Mating and Parental Care in Lake Tanganyika's Cichlids

Cichlid fishes of Lake Tanganyika display a variety of mating and parental care behaviors, including polygamous and monogamous mouthbrooding and substrate breeding, cooperative breeding, as well as various alternative reproductive tactics such as sneaking and piracy. Moreover, reproductive behaviors sometimes vary within species both in space and in time. Here, I survey reports on mating and parenting behaviors of Lake Tanganyika cichlid species and address the evolution of mating and parental care patterns and sexual dimorphism. Notes on measures of sexual selection intensity and the difficulties of defining mating systems and estimating selection intensities at species level conclude the essay.

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title: Parental investment, sexual selection and sex ratios
abstract: Conventional sex roles imply caring females and competitive males. The evolution of sex role divergence is widely attributed to anisogamy initiating a self‐reinforcing process. The initial asymmetry in pre‐mating parental investment (eggs vs. sperm) is assumed to promote even greater divergence in post‐mating parental investment (parental care). But do we really understand the process? Trivers [Sexual Selection and the Descent of Man 1871–1971 (1972), Aldine Press, Chicago] introduced two arguments with a female and male perspective on whether to care for offspring that try to link pre‐mating and post‐mating investment. Here we review their merits and subsequent theoretical developments. The first argument is that females are more committed than males to providing care because they stand to lose a greater initial investment. This, however, commits the ‘Concorde Fallacy’ as optimal decisions should depend on future pay‐offs not past costs. Although the argument can be rephrased in terms of residual reproductive value when past investment affects future pay‐offs, it remains weak. The factors likely to change future pay‐offs seem to work against females providing more care than males. The second argument takes the reasonable premise that anisogamy produces a male‐biased operational sex ratio (OSR) leading to males competing for mates. Male care is then predicted to be less likely to evolve as it consumes resources that could otherwise be used to increase competitiveness. However, given each offspring has precisely two genetic parents (the Fisher condition), a biased OSR generates frequency‐dependent selection, analogous to Fisherian sex ratio selection, that favours increased parental investment by whichever sex faces more intense competition. Sex role divergence is therefore still an evolutionary conundrum. Here we review some possible solutions. Factors that promote conventional sex roles are sexual selection on males (but non‐random variance in male mating success must be high to override the Fisher condition), loss of paternity because of female multiple mating or group spawning and patterns of mortality that generate female‐biased adult sex ratios (ASR). We present an integrative model that shows how these factors interact to generate sex roles. We emphasize the need to distinguish between the ASR and the operational sex ratio (OSR). If mortality is higher when caring than competing this diminishes the likelihood of sex role divergence because this strongly limits the mating success of the earlier deserting sex. We illustrate this in a model where a change in relative mortality rates while caring and competing generates a shift from a mammalian type breeding system (female‐only care, male‐biased OSR and female‐biased ASR) to an avian type system (biparental care and a male‐biased OSR and ASR).

title: Sexual Selection Determines Parental Care Patterns in Cichlid Fishes
abstract: Despite a massive research effort, our understanding of why, in most vertebrates, males compete for mates and females care for offspring remains incomplete. Two alternative hypotheses have been proposed to explain the direction of causality between parental care and sexual selection. Traditionally, sexual selection has been explained as a consequence of relative parental investment, where the sex investing less will compete for the sex investing more. However, a more recent model suggests that parental care patterns result from sexual selection acting on one sex favoring mating competition and lower parental investment. Using species-level comparative analyses on Tanganyikan cichlid fishes we tested these alternative hypotheses employing a proxy of sexual selection based on mating system, sexual dichromatism, and dimorphism data. First, while controlling for female reproductive investment, we found that species with intense sexual selection were associated with female-only care whereas species with moderate sexual selection were associated with biparental care. Second, using contingency analyses, we found that, contrary to the traditional view, evolutionary changes in parental care type are dependent on the intensity of sexual selection. Hence, our results support the hypothesis that sexual selection determines parental care patterns in Tanganyikan cichlid fishes.

title: Environmental Complexity and Social Organization Sculpt the Brain in Lake Tanganyikan Cichlid Fish
abstract: Complex brains and behaviors have occurred repeatedly within vertebrate classes throughout evolution. What adaptive pressures drive such changes? Both environmental and social features have been implicated in the expansion of select brain structures, particularly the telencephalon. East African cichlid fishes provide a superb opportunity to analyze the social and ecological correlates of neural phenotypes and their evolution. As a result of rapid, recent, and repeated radiations, there are hundreds of closely-related species available for study, with an astonishing diversity in habitat preferences and social behaviors. In this study, we present quantitative ecological, social, and neuroanatomical data for closely-related species from the (monophyletic) Ectodini clade of Lake Tanganyikan cichlid fish. The species differed either in habitat preference or social organization. After accounting for phylogeny with independent contrasts, we find that environmental and social factors differentially affect the brain, with environmental factors showing a broader effect on a range of brain structures compared to social factors. Five out of seven of the brain measures show a relationship with habitat measures. Brain size and cerebellar size are positively correlated with species number (which is correlated with habitat complexity); the medulla and olfactory bulb are negatively correlated with habitat measures. The telencephalon shows a trend toward a positive correlation with rock size. In contrast, only two brain structures, the telencephalon and hypothalamus, are correlated with social factors. Telencephalic size is larger in monogamous species compared to polygamous species, as well as with increased numbers of individuals; monogamy is also associated with smaller hypothalamic size. Our results suggest that selection or drift can act independently on different brain regions as the species diverge into different habitats and social systems.

title: Why do females care more than males?
abstract: Females tend to provide more parental care than males. Previous efforts to account for this have been confused because it is difficult to express the costs of care for males and females in the same currency. Here I propose a null model that does so, using the Fisherian constraint that total male and female reproduction must be equal. The model shows that, contrary to a number of recent analyses, lower probability of parentage for males does tend to make males less likely than females to provide care. It also shows how sexual selection stemming from premating asymmetries in investment promotes similar post–mating asymmetries.

title: Male “Mixed” Reproductive Strategies in Biparental Species: Trivers Was Probably Right, but Why?
abstract: Trivers proposed that, if parental care by both sexes is advantageous, males should practice a “mixed” strategy of seeking extrapair copulations, while restricting their parental investment to offspring of social mates. We explore circumstances under which males should limit their parental care in the predicted manner. We find that Trivers’s “mixed” strategy will generally be evolutionarily stable so long as either socially monogamous or polygynous males usually sire more offspring per brood from a social mate than they typically sire in broods of extrapair mates. Polygynous males should spread investment across their home nests unless the expected number of chicks sired in them differs widely. Whether polygynous males should restrict paternal care to social mates’ offspring hinges additionally on resident male investment in broods containing extrapair young: if resident males contribute minimally, some investment by a polygynous extrapair male becomes more advantageous. Recently reviewed data on extrapair fertilization distributions within monogamous and polygynous passerines suggest that extrapair offspring often predominate numerically within their broods, consistent with sperm expenditure theory. Nevertheless, most species conform to the model’s criterion regarding relative parentage levels in broods of social versus extrapair mates. Patterns of extrapair parentage thus appear sufficient to stabilize biparental care systems.

title: Experimental evolution of a sexually selected display in yeast
abstract: The fundamental principle underlying sexual selection theory is that an allele conferring an advantage in the competition for mates will spread through a population. Remarkably, this has never been demonstrated empirically. We have developed an experimental system using yeast for testing genetic models of sexual selection. Yeast signal to potential partners by producing an attractive pheromone; stronger signallers are preferred as mates. We tested the effect of high and low levels of sexual selection on the evolution of a gene determining the strength of this signal. Under high sexual selection, an allele encoding a stronger signal was able to invade a population of weak signallers, and we observed a corresponding increase in the amount of pheromone produced. By contrast, the strong signalling allele failed to invade under low sexual selection. Our results demonstrate, for the first time, the spread of a sexually selected allele through a population, confirming the central assumption of sexual selection theory. Our yeast system is a powerful tool for investigating the genetics of sexual selection.

title: A New Viewpoint on the Evolution of Sexually Dimorphic Human Faces
abstract: Human faces show marked sexual shape dimorphism, and this affects their attractiveness. Humans also show marked height dimorphism, which means that men typically view women's faces from slightly above and women typically view men's faces from slightly below. We tested the idea that this perspective difference may be the evolutionary origin of the face shape dimorphism by having males and females rate the masculinity/femininity and attractiveness of male and female faces that had been manipulated in pitch (forward or backward tilt), simulating viewing the face from slightly above or below. As predicted, tilting female faces upwards decreased their perceived femininity and attractiveness, whereas tilting them downwards increased their perceived femininity and attractiveness. Male faces tilted up were judged to be more masculine, and tilted down judged to be less masculine. This suggests that sexual selection may have embodied this viewpoint difference into the actual facial proportions of men and women.

title: Fishes as models in studies of sexual selection and parental care
abstract: Fishes are by far the most diverse group of vertebrates. This fact is in no way, however, reflected in their use as model organisms for understanding sexual selection or parental care. Why is this so? Is it because fishes are actually poor models? The usefulness of fishes as models for sexual selection and parental care is discussed by emphasizing some problems inherent in fish studies, along with a number of reasons why fishes are indeed excellently suited. The pros and cons of fishes as models are discussed mainly by comparison with birds, the most popular model organisms in animal behaviour. Difficulties include a lack of background knowledge for many species, and the problems of marking and observing fishes in their natural environment. Positive attributes include the diversity of lifestyles among fishes, and the ease with which they can be studied experimentally in the laboratory. How useful fish models can be is briefly illustrated by the impressive and broadly relevant advances derived from studies of guppies Poecilia reticulata and three-spined sticklebacks Gasterosteus aculeatus. A selection of topics is highlighted where fish studies have either advanced or could greatly enhance, the understanding of processes fundamental to animal reproductive dynamics. Such topics include sex role dynamics, the evolution of female ornamentation and mate choice copying. Finally, a number of potential pitfalls in the future use of fish as models for sexual selection and parental care are discussed. Researchers interested in these issues are recommended to make much more extensive use of fish models, but also to adopt a wider range of models among fishes.

title: Behaviorally Induced Sex Reversal of Metriaclima cf. livingstoni (Cichlidae) from Lake Malawi
abstract: Sex reversal in the Lake Malawi cichlid, Metriaclima cf. livingstoni, was documented. A functional male in an aquarium was separated from three functional females by a divider of clear Plexiglas® sealed to tank walls. After the male was removed, one of the functional females developed male secondary sexual characteristics and fertilized eggs produced by the other females. In a second experiment, 6–7 functional females were placed in tanks without males. Again, sex change was noted with one of the former females fertilizing several broods.

title: The Lake Tanganyika cichlid species assemblage: recent advances in molecular phylogenetics
abstract: Tanganyika is not the most species-rich of the Great East African Lakes, but comprises the greatest diversity of cichlid fishes in terms of morphology, ecology, and breeding styles. The lake contains a polyphyletic assemblage of cichlid lineages, which evolved from several ancient species that colonized the emerging lake some 9–12 million years ago. Based on morphological characteristics, the Tanganyikan cichlids have been classified into 12, or, more recently, 16 tribes, which are largely supported by molecular data. The radiations of East African cichlids are believed to be driven by complex interactions between extrinsic factors, such as climatic changes and geological processes, and intrinsic biological characteristics of the involved organisms. Diversification within different lineages occurred simultaneously in response to drastic habitat changes such as the establishment of lacustrine deep-water conditions 5–6 MYA and subsequent major lake-level fluctuations. This seems particularly true for the mouthbrooding lineages whereas the substrate breeders underwent a more gradual process of diversification. This review presents an account of the taxonomy and phylogeny of the Lake Tanganyika cichlid species assemblage, its relationship to the African cichlid fauna, key factors leading to the astonishing diversity and discusses recently proposed alternative age estimates for the Lake Tanganyika cichlid species assemblage.

title: Life history evolution in cichlids 1: revisiting the evolution of life histories in relation to parental care
abstract: Empirical links between egg size and duration of parental care in fishes have generated a considerable amount of theory concerning life history evolution. However, to date, this link has not been investigated in relation to other important life‐history traits such as clutch size and body size, or while controlling for shared ancestry between species. We provide the first phylogenetically based tests using a database with information on egg size, clutch size, body size and care duration in cichlid fishes (Cichlidae). Multiple regression analyses, based on independent contrasts on both the species and the genus level, showed that clutch size is the variable most closely related to duration of care. This pattern appeared to be driven by post‐hatch care relationships. Our results show that, contrary to expectation, there is no positive link between egg size and care duration in Cichlidae. Instead, greater reproductive output through increased clutch size investment appears to have coevolved with greater care of offspring. We suggest that re‐evaluation of the generality of current models of the evolution of egg size under parental care in fishes is needed.

title: Female promiscuity promotes the evolution of faster sperm in cichlid fishes
abstract: Sperm competition, the contest among ejaculates from rival males to fertilize ova of a female, is a common and powerful evolutionary force influencing ejaculate traits. During competitive interactions between ejaculates, longer and faster spermatozoa are expected to have an edge; however, to date, there has been mixed support for this key prediction from sperm competition theory. Here, we use the spectacular radiation of cichlid fishes from Lake Tanganyika to examine sperm characteristics in 29 closely related species. We provide phylogenetically robust evidence that species experiencing greater levels of sperm competition have faster-swimming sperm. We also show that sperm competition selects for increases in the number, size, and longevity of spermatozoa in the ejaculate of a male, and, contrary to expectations from theory, we find no evidence of trade-offs among sperm traits in an interspecific analysis. Also, sperm swimming speed is positively correlated with sperm length among, but not within, species. These different responses to sperm competition at intra- and interspecific levels provide a simple, powerful explanation for equivocal results from previous studies. Using phylogenetic analyses, we also reconstructed the probable evolutionary route of trait evolution in this taxon, and show that, in response to increases in the magnitude of sperm competition, the evolution of sperm traits in this clade began with the evolution of faster (thus, more competitive) sperm.

title: Parallel life history evolution in mouthbrooding cichlids from the African Great Lakes
abstract: The existence of ancient deep-water lakes provides an opportunity to study the independent adaptation of aquatic organisms to pelagic, benthic, and rocky shore habitats. With improving resolution of their phylogenetic relationships, the many cichlid fish species endemic to the African Great Lakes Malawi, Tanganyika, and Victoria provide a significant resource for the comparative study of such evolutionary processes. Here, we show that cichlid lineages colonizing rocky shores and pelagic habitats in the different lakes have independently evolved larger eggs and lower fecundities than benthic lineages, suggesting parallel adaptive life-history evolution. By contrast, other pelagic teleost fishes in both marine and freshwater habitats, including African lakes, typically produce large numbers of very small eggs. Our results also suggest that decreased fecundity and increased egg size not only occurred independently in each lake but occurred independently in the colonization of rocky and pelagic habitats.

title: Sperm size of African cichlids in relation to sperm competition
abstract: We compared pairs of closely related taxa of cichlid fishes from Lake Tanganyika to examine the relationship between sperm size and the presumed intensity of sperm competition. In contrast to previous reports of relatively short sperm in polygamous fishes across a variety of taxa, we found that polygamous cichlids had significantly longer sperm than their closest monogamous relatives. In addition, sperm length was significantly related to relative testis size (controlling for body size and phylogeny). The site of fertilization may also be correlated with sperm length, as species that fertilize in the female’s buccal cavity had significantly shorter sperm than those that fertilized eggs on the substrate. Assuming that relatively large testes and polygamous mating are indicative of more intense sperm competition, these results indicate that sperm length is related to the intensity of sperm competition in this clade of cichlids, as has been found previously in insects, birds, and mammals.

