP) in Neochromis greenwoodi , a rock-dwelling haplochromine cichlid from Lake Victoria. Adaptive phenotypic plasticity will enhance the chance of survival and reproduction in changing or novel environments. For rock-dwelling cichlids it may contribute to the success of settlement after migration between rocky outcrops or islands, since newly colonized patches of rock may offer different food regimes and different competitors. In our experiments, we simulated such situations by raising fry on different diets and with different competitors. We chose diet treatments in such a way that we could predict the direction of anatomical changes from functional demands. We expected an indirect effect of interspecific competition: selectively removing one prey-type by the competing species would leave N. greenwoodi with the other. We found the predicted phenotypic responses to food in trophic traits of N. greenwoodi . However, in the current experimental set-up we did not find a differentiating effect of species of competitor. We discuss the possible role of PP in allopatric and sympatric speciation.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 185−192.     

A revision of the genus Labidochromis (Teleostei: Cichlidae) from Lake Malawi

The diagnosis of the genus Labidochromis is revised to include species with bicuspid outer teeth. Sixteen species, of which 13 are new, are described and illustrated and the validity of five others is discussed. The interrelationships between members of the genus and the relationship of the genus to other genera are discussed. No clearly defined sister group to the genus has been found either within or outside the Lake Malawi cichlid species flock. A key based on the coloration of known species is included.     

Midwater spawning inHaplochromis chrysonotus (Boulenger) (Teleostei: Cichlidae) in Lake Malawi

Synopsis Haplochromis chrysonotus, a semipelagic shoaling cichlid from Lake Malawi, has been observed spawning in open water up to ten metres above the substratum. It is suggested that open water spawning without the establishment of a substratum-based territory eliminates competition with other species for breeding space and that it may be a stage in the evolution of a totally pelagic mode of life.     

Foraging behavior of the scale-eater Plecodus straeleni (Cichlidae,Teleostei) in Lake Tanganyika,Africa

Synopsis Sixty-nine individuals of Plecodus straeleni were followed for 1 h each in the field with the aid of SCUBA, and time budgets, hunting techniques and prey selection were investigated in relation to sex and body size. The time of cruising in midwater and on the substrate amounted to 3/4 of the total time. The rest of the time was mainly spent on five hunting techniques named pursuing, waiting, mingling, aiming and stealthy approaching. Pursuing (following a flying prey at high speed) was frequently used by adults, especially males, mainly to attack the spiny eel Afromastacembelus moorii. Waiting (keeping motionless on the substrate, waiting for a known prey) was used by some adult females when they tried to steal eggs of the mouthbrooder Cyathopharynx furcifer on the bower and by adult males when they targeted an eel having hidden under a rock. Mingling (mixing in a school of prey to attack school members) was a favorite tactic of subadults to attack plankton-feeders. Aiming (directing the head to a target fish for a moment) commonly occurred when both adults and subadults attacked solitary fishes. In stealthy approaching, the scale-eater approached an unwary prey from behind or sideway. Attacks by these hunting techniques amounted to 97% of the total attacks, which were made on 38 cichlid species and 7 non-cichlid species. Hunting techniques and prey preference varied not only with sex and size but even among consexuals of similar sizes. A number of individuals successively attacked only one or a few prey species in 1 h. Food specialization among individuals was attributed to their learning of the behavior and life style of preferred prey species.     

Grasping ecological opportunities: not one but five paedophagous species of Haplochromis (Teleostei: Cichlidae) in the Lake Edward system

Hydrobiologia - The Lake Edward system in Eastern Central Africa, including Lakes Edward and George and their associated rivers, is home to a species assemblage of endemic haplochromine cichlids....     

