Neolamprologus cancellatus, a new cichlid fish from Lake Tanganyika, Africa

Neolamprologus cancellatus, a new cichlid species, is described on the basis of eight specimens from the Zambian coast of Lake Tanganyika. The new species is characterized by a subtruncate caudal fin and a body that is slender (depth 22.3–25.2% in standard length) and easily distinguishable from its congeners by having 7–8 anal fin spines, 34–37 scales in longitudinal line, 33 total vertebrae, and a gridlike body pattern on a pale brownish body color. This species is only known to inhabit the shallow (2–7 m depth), rocky bottom of Wonzye Point in the southern part of the lake.     

Genetic population structure as indirect measure of dispersal ability in a Lake Tanganyika cichlid

Diversification and speciation processes are influenced by intrinsic (ecological specialization, dispersal) and extrinsic (habitat structure and instability) factors, but the effect of ecological characteristics on dispersal is difficult to assess. This study uses mitochondrial control region sequences to investigate the population structure and demographic history of the endemic Lake Tanganyika cichlid Neolamprologus caudopunctatus with a preference for the rock-sand interface along two stretches of continuous, rocky shoreline, and across a sandy bay representing a potential dispersal barrier. Populations along uninterrupted habitat were not differentiated; whereas, the sandy bay separated two reciprocally monophyletic clades. The split between the two clades between 170,000 and 260,000 years BP coincides with a period of rising water level following a major lowstand, and indicates that clades remained isolated throughout subsequent lake level fluctuations. Low long-term effective population sizes were inferred from modest genetic diversity estimates, and may be due to recent population expansions starting from small population sizes 45,000–60,000 years BP. Comparisons with available data from specialized rock-dwelling species of the␣same area suggest that habitat structure and lake level fluctuations determine phylogeographic patterns on large scales, while fine-scale population structure and demography are modulated by species-specific ecologies.     

Widespread geographical distribution of mitochondrial haplotypes in rock-dwelling cichlid fishes from Lake Tanganyika

The spectacularly diverse cichlid fish species flocks of the East African Rift Lakes have elicited much debate on the potential evolutionary mechanisms responsible for the origin of these adaptive radiations. An historical perspective on population structure may offer insights into the processes driving population differentiation and possibly speciation. Here, we examine mitochondrial DNA (mtDNA) sequence variation in two endemic species of rock-dwelling cichlids, Simochromis babaulti and S. diagramma , from Lake Tanganyika. Phylogeographic analyses were used to infer what factors might have been important in the genetic structuring of Simochromis populations. Patterns of mtDNA differentiation in Simochromis were compared to those of other rock-dwelling cichlids to distinguish between competing hypotheses concerning the processes underlying their evolution. In striking contrast to previous findings, populations of Simochromis , even those separated by up to 300 km, were found to share mitochondrial DNA haplotypes. There is no correspondence between mtDNA genealogies and the geographical distribution of populations. Only S. babaulti , but not S. diagramma was found to have a significant association between genetic and geographical distance. These phylogeographic patterns suggest that the evolutionary effects of abiotic and biotic factors shaping population genetic structure may differ substantially even among closely related species of rock-dwelling cichlids. Physical events and barriers to gene flow that are believed to have had a major impact on the geographical distribution and intralacustrine speciation of Tropheus do not seem to have equally strongly affected its close relative Simochromis . These findings emphasize that no single mechanism can be responsible for the formation of population structure, speciation, and the adaptive radiation of all cichlid fishes.     

Sexual conflict over breeding substrate causes female expulsion and offspring loss in a cichlid fish

Females of the Lake Tanganyika cichlid Lamprologus callipterusexclusively breed in empty snail shells that males collect intheir territories. Male–male competition for shells issevere, leading to frequent shell stealing and territory takeover.As a consequence, males have breeding females in their shellsthat spawned with competitors. In this field study, we investigatedboth naturally occurring and experimentally induced encountersof territorial males with females that had spawned with othermales. We found that the breeding success of females that weretaken over by a different male was significantly reduced. Behavioralobservations after experimental shell relocation further showedthat males recognized females that they had not spawned with:males directed more exploration and manipulation behavior towardsuch shells compared with controls. Reoccupation rate of emptiedexperimental shells was significantly higher than that of unmanipulatedempty shells. This indicates that shell stealing and nest takeover,followed by female expulsion, contribute to the reproductivesuccess of L. callipterus males. We also found that female matechoice reduces expulsion risk: females preferred to mate withlarge males, and male size correlated with dominance. We concludethat the limited availability of breeding substrate is a keydeterminant of both intrasexual competition and intersexualconflict in this species.     

Dynamics of sibling aggression of a cichlid fish in Lake Tanganyika

Hydrobiologia - Siblings often compete for limited resources, particularly food provided by their parents. Such competition is usually nonviolent, but direct aggression has evolved in some species....     

Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika

We studied foraging site partitioning between the sexes in Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid with harem-polygyny. Females maintained small territories against heterospecific food competitors within large territories of males, foraging exclusively at the inner side of their own territories (foraging areas). Males fed as frequently as females in their own territories, but mostly outside female foraging areas, although they frequently entered female territories and repelled food competitors from the territories. Soon after removal of the resident females, however, harem males, as well as many food competitors, invaded the vacant territories and intensively devoured prey of female foraging areas. This indicates that although female foraging areas appear to contain more food than outside the areas, harem males refrained from foraging there when the resident females were present. We suggest that harem males will attempt to keep female foraging areas in good condition, whereby they may get females to reside in male territories and/or promote female gonadal maturation.     

Lateralized kinematics of predation behavior in a Lake Tanganyika scale-eating cichlid fish

Behavioral lateralization has been documented in many vertebrates. The scale-eating cichlid fish Perissodus microlepis is well known for exhibiting lateral dimorphism in its mouth morphology and lateralized behavior in robbing scales from prey fish. A previous field study indicated that this mouth asymmetry closely correlates with the side on which prey is attacked, but details of this species' predation behavior have not been previously analyzed because of the rapidity of the movements. Here, we studied scale-eating behavior in cichlids in a tank through high-speed video monitoring and quantitative assessment of behavioral laterality and kinematics. The fish observed showed a clear bias toward striking on one side, which closely correlated with their asymmetric mouth morphologies. Furthermore, the maximum angular velocity and amplitude of body flexion were significantly larger during attacks on the preferred side compared to those on the nonpreferred side, permitting increased predation success. In contrast, no such lateral difference in movement elements was observed in acoustically evoked flexion during the escape response, which is similar to flexion during scale eating and suggests that they share a common motor control pathway. Thus the neuronal circuits controlling body flexion during scale eating may be functionally lateralized upstream of this common motor pathway.     

Distinct population structure in a phenotypically homogeneous rock-dwelling cichlid fish from Lake Tanganyika

Several lineages of cichlid fishes in the East African Great Lakes display stunning levels of morphological diversification. The rapid evolution of rock-dwelling polygynous mouthbrooders in Lake Malawi, for example, was in part ascribed to their allopatric distribution on disjunct stretches of rocky coast, where even short habitat discontinuities reduce gene flow effectively. However, as seen in other cichlids, ecological barriers do not always prevent gene flow, whereas genetic structure can develop along continuous habitat, and morphological diversification does not necessarily accompany genetic differentiation. The present study investigates the population structure of Variabilichromis moorii, a monogamous substrate-brooding lamprologine of rocky coasts in Lake Tanganyika, which occurs over about 1000 km of shoreline almost without phenotypic variation. Phylogeographic analyses of mitochondrial DNA sequences indicated that dispersal is infrequent and generally occurs between adjacent locations only. Exceptions to this pattern are closely related haplotypes from certain locations on opposite lakeshores, a phenomenon which has been observed in other species and is thought to reflect lake crossing along an underwater ridge in times of low water level. Genetic population differentiation, estimated from mitochondrial DNA and microsatellite data in six adjacent populations, was equally high across localities separated by sandy shores and along uninterrupted stretches of rocky shore. Our results suggest that ecological barriers are not required to induce philopatric behavior in Variabilichromis, and that morphological stasis persists in the face of high levels of neutral genetic differentiation.     

Mating system and kin relationship between adults and young in the shell-brooding cichlid fish <Emphasis Type="Italic">Neolamprologus meeli</Emphasis> in Lake Tanganyika

Mating system and parental behavior of ten monogamous pairs and two polygynous groups of the Tanganyikan cichlid Neolamprologus meeli were observed in their natural habitat. The home ranges of males and females overlapped with each other. Most groups included one to six young. Paternal and maternal relationships were determined for 22 young from DNA microsatellite markers. Three types of kinship were found: (I) kinship to both the male and female; (II) kinship to females only; and (III) non-kinship to both sexes. In the groups with type II young, step-fathering or sneaking may have occurred. Type III young were larger than type I, suggesting that the former were of sufficient size to leave their birth nest and settle in the territories of foster parents. Both males and females drove out potential predators of young (including three species of Lepidiolamprologus) as a parental behavior. Adults with type III young attacked approaching predators with as much frequency as those with type I young only, indicating that they provided alloparental care. Adults and young swam together, but, a significant difference existed in the frequencies of interactions between adults versus kin young and adults versus non-kin young. The results suggest that both adults and young recognized kin. Electronic Publication     