title: Social fishes and single mothers: brain evolution in African cichlids
abstract: As with any organ, differences in brain size—after adequate control of allometry—are assumed to be a response to selection. With over 200 species and an astonishing diversity in niche preferences and social organization, Tanganyikan cichlids present an excellent opportunity to study brain evolution. We used phylogenetic comparative analyses of sexed adults from 39 Tanganyikan cichlid species in a multiple regression framework to investigate the influence of ecology, sexual selection and parental care patterns on whole brain size, as well as to analyse sex-specific effects. First, using species-specific measures, we analysed the influence of diet, habitat, form of care (mouthbrooding or substrate guarding), care type (biparental or female only) and intensity of sexual selection on brain size, while controlling for body size. Then, we repeated the analyses for male and female brain size separately. Type of diet and care type were significantly correlated with whole brain size. Sex-specific analyses showed that female brain size correlated significantly with care type while male brain size was uncorrelated with care type. Our results suggest that more complex social interactions associated with diet select for larger brains and further that the burden of uniparental care exerts high cognitive demands on females.

title: Distinct Evolutionary Patterns of Brain and Body Size During Adaptive Radiation
abstract: Morphological traits are often genetically and/or phenotypically correlated with each other and such covariation can have an important influence on the evolution of individual traits. The strong positive relationship between brain size and body size in vertebrates has attracted a lot of interest, and much debate has surrounded the study of the factors responsible for the allometric relationship between these two traits. Here, we use comparative analyses of the Tanganyikan cichlid adaptive radiation to investigate the patterns of evolution for brain size and body size separately. We found that body size exhibited recent bursts of rapid evolution, a pattern that is consistent with divergence linked to ecological specialization. Brain weight on the other hand, showed no bursts of divergence but rather evolved in a gradual manner. Our results thus show that even highly genetically correlated traits can present markedly different patterns of evolution, hence interpreting patterns of evolution of traits from correlations in extant taxa can be misleading. Furthermore, our results suggest, contrary to expectations from theory, that brain size does not play a key role during adaptive radiation.

title: Brain structure evolution in a basal vertebrate clade: evidence from phylogenetic comparative analysis of cichlid fishes
abstract: BackgroundThe vertebrate brain is composed of several interconnected, functionally distinct structures and much debate has surrounded the basic question of how these structures evolve. On the one hand, according to the 'mosaic evolution hypothesis', because of the elevated metabolic cost of brain tissue, selection is expected to target specific structures mediating the cognitive abilities which are being favored. On the other hand, the 'concerted evolution hypothesis' argues that developmental constraints limit such mosaic evolution and instead the size of the entire brain varies in response to selection on any of its constituent parts. To date, analyses of these hypotheses of brain evolution have been limited to mammals and birds; excluding Actinopterygii, the basal and most diverse class of vertebrates. Using a combination of recently developed phylogenetic multivariate allometry analyses and comparative methods that can identify distinct rates of evolution, even in highly correlated traits, we studied brain structure evolution in a highly variable clade of ray-finned fishes; the Tanganyikan cichlids.ResultsTotal brain size explained 86% of the variance in brain structure volume in cichlids, a lower proportion than what has previously been reported for mammals. Brain structures showed variation in pair-wise allometry suggesting some degree of independence in evolutionary changes in size. This result is supported by variation among structures on the strength of their loadings on the principal size axis of the allometric analysis. The rate of evolution analyses generally supported the results of the multivariate allometry analyses, showing variation among several structures in their evolutionary patterns. The olfactory bulbs and hypothalamus were found to evolve faster than other structures while the dorsal medulla presented the slowest evolutionary rate.ConclusionOur results favor a mosaic model of brain evolution, as certain structures are evolving in a modular fashion, with a small but non-negligible influence of concerted evolution in cichlid fishes. Interestingly, one of the structures presenting distinct evolutionary patterns within cichlids, the olfactory bulbs, has also been shown to evolve differently from other structures in mammals. Hence, our results for a basal vertebrate clade also point towards a conserved developmental plan for all vertebrates.

title: Parallel evolution of opsin gene expression in African cichlid fishes.
abstract: Phenotypic evolution may occur either through alterations to the structure of protein-coding genes or their expression. Evidence for which of these two mechanisms more commonly contribute to the evolution of a phenotype can be garnered from examples of parallel and convergent evolution. The visual system of East African cichlid fishes is an excellent system with which to address this question. Cichlid fishes from Lakes Malawi (LM) and Victoria together exhibit three diverse palettes of coexpressed opsins and several important protein-coding mutations that both shift spectral sensitivity. Here we assess both opsin expression and protein-coding diversity among cichlids from a third rift lake, Lake Tanganyika (LT). We found that Tanganyikan cichlids exhibit three palettes of coexpressed opsins that largely overlap the short-, middle-, and long-wavelength-sensitive palettes of LM cichlids. Bayesian phenotypic clustering and ancestral state reconstructions both support the parallel evolution of the short- and middle-wavelength palettes among cichlids from LT and LM. In each case, these transitions occurred from different ancestors that expressed the same long-wavelength palette. We also identified similar but distinct patterns of correlated evolution between opsin expression, diet, and lens transmittance among cichlids from LT and LM as well. In contrast to regulatory changes, we identified few functional or potentially functional mutations in the protein-coding sequences of three variable opsins, with the possible exception of the SWS1 (ultraviolet) opsin. These results underscore the important contribution that gene regulation can make to rapid phenotypic evolution and adaptation.

title: Resource defence or exploded lek? - A question of perspective
abstract: It becomes increasingly obvious that animal mating systems cannot be classified into distinct categories, but transitions between mating system classes are continuous. Positioning a certain mating system at this continuum is often not straightforward, however. Depending on which characteristic is considered, a mating system may end up at very different positions on this gradient. Here, we explore the potential conflict between mating system classifications that may arise when they are based on different criteria by investigating the mating system of the cichlid fish Simochromis pleurospilus in which males defend small patches of homogeneously distributed food resources (turf algae) vigorously against food competitors, but they allow specific females to use them. We hypothesized that male defence may generate high-quality feeding patches serving to attract females, and hence male territoriality constitutes a form of courtship. Our field data show that males selectively allow approximately one-third of the visiting females to feed on their territory and that females preferentially feed in male territories and usually sample several territories successively. As males protect food patches against other algae grazers and guard females from harassment by food competitors, females gain nutritional benefits from visiting male territories. Hence, males appear to generate essential resources for females, which is the key feature of resource-defence mating systems, although the distributions of resources and of males and females are characteristics of an exploded lek.

title: Survival of broods under parental care and parental roles of the cichlid fish,Lamprologus toae, in Lake Tanganyika
abstract: The survival of offspring guarded by both parents and by only the female parent who remained after experimental removal of the male parent was observed in the cichlid fishLamprologus toae in its natural habitat. The number of offspring guarded by both parents abruptly decreased during the period from eggs through postlarvae. It became stable for low rate of mortality when the offspring were between 0.5 and 1.8 cm in body length. The number again decreased this time gradually after attaining 1.8 cm in body length. Mortality seemed to be caused by predation. Experimental removal of the male parent at any larval stage caused the decrease of offspring by predatory fishes. It seems that both parents are indispensable to protect the offspring against predators under natural conditions.

title: Lunar synchronization of spawning in cichlid fishes of the tribe Lamprologini in Lake Tanganyika
abstract: Lunar synchronization of spawning was investigated for eight substrate brooding cichlid fishes belonging to the tribe Lamprologini. Their spawning activities all peaked during the second quarter of the lunar cycle. Comparison between their breeding styles and the degrees of spawning synchronization suggested that the nocturnal guarding-efficiency of eggs, especially exposed ones, is improved by the maximal amount of moonlight during full moon and, in species whose young leave the breeding sites about 2 weeks after spawning, the survival of dispersing young is enhanced by the darkness of moonless nights.

title: Brood protection at a cost: mouth brooding under hypoxia in an African cichlid
abstract: This study quantifies the behavioral response of the widespread mouth brooding African cichlid Pseudocrenilabrus multicolor victoriae to progressive hypoxia. We exposed four gender/stage classes of P. multicolor (males, brooding females, females that had just released young, and non-brooding females) to progressive hypoxia and recorded the percent time spent using aquatic surface respiration (surface skimming, ASR) and gill ventilation rates. This was done for fish collected from three sites in Uganda (lake, swamp, and river) after long-term acclimation to normoxia. There was no effect of site of origin on response to hypoxia, but ASR thresholds did differ between gender/stage classes. The oxygen level (threshold) at which spent 10, 50, and 90% of their time at the surface using ASR was much higher for brooding females than for males, whereas ASR thresholds did not differ between non-brooding females and males. Similarly, the level at which ASR was initiated was much higher in brooding females than males, but did not differ between males and non-brooders, or between males and females than had just released young. The rate of gill ventilation dropped significantly in males and all stages of females after initiation of ASR, suggesting that surface skimming increases efficiency of oxygen acquisition. These results suggest that mouth brooding in female P. multicolor ASR improves oxygen uptake but imposes a cost in terms of time spent at the water surface, and this may affect maternal predation risk in low-oxygen habitats.

title: Mouthbreeding in fishes.
abstract: Mouthbreeding occurs in at least fifty-three genera of fishes (four orders, eight families) that inhabit marine, brackish and fresh water. It possibly developed in some Betta species from the habit of orally placing eggs in a bubble nest, whereas in cichlids it possibly evolved from mouthing and transferring behaviour. Invasion of new habitats, high competition for or lack of nest sites, new courtship patterns, and predation may be some of the factors involved in the selection for this advanced type of parental care. Morphological adaptations for either herbivorous or carnivorous feeding can be advantageous for mouthbreeding, and apparently no special adaptations have evolved in the mouth for this behaviour. Increased efficiency in fertilization and/or pick up of the eggs has been brought about in some species by the possession of colourful genital tassels, false eggs on the anal fin, and by enlarged pelvic fins. Mouthbreeders tend to have a small number of large eggs without adhesive areas that are released in batches during spawning. The greater the amount of yolk in the eggs, the less likely it is that parental care will be continued after the fry are released. The incubating parent usually goes without food, although some fry may feed while in the parent's mouth. Coughing, yawning and swallowing appear to be carefully controlled during the parental period. Churning movements of the mouth are functionally analogous to fanning movements of the pectoral fins in substrate-breeders. Some of the changes in the amounts and rates of churning and respiration are associated with the metabolic development of the young, whereas others appear to be involved with bouyancy adjustments by the parent.

title: Intra-buccal feeding of young in an undescribed Tanganyikan cichlid Microdontochromis sp.
abstract: SynopsisThe zooplanktivorous cichlid Microdontochromis sp. formed large stationary schools in midwater consisting of mouthbrooding and nonbrooding individuals. Early young smaller than 6.0 mm in standard length were mouthbrooded solely by females, but large young, up to 17.4 mm, were mouthbrooded by both females and males. Mouthbrooding fish took food as actively as nonbrooding fish to nourish the young and themselves. Eggs were 1.9 mm in maximum diameter, among the smallest known for mouthbrooding cichlid fishes. Young which ate food within the parent's buccal cavity showed a 10-fold increase in dry weight during the mouthbrooding period. Data suggested that parents finally farmed out their young into broods of other cichlid fishes.

title: Probable evolution of the coelacanth’s reproductive style: lecithotrophy and orally feeding embryos in cichlid fishes and in Latimeria chalumnae
abstract: The living coelacanth is a livebearer. Yolk seems to be the main source of nutrients and of oxygen to the embryo (fetus). Long before birth, young may also possibly feed orally on histotrophe secretion and egg debris. This type of reproduction evolved, as in most other fishes, from oviparity. The Carboniferous coelacanth Rhabdoderma exiguum had eggs of much lesser yolk volume and may represent an earlier form of oviparity with hiding, guarding or brooding type of parental care. The Jurassic coelacanth Holophagus (Undina) and the Cretaceous Axelrodichthys appear to have already evolved the internal-bearing style. Much of this evolutionary sequence is similar to that in cichlids. Ancestral cichlids are substrate tenders and nesters, with small eggs, little yolk and a feeding larva with indirect development. Mouthbrooding cichlids evolved a few, large eggs with denser yolk, direct development and, ultimately, orally feeding embryos while yolk is still in ample supply. Mixed feeding from yolk and orally ingested food in cichlids and in coelacanths is shown to be an enhanced mode of food delivery to the embryos over that from each source separately, in order to produce directly a better developed or larger young at the time of release, i.e. independence. Increase in egg size is regarded as an environmentally induced, altered pattern of yolk synthesis and an initial component of the epigenetic mechanism leading towards greater specialization. Carotenoids are incorporated within the yolk to assist the oxidative metabolism of the developing embryo.

title: Active browsing by mouthbrooding females of Tropheus duboisi and Tropheus moorii (Cichlidae) to feed the young and/or themselves
abstract: SynopsisFishes usually do not eat while brooding offspring in their mouths. In two epilithic algal eaters Tropheus duboisi and T. moorii in Lake Tanganyika, however, mouthbrooding females exhibited feeding actions. In T. duboisi, the feeding rate of mouthbrooding females was 80 percent of that of males and non brooding females irrespective of the developmental state of their offspring. In T. moori, females brooding early embryos rarely fed but their feeding rate increased with development of offspring. An examination of specimens revealed that such females took food for nourishment of themselves and the young in the former species but for nourishment of only the young in the latter.

title: Social organization of a polygynous CichlidLamprologns furcifer in Lake Tanganyika
abstract: Lamprologus furcifer inhabited vertical or overhanging rock surfaces. Females persistently stayed at particular sites and singly guarded the offspring there. Males also stayed at particular sites but often left them to visit females. A dominant male controlled the territories of several breeding females at a time. Schools of young under maternal care spread horizontally on the rock surface as they developed and fed mainly on benthic organisms such as atyid shrimps. It is suggested that the emancipation of males from brood care is attributed to the benthic life of young.

title: Lunar spawning synchroneity in a freshwater fish
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title: Lunar cyclic spawning of a mouthbrooding cichlid,cyprichromis leptosoma, in lake tanganyika
abstract: L unar reproductive cycles have rarely been documented in freshwater fishes (Schwanck, 1987; Nakai et al., 19901, although common among marine reef fishes (Robertson et al., 19901. Several hypotheses for lunar synchronous spawning have been proposed, based on the effects of the tidal regime on planktonic egg dispersal or hatching from adult habitats (Robertson et at., 19901. However, these hypotheses cannot be applied to fishes inhabiting freshwater bodies, where the lunar cycle never causes tidal fluctuations. However, some authors have proposed several explanations for the adaptive significance of lunar cyclic reproductive activity in freshwater fishes, based on the effects of moonlight. (l) Moonlight prorides a cue for pairs to spawn at the same time (Schwanck, 19871. (2) Spawning prior to a full moon enhances the effectiveness of nocturnal parental care of the brood (Schwanck, 1987; Nakai et al., 1990; Rossiter, 1991). (3) Dispersal of young during the fourth quarter of the lunar cycle and the new moon improves the survival of young dispersing under the cover of darkness (Nakai et al., 1990). Cichlid fishes that dominate the coastal fish communities in Lake Tanganyika have two well-developed patterns of parental care, substrate brooding and mouthbrooding (Coulter, 1991; Keenleyside, 199 t). In this take, about one third of the cichlid species are substrate brooders and the remainder mouthbrooders (Poll, 19861. Lunar cyclic spawning has been reported in some substrate brooders, but never in mouthbrooders. However, the spawning activity of a mouthbrooding cichlid, Q>prichromis leptosoma, in fact follows the lunar cycle at the southern end of the take. I examined whether or not any of the three explanations for lunar spawning synchroneity are applicable to this species, and herein propose an alternative explanation that considers interspecific relationships between juvenile C. leptosoma and substrate brooding species cohabiting the area.