The feeding habits of a group of herbivorous rock-dwelling cichlid fishes (Cichlidae: Perciformes) from Lake Malawi,Africa

Synopsis The feeding; habits of a group of tropical herbivorous rock-dwelling cichlid fishes from Lake Malawi, Africa, are investigated using stomach content analyses. The various species fed selectively on the periphyton of the rocky shores. Blue-green alga of the genus Calothrix was the most common item ingested by the group. Diatoms (Chrysophyta) also were abundant food items. Discriminant analysis showed that dietary items were good variables to identify species. Interspecific dietary differences showed a continuum from those species feeding primarily on Calothrix to those feeding primarily on diatoms. Algal resources exhibit distinct patterns of spatial variation. Diet was correlated with foraging behavior and trophic morphology. Interspecific differences in diet could possibly facilitate ecological coexistence among various species. Such coexistence would contribute to the maintenance of the high diversity fish faunas characteristic of the Great Rift Lakes of Africa.     

Homing and home-site fidelity in rock-dwelling cichlids (Pisces: Teleostei) of Lake Malawi,Africa

Synopsis Existence of home site fidelity and homing ability was established for Pseudotropheus aurora. Territorial males stayed up to 1 1/2 years within the same territories. Females showed long-term fidelity to certain places. Transplantation experiments revealed that P. aurora males home over distances of up to at least 2500 m but would not cross deep water. The territorial males of five other sympartic species returned from 1000 m to their territories. The mechanisms and functions of homing in cichlids are discussed.     

A sympatric sibling species complex of Petrotilapia Trewavas from Lake Malawi analysed by enzyme electrophoresis (Pisces: Cichlidae)

The protein electromorphs of three taxa of Petrotilapia , referred to in earlier works as sibling species, were examined by starch-gel electrophoresis. The results indicated that, for the three taxa, no alternative fixed alleles occurred for any of the 25 loci (enzymes) examined. However, heterogeneous gene frequencies were indicated at seven polymorphic loci. This suggests that either the taxa are isolated 'sibling' species which recently diverged, or that they are 'incipient' species with minimal gene flow between morphs. The genetic and field data lend support to the hypothesis that sympatric splitting of morphs could be important in the explosive radiation of the cichlidae.     

Notograptidae, sister to Acanthoplesiops Regan (Teleostei: Plesiopidae: Acanthoclininae), with comments on biogeography, diet and morphological convergence with Congrogadinae (Teleostei: Pseudochromidae)

The Notograptidae contains one genus, Notograptus Günther, and five nominal species from northern Australia and southern New Guinea. Morphological evidence places Notograptus among acanthoclinine plesiopids (continuous free margin of lower lip; head naked; dorsal and anal fins with many spines and few segmented rays; no extensor proprius; reduced number of caudal-fin rays) and supports a sister relationship with Acanthoplesiops (symphyseal flap on lower lip; reduced hypural 5; reduced hypurapophysis). This hypothesis resolves the relationships within Acanthoplesiops , clarifying the polarity of autogenous middle radials of dorsal- and anal-fin pterygiophores. The proposed relationships among acanthoclinines are: Acanthoclinus ( Belonepterygion ( Beliops ( Notograptus ( Acanthoplesiops hiatti ( A. indicus ( A. psilogaster ( A. echinatus ))))))). The distribution of Notograptus compliments that of its proposed sister clade in that Acanthoplesiops is unknown from northern Australia or southern New Guinea. There are repeated geographical patterns among several groups suggesting that Australia is a basal area to a broader Indo-Pacific region. Similarity between the Congrogadinae (Pseudochromidae) and Notograptus has long been noted, both having a loosely connected suspensorium and elongate body which were mistakenly considered indicators of relationship; we add reduced branchial arches, straight, tube-like gut and highly expandable anus. We examine these similarities as an indication of a shared specialized feeding habit. Notograptus is an alpheid shrimp predator, able to swallow its large prey whole. Most species of congrogadines eat whole, large crustaceans. This is probably an example of convergent adaptation to a particular selective regime.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 179–205.     

The evolution of the fishery of Oreochromis niloticus (Pisces: Cichlidae) in Lake Victoria

The Lake Victoria ecosystem has experienced changes associated with fishing levels, a rise in lake level in the 1960s, fish introductions, and human activities in the drainage basin. Following the fish introductions of the 1950s and early 1960s, Oreochromis niloticus has become the most abundant and commercially important species among the tilapiines, and the only species which has managed to co-exist with the Nile perch in Lakes Victoria and Kyoga. There is, however, little published information on the biology and ecology of the specie in the new habitats. It has therefore been found necessary to initiate studies on the characteristics of O. niloticus in Lake Victoria.     