Body size evolution of a shell‐brooding cichlid fish from Lake Tanganyika

The substrate‐brooding cichlid fish Telmatochromis temporalis in Lake Tanganyika demonstrates a simple example of ecological speciation between normal and dwarf morphs through divergent natural selection on body size. The dwarf morph most likely evolved from the ancestral normal morph; therefore, elucidating the evolution of its small body size is a key to understanding this ecological speciation event. Previous studies suggest that the small body size of the dwarf morph is an adaptation to the use of empty snail shells as shelters (males) and spawning sites (females), but this idea has not been fully evaluated. Combining original and previously published information, this study compared likelihood values to determine the primary factor that would be responsible for regulating the body size of the dwarf morph. Male body size is most likely regulated by the ability to turn within shells, which may influence the predation avoidance of adult fish. Females are smaller than males, and their body size is most likely regulated by the ability to lay eggs in the small spaces within shells close to the shell apices where predation risk on eggs is lower. This study provides new evidence supporting the hypothesis that different natural selection factors affected body size of the different sexes of the dwarf morph, which has not been reported in other animal species.     

Genomics of speciation and introgression in Princess cichlid fishes from Lake Tanganyika

How variation in the genome translates into biological diversity and new species originate has endured as the mystery of mysteries in evolutionary biology. African cichlid fishes are prime model systems to address speciation‐related questions for their remarkable taxonomic and phenotypic diversity, and the possible role of gene flow in this process. Here, we capitalize on genome sequencing and phylogenomic analyses to address the relative impacts of incomplete lineage sorting, introgression and hybrid speciation in the Neolamprologus savoryi‐complex (the ‘Princess cichlids’) from Lake Tanganyika. We present a time‐calibrated species tree based on whole‐genome sequences and provide strong evidence for incomplete lineage sorting in the early phases of diversification and multiple introgression events affecting different stages. Importantly, we find that the Neolamprologus chromosomes show centre‐to‐periphery biases in nucleotide diversity, sequence divergence, GC content, incomplete lineage sorting and rates of introgression, which are likely modulated by recombination density and linked selection. The detection of heterogeneous genomic landscapes has strong implications on the genomic mechanisms involved in speciation. Collinear chromosomal regions can be protected from gene flow and harbour incompatibility genes if they reside in lowly recombining regions, and coupling can evolve between nonphysically linked genomic regions (chromosome centres in particular). Simultaneously, higher recombination towards chromosome peripheries makes these more dynamic, evolvable regions where adaptation polymorphisms have a fertile ground. Hence, differences in genome architecture could explain the levels of taxonomic and phenotypic diversity seen in taxa with collinear genomes and might have contributed to the spectacular cichlid diversity observed today.     

Overlapping territory of the benthophagous cichlid fish, Lobochilotes labiatus, in Lake Tanganyika

Synopsis Feeding territories do not usually overlap in the same fish species. We studied overlapping territories and feeding behaviours of a benthophagous cichlid fish Lobochilotes labiatus Boulenger in a coastal area of Lake Tanganyika. Forty-four individuals of both sexes, 6–31 cm in total length, maintained territories in a 21 m × 25 m area. These could be regarded as feeding territories. The territories of similar-sized owners (usually different by less than 5 cm) did not overlap each other, but those of different-sized owners overlapped widely. Aggressive interactions were rarely observed between fish distinctly different in body size. Both large and small fish took benthic animals of the same kinds and of similar size. In contrast, the size of foraging sites (crevices) was different between them: the large fish foraged at large crevices and small fish at small crevices. This partitioning of foraging sites may enable the overlap of territories in this benthivore.     

Microdontochromis rotundiventralis, a new cichlid fish (perciformes: Cichlidae) from Lake Tanganyika

Microdontochromis rotundiventralis, a new species of cichlid fish, is described on the basis of 13 specimens from Nkumbula Island, Lake Tanganyika (Zambia). It is distinct from its only congener,M. tenuidentatus, in having two (rarely one) rows of teeth on both jaws, a rounded distal margin on the pelvic fin, the outermost pelvic fin soft ray length 1.23–1.43 times the innermost ray, a deeper body (depth 26.8–29.1% standard length) and the anal fin with 9 (rarely 8 or 10) soft rays.     

Group foraging by the herbivorous cichlid fish,Petrochromis fasciolatus,in Lake Tanganyika

Group foraging of the algae eating cichlid,Petrochromis fasciolatus, was studied along a rocky shore in Lake Tanganyika, in an area occupied by the feeding territories of several herbivorous cichlid species. SolitaryP. fasciolatus or schools comprising a few members only were nearly always repelled from such territories, although larger schools (40–250 individuals) ofP. fasciolatus were able to overwhelm the defenses of territory owners and enable subsequent foraging. The frequency and duration of individual foraging sessions on the algal mats were positively correlated with group size, probably owing to individuals in large schools suffering fewer attacks from territory owners. Almost all of the foraging sites ofP. fasciolatus were inside the territories of other herbivorous cichlids, especially those ofNeolamprologus moorii (more than 80% of total), in which the algal density was more than 10 times that in territories of the other species.N. moorii territories included only 4% rocky substrate, indicating that schools ofP. fasciolatus selectively chose territories containing rich resources.     