title: Female-to-male shift of mouthbrooding in a cichlid fish, Tanganicodus irsacae, with notes on breeding habits of two related species in Lake Tanganyika
abstract: SynopsisParental-care patterns and mating systems of three goby-like cichlids in Lake Tanganyika were investigated. In Tanganicodus irsacae females mouthbrooded eggs and small young for about two weeks and then males took over the role for about one week. Field observations of tagged fish suggest that this species is monogamous: a male's home range largely overlapped with that of its mate, while their home ranges were segregated from those of similar-sized consexual adults. Eretmodus cyanostictus also performed female-to-male shift of mouthbrooding and appeared to be monogamous. The third species, Spathodus marlieri, however, exhibited exclusively maternal mouthbrooding. The differences in parental care and mating system among the three species are discussed in relation to their feeding habits, and the pattern of monogamy in the goby-like cichlids is compared with those of other fishes.

title: Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika
abstract: 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.

title: The social and mating system of the maternal mouthbrooderTropheus moorii (Cichlidae) in Lake Tanganyika
abstract: Territorial behaviour, reproduction and migration of the epilithic algal eater,Tropheus moorii, were investigated in Lake Tanganyika, Africa. Adults of both sexes had individual feeding territories which adjointed each other. Males, who occupied higher rocks than females, usually stayed at the same sites for more than 5 months. Females left their territories to pair with males in the males’ territories. Paired females actively foraged under the protection of their mates for up to 3 weeks prior to spawning. After spawning, females usually settled in a site unoccupied by territory-holders to mouthbrood the offspring for a month. An examination of the ovaries and a removal experiment of dominant males suggest that females cannot attain fully mature ovaries in their own territories and choose males whose territories can provide enough food to satisfy their nutritive demand. The evolution of a number of local colour morphs in this fish is briefly discussed in relation to social selection.

title: Nuclear and mitochondrial data reveal different evolutionary processes in the Lake Tanganyika cichlid genus Tropheus
abstract: BackgroundCichlid fishes are notorious for their wealth of intra- and interspecific colour pattern diversity. In Lake Tanganyika, the endemic genus Tropheus represents the most impressive example for geographic variation in the pattern and hue of integument colouration, but the taxonomy of the over 100 mostly allopatric colour morphs remains to a large degree unresolved. Previous studies of mitochondrial DNA sequence data revealed polyphyly of the six nominally described species and complex phylogeographic patterns influenced by lake level fluctuations and population admixture, and suggested the parallel evolution of similar colour patterns in divergent evolutionary lineages. A gene tree of a rapidly radiating group may be subject to incomplete and stochastic lineage sorting, and to overcome this problem we used multi-locus, nuclear AFLP data in comparison with mtDNA sequences to study diversification, migration and introgression in Tropheus colour morphs in Lake Tanganyika.ResultsSignificant incongruence between phylogenetic reconstructions from mitochondrial and AFLP data suggested incomplete sorting of mitochondrial haplotypes as well as frequent introgression between differentiated lineages. In contrast to the mitochondrial phylogeny, the AFLP phenogram was largely congruent with species classifications, colour pattern similarities, and in many cases also with the current geographic distribution of populations, and did not produce evidence of convergent colour pattern evolution. Homoplasy in the AFLP data was used to identify populations that were strongly affected by introgression.ConclusionDifferent evolutionary processes were distinguished by the combination of mitochondrial and AFLP data. Mitochondrial phylogeographic patterns retained signals of large-scale migration events triggered by historical, major lake level fluctuations, whereas AFLP data indicated genetic cohesion among local groups of populations resulting from secondary contact of adjacent populations in the course of the more frequently occurring, minor lake level fluctuations. There was no support for the parallel evolution of similar colour patterns in the AFLP data. Genetic signatures of introgression and hybridisation detected in several populations suggest that lake level fluctuations drove the stunning diversification of Tropheus morphs not only through population fragmentation, but also by promoting hybridisation between differentiated morphs in secondary contact.

title: Polychromatism of the scale-eaterPerissodus microlepis (Cichlidae, Teleostei) in relation to foraging behavior
abstract: Variation in foraging behavior in a population of the scale-eaterPerissodus microlepis was studied on the northwest coast of Lake Tanganyika. Differences in body coloration were found among adult and subadult individuals, which were classified into 4 color morphs designated as Beige, Dark, Grey and Stripe. These color morphs were not strictly related to either sex or size. Each morph spent much time in specific microhabitats and had a major hunting technique that differed from other morphs. Beige morph. which predominated in number, ambushed prey at open surfaces of the substrate, whereas Dark morph used the shade of rock as an ambush site. Grey morph mixed in schools of fisheds hovering in midwater to attack school members, and Stripe morph cruised in the water column and stooped mainly at bottom-fishes. Prey preference differed among the morphs corresponding to their hunting techniques but successful attack rates were similar among them. Observations of marked individuals demonstrated adherence to particular hunting techniques and, in some cases, to particular hunting sites. Intraspecific foraging specialization is discussed in relation to the function of body color and diversity of life styles of prey fishes.

title: Influence of brood size and offspring size on parental investment in a biparental cichlid fish,Neolamprologus moorii
abstract: Both males and females ofNeolamprologus moorii, a substrate-spawning cichlid fish in Lake Tanganyika, tend their eggs and fry. In this study, the influence of brood size and fry size on parental investment of this species was examined under natural conditions. Breeding parents intensively attacked fry-eating fishes that approached their broods. The attack rate by the parents against the approaching fry-eaters was positively correlated with brood size and negatively with fry size, but was much more strongly related to brood size. This suggests that the parental decision for brood defense appeared to be primarily determined by brood size. The reason for fry size being less important in the parental decision may be that even large fry have a low survival ability on their own under the high predation pressure in the study sites.

title: Variation of territory size and defense behavior in breeding pairs of the endemic Lake Tanganyika cichlid fish Variabilichromis moorii
abstract: Tanganyika harbors the ecologically, morphologically, and behaviorally most diverse species flock of cichlid fishes. It is comprised by substrate breeding and mouthbrooding species, most of which live in littoral habitats. Species communities are characterized by complex behavioral and trophic interactions, resulting in a dense pattern of partially overlapping territories, depending on the degree of ecological distinctness. We studied territorial behavior of breeding pairs in a substrate breeding species, with respect to territory size and defense behavior. The study species Variabilichromis moorii belongs to the tribe Lamprologini, the most species rich tribe of cichlids in Lake Tanganyika. Our study shows that breeding pairs of V. moorii can have highly complex territories, in which both parents hold separate sub-territories which are shifted slightly according to the movements of the fry, but the outer borders are conjointly defended. The size of the total defended territory varied from <1 to almost 4 m2, averaging at about 2 m2. Depending on presence of competitors or fry-predators evoking agonistic interactions, the territory size varied quite substantially over the day. Attack rates and size of the defended area decreased with water depth. Agonistic behavior was observed toward heterospecifics as well as conspecifics, with heterospecific attacks mostly concerning territorial neighbors and potential fry-predators in about equal frequencies.

title: PARENTAL ROLES AND THE AMOUNT OF CARE IN A BI-PARENTAL SUBSTRATE BROODING CICHLID: THE EFFECT OF SIZE DIFFERENCES WITHIN PAIRS
abstract: Parental roles and the amount of care in bi-parental fish have been assumed to be determined by sex. We studied the parental behaviour in a substrate brooding cichlid Julidochromis ornatus (40-90 mm in total length) in which both parents participate in care of eggs and young. In the study population, ca 80% of paired females were larger than their partners and pairs mated assortatively for size. Males spent more time with their offspring in female-largest pairs, whereas the opposite was found for male-largest pairs. These differences in the amount of care were more conspicuous when differences in body size were greater, whereas similar sized pairs shared parental tasks. These results suggest that the amount of parental care is largely affected by the relative size within pairs independent of sex in J. ornatus .H owever, the frequencies of defensive behaviours were not different in both female-largest and malelargest pairs. This indicates that parental roles would not completely change as the change of the body size difference within pairs. The larger parents were socially dominant over the partners regardless of sex, and observations of aggressive behaviours within pairs suggest that the larger fish are likely to make the partners perform parental care. Higher frequencies of aggressive encounters were observed in the similar-sized pairs than in the different-sized ones. Higher costs associated with frequent aggressive behaviours in the similar-sized pairs may be related to their small brood size, and may be partly responsible for the size-assortative mating with sexual size difference in this fish.

title: Alternative Male Mating Tactics of the Substrate Brooding Cichlid Telmatochromis temporalis in Lake Tanganyika
abstract: Telmatochromis temporalis is a bi-parental substrate brooding cichlid endemic to Lake Tanganyika. Paired males were always larger than their mates and had territories around nests against con-specific males. However, males smaller than the paired females were found in 18% of the nests. Here we report a reproductive tactic of these small males. The small males had as heavy gonads as paired males, and the gonad somatic index (GSI) of the small males was much higher than that of the latter. The examinations of the paternity and maternity using microsatellite-DNA as a genetic marker revealed that the small males were not genetically related to the pair members, and sired some young in 3 of 5 nests. These small males did not guard the broods, suggesting that they are likely to perform reproductive parasitism as sneakers. Paired males could not enter their spawning nests due to their large size, which made it difficult to chase out sneakers once they entered the nest. Some males as small as the sneakers were found outside the territories of paired males, and their gonads were quite small. Circumstantial evidence suggests that small males have two alternative investment patterns: investing in gonad to be sneakers, and investing in growth to probably be territorial males.

title: Active accumulation of spawning substrate: a determinant of extreme polygyny in a shell-brooding cichlid fish
abstract: Observations on the reproductive behaviour and polygynous mating system of an endemic shell-brooding cichlid fish, Lamprologus callipterus , revealed that territorial males actively transported and accumulated spawning substrate (empty gastropod shells) to make nests. In contrast to conventional resource defence polygyny, males of this species can therefore create a defensible resource distribution. Females successively visited the nests, chose one shell, spawned the whole clutch inside the shell and stayed inside until the young became independent. Males were freed from direct parental care but defended the territory. Nest usurpation by larger males and frequent shell stealing among males were observed; these resulted in cannibalism of the young fathered by the former males. Large males accumulated more shells and consequently monopolized more females than did smaller males. Maximum size of females was apparently limited by the size of available shells, resulting in extreme sexual dimorphism. Extreme polygyny in this species, associated with intense intra-sexual competition and extreme sexual dimorphism, was based mainly on transportability of spawning substrate and the specific behavioural competence of males to use it.

title: Social status-dependent nest choice of territorial males under reproductive parasitism in a Lake Tanganyika cichlid Telmatochromis vittatus.
abstract: Field and laboratory studies were conducted to examine how territorial males of a Lake Tanganyika cichlid Telmatochromis vittatus balance the conflicting demands on nest choice between occupying large nests with more females and avoiding reproductive parasitism (nest piracy, which is adopted by the largest males in the population). Pirates less frequently intruded the nests farther from neighbours, perhaps due to the costs associated with travelling between nests. The field experiment showed that territorial male T. vittatus sacrificed the fitness benefits that large nests offer and instead prioritized occupying the nests farther from neighbours on which fewer pirates intruded. The field observations suggested that they adopt different strategies for nest choice according to their relative competitive ability to pirates; the large territorial males, who are size-matched to pirates and can defend their nests against them, compete for larger nests among the more-isolated nests, whereas subordinate territorial males, which are smaller than pirates and thus inferior to them, compete for the more-isolated nests among the less-isolated nests. These findings suggest that the territorial male T. vittatus chooses the more-isolated nests to avoid pirate males at the expense of occupying large nests.

title: Genetic mating systems and reproductive natural histories of fishes: lessons for ecology and evolution.
abstract: Fish species have diverse breeding behaviors that make them valuable for testing theories on genetic mating systems and reproductive tactics. Here we review genetic appraisals of paternity and maternity in wild fish populations. Behavioral phenomena quantified by genetic markers in various species include patterns of multiple mating by both sexes; frequent cuckoldry by males and rare cuckoldry by females in nest-tending species; additional routes to surrogate parentage via nest piracy and egg-thievery; egg mimicry by nest-tending males; brood parasitism by helper males in cooperative breeders; clutch mixing in oral brooders; kinship in schooling fry of broadcast spawners; sperm storage by dams in female-pregnant species; and sex-role reversal, polyandry, and strong sexual selection on females in some male-pregnant species. Additional phenomena addressed by genetic parentage analyses in fishes include clustered mutations, filial cannibalism, and local population size. All results are discussed in the context of relevant behavioral and evolutionary theory.

title: High frequency of multiple paternity in broods of a socially monogamous cichlid fish with biparental nest defence
abstract: In several animal taxa, genetic analyses have demonstrated that social monogamy and biparental brood care do not preclude polygamous reproduction. Few studies have been conducted in fish, but in fish species without alternative reproductive phenotypes, social monogamy was largely congruent with genetic parentage. In contrast to these findings, we report an exceptionally high level of multiple paternity in a socially monogamous cichlid fish with biparental nest defence (Variabilichromis moorii), inferred from microsatellite and mitochondrial data of 10 broods. Whereas all offspring in a nest shared a common mother, each brood was sired by 2 to > 10 males. None of the inferred sires was assigned a large proportion of the brood. Paternity was estimated as the minimum number of sires required to explain multilocus offspring genotypes, and as the maximum‐likelihood number of sires given population allele frequencies. Analysis of simulated brood genotypes suggested that, although these two methods tend to under‐ and overestimate, respectively, the true number of sires, primary sires with many offspring in a brood would have been detected. Hence, the genetic data indicate that the nest tending males suffer substantial cuckoldry and provide alloparental care for a large number of unrelated fry. We have no data on the social status of the cuckolding males, but due to synchronous spawning of pairs and commitment to brood care of paired males, it is possible that most of the parasitic spawners are solitary males.

title: Morphological and genetic distinctness of rock and shell-bed dwelling Telmatochromis(Teleostei, Cichlidae) in the south of Lake Tanganyika suggest the existence of two species
abstract: Two populations from Wonzye and Nkumbula Island, Lake Tanganyika, containing both rock and shell-bed dwelling individuals of the genus Telmatochromis were analysed by means of comparative morphological and DNA haplotype data from the mitochondrial control region. The data on individuals from rock v. shell-bed habitats allowed a clear discrimination to be made between habitats, while there was no distinction among individuals of the same habitat from Wonzye and Nkumbula. The shell-bed dwellers had small body size, including smaller size at sexual maturity and fewer eggs than rock dwellers. The existence of two distinct species that diversified according to habitat choice is suggested.

title: Evidence for divergent natural selection of a Lake Tanganyika cichlid inferred from repeated radiations in body size
abstract: Divergent natural selection is thought to play a vital role in speciation, but clear, measurable examples from nature are still few. Among the many possible sources of divergent natural selection, predation pressure may be important because predators are ubiquitous in food webs. Here, we show evidence for divergent natural selection in a Lake Tanganyika cichlid, Telmatochromis temporalis, which uses burrows under stones or empty snail shells as shelters. This species contains normal and dwarf morphs at several localities. The normal morph inhabits rocky shorelines, whereas the dwarf morph invariably inhabits shell beds, where empty snail shells densely cover the lake bottom. Genetic evidence suggested that the dwarf morph evolved independently from the normal morph at two areas, and morphological analysis and evaluation of habitat structure revealed that the body sizes of morphs closely matched the available shelter sizes in their habitats. These findings suggest that the two morphs repeatedly evolved through divergent natural selection associated with the strategy for sheltering from predators.