Morphological analyses of the neurocranium of a group of rock-dwelling cichlid fishes (Cichlidae: Perciformes) from Lake Malawi,Africa

Variation in the size and the shape of neurocrania of a group of rock-dwelling cichlid fishes (mbuna; from Lake Malawi, Africa was analysed using principal components and regression analyses on a set of 24 neurocranial measurements of 86 individuals from 11 species. The results reveal effective morphological discrimination between species. Most of the structural variation between species demonstrates positive allometry and represents size-dependent species differences. Thus, morphological differentiation in these fish may be largely the result of heterochronic changes in relative growth rates. The vomerine region, however, exhibited shape differentiation. Distinct groupings, based on vomer shape, are identified. Foraging behaviour and diet are found to be correlated with this region of the neurocrania. The correlation of the shape of the vomer and fine-scale resource partitioning suggests that this structure may be important in the evolutionary success of the group. Also, most structural shape variation in neurocranial morphology is associated with structural variation in buccal jaws.     

Comparative ecology of Serranochromis species (Teleostei: Cichlidae) in the Upper Zambezi River floodplain

The Upper Zambezi River drainage of Zambia contains six Serrunochromis species belonging to the largemouth, piscivorous subgenus Serranochromis and three smallmouth, invertebrate-feeding species of the subgenus Sargochromis. A survey of the Barotse floodplain and associateddrainages revealed that two species, S. (S.) longimanus (Boulenger) and S. (S.) thumbergi (Castelnau), were uncommon on the floodplain. S. thumbergi appeared to be more abundant in the Kafue, Chobe, and Kabompo rivers than the Upper Zambezi proper. Adults of the three largest species (i.e. S. (S) robustus (Günther), S. (S.) altus (Winemiller & Kelso-Winemiller), and S. (Sar.) giurdi (Pellegrin)) inhabit the main channel of the Zambezi River during low water, whereas most adults of the other four species (i.e., S. (S.) macrocephalus (Boulenger), S. (S.)ungusticeps (Boulenger), S. (Sar.) codringtoni (Boulenger), and S. (Sar.) carlottae (Boulenger)) were collected from lagoons, canals and backwaters. Comparisons of population size structure and gonad condition during the falling water (May-August) and low water (September- December) seasons indicated that the seven common Serranochromis initiate spawning near the end of the low water period. Examination of gonad condition during the pre-spawning period indicated that Serrunochromis attain maturity at the following sizes: S. robustus, males (275-300 mm s.l .), females (250-275 mm); S. macrocephalus, both sexes (150-200 mm); S. altus, males (275-325 mm), females (250-300 mm);S. angusticeps, males (250-275 mm), females (175-225 mm); S. giardi, males (275-300 mm), females (225-250 mm); S. codringtoni, males (150 mm), females (125-150 mm); S. carlottue, males (150-175 mm), females (100-125 mm). Age was estimated from scale annuli comprised of closely spaced circuli that corresponded to the period of rising water (December- March). S. robustus, S. altus, S. angusticeps, and S. giurdi matured at approximately 3 years, and S. macrocephalus, S. codringtoni, and S. curlottae matured at approximately 2 years in the Barotse floodplain. The average numbers of mature oocytes per female were as follows: S. robustus- 1165; S. macrocephalus-810; S. altus-920; S. angusticeps-627; S. codringtoni-580; S. carlottue-362. Fecundity exhibited a weak positive relationship with female size. Analysis of stomach contents showed that interspecific diet overlap was low among both immature and mature size classes of Serrunochromis. Adult size classes of S. robustus fed primarily on small Synodontis catfishes, S. mucrocephalus fed mostly on mormyrid and cichlid fishes, S. altus specialized on mormyrids, and S. angusticeps fed primarily on small cyprinid and characid fishes. The three members of the subgenus Sargochromis consumed mostly invertebrates. Molluscs were the most important item in the diet of S. giardi, and aquatic insects and fish scales were predominant in the diet of S. carlottae. Larger size classes of S. codringtoni consumed large amounts of seeds (Graminaceae). If habitat and diet are considered together, niche separation among Upper Zambezi Serrunochromis species appears to be nearly complete.     