Function of nuchal humps of a cichlid fish from Lake Tanganyika: inferences from morphological data

Nuchal humps occur in various fish groups, but their functions are rarely studied. Both sexes of Cyphotilapia gibberosa, a cichlid fish endemic to Lake Tanganyika, possess nuchal humps. This study morphologically analysed the humps of this species to assess the primary factors responsible for hump development. Semi-landmarks showed that the size of male humps positively correlated with body condition, and thicknesses of hypodermises implied that the humps were larger in males than in females. These facts suggest that sexual selection on male humps is intense. Semi-landmarks showed that the humps were less prominent in females than in males, and that the transformation of nuchal humps with growth (becoming more prominent) ceased at medium body size in females. These facts suggest that sex recognition likely plays a role in the evolution of the uniform humps of large females, because, according to this hypothesis, a morphologically moderate hump may be favoured. If male humps also function as a sex recognition trait, the transformation of nuchal humps is expected to cease at a certain body size, as did female humps. However, the male humps became continuously more prominent from the smallest to the largest individuals examined. The body size at which hump transformation stops or at which extreme hump shape interferes with sex recognition may be outside of the size range of the present samples. A prominent nuchal hump may also function as a species recognition trait, because sympatric cichlid species do not develop nuchal humps as prominent as this species. In short, the present morphological analyses do not contradict the hypotheses that C. gibberosa individuals recognise conspecifics and their sex from the shapes of the nuchal humps, and that females prefer males with larger humps.     

Mouth dimorphism in scale‐eating cichlid fish from Lake Tanganyika advances individual fitness

Random asymmetry, that is the coexistence of left‐ and right‐sided (or ‐handed) individuals within a population, is a particular case of natural variation; what triggers and maintains such dimorphisms remains unknown in most cases. Here, we report a field‐based cage experiment in the scale‐eating Tanganyikan cichlid Perissodus microlepis, which occurs in two morphs in nature: left‐skewed and right‐skewed individuals with respect to mouth orientation. Using underwater cages stocked with scale‐eaters and natural prey fish, we first confirm that, under semi‐natural conditions, left‐skewed scale‐eaters preferentially attack the right flank of their prey, whereas right‐skewed individuals feed predominantly from the left side. We then demonstrate that scale‐eaters have a higher probability for successful attacks when kept in dimorphic experimental populations (left‐ and right‐skewed morphs together) as compared to monomorphic populations (left‐ or right‐skewed morphs), most likely because prey fishes fail to accustom to strikes from both sides. The significantly increased probability for attacks appears to be the selective agent responsible for the evolution and maintenance of mouth dimorphism in P. microlepis, lending further support to the hypothesis that negative frequency‐dependent selection is the stabilizing force balancing the mouth dimorphism at quasi‐equal ratios in scale‐eating cichlids.     

Differential gene flow of mitochondrial and nuclear DNA markers among chromosomal races of Australian morabine grasshoppers (Vandiemenella, viatica species group)

Recent theoretical developments have led to a renewed interest in the potential role of chromosomal rearrangements in speciation. Australian morabine grasshoppers (genus Vandiemenella, viatica species group) provide an excellent study system to test this potential role of chromosomal rearrangements because they show extensive chromosomal variation and formed the basis of a classic chromosomal speciation model. There are three chromosomal races, viatica19, viatica17, and P24(XY), on Kangaroo Island, South Australia, forming five parapatric populations with four putative contact zones among them. We investigate the extent to which chromosomal variation among these populations may be associated with barriers to gene flow. Population genetic and phylogeographical analyses using 15 variable allozyme loci and the elongation factor-1alpha (EF-1alpha) gene indicate that the three races represent genetically distinct taxa. In contrast, analyses of the mitochondrial cytochrome c oxidase subunit I (COI) gene show the presence of three distinctive and geographically localized groups that do not correspond with the distribution of the chromosomal races. These discordant population genetic patterns are likely to result from introgressive hybridization between the chromosomal races and range expansions/contractions. Overall, these results suggest that reduction of nuclear gene flow may be associated with chromosomal variation, or underlying genetic variation linked with chromosomal variation, whereas mitochondrial gene flow appears to be independent of this variation in these morabine grasshoppers. The identification of an intact contact zone between P24(XY) and viatica17 offers considerable potential for further investigation of molecular mechanisms that maintain distinct nuclear genomes among the chromosomal races.