title: Reticulate phylogeny of gastropod-shell-breeding cichlids from Lake Tanganyika – the result of repeated introgressive hybridization
abstract: BackgroundThe tribe Lamprologini is the major substrate breeding lineage of Lake Tanganyika's cichlid species flock. Among several different life history strategies found in lamprologines, the adaptation to live and breed in empty gastropod shells is probably the most peculiar. Although shell-breeding arose several times in the evolutionary history of the lamprologines, all obligatory and most facultative shell-breeders belong to the so called "ossified group", a monophyletic lineage within the lamprologine cichlids. Since their distinctive life style enables these species to live and breed in closest vicinity, we hypothesized that these cichlids might be particularly prone to accidental hybridization, and that introgression might have affected the evolutionary history of this cichlid lineage.ResultsOur analyses revealed discrepancies between phylogenetic hypotheses based on mitochondrial and nuclear (AFLP) data. While the nuclear phylogeny was congruent with morphological, behavioral and ecological characteristics, several species – usually highly specialized shell-breeders – were placed at contradicting positions in the mitochondrial phylogeny. The discordant phylogenies strongly suggest repeated incidents of introgressive hybridization between several distantly related shell-breeding species, which reticulated the phylogeny of this group of cichlids. Long interior branches and high bootstrap support for many interior nodes in the mitochondrial phylogeny argue against a major effect of ancient incomplete lineage sorting on the phylogenetic reconstruction. Moreover, we provide morphological and genetic (mtDNA and microsatellites) evidence for ongoing hybridization among distantly related shell-breeders. In these cases, the territorial males of the inferred paternal species are too large to enter the shells of their mate, such that they have to release their sperm over the entrance of the shell to fertilize the eggs. With sperm dispersal by water currents and wave action, trans-specific fertilization of clutches in neighboring shells seem inevitable, when post-zygotic isolation is incomplete.ConclusionFrom the direct observation of hybrids we conclude that hybridization between distantly related gastropod-shell-breeding cichlids of Lake Tanganyika follows inevitably from their ecological specialization. Moreover, the observed incongruence between mtDNA and nuclear multilocus phylogeny suggests that repeated hybridization events among quite distantly related taxa affected the diversification of this group, and introduced reticulation into their phylogeny.

title: Sperm competition in fish: `bourgeois' males and parasitic spawning.
abstract: Fish exhibit an enormous variety of reproductive patterns. There is external and internal fertilization, simultaneous and sequential hermaphroditism as well as gonochorism, and an extremely widespread occurrence of parasitic reproductive behaviour among males. In most fish species there is a great size range of reproductive males, setting the stage for divergent, intraspecific reproductive patterns and an unparalleled concentration of alternative male reproductive phenotypes. Recent theoretical, empirical and comparative evidence suggests that adaptations to sperm competition in fish might be responsible for some of the most intriguing examples of reproductive design known.

title: Giant males or dwarf females : what determines the extreme sexual size dimorphism in Lamprologus callipterus?
abstract: In the Lake Tanganyika cichlid Lamprologus callipterus, males were >12 times heavier than females, the most extreme sexual size dimorphism in this direction among animals. L. callipterus males construct nests of empty snail shells in which the females breed. If the ancestors of L. callipterus were small cichlids, both sexes may have used shells for shelter. If the ancestors were larger, snail shells could not be used as shelters and would be important only for reproduction. In the field and the laboratory, females bred only in shells and the largest spawned in the largest shells. In the field, females chose larger shells than the average available in males' territories and did not copy the mate choice of other females. They never hid from predators in snail shells and they occurred commonly in areas without any potential shell shelters. In laboratory experiments females used shells only for reproduction and hardly for shelter. Therefore, it seems unlikely that L. callipterus descended from small shell-brooding cichlids which used shells for shelter, but more likely that the ancestors were of large or intermediate size, and that female size is constrained by the sizes of snail shells, which appear to be optimal breeding substrata.

title: The evolution of bourgeois, parasitic, and cooperative reproductive behaviors in fishes.
abstract: Among vertebrate classes, fishes exhibit by far the greatest variability in competitive and cooperative behaviors in male reproduction. Scramble competition between reproductive males is one possibility. Another possibility occurs when resources, mates, or locations can be monopolized, in which case males may invest in primary access to fertilizations by adopting a "bourgeois" strategy, or they may employ alternative mating tactics to evade the reproductive monopoly of other males. Adaptations in morphology, physiology, and behavior to bourgeois and alternative phenotypes are highly divergent. Here I review the functional characteristics that differ between bourgeois and parasitic phenotypes, and discuss the variability of alternative reproductive tactics at the levels of plasticity, determination, and selection. Examples will illustrate the importance of ecology, and will suggest that variation in reproductive tactics is largely adaptive. Behavioral solutions to competition for mates and fertilizations often involve agonistic behavior and conflict, but also cooperation among competitors (e.g., when subordinate males pay a price to bourgeois males for gaining access to fertilizable eggs). Application of molecular genetic tools has helped to uncover intricate sexual and social relationships in various fish species, including species that display some of the most complex reproductive and social patterns known among the vertebrates.

title: Nest use by territorial males in a shell-brooding cichlid: the effect of reproductive parasitism
abstract: Male Telmatochromis vittatus, a substrate-brooding Tanganyikan cichlid, exhibit two parasitic reproductive tactics: takeover of spawning by larger males (“pirates”) and sneaking by smaller males (“sneakers”). Medium-sized males are territorial and pair-spawn within nests of clumped shells that harbor several resident females that are potential mates of the territory owner. To study nest use by territorial males, we analyzed the relationship between the body size of territorial males and nest quality, the number of females per nest, the distance between nests, and the frequency and intensity of reproductive parasitism. The size of males was not correlated with nest quality, the number of females, or the number of sneakers, but was negatively correlated with the frequency of intrusion by pirates and was positively correlated with the distance between nests. Territorial males effectively defended nests against sneakers but failed to defend against pirates. These results suggest that larger territorial males selected nests that have a lower risk of usurpation of spawning. We hypothesize that the risk of intrusion by pirate males affects the selection of nests by territorial males in this species.

title: Female-biased immigration and male peace-keeping in groups of the shell-dwelling cichlid fish Neolamprologus multifasciatus
abstract: The small cichlid fish Neolamprologus multifasciatus is endemic to Lake Tanganyika and is unique among fish because it lives in complex social groups with several reproductive males and females, all of which participate in defending their territory against neighbors and intruders. Individuals use empty snail shells for breeding and shelter. Previous parentage analysis using microsatellites suggested occasional exchange of individuals between groups. In field experiments, we found that females showed a higher tendency than males to migrate into territories already occupied by a resident pair. The phenomenon and causes of female-biased immigration were further investigated in aquarium experiments: Nine of 15 females, but only 2 of 15 equally sized males, settled in territories of established pairs. Territorial males exhibited more aggression toward strange males (potential reproductive competitors) than toward strange females (potential additional mates); their females were more aggressive toward strange females (probably competitors for shells) than toward strange males. Apparently, a conflict exists between the sexes regarding the immigration of additional females. This conflict seems to be the selective basis for observed active male interference in aggressive disputes between females in the territory. Interfemale tolerance in a group was greater in the male’s presence than in his absence.

title: Female aggression and male peace-keeping in a cichlid fish harem: conflict between and within the sexes in Lamprologus ocellatus
abstract: SummaryConflicts of interest within and between the sexes are important processes leading to variability in mating systems. The behavioral interactions mediating conflict are little documented. We studied pairs and harems of the snail-shell inhabiting cichlid fish Lamprologus ocellatus in the laboratory. Due to their larger size, males controlled the resource that limited breeding: snail shells. Males were able to choose among females ready to spawn. Females were only accepted if they produced a clutch within a few days of settling. When several females attempted to settle simultaneously the larger female settled first. Females were least aggressive when guarding eggs. Secondary females were more likely to settle when the primary female was guarding eggs. In established harems females continued to be aggressive against each other. The male intervened in about 80% of female aggressive interactions. Male intervention activity correlated with the frequency of aggression among the females in his harem. The male usually attacked the aggressor and chased her back to her own snail shell. When a male was removed from his harem, aggression between females increased immediately and usually the secondary female was expelled by the primary female within a few days. Time to harem break-up was shorter the more mobile the primary females' young were and did not correlate with the size difference between harem females. Male L. ocellatus interfere actively in female conflict and keep the harem together against female interests. Female conflict presumably relates to the cost of sharing male parental investment and to the potential of predation by another female's large juveniles on a female's own small juveniles.

title: Phylogenetic relationships of the lamprologine cichlid genus Lepidiolamprologus (Teleostei: Perciformes) based on mitochondrial and nuclear sequences, suggesting introgressive hybridization.
abstract: Using sequences of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2, 1047bp) and a segment of the non-coding mitochondrial control region, as well as nuclear sequences including two introns from the S7 ribosomal protein and the loci TmoM25, TmoM27, and UME002, we explore the phylogenetic relationships of Lepidiolamprologus, one of seven lamprologine cichlid genera in Lake Tanganyika, East Africa. Analyses consisted of direct optimization using POY, including a parsimony sensitivity analysis, and maximum likelihood and Bayesian inference for comparison. With respect to Lepidiolamprologus, the results based on the mitochondrial dataset were robust to parameter variation in POY. Lepidiolamprologus cunningtoni was resolved in a large clade sister to ossified group lamprologines, among which the remaining Lepidiolamprologus were nested. In addition to L. attenuatus, L. elongatus, L. kendalli, and L. profundicola, Neolamprologus meeli, N. hecqui, N. boulengeri, N. variostigma, and two undescribed species were resolved in a two-pore Lepidiolamprologus clade sister to Lamprologus callipterus and two species of Altolamprologus. Lepidiolamprologus nkambae, in marked conflict with morphological and nuclear DNA evidence, nested outside of the two-pore Lepidiolamprologus clade, suggesting that the mtDNA signal has been convoluted by introgressive hybridization.

title: Mating system and kin relationship between adults and young in the shell-brooding cichlid fish Neolamprologus meeli in Lake Tanganyika
abstract: 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.

title: Mothers determine offspring size in response to own juvenile growth conditions
abstract: Through non-genetic maternal effects, mothers can tailor offspring phenotype to the environment in which young will grow up. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult environment of mothers. In this case maternal decisions about investment in offspring quality should already be determined during the juvenile phase of mothers. I tested this hypothesis by manipulating juvenile and adult maternal environments independently in a cichlid fish. Females raised in a poor environment produced larger young than females raised without food limitations, irrespective of the feeding conditions experienced during adulthood. This maternal boost was due to a higher investment in eggs and to faster larval growth. Apparently, mothers prepare their offspring for similar environmental conditions to those they encountered as juveniles. This explanation is supported by the distribution of these fishes under natural conditions. Juveniles live in a different and much narrower range of habitats than adults. Therefore, the habitat mothers experienced as juveniles will allow them to predict their offspring's environment better than the conditions in the adult home range.

title: Hypoxia and energetics of mouth brooding: is parental care a costly affair?
abstract: This study explores costs of mouth brooding and the response of parent and offspring to brooding under hypoxia in the maternal African mouth brooder Pseudocrenilabrus multicolor. Fish of swamp origin were acclimated to hypoxia (1.42 mg oxygen L(-1)) or normoxia (8.00 mg oxygen L(-1)) for a minimum of 6 months prior to measures of metabolic rate and embryo traits. Regardless of brooding stage, standard metabolic rates were lower in females acclimated to low dissolved oxygen (DO) compared to high-DO acclimated females. Regardless of DO acclimation treatment, standard metabolic rates were approximately 48% higher in brooding females (with the estimated metabolic rate of the brood removed) compared to post-brooding females. There was no difference in brood maintenance metabolism, female relative condition, embryo size, and embryo number between fish acclimated to low DO vs. high DO. However, the length of the brooding period (from egg-laying to release of fry) was approximately 27% shorter in females acclimated to low DO compared to females acclimated to high DO suggesting accelerated development in offspring brooded under hypoxia. These findings demonstrate a cost to mouth brooding and provide evidence for a parental strategy to deal with the expense of providing offspring care under hypoxia.

title: Food intake by mouthbrooding females ofCyphotilapia frontosa (Cichlidae) to feed both themselves and their young
abstract: SynopsisFeeding behaviour of mouthbrooding females ofCyphotilapia frontosa was observed in their natural habitat, and specimens of mouthbrooding females and the young in their mouths were examined in the laboratory. Mouthbrooding females exhibited feeding actions and their guts contained about one quarter as much food as those of nonbrooding adults. A substantial amount of food was found in young 12.5 mm TL who retained a large quantity of yolk, and gut fullness of young increased as they grew. Weight changes of the young suggested that the buccal feeding augmented their growth.

title: Reproductive biology of Eretmodus cyanostictus, a cichlid fish from Lake Tanganyika
abstract: One of the most widely accepted explanations for monogamy is the need for biparental care. However, the occurrence of monogamy combined with biparental care is extremely rare in oral incubating (mouthbrooding) cichlid fishes. Few studies have been performed on cichlid species that exhibit this behaviour, and therefore the ecological factors that favour monogamy in these cases remain obscure. Here we present new information on the natural history and reproductive biology of Eretmodus cyanostictus (Boulenger 1898), a monogamous biparental mouthbrooder from Lake Tanganyika. The populations studied consisted of territorial pairs and a male-biased non-territorial population of smaller 'floater' individuals. We present the first detailed description of spawning in this species, show that breeding does not appear to be synchronised within the population, and provide evidence that parental care is costly. We discuss the implications of this information for our understanding of monogamy.

title: The functional significance of buccal feeding in the mouthbrooding cichlid Tropheus moorii
abstract: The mouthbrooding cichlid Tropheus moorii exhibits an exceptional degree of maternal investment. Females produce very large eggs, which are incubated for a period of six weeks, and they feed their young in the mouth while starving themselves. Only very few fish species are known to feed their young. We hypothesized that feeding may either (1) benefit females directly if young develop faster, or (2) provide a size advantage to young. To distinguish between these mutually non-exclusive hypotheses, we measured costs and benefits arising for mouthbrooding females when feeding their young. Field observations revealed that mouthbrooding females reduced bite rates, locomotion and territorial defence compared to non-brooding adults. Feeding rates correlated positively with locomotion, as fish moving around more also spent more time with territory defence and other social interactions. This suggests that buccal feeding is costly when compared to mere incubation without feeding the young. In an experiment, in which we controlled the access of females to food, we showed that these costs are apparently not counterbalanced by a benefit to females through a shorter incubation duration. Rather, fed young were larger, heavier and had higher burst-swimming speeds. The extreme maternal investment of T. moorii appears to yield fitness benefits to females by producing larger and stronger young, which consequently should have better survival chances in their natural environment.

title: Long-term territory maintenance by female Tropheus duboisi (Cichlidae) involving foraging during the mouth-brooding period
abstract: The herbivorous cichlid fish Tropheus duboisi in Lake Tanganyika held feeding territories that were defended against individuals of the same but not the opposite sex. The male home ranges were 3 times as large as female home ranges, each overlapping with the ranges of 2–5 females. After spawning in the male's territory, each female foraged actively within her own territory while brooding the young in the buccal cavity. Both males and females rarely shifted their territories during the 6-month observation period. The social organization of this fish was compared with that of a congener, Tropheus moorii.

title: Systematics of Tanganyikan cichlid fishes (Teleostei: Perciformes)
abstract: The relationships among 53 genera of Tanganyikan cichlid fishes were analyzed based on internal and external morphological features. Comparison of the morphological cladistic tree with a previously proposed classification showed 5 of 12 tribes to be nonmonophyletic. Sixteen tribes were recognized, the changes in classification being that Trematocarini was treated as a junior synonym of Bathybatini; 5 new tribes were established for each of the following genera, Benthochromis, Boulengerochromis, “Ctenochromis” benthicola, Cyphotilapia, and Greenwoodochromis; “Ctenochromis” horei was transferred from Haplochromini to Tropheini; and “Gnathochromis” pfefferi was transferred from Limnochromini to Tropheini. The revised classification was supported by previously proposed molecular trees.