A new species and geographical variation in the Telmatochromis temporalis complex (Teleostei, Cichlidae) from Lake Tanganyika

Telmatochromis brachygnathus sp.n. is described from the southern and central parts of Lake Tanganyika. It can be distinguished from the similar T. temporalis mainly by its smaller mouth. Morphological distinct populations were found in both species.     

From conical to spatulate: intra- and interspecific changes in tooth shape in closely related cichlids (Teleostei; Cichlidae: Eretmodini)

The Eretmodini are closely related cichlids endemic to Lake Tanganyika with very divergent oral tooth shapes, ranging from spatulate in Eretmodus to conical in Tanganicodus. To study how closely related cichlids can generate such divergent tooth shapes, we investigated how the enamel organ directs the development of spatulate teeth in Eretmodus cf. cyanostictus (lineage A), both in ontogeny and in adults, and of conical teeth in adult Tanganicodus cf. irsacae, using 3D-reconstructions from serially sectioned tooth germs. The spatulate oral tooth shape that characterizes adult E. cf. cyanostictus (lineage A) is preceded early in ontogeny by a conical tooth shape. We propose two possible hypotheses to account for changes in the folding of the enamel organ (in particular its epithelio-mesenchymal boundary) capable of generating such distinct tooth shapes. Different arguments lead us to favor the hypothesis of an asymmetric growth and differentiation of the enamel organ, such that the tip of a conical tooth corresponds to one "corner" of a spatulate tooth. Applying current molecular models of tooth shape variation, this would imply the existence of asymmetric fields of inhibition. Whether such asymmetric growth reflects the reutilization of a simple mechanism operating in ontogeny has to be clarified.     

Ontogenetic shape changes in Pomacentridae (Teleostei,Perciformes) and their relationships with feeding strategies: a geometric morphometric approach

The present study explores the shape changes of cranial structures directly involved in food capturing during growth after reef settlement in two species of Pomacentridae (Dascyllus aruanus and Pomacentrus pavo). Landmark‐based geometric morphometrics were used to study allometric patterns and related shape changes in four skeletal units: neurocranium, suspensorium and opercle, mandible and premaxilla. At settlement, the larvae of both species have a relatively similar morphology, especially with respect to the mandible. Their shapes suggest a feeding mode defined as ram/suction‐feeding. Ontogenetic shape changes show a shift to a suction feeding mode of prey capture. The main transformations involved are an increase in height of the suspensorium and the opercle, an elevation of the supraoccipital crest, a relative shortening of the mandible, and a lengthening of the ascending process of the premaxilla. Shape changes of the mandible in the two studied species also reflect an increase of biting capacities. The high disparity between adult shape results from differences in the rate and in the length of ontogenetic trajectories, from divergence of the ontogenetic trajectories (neurocranium, mandible, and premaxilla) and parallel shifts of the trajectories in the size‐shape space (suspensorium and opercle). In an evolutionary context, allometric heterochronies during ontogeny of different skeletal unit of the head may be considered as a basis for the explanation of the diversity of damselfishes. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 92–105.     

Progressive invasion and allopatric speciation can also explain distribution patterns of rock-dwelling cichlids from southern Lake Victoria: a comment on Seehausen and van Alphen (1999)

I show that distribution patterns of rock-dwelling haplochromine cichlids in the south-eastern part of Lake Victoria can be explained by allopatric speciation alone. Observed patterns are consistent with a progressive invasion of different lineages from older ( = deeper) parts of the lake, after which some lineages speciate more frequently than others. Based on a quantitative analysis of distribution data, Seehausen and van Alphen conclude that a high percentage of sympatric speciation (17.5%) is essential to explain the patterns. In my view this conclusion has its origins in an unjustifiable assumption. Seehausen and van Alphen assume that the unresolved phylogeny is balanced. However, allopatric speciation would typically result in imbalanced phylogenies.