title: Biparental mouthbrooding and guarding in a Tanganyikan CichlidHaplotaxodon microlepis
abstract: Differences between parental roles of males and females inHaplotaxodon microlepis (Cichlidae) were investigated in Lake Tanganyika, and the early ontogeny and growth of the species were studied in the aquarium. Eggs were mouthbrooded by the female, and it is suggested that small larvae (<9 mm in total length) were also mouthbrooded by females though such samples were not collected. Above this size the larvae began to feed, and parents jointly performed mouthbrooding and guarding until the young grew to 25–30 mm, nearly 2 months after spawning. Males and females mouthbrooded to the same extent, but when a part of the brood was released, females mainly guarded the released brood and males took the mouthbrooding role. Differences in parentalcare patterns betweenH. microlepis and other monogamous mouthbrooding cichlids are discussed.

title: Parental care in a monogamous mouthbrooding cichlidXenotilapia flavipinnis in Lake Tanganyika
abstract: Breeding pairs ofXenotilapia flavipinnis held their territories on the sandy bottom and repeated several breeding cycles. Females mouthbrooded the eggs and early larvae but afterward males took over the mouthbrooding role. When the young became free swimming, they were guarded by both parents and remained in the spawning territory. Males played a leading role in the guarding, while females were more active in foraging during the guarding period. It was concluded that males’ active participation in the parental care could accelerate the gonadal recovery of females and consequently could maximize the fecundity of serially monogamous pairs.

title: Additions and amendments to the classification of reproductive styles in fishes
abstract: SynopsisThe guild concept is explained to represent peaks in the landscape of reproductive styles, which respond to the ‘altricial ⇄ precocial principle’ caused by the processes of heterochrony and saltation. Guilds represent evolutionary trends but not units. New evidence for a specialization within the mouthbrooding external bearers is reviewed, e.g. the unique specialization of juvenile feeding inside the female's buccal cavity. Ability to withstand extreme environments is revealed in some water-edge spawners and a new guild of nonguarding aerophils is distinguished. The walleye is placed into another guild and the significance of spawning inside of sponges discussed. Due to a better understanding of reproductive styles used by some external bearers, termed earlier as forehead, skin and possibly armpit and backpack brooders, a single guild of auxiliary brooders is named. The classification of internal bearers is revised into four redefined and renamed guilds — facultative internal bearers, obligate lecithotrophic livebearers, matrotrophous oophages and adephophages, and viviparous trophoderms.

title: Female mouthbrooders adjust incubation duration to perceived risk of predation
abstract: The protection of young from predators is an important and costly component of parental care, and this investment should be adjusted to the predation risk for the offspring. Parents are known to fine-tune defence intensity to the perceived need of protection, yet little attention has been paid to individual variation in the duration of brood defence. To optimize parental investment it may be beneficial to prolong brood care when the predation risk for the young is high. We tested whether cichlids of the species Ctenochromis horei adjust mouthbrooding duration to the perceived predation risk of their offspring. After spawning, females were transferred to an experimental tank, where they incubated the clutch either alone or in the presence of a natural predator of young. Females extended their incubation period by 4.3 days on average when the predator was present. During this prolonged incubation, young continued to grow in the female's buccal cavity. Interspawning interval was longer for females that incubated eggs than for those that did not, although incubation had no long-term negative effects on female condition or egg number or size. Incubating females may thus breed less frequently. Brood care duration should hence be considered as a flexible and potentially costly trait when investigating costs and benefits of parental care.

title: The influence of juvenile and adult environments on life-history trajectories
abstract: There is increasing evidence that the environment experienced early in life can strongly influence adult life histories. It is largely unknown, however, how past and present conditions influence suites of life-history traits regarding major life-history trade-offs. Especially in animals with indeterminate growth, we may expect that environmental conditions of juveniles and adults independently or interactively influence the life-history trade-off between growth and reproduction after maturation. Juvenile growth conditions may initiate a feedback loop determining adult allocation patterns, triggered by size-dependent mortality risk. I tested this possibility in a long-term growth experiment with mouthbrooding cichlids. Females were raised either on a high-food or low-food diet. After maturation half of them were switched to the opposite treatment, while the other half remained unchanged. Adult growth was determined by current resource availability, but key reproductive traits like reproductive rate and offspring size were only influenced by juvenile growth conditions, irrespective of the ration received as adults. Moreover, the allocation of resources to growth versus reproduction and to offspring number versus size were shaped by juvenile rather than adult ecology. These results indicate that early individual history must be considered when analysing causes of life-history variation in natural populations.

title: Egg size and food abundance interactively affect juvenile growth and behaviour
abstract: Comparative evidence from several animal taxa suggests that juveniles hatching from larger eggs have fitness benefits when growing up in a harsh environment, whereas under benign conditions egg size should be of less importance. However, the physiological and behavioural mechanisms responsible for these context-dependent fitness differences are as yet poorly understood. 2. We studied the interactions between the phenotype of developing offspring and their environment in the mouthbrooding cichlid Simochromis pleurospilus. We hand-raised young from large and small eggs, and measured their initial body size and burst swimming speed. Thereafter we raised half of each egg-size class on high ration and half on low food ration and followed their growth trajectories and behavioural development until the age of 12 weeks. 3. We found that larger eggs gave rise to larger young that had a higher burst swimming speed. Food ration greatly influenced long-term growth, while egg size predominantly affected fish size during the first 2 weeks of life. However, large egg size caused a size advantage of juveniles persisting throughout the experimental period. 4. Egg size and food ration interactively affected the hiding and foraging behaviour of young. In the low-food treatment, individuals from small eggs spent less time in shelter and showed a higher commitment to foraging than individuals from large eggs. In a natural setting, this should markedly increase predation risk of young originating from small eggs, particularly in poor environments. In contrast, when food was plentiful juveniles behaved similarly, irrespective of egg size. 5. Our results show that egg size affects juvenile growth trajectories and behaviour differently in different environments. While it is well-established that a large egg size raises offspring fitness particularly in harsh environments, our study suggests that this advantage arises through riskaverse behaviour being tightly linked to offspring size.

title: Mating system variability in a mouthbrooding cichlid fish from a tropical lake
abstract: Intraspecific variability in mating behaviour and disparities between social and reproductive behaviour add complexity to the description of animal mating systems. A previously published field study on a population of the maternally mouthbrooding cichlid fish Ctenochromis horei in the north of Lake Tanganyika suggested mate monopolization by the most dominant male. In the present study, genetic reconstructions of paternity in a population in the south of the lake provided no evidence for male mate monopolization, as none of the inferred sires had offspring in more than one brood. The ability to reconstruct sire genotypes from offspring alleles was confirmed by computer simulations. Multiple paternity occurred in several broods and was significantly more frequent in the sample taken in the rainy season than in the dry season sample (100% of broods vs. 14%, respectively; P = 0.005). The data suggest geographical and temporal variation in the mating behaviour of C. horei despite its continuous year‐round reproduction, invariable brood care behaviour and habitat‐specific distribution in the relatively constant and confined environment of a tropical lake. Moreover, our data show that inferences on the distribution of male reproductive success, if based on each geographical or temporal data set alone, would fail to describe the potential for sexual selection in this species.

title: Mating systems of two midwater-spawning cichlids,Cyprichromis microlepidotus andParacyprichromis brieni, in Lake Tanganyika
abstract: The breeding habits of 2 maternal mouthbrooding cichlids,Cyprichromis microlepidotus andParacyprichromis brieni, were investigated in Lake Tanganyika. Although spawning on the substrate in the male's mating territory is prevalent in maternal mouthbrooders, bothC. microlepidotus andP. brieni spawned in the water column. MaleC. microlepidotus established their mating territories in the open water column, while maleP. brieni did so around fixed spawning sites near a vertical rock surface. In both species, females visited male mating territories, departing soon after spawning and collecting the eggs. Sneaking, which was observed only inP. brieni, may be attributed to the presence of refuges for sneakers in this species. FemaleC. microlepidotus deposited their entire clutch of about 9 eggs in one male territory. In contrast, femaleP. brieni divided their clutch of about 11 eggs among several males. After the final egg-release, femaleC. microlepidotus repeatedly approached their mate, with the mouth near the abdomen of the latter (nuzzling), but femaleP. brieni often departed without nuzzling. Males may eject sperm during nuzzling to fertilize eggs inside the female's mouth. However, maleP. brieni is also known to eject sperm near spawning females when the females are not nuzzling. Such behavior seems to be a male countermeasure against female mate infidelity, because males could not ensure paternity of eggs by ejecting sperm only during female nuzzling.

title: Female mate preference for males having long and symmetric fins in the bower-holding cichlid Cyathopharynx furcifer
abstract: Female mate preference in a bower-holding cichlid, Cyathopharynx furcifer, was studied in Lake Tanganyika. Most males held territories with crater-shaped bowers in sand, but some males held territories without bowers. Territories were distributed adjacently and females visited them to spawn. After engaging in circling behaviour with the male, a female deposited eggs in the bower. Soon after spawning, the female picked the eggs up into her mouth and brooded them in places away from male territories. Female mate choice appeared to follow three steps: 1) females visited only bower-holding male territories, and more frequently visited territories of males that performed courtship displays at a higher frequency and had longer pelvic fins; 2) females preferred to start circling with males having longer and more symmetrical pelvic fins; 3) females chose males with more symmetrical pelvic fins as their mates. Less than 7% of females that visited male territories spawned eggs in the bowers. In contrast to other bower-holding species, bower size did not correlate with male reproductive success in C. furcifer. Bowers may therefore be essential as spawning sites or may function as a species recognition character for females. Female choice may be dependent instead on males having long and symmetrical pelvic fins apparent during the circling behaviour carried out in the bowers.

title: Female choice of a non-bodily ornament: an experimental study of cichlid sand craters in Cyathopharynx furcifer
abstract: In many species of East African cichlids, males build sand craters or “bowers” to attract females for spawning. It is commonly assumed that these exaggerated sexual traits have a function similar to bodily ornaments. In non-bodily ornaments, however, the behaviour creating the structure may serve as an additional source of information to potential partners, providing multiple signals for mate choice. We tested whether and to what extent females use these signals for choosing males by observing the individual sampling behaviour of female Cyathopharynx furcifer in the field. In addition, we experimentally manipulated crater characteristics in the field and laboratory. We found that females spawn preferably with owners of large and well-maintained craters, but when crater size was enlarged or reduced, the resulting building activity of crater owners affected female choice more strongly than the manipulated crater size per se. We discuss the importance of multiple signals in species constructing extended phenotypes.

title: Sequential polyandry affords post-mating sexual selection in the mouths of cichlid females
abstract: Females mating with multiple males may obtain direct benefits such as nuptial gifts or paternal care or indirect (i.e. genetic) benefits resulting in higher-quality offspring. While direct benefits are easily identified, it is difficult to determine indirect benefits, and it is hence largely unclear how they are obtained. This is particularly true in species with external fertilisation, where females seem to have little control over fertilisation. In cichlids, most maternal mouthbrooders show sequential multiple mating, where females visit several males for egg deposition. Genetic data revealed that multiple paternity of eggs and young in the mouth of females is common, but behavioural data of female spawning decisions are missing. Here, we test four hypotheses to explain female multiple mating in the maternally mouthbrooding cichlid, Ophthalmotilapia ventralis: (1) fertilisation insurance, (2) genetic bet-hedging, (3) female choice and (4) ‘sperm shopping’ (i.e. induction of sperm competition resulting in sexually selected sperm). Detailed observations of spawning behaviour in the field combined with histological analyses of the male reproductive organs suggest that fertilisation insurance, genetic bet-hedging and pre-mating female choice are unlikely to explain the sequential female multiple mating in O. ventralis. Instead, cryptic female choice by sperm shopping, i.e. post-mating sexual selection, is most compatible with our data and might be the major ultimate cause of multiple mating in females of this species and of mouthbrooding cichlids with maternal care in general. Our study provides new insight into ultimate causes of sequential polyandry in species with external fertilisation, as hitherto post-mating sexual selection by cryptic female choice has been assumed to be incompatible with external fertilisation mechanisms except by components of the ovarian fluid.

title: Reproductive parasitism: male and female responses to conspecific and heterospecific intrusions at spawning in a mouth-brooding cichlid Ophthalmotilapia ventralis.
abstract: A rare form of alternative reproductive behaviour without simultaneous parasitic spawning was observed in Ophthalmotilapia ventralis, a lekking mouth-brooding cichlid from Lake Tanganyika. Floater males attempted to sneak opportunistically into the territory to actively court the female, while the owner (bourgeois male) defended the territory against other potential intruders. Floater males had more body fat than territory owners and generally higher condition factors. In field experiments, the response of bourgeois males and courted females was tested towards floaters and egg predators (a catfish Synodontis multipunctatus) present in the territories. Territory owners responded aggressively particularly to floaters, and female responsiveness to bourgeois male courtship tended to decline when floaters were present. The potential influence of reproductive parasitism on sexual selection in mouth-brooding cichlids is discussed.

title: Monogamy in the maternally mouthbrooding Lake Tanganyika cichlid fish Tropheus moorii
abstract: Supported by evidence for assortative mating and polygynandry, sexual selection through mate choice was suggested as the main force driving the evolution of colour diversity of haplochromine cichlids in Lakes Malawi and Victoria. The phylogenetically closely related tribe Tropheini of Lake Tanganyika includes the genus Tropheus, which comprises over 100 colour variants currently classified into six morphologically similar, polyphyletic species. To assess the potential for sexual selection in this sexually monochromatic maternal mouthbrooder, we used microsatellite-based paternity inference to investigate the mating system of Tropheus moorii. In contrast to haplochromines in Lake Malawi, multiple paternity is rare or even absent in broods of T. moorii. Eighteen of the 19 analysed families were consistent with genetic monogamy, while either a mutation or more than one sire explained the genotype of one offspring in another brood. We discuss the differences in breeding behaviour between T. moorii and the Lake Malawi haplochromines, and evaluate additional factors or alternatives to sexual selection as promoters of colour diversification. A preliminary survey of other Tropheini species suggested that multiple paternity is infrequent in the entire tribe.

title: Faithful fish: territory and mate defence favour monogamy in an African cichlid fish
abstract: Monogamy is thought to occur when the opportunities for polygyny are constrained by strong intra-sexual competition, or when two individuals are needed to defend a territory, or to feed or defend their young. In Eretmodus cyanostictus, a monogamous cichlid from Lake Tanganyika, parents do not feed young and each sex is capable of caring for all the offspring alone. We used a series of field-based removal experiments to investigate what other factors may select for monogamy. We found that single individuals usually retained their territory and gained new mates quickly. Thus, it does not seem to be mate availability or the need for a pair to defend a territory per se that maintains monogamy in this species. However, the new mates of widowed individuals were smaller than the original mates had been, and single fish were not capable of taking over empty territories. Single females remated faster than males and had to significantly increase their defence behaviour as singletons. Thus we argue that monogamy results from the need to guarantee a high quality mate and territory in a competitive environment.

title: A field survey of the bredding habits of Eretmodus cyanostictus, a biparental mouthbrooding cichlid in Lake Tanganyika
abstract: Oral incubation of young or ‘mouthbrooding’ reduces the selective advantages of care by two parents and thus biparental care is rare among ‘mouthbrooding’ fish. We surveyed the breeding biology of Eretmodus cyanostictus, a biparental mouthbrooder from Lake Tanganyika, to understand what factors maintain biparental care. We found larger males than females, a male-biased sex ration and indications that spawning is synchronized around the full moon. These preliminary findings suggest that the benefits of desertion for males are low; males may maximize their reproductive success by helping raise young while females regain reproductive condition.

title: 
Evolutionary Relationships in the Sand-Dwelling Cichlid Lineage of Lake Tanganyika Suggest Multiple Colonization of Rocky Habitats and Convergent Origin of Biparental Mouthbrooding

abstract: The cichlid species flock of Lake Tanganyika is comprised of seven seeding lineages that evolved in step with changes of the lake environment. One seeding lineage diversified into at least six lineages within a short period of time. Our study focuses on the diversification of one of these lineages, the Ectodini, comprising highly specialized, sand- and rock-dwelling species. They display two distinct breeding styles: maternal and biparental mouthbrooding. By analyzing three mtDNA gene segments in 30 species representing all 13 described genera, we show that the Ectodini rapidly diversified into four clades at the onset of their radiation. The monotypic genus Grammatotria is likely to represent the most ancestral split, followed by the almost contemporary origin of three additional clades, the first comprising the benthic genus Callochromis, the second comprising the benthic genera Asprotilapia, Xenotilapia, Enantiopus, and Microdontochromis, and the third comprising the semi-pelagic genera Ophthalmotilapia, Cardiopharynx, Cyathopharynx, Ectodus, Aulonocranus, Lestradea, and Cunningtonia. Our study confirms the benthic and sand-dwelling life-style as ancestral. Rocky habitats were colonized independently in the Xenotilapia- and Ophthalmotilapia-clade. The Xenotilapia-clade comprises both maternal and biparental mouthbrooders. Their mode of breeding appears to be highly plastic: biparental mouthbrooding either evolved once in the common ancestor of the clade, to be reverted at least three times, or evolved at least five times independently from a maternally mouthbrooding ancestor. Furthermore, the genera Xenotilapia, Microdontochromis, Lestradea, and Ophthalmotilapia appeared paraphyletic in our analyses, suggesting the need of taxonomic revision.

title: Repeated Parallel Evolution of Parental Care Strategies within Xenotilapia, a Genus of Cichlid Fishes from Lake Tanganyika
abstract: The factors promoting the evolution of parental care strategies have been extensively studied in experiment and theory. However, most attempts to examine parental care in an evolutionary context have evaluated broad taxonomic categories. The explosive and recent diversifications of East African cichlid fishes offer exceptional opportunities to study the evolution of various life history traits based on species-level phylogenies. The Xenotilapia lineage within the endemic Lake Tanganyika cichlid tribe Ectodini comprises species that display either biparental or maternal only brood care and hence offers a unique opportunity to study the evolution of distinct parental care strategies in a phylogenetic framework. In order to reconstruct the evolutionary relationships among 16 species of this lineage we scored 2,478 Amplified Fragment Length Polymorphisms (AFLPs) across the genome. We find that the Ectodini genus Enantiopus is embedded within the genus Xenotilapia and that during 2.5 to 3 million years of evolution within the Xenotilapia clade there have been 3–5 transitions from maternal only to biparental care. While most previous models suggest that uniparental care (maternal or paternal) arose from biparental care, we conclude from our species-level analysis that the evolution of parental care strategies is not only remarkably fast, but much more labile than previously expected.

title: Evidence for genetic monogamy and female‐biased dispersal in the biparental mouthbrooding cichlid Eretmodus cyanostictus from Lake Tanganyika
abstract: In this study, we investigate whether apparent social monogamy (where a species forms a pair bond but may participate in copulations outside the pair bond) corresponds with genetic monogamy (where individuals participate only in copulations within a pair bond) in a biparental mouthbrooding cichlid fish, Eretmodus cyanostictus, from Lake Tanganyika, Africa. Our findings suggest that E. cyanostictus is both socially and genetically monogamous and that monogamy may result from limited opportunities for polygyny, rather than from reproductive benefits of monogamy. Mating systems are believed to influence the relative rate of dispersal of the sexes, and our results suggest that E. cyanostictus displays female‐biased dispersal, providing some support for the ‘resource competition’ hypothesis driving sex‐biased dispersal.

title: Mouthbrooding and biparental care: an unexpected combination, but male brood care pays
abstract: Biparental care is expected to occur if (1) the costs of desertion for a parent are high, because of greatly reduced survival prospects of offspring, or (2) the benefits of desertion are low. Among mouthbrooding fish, biparental care is rare because the mouth cavity provides a safe brooding site, thus reducing the selective advantages of shared brood care. Eretmodus cyanostictus is a monogamous mouthbrooding cichlid in which the entire clutch is brooded first by the female and then by the male. To test the hypothesis that females alone can produce viable young, we designed an experiment in which females were separated from their mates. Unassisted females prolonged incubation but released as many young as females assisted by males. However, they compensated only partially for male incubation and released smaller and less-developed young. This may substantially reduce offspring survival chances in the wild. The body condition of single females decreased more during incubation and they had a prolonged interspawning interval, but produced similar egg numbers and weights in the next clutch. Our results suggest that the male's brood care effort is an important cause of the maintenance of biparental care and monogamy in E. cyanostictus.

title: Sex ratio and the sexual conflict about brood care in a biparental mouthbrooder
abstract: The pay-off of deserting and leaving a mate to care for the offspring alone is generally assumed to depend mainly on the availability of alternative mating partners and on the potential spawning rate of males and females. Eretmodus cyanostictus is a monogamous mouthbrooding cichlid in which the clutch is successively incubated first by the female and then by the male. It has been suggested that parents are constrained to monogamy due to low remating probabilities for both sexes. We tested this hypothesis by varying the sex ratio experimentally. Mate desertion by either sex was not significantly higher when additional potential mates were present (males: 8.3%, females: 0%) than when there were no other same-sex conspecifics present (males: 0%, females: 0%). Males lost their mate to a male intruder during their incubation in 26.7% of cases. Pair members were more active and showed more aggression when same-sex conspecifics were present. Behavioural differences between treatments were strongest during the incubation period of a given sex. If no desertion takes place, sexual conflict may be expressed also on a second level, the amount of parental care each parent provides. Indeed, males took the offspring later when additional females were present, although male incubation time did not differ between treatments. A hitherto undescribed display behaviour of females was clear evidence of a conflict about the timing of shift of young. In conclusion, offering alternative mating opportunities did not strongly favour mate desertion in E. cyanostictus. It rather revealed a conflict between mates about when to shift the young.

title: Asymmetric sexual conflict over parental care in a biparental cichlid
abstract: Theoretical models predict that parents should adjust the amount of care both to their own and their partner’s body condition. In most biparental species, parental duties are switched repeatedly allowing for repeated mutual adjustment of the amount of care. In the mouthbrooding cichlid Eretmodus cyanostictus, terms are switched only once with females taking the first share. The timing of the shift of the clutch between mates strongly determines both partners’ brooding period and thereby their parental investment. Females signal their readiness to transfer the young several days before the male finally takes them, suggesting sexual conflict over the timing of the shift. In a lab experiment, we reduced the body condition of either the female or the male of a pair to test whether energy reserves affect the timing of the shift and whether female signalling behaviour depends on energetic state. Males with a lowered condition took the young later and incubated for a shorter period, which prolonged the incubation time of their female partners. When female condition was lowered, female and male incubation durations remained unchanged, although females signalled their readiness to shift more intensely. Our results suggest that males adjust their parental investment to own energy reserves but are unresponsive to their mate’s condition. Females appear to carry the entire costs for the male’s adjustment of care. We propose that intrinsic asymmetries in the scope for mutual adjustment of parental investment and the costs of negotiation crucially influence solutions of the conflict between sexes over care.

title: Sneakers, Satellites, and Helpers: Parasitic and Cooperative Behavior in Fish Reproduction
abstract: Organisms compete for various resources in the course of sexual reproduction. First, there is intrasexual competition for obtaining mates. Then, there is the need to exclude reproductive competitors who might displace or affect the individual's own gametes (e.g., sperm competition, egg dumping). There is competition for sites that are used to raise progeny andtor that will optimally support them, and it may be highly advantageous to monopolize the resources that are essential for offspring survival and development. In this chapter I review our current knowledge of the ways in which fish compete at these different levels. I further describe how competition for resources may lead to cooperative behavior, even between the competitors themselves. Finally, I draw attention to the model character of fish social systems and suggest crucial directions for future research.

title: Parental strategy of the cichlid fish Perissodus microlepis, with particular reference to intraspecific brood ‘farming out’
abstract: SynopsisOne parent was experimentally removed from brooding pairs of Perissodus microlepis in the field. The removal elicited drastic behavioural changes in the remaining parent and young. The remaining parent sometimes showed a sequence of peculiar behaviours which were not observed when guarding the young with its mate. Some left the brooding site with young in their mouths and then put the young under the care of another pair of brooding parents.

title: Interspecific brood-mixing in Tanganyikan cichlids
abstract: SynopsisWe collected schools of young, guarded by parents, of six common cichlid species to investigate the frequency and origin of interspecific brood-mixing. The main host species were a piscivore Lepidiolamprologus elongatus and a scale-eater Perissodus microlepis; more than half of their schools included heterospecific young, accounting for 20–40% of the total young. Most of the foreign young belonged to four biparental mouth-brooders whose parents have a habit of carrying their young in their mouths. Many of these young were smaller than the largest young brooded by their own parents. We concluded that adoption of young before independence results from farming-out, a behavior by which parents actively transfer their young to foster parents.

title: The Nesting Site of a Piscivorous Cichlid Lepidiolamprologus profundicola as a Safety Zone for Juveniles of a Zooplanktivorous Cichlid Cyprichromis leptosoma in Lake Tanganyika
abstract: Juvenile Cyprichromis leptosoma, an endemic cichlid fish in Lake Tanganyika, form large schools near the nesting sites of the piscivorous cichlid Lepidiolamprologus profundicola. The female L. profundicola guarding the nesting site drove away piscivorous fish that approached the nest. After the disappearance of the guarding L. profundicola, the frequency of predatory dashes toward the school of juvenile C. leptosoma near the nesting site increased markedly. This suggests that juvenile C. leptosoma use the breeding territory of L. profundicola as a safety zone from their potential predators. The relationship between C. leptosoma and L. profundicola should be regarded as a type of commensalism.

title: Social system and reproduction of helpers in a cooperatively breeding cichlid fish (Julidochromis ornatus) in Lake Tanganyika: field observations and parentage analyses
abstract: It is suggested that some fish of the genus Julidochromis, substrate-brooding Tanganyikan cichlids with biparental care, breed cooperatively with helpers. We studied the social system of J. ornatus in the wild and analysed genetic parentage using microsatellites. Within the studied population three patterns of social system were identified: monogamous pairs (61%, 44 of 72 groups), pairs with helpers (29%, 21), and polygamous harems with helpers (controlled by either a large female or large male owner; 10%, 7). In cooperatively breeding groups, the number of helpers at each nest ranged from 1 to 6 (median 1), and male helpers were more numerous than female helpers. In both sexes, the body size was different among individuals of different social status (harem owner > breeder > helper). Helpers and harem owners of both sexes exhibited brood defence although its frequency was low. The molecular analysis revealed that (1) the helpers were mostly unrelated to dominant breeders, (2) many helpers of both sexes contributed genetically to the next generations, (3) male helpers had high siring success (41% of young in total), and (4) large young unrelated to group members were detected at 30% of observed nests, which may be due to breeder (or helper) replacements and immigration of young. We discuss the implications of these results for understanding the complex social system of this species, especially the low reproductive skew in comparison with other cooperatively breeding cichlids.

title: Cooperative Breeding in the Lake Tanganyika Cichlid Julidochromis ornatus
abstract: Cooperative breeding has been described for several cichlids from the genus Julidochromis (Perciformes: Cichlidae) under laboratory conditions, but field evidence is scarce. Here we describe the breeding system of the cichlid Julidochromis ornatus (Boulenger) in Lake Tanganyika (Zambia). Groups defended a breeding shelter under a large flat stone. Smaller group members stayed and fed under or close to the stone, actively guarded by the larger group members. Six out of 28 groups were newly established by breeders, joined by subordinates from a large pool of independent fish (comprising 50–70% of the total population), and four groups were seen to dissolve during a total of 77 observation days. Breeding groups consisted of a large breeding male and female with zero to five smaller subordinates (average 2). Larger breeders and subordinates were found in larger groups. All group members participated in territory defence and -maintenance, but the breeders were only present at the shelter 48% of the time, in contrast to the subordinates which guarded the breeding shelter 94% of the time. Smaller group members showed submissive behaviours to larger group members. We conclude subordinates in J. ornatus are helpers, but we did not find evidence that helpers increased the group’s current reproductive success. Personal observations combined with a literature review revealed at least 19 species of Lake Tanganyika cichlids show evidence of cooperative breeding, entirely confined to the substrate breeding tribe of the Lamprologini (24% of 80 species in total): 2 Chalinochromis spp., 5 Julidochromis spp., 12 Neolamprologus spp. More effort should be put into detecting cooperative breeding in American and Asian substrate breeding cichlid species.

title: Evolutionary history of the Lake Tanganyika cichlid tribe Lamprologini (Teleostei: Perciformes) derived from mitochondrial and nuclear DNA data
abstract: Tanganyika comprises a cichlid species flock with substrate-breeding and mouthbrooding lineages. While sexual selection via mate choice on male mating color is thought to boost speciation rates in mouthbrooding cichlids, this is not the case in substrate-breeding lamprologines, which mostly form stable pairs and lack sexual dichromatism. We present a comprehensive reconstruction of the evolution of the cichlid tribe Lamprologini, based upon mtDNA sequences and multilocus nuclear DNA (AFLP) markers. Twelve mtDNA clades were identified, seven of which were corroborated by the AFLP tree. The radiation is likely to have started about 5.3 MYA, contemporarily with that of the mouthbrooding C-lineage, and probably triggered by the onset of deep-water conditions in Lake Tanganyika. Neither the Congo- nor the Malagarazi River species form the most ancestral branch. Several conflicts in the mtDNA phylogeny with taxonomic assignments based upon color, eco-morphology and behavior could be resolved and complemented by the AFLP analysis. Introgressive hybridization upon secondary contact seems to be the most likely cause for paraphyly of taxa due to mtDNA capture in species involving brood-care helpers, while accidental hybridization best explains the para- or polyphyly of several gastropod shell breeders. Taxonomic error or paraphyly due to the survival of ancestral lineages appear responsible for inconsistencies in the genera Lamprologus and Neolamprologus.

title: The evolution of cooperative breeding in the African cichlid fish, Neolamprologus pulcher
abstract: The conundrum of why subordinate individuals assist dominants at the expense of their own direct reproduction has received much theoretical and empirical attention over the last 50 years. During this time, birds and mammals have taken centre stage as model vertebrate systems for exploring why helpers help. However, fish have great potential for enhancing our understanding of the generality and adaptiveness of helping behaviour because of the ease with which they can be experimentally manipulated under controlled laboratory and field conditions. In particular, the freshwater African cichlid, Neolamprologus pulcher, has emerged as a promising model species for investigating the evolution of cooperative breeding, with 64 papers published on this species over the past 27 years. Here we clarify current knowledge pertaining to the costs and benefits of helping in N. pulcher by critically assessing the existing empirical evidence. We then provide a comprehensive examination of the evidence pertaining to four key hypotheses for why helpers might help: (1) kin selection; (2) pay‐to‐stay; (3) signals of prestige; and (4) group augmentation. For each hypothesis, we outline the underlying theory, address the appropriateness of N. pulcher as a model species and describe the key predictions and associated empirical tests. For N. pulcher, we demonstrate that the kin selection and group augmentation hypotheses have received partial support. One of the key predictions of the pay‐to‐stay hypothesis has failed to receive any support despite numerous laboratory and field studies; thus as it stands, the evidence for this hypothesis is weak. There have been no empirical investigations addressing the key predictions of the signals of prestige hypothesis. By outlining the key predictions of the various hypotheses, and highlighting how many of these remain to be tested explicitly, our review can be regarded as a roadmap in which potential paths for future empirical research into the evolution of cooperative breeding are proposed. Overall, we clarify what is currently known about cooperative breeding in N. pulcher, address discrepancies among studies, caution against incorrect inferences that have been drawn over the years and suggest promising avenues for future research in fishes and other taxonomic groups.

title: Parallel evolution of facial stripe patterns in the Neolamprologus brichardi/pulcher species complex endemic to Lake Tanganyika.
abstract: Colour pattern diversity can be due to random processes or to natural or sexual selection. Consequently, similarities in colour patterns are not always correlated with common ancestry, but may result from convergent evolution under shared selection pressures or drift. Neolamprologus brichardi and Neolamprologus pulcher have been described as two distinct species based on differences in the arrangement of two dark bars on the operculum. Our study uses DNA sequences of the mitochondrial control region to show that relatedness of haplotypes disagrees with species assignment based on head colour pattern. This suggests repeated parallel evolution of particular stripe patterns. The complete lack of shared haplotypes between populations of the same or different phenotypes reflects strong philopatric behaviour, possibly induced by the cooperative breeding mode in which offspring remain in their natal territory and serve as helpers until they disperse to nearby territories or take over a breeding position. Concordant phylogeographic patterns between N. brichardi/N. pulcher populations and other rock-dwelling cichlids suggest that the same colonization routes have been taken by sympatric species and that these routes were affected by lake level fluctuations in the past.

title: Broodcare helpers in the cichlid fish Lamprologus brichardi: Their costs and benefits
abstract: Helping' in birds and mammals involves seemingly altruistic behaviour. In the cichlid fish Lamprologus birchardi helpers are usually young of former broods staying in their parents' territories and participating in all kinds of parental duties (broodcare, territory maintenance and defence). The discovery of helpers in fish offered the chance of attempting an extensive analysis of potential costs and benefits influencing the evolution of helpers in a vertebrate. Three factors proved to be of major impor- tance in the cost-benefit analysis of helping as opposed to leaving for family-independent non- reproductive aggregations. Due to investment and to their rank within a family's hierarchy, helpers grow at a slower rate than non-helpers. This cost is compensated for by (i) a lower mortality risk to helpers caused by their access to a defended shelter and by protection afforded by bigger family members, and (ii) a positive contribution by helpers to the future reproductive success of their parents: females with helpers produce bigger clutches and consequently more free-swimming fry (=siblings). Other variables, such as the helpers' influence on the relative breeding success of their parenis, broodcare experience through helping, the chances of territory take-over, parasitism of parents' reproduction and cannibalism are of minor importance. Similar social organizations in other fish are discussed with respect to their ecology and are compared with cooperatively breeding birds and mammals.

title: Helpers in fish
abstract: SummaryField data show that in the cichlid fish Lamprologus brichardi conspecifics other than the reproducing pair help in brood care and territory maintenance. The expected degree of relatedness between helpers and the eggs or larvae they tend lies between 0.25 and 0.5, decreasing with the helper's age. This decrease might influence the point of time at which helpers depart. Five other endemic Lake Tanganyika cichlids showing rather similar helping behaviour are described.

title: Group composition affects male reproductive partitioning in a cooperatively breeding cichlid
abstract: Individuals within groups of cooperatively breeding species may partition reproduction, with the dominant pair often taking the largest share. The dominant's ability to reproductively control subordinates may depend on differences in competitive ability, due to, e.g. body size differences, but may also depend on the number of same‐sex competitors inside the group. We tested experimentally whether subordinates reproduce more when these subordinates are large or when a second subordinate of the same sex need to be controlled by the dominants, using the cooperatively breeding cichlid Neolamprologus pulcher. Dominant pairs were assisted by a large and a small unrelated subordinate; sexes of these fish were varied in a full‐factorial design (giving four treatments). Dominant males lost significantly more parentage to the large subordinate male when a small subordinate male was also present, compared to when a small subordinate female was present. However, subordinate paternity was generally low and did not significantly curb total dominant male reproductive output, which was more affected by the sizes and numbers of reproductive females present inside his group. Dominant female maternity, clutch sizes and total output did not depend on the treatments. Subordinate–subordinate reproduction was virtually absent (one out of 874 offspring). Female subordinates were more likely to provide care for their own broods. In contrast, male subordinates did not adjust their level of care to their parentage. Variability in female subordinate alloparental brood care was particularly high, with females showing more care than males in general. We also detected effects of growth rate and food ration on parentage independent of the treatments, most notably: (i) a trade‐off between dominant male growth rate and paternity; (ii) a decrease in dominant male paternity with increasing food ration; (iii) a positive effect of growth rate on paternity in small males. We conclude that dominant males should be sensitive to the number and sizes of subordinate males present in their group, particularly when these subordinates are not helpful or grow fast, and food is plentiful. Dominant females should be less sensitive, because female subordinates do not appear to impose reproductive costs and can be helpful through alloparental brood care.

title: Reproductive suppression in female cooperatively breeding cichlids
abstract: Suppression by dominants of female subordinate reproduction has been found in many vertebrate social groups, but has rarely been shown experimentally. Here experimental evidence is provided for reproductive suppression in the group-living Lake Tanganyika cichlid Neolamprologus pulcher. Within groups of three unrelated females, suppression was due to medium- and small-sized females laying less frequently compared with large females, and compared with medium females in control pairs. Clutch size and average egg mass of all females depended on body size, but not on rank. In a second step, a large female was removed from the group and a very small female was added to keep the group size constant. The medium females immediately seized the dominant breeding position in the group and started to reproduce as frequently as control pairs, whereas clutch size and egg mass did not change. These results show that female subordinate cichlids are reproductively capable, but apparently suppressed with respect to egg laying. Nevertheless, some reproduction is tolerated, possibly to ensure continued alloparental care by subordinate females.

title: Costs and benefits of polygyny in the cichlid Neolamprologus pulcher

abstract: Polygyny is regarded as a beneficial strategy for males, whereas females mated with polygynous males (males simultaneously paired to more than one female) often suffer a reduction in pair male contributions. This study examined the costs and benefits associated with polygyny in the cooperatively breeding cichlid Neolamprologus pulcher. In this species, males are facultatively polygynous; some males hold only one territory with one breeding female whereas other males hold multiple territories, each with its own breeding female. Polygynous males were larger than monogamous males and body-scraped less, a behaviour often associated with ectoparasite loads. Polygynous males also had larger testes (controlling for body mass) and higher circulating 11-ketotestosterone levels than monogamous males. Paradoxically, monogamous males occupied higher-quality territories with more shelter and fewer predators. Monogamous males also provided more parental care than polygynous males but the number and survival of young did not vary according to male mating behaviour. The results of our study suggest that females trade-off between male genetic quality and resources in N. pulcher. Our results imply that males holding only one territory may provide their mates with significant assets but may not be able to outcompete neighbours for additional breeding positions because of their small body size and possible higher parasite load. The lack of differences between monogamous and polygynous groups in terms of offspring survival (a measure of reproductive success) suggests that there may be few if any fitness consequences of polygynous pairing for females.

title: Dispersal patterns and status change in a co‐operatively breeding cichlid Neolamprologus pulcher: evidence from microsatellite analyses and behavioural observations
abstract: Genetic techniques and long-term behavioural observations were combined to investigate dispersal patterns and changes in social position in Neolamprologus pulcher, a co-operatively breeding cichlid from Lake Tanganyika. Comparisons of genetic variance (FsT) across sub-populations demonstrated that fish were genetically more similar to individuals from proximate sub-populations compared to individuals from distant sub-populations. Microsatellite analyses revealed year-long philopatry for some individuals and that other individuals dispersed to new territories and sub-populations. Individuals appeared to disperse farther (across many territories in a sub-population or to new sub-populations) to achieve breeding status. Non-breeding group members (or helpers) were observed to inherit breeding positions and male breeders were replaced faster than female breeders. These results demonstrate that important and difficult to obtain life-history information can be obtained from genetic sampling.

title: Genetic relatedness in groups is sex-specific and declines with age of helpers in a cooperatively breeding cichlid.
abstract: Kin selection can explain the evolution of cooperative breeding and the distribution of relatives within a population may influence the benefits of cooperative behaviour. We provide genetic data on relatedness in the cooperatively breeding cichlid Neolamprologus pulcher. Helper to breeder relatedness decreased steeply with increasing helper age, particularly to the breeding males. Helper to helper relatedness was age-assortative and also declined with age. These patterns of relatedness could be attributed to territory take-overs by outsiders when breeders had disappeared (more in breeding males), between-group dispersal of helpers and reproductive parasitism. In six of 31 groups females inherited the breeding position of their mother or sister. These matrilines were more likely to occur in large groups. We conclude that the relative fitness benefits of helping gained through kin selection vs. those gained through direct selection depend on helper age and sex.

title: Extended safe havens and between-group dispersal of helpers in a cooperatively breeding cichlid
abstract: In cooperative breeders, between-group dispersal of helpers is expected to occur if it increases their fitness. Genetic data suggest that helpers in the cooperatively breeding Lake Tanganyika cichlid Neolamprologus pulcher occasionally migrate into nearby groups where they again become helpers. We studied in the field how and why helpers migrate between groups by recording their ranging and social behaviours. We found that helpers spent 5.3% of their time visiting other groups, where they received similar low levels of aggression as within their home group. Large helpers visited other groups more often than small helpers and helpers visited other groups more frequently when the queue in their home group was large, suggesting that helpers with low chances to inherit the territory search for alternatives. Our data show that helpers may use other groups' territories as a refuge, as helpers actively sought shelter within territories of neighbouring groups when we experimentally increased the perceived risk of staying in their home territory. We observed two attempted and one successful case of 'voluntary' (i.e., strategic) between-group dispersal, and experimentally induced three helpers to disperse into other groups. By regular visits, helpers appear to establish familiarity and social relationships with nearby groups, which serve as 'extended safe havens' to hide from predators. In the long run frequent visiting behaviour may facilitate between group dispersal.

title: Sex differences in rates of territory joining and inheritance in a cooperatively breeding cichlid fish
abstract: In social groups, subordinates may gain dominant breeding status either by inheriting the top position in their current group or by dispersing to join a new group. The pathway to breeder status is likely to vary between males and females as a result of sex differences in the costs of dispersal and inbreeding. We report results from a field study conducted to explore sex differences in the rates of territory joining and inheritance in a cooperatively breeding cichlid, Neolamprologus pulcher. We experimentally created 34 breeding vacancies and investigated the effects of breeder removal on the degree of cooperative behaviour and how changes in status influenced reproductive physiology. We found that 71% of male vacancies were filled by joiners (incoming fish, not previously members of the group) entering the territory. In contrast, only 15% of female vacancies were filled by joiners entering the group from elsewhere. Helpers increased their frequency of cooperative behaviour following the removal of a female breeder, but not following removal of a male breeder. Our results suggest that female breeder vacancies are typically filled by subordinate helpers who inherit from within the group and that male breeder vacancies are commonly filled by joining individuals (existing breeders or former helpers from other groups). Male social status and gonadal investment were positively correlated. This study represents one of the first experimental attempts to examine sexual differences in the pathway to breeding status in a cooperatively breeding species.

title: Evidence for size and sex‐specific dispersal in a cooperatively breeding cichlid fish
abstract: African Great Lake cichlid populations are divided into thousands of genetic subpopulations. The low gene flow between these subpopulations is thought to result from high degrees of natal philopatry, heavy predation pressure, and a patchy distribution of preferred habitats. While predation pressure and habitat distribution are fairly straightforward to assess, data on dispersal distances and rates are scarce. In fishes, direct observations of dispersal events are unlikely, but dispersal can be studied using molecular markers. Using seven microsatellite loci, we examined dispersal in the cooperatively breeding cichlid fish, Neolamprologus pulcher. As this species is found in well‐defined groups clustered into subpopulations, we could assess dispersal on a narrow (within subpopulation) and broad (between subpopulation) scale. While fish were generally more related to others in their own subpopulation than they were to fish from other subpopulations, large males diverged from this pattern. Large males were more related to other large males from different subpopulations than they were to large males from their own subpopulation, suggesting more frequent dispersal by large males. Across subpopulations, relatedness between large males was higher than the relatedness among large females; this pattern was not detected in small males and small females. Within a subpopulation, individuals appeared to be preferentially moving away from relatives, and movement was unrestricted by the physical distance between groups. Our results highlight the importance of examining multiple spatial scales when studying individual dispersal biases.

title: The role of genetic relatedness among social mates in a cooperative breeder
abstract: Breeding with relatives can have severe fitness consequences, so avoiding these costs is often evolutionarily favored. There are a number of mechanisms that reduce the likelihood of mating with relatives, including avoiding relatives as mates (through sex-biased dispersal and mate choice) and delayed sexual maturity in the presence of relatives. Here, we examine these mechanisms in Neolamprologus pulcher, a group-living cichlid fish that exhibits male-biased dispersal. Despite sex-biased dispersal in this species, mean relatedness between social mates was not different from that expected if pairs had formed randomly, suggesting individuals neither actively avoid nor prefer pairing with relatives. Furthermore, gonadal investment of subordinates living in social groups was not correlated with their relatedness to the opposite-sex dominant breeder in the group, suggesting that sexual maturation does not depend on the presence or absence of a relative. Highly related social pairs showed higher rates of within-pair aggression and lower rates of nonaggressive social affiliation than less-related social pairs. Breeder investment and indicators of female breeder and group quality were not correlated with relatedness values between social mates. However, scraping rates (a potential quality indicator) were lower in males paired with more closely related females. We consider whether the apparent lack of inbreeding avoidance reflects an evolutionary history of limited breeding opportunities in N. pulcher or a facultative strategy of more-fit individuals and discuss the behavioral results in light of the suggested nonassortative mating with regard to relatedness.

title: Helpful Female Subordinate Cichlids Are More Likely to Reproduce
abstract: In many cooperatively breeding vertebrates, subordinates assist a dominant pair to raise the dominants' offspring. Previously, it has been suggested that subordinates may help in payment for continued residency on the territory (the ‘pay-to-stay hypothesis’), but payment might also be reciprocated or might allow subordinates access to reproductive opportunities.  We measured dominant and subordinate female alloparental brood care and reproductive success in four separate experiments and show that unrelated female dominant and subordinate cichlid fish care for each other's broods (alloparental brood care), but that there is no evidence for reciprocal ‘altruism’ (no correlation between alloparental care received and given). Instead, subordinate females appear to pay with alloparental care for own direct reproduction.  Our results suggest subordinate females pay with alloparental care to ensure access to the breeding substrate and thereby increase their opportunities to lay their own clutches. Subordinates' eggs are laid, on average, five days after the dominant female has produced her first brood. We suggest that immediate reproductive benefits need to be considered in tests of the pay-to-stay hypothesis.

title: Cichlids do not adjust reproductive skew to the availability of independent breeding options
abstract: Helpers in cooperatively breeding species forego all or part of their reproduction when remaining at home and assisting breeders to raise offspring. Different models of reproductive skew generate alternative predictions about the share of reproduction unrelated subordinates will get depending on the degree of ecological constraints. Concession models predict a larger share when independent breeding options are good, whereas restraint and tug-of-war models predict no effects on reproductive skew. We tested these predictions by determining the share of reproduction by unrelated male and female helpers in the Lake Tanganyika cichlid Neolamprologus pulcher depending on experimentally manipulated possibilities for helper dispersal and independent breeding and depending on helper size and sex. We created 32 breeding groups in the laboratory, consisting of two breeders and two helpers each, where only the helpers had access to a nearby dispersal compartment with (treatment) or without (control) breeding substrate, using a repeated measures design. We determined the paternity and maternity of 1185 offspring from 47 broods using five to nine DNA microsatellite loci and found that: (1) helpers participated in reproduction equally across the treatments, (2) large male helpers were significantly more likely to reproduce than small helpers, and (3) male helpers engaged in significantly more reproduction than female helpers. Interestingly, in four broods, extragroup helper males had fertilized part of the brood. No helper evictions from the group after helper reproduction were observed. Our results suggest that tug-of-war models based on competition over reproduction within groups describe best the reproductive skew observed in our study system. Female breeders produced larger clutches in the treatment compared to the control situation when the large helpers were males. This suggests that male breeder-male helper reproductive conflicts may be alleviated by females producing larger clutches with helpers around.

title: Habitat saturation, benefits of philopatry, relatedness, and the extent of co-operative breeding in a cichlid
abstract: Co-operative breeding in vertebrates may emerge due to subordinates delaying dispersal when free breeding habitat is not available ('habitat saturation' hypothesis, HS). However, delayed dispersal might also be due to younger individuals postponing dispersal to when they are more competitively able or have more to gain from breeding independently ("benefits-of-philopatry" hypothesis, BP) or to when inclusive fitness benefits no longer outweigh the benefits from independent breeding ("kin selection" hypothesis, KS). Here, we show in three experiments that both HS and BP determine the extent of co-operative breeding in the cichlid Neolamprologus pulcher. Contrary to the KS, individuals significantly avoided settlement with related individuals, and an additional settlement experiment confirmed this result. Our results suggest that kin structure in these cichlids emerges from limits on dispersal, but if such barriers are absent, cichlids prefer to settle with unrelated individuals to maximize the benefits of direct reproductive participation.

title: Reproductive parasitism of broodcare helpers in a cooperatively breeding fish
abstract: Large male helpers in the cooperatively breeding cichlid Neolamprologus pulcher gain reproductive success by parasitizing the reproductive effort of male territory owners. Under controlled, experimental conditions we examined the genetic relatedness between the members of brood pairs (n 5 14), their male helpers (n 5 8), and offspring (n 5 292) in seven families. We used multilocus DNA fingerprinting to check for potential reproductive parasitism by male helpers and to assess their fertilization success. Of offspring produced in these families, 10.3% were sired by helpers. In parasitized broods, helper fertilization success varied between 12.5% and 35.8%. Male helpers parasitized parental reproduction when their body size exceeded 4.5 cm standard length (SL), even though sexual maturity may be reached much earlier (3.5 cm SL). Two of three parasitic helpers were punished by severe aggressive attacks when parasitizing the reproduction of breeders, which led to their expulsion from the territory. This study demonstrates a potential fitness benefit to broodcare helpers that is often neglected. It also points to the delicate balance that may exist between cooperative and competitive behavior in cooperatively breeding species.  Lake Tanganyika, cichlids, cooperative breeding, DNA fingerprinting, Neolamprologus pulcher, N. brichardi, reproductive competition.

title: Multiple paternity in the cooperatively breeding fish Neolamprologus pulcher
abstract: In cooperative breeders, mature males may compete for fertilizations. In this study, we measured the degree of multiple paternity in a natural population of a cooperatively breeding fish. Neolamprologus pulcher (Perciformes: Cichlidae) is a highly social cichlid endemic to Lake Tanganyika. We used highly variable microsatellite loci to survey 12 groups with an average number of 10.6 brood care helpers per group and a total of 43 offspring (mean 3.6 per brood). In 11 of 12 groups, all young were assigned to the dominant female. The dominant male sired all offspring in three groups, part of the offspring in four groups, and in five groups, he had no paternity at all. In total, 44.2% of young were not fathered by the current male territory owner. Multiple paternity was found in 5 of 12 broods (41.7 %), with 8 of 35 young (22.9 %) being sired by males other than the respective territory owners. This is an exceptionally high rate of extra-pair paternity among cooperatively breeding vertebrates. Neither helpers present in these territories during collection nor neighbouring males were unequivocally assigned to have sired these extra-pair young. However, behavioural observations suggest that male helpers may have produced these young before being expelled from the territory in response to this reproductive parasitism. We discuss these results in the light of reproductive skew theory, cooperative breeding in vertebrates and alternative reproductive tactics in fish.

title: Male reproductive suppression in the cooperatively breeding fish Neolamprologus pulcher
abstract: In most cooperative breeders, dominants suppress the reproduction of subordinates. However, two previous studies of Neolamprologus pulcher, a cooperatively breeding cichlid fish, have suggested that socially subordinate helper males sneak fertilizations from dominant breeding males. If such sneaking does occur, both theoretical work and empirical studies of other fish species suggest that sperm competition will select for increased reproductive investment by sneaker males, relative to more dominant males. To address these issues, we quantified gonadal investment and sperm characteristics of 41 N. pulcher male breeders and 62 male helpers from 55 groups in Lake Tanganyika. Gonadal investment followed patterns consistent with reproductive suppression, with breeders having considerably larger testes masses than helpers. Breeders also had faster and longer swimming sperm and a higher percentage of motile sperm compared to helpers. However, sperm characteristics of large helpers were similar to those of breeders, but these same helpers had lower testes masses. Thus, large helpers had sperm that were physiologically equivalent to that of breeders, but their relatively small gonads imply that they were reproductively suppressed.

title: The relationship between social status, behaviour, growth and steroids in male helpers and breeders of a cooperatively breeding cichlid
abstract: We tested whether subordinate helper males of the Lake Tanganyika cooperatively breeding cichlid Neolamprologus pulcher show elevated excretion levels of the stress hormone cortisol and reduced levels of 11-ketotestosterone and testosterone when living in groups with a small breeder male, compared to similar helper males living in groups with a large breeder male, in a full-factorial repeated measures experimental design. We also measured the same hormones in breeder males with and without helper males. Previous research showed that the size difference between large male helpers and male breeders in groups of this species influences behaviour and growth decisions. Contrary to our expectation, no effect of the size-difference between helper males and breeder males on helper hormone levels was detected. Furthermore, helper males had similar hormone excretion levels to those of size-matched breeder males without helpers, and to small breeder males. There was no influence of egg laying on breeder male and helper hormone levels during the experiment. Interestingly, all three hormone levels were significantly lower in helpers showing elevated levels of submissive behaviour towards the breeders, independently of the size of the breeder males. The low cortisol levels suggest that helper males can successfully reduce stress by appeasing breeder males through submission. Furthermore, helper males showing a high level of submissive behaviour had lower levels of androgens than less submissive helpers, suggesting a lower reproductive potential in submissive helpers. We propose that helper submission may be used as an honest signal of reduced interest in reproduction towards the breeder male in this species.

title: Tug-of-war over reproduction in a cooperatively breeding cichlid
abstract: In group-living animals, dominants may suppress subordinate reproduction directly and indirectly, thereby skewing reproduction in their favour. In this study, we show experimentally that this ability (‘power’) is influenced by resource distribution and the body size difference between unrelated dominants and subordinates in the cichlid Neolamprologus pulcher. Reproduction was strongly skewed towards the dominant female, due to these females producing more and larger clutches and those clutches surviving egg eating better than those of subordinate females, but was not so when subordinates defended a patch. If breeding shelters were provided in two patches, subordinate females were more likely to exclusively defend a patch against the dominant female and breed, compared to when the same breeding resource was provided in one patch. Relatively large subordinate females were more likely to defend a patch and reproduce. Females also directly interfered with each other’s reproduction by eating the competitors’ eggs, at which dominants were more successful. Although dominant females benefited from subordinate females due to alloparental care and an increase in egg mass, they also showed costs due to reduced growth in the presence of subordinates. The results support the view that the dominant’s power to control subordinate reproduction determines reproductive partitioning, in agreement with the predictions from tug-of-war models of reproductive skew.

title: Cooperative Breeding and Group Structure in the Lake Tanganyika Cichlid Neolamprologus savoryi
abstract: As yet, cooperative breeding has been described only for some fish species. However, evidence is accumulating that it is widespread among Lake Tanganyika cichlids. We studied the cooperative breeding system of the substrate breeding cichlid Neolamprologus savoryi. Breeding groups typically consisted of a large breeding male with one to four breeding females and three to 33 helpers (mean group size: 14.3 members). Group size was significantly related to breeding male and female body sizes, and larger males had more breeding females and larger sized male helpers. The size of the largest female in the group was positively related to the number and sizes of secondary breeding females and female helpers. In case of multiple breeding females, these females usually divided the group's territory into sub-territories, each with its own helpers (subgroups). Interspersed between groups, independent fish were detected defending an individual shelter (4.4% of all fish). In 9% of the groups no breeding female was present. All group members participated in territory defence and maintenance, and showed submissive behaviours to larger group members. As expected, the level of between-subgroup conflicts was high compared with the level of within-subgroup conflicts. We compare these results with data available from other cooperatively breeding fishes.

title: Causes of Female Emigration in the Group-Living Cichlid Fish Neolamprologus multifasciatus
abstract: Neolamprologus multifasciatus is a small cichlid breeding in empty gastropod shells that forms complex family groups with several reproductive males and females, presumably due to dispersal constraints. Nevertheless, some exchange of females between groups has been demonstrated. We investigated experimentally emigration patterns of female group members to elucidate which factors influence the decision of adult females either to stay or to emigrate into another territory. Four females resident in a large (13 shells) male territory were given the opportunity to stay there or to emigrate to another large (13 shells) or a small (four shells) male territory. In six replicates we found that (1) the most aggressive females had the largest subterritories (most shells); (2) invariably, the two females with the fewest shells emigrated; (3) both the emigrating and the staying females had more shells after emigration than before. Apparently, females explore their options in neighboring territories and decide accordingly whether to stay or emigrate.

title: Sex-reversed dominance and aggression in the cichlid fish Julidochromis marlieri
abstract: The plesiomorphic breeding system of the large fish family Cichlidae is a monogamous, biparental pair that cares for eggs and extends care to shepherding and protecting tiny fry. Typically, the male is larger than his mate and dominates her. Julidochromis marlieri, an African cichlid from Lake Tanganyika, follows this pattern but with an important difference: the female is larger than her mate; one female was reported mated to two males simultaneously (Yamagishi & Kohda 1996). We asked whether the female also dominates her mate, as in polyandrous, sex-role reversed birds. Additionally, documenting an inherent difference in aggressiveness would clarify the behavioral mechanisms that support pair-bonding in monogamous species. We staged contests between males and females of various relative sizes. When equal in size, females regularly won contests. Females were also more likely to initiate advancement to more aggressive stages of the fight. Thus dominance and aggression are sex-reversed in Julidochromis marlieri.

title: Testis size depends on social status and the presence of male helpers in the cooperatively breeding cichlid Julidochromis ornatus
abstract: Cooperatively breeding animals, in which helpers may participate in reproduction with dominant breeders, are ideal species for examining intraspecific variation in testis size because they often exhibit both monogamous breeding (low risk of sperm competition) and polyandrous breeding (high risk) within a population. However, little is known about testis investment as a result of sperm competition in these animals. The substrate-brooding cichlid fish Julidochromis ornatus has a cooperatively breeding system, in which some males mate monogamously and other males reproduce as dominant breeders or helpers within cooperatively breeding groups, in which male helpers frequently sire young. We examined the relationship between testis investment and male social status in relation to the risk of sperm competition. As predicted from sperm competition models, in groups with male helpers, both the male breeders and the male helpers invested more in testes mass, compared to breeding males without male helpers. We also found a positive relationship between the testes mass of male breeders and their male helpers, suggesting that males increase their investment in reproductive capability under the risk of sperm competition. Sperm competition models also predict that larger testes are associated with increased siring success. Our paternity analysis supported this prediction; we found a positive relationship between testis investment by male helpers and the number of offspring they sired.

title: Cooperatively breeding cichlid fish adjust their testis size but not sperm traits in relation to sperm competition risk
abstract: Theoretical models predict that in external fertilisers, sperm competition risk selects for larger testes and larger sperm with higher velocity, at the expense of sperm longevity. Although several empirical studies have tested this prediction within a fish species, there has been little evidence that an individual male can adjust his testis size and sperm traits according to the risk of sperm competition. Using the cooperatively breeding cichlid Julidochromis transcriptus, we examined testis size and sperm traits (sperm size, velocity and longevity) of males that were assigned to monogamous pairs (without risk of sperm competition) or cooperatively polyandrous trios composed of two males and one female (with risk); the groups were reared for breeding for 2 or more months. Our experiments showed that the polyandrous males, both siring eggs in a clutch, had larger relative testis mass than the monogamous males. However, sperm traits were unrelated to sperm competition. These results suggest that individual males of J. transcriptus can adjust their testis size but not sperm traits in response to sperm competition risk. We also examined the effect of testis investment on growth rate and the somatic condition of males. Increased investment in testes slowed growth rates, whereas testis investment did not affect somatic condition, indicating a trade-off between investment in testes and growth. We conclude that testis investment, in response to perceived sperm competition risks, occurs at the expense of growth and may be a conditional reproductive strategy that is determined by the social status (monogamous or polyandrous) of each J. transcriptus male.

title: Living on the wedge: female control of paternity in a cooperatively polyandrous cichlid
abstract: Theories suggest that, in cooperatively breeding species, female control over paternity and reproductive output may affect male reproductive skew and group stability. Female paternity control may come about through cryptic female choice or female reproductive behaviour, but experimental studies are scarce. Here, we show a new form of female paternity control in a cooperatively polyandrous cichlid fish (Julidochromis transcriptus), in which females prefer wedge-shaped nesting sites. Wedge-shaped sites allowed females to manipulate the siring success of the group member males by spawning the clutch at the spot where the large males were just able to enter and fertilize the outer part of the clutch. Small males fertilized the inner part of the clutch, protected from the large aggressive males, leading to low male reproductive skew. Small males provided more brood care than large males. Multiple paternity induced both males to provide brood care and reduced female brood care accordingly. This is, to our knowledge, the first documented case in a species with external fertilization showing female mating behaviour leading to multiple male paternity and increased male brood care as a result.

title: The evolution of alternative reproductive tactics: concepts and questions
abstract: Here we outline the meaning of the term alternative reproductive tactics, or ARTs, and discuss why the existence of ARTs is so widespread in animals. We ask what we need to know to understand the evolution of ARTs and the importance of general principles such as frequency dependence, density dependence, and condition dependence, and what we need to know about proximate mechanisms involved in the regulation of ARTs to comprehend evolutionary patterns. We discuss current issues in the study of ARTs and list 12 questions that we think need particular attention. Throughout we shall provide representative examples of ARTs in animals to illustrate the ubiquitous nature of this phenomenon.

title: Sexual conflict over breeding substrate causes female expulsion and offspring loss in a cichlid fish
abstract: Females of the Lake Tanganyika cichlid Lamprologus callipterus exclusively breed in empty snail shells that males collect in their territories. Male--male competition for shells is severe, leading to frequent shell stealing and territory takeover. As a consequence, males have breeding females in their shells that spawned with competitors. In this field study, we investigated both naturally occurring and experimentally induced encounters of territorial males with females that had spawned with other males. We found that the breeding success of females that were taken over by a different male was significantly reduced. Behavioral observations after experimental shell relocation further showed that males recognized females that they had not spawned with: males directed more exploration and manipulation behavior toward such shells compared with controls. Reoccupation rate of emptied experimental shells was significantly higher than that of unmanipulated empty shells. This indicates that shell stealing and nest takeover, followed by female expulsion, contribute to the reproductive success of L. callipterus males. We also found that female mate choice reduces expulsion risk: females preferred to mate with large males, and male size correlated with dominance. We conclude that the limited availability of breeding substrate is a key determinant of both intrasexual competition and intersexual conflict in this species.

title: Reproductive investment of giants and dwarfs: specialized tactics in a cichlid fish with alternative male morphs
abstract: 1. Adopting alternative reproductive tactics may require divergent solutions to reproductive competition among individuals of a population. Often investment in reproduction differs substantially between individuals pursuing bourgeois and parasitic tactics, which may result in different trade-offs and limitations. 
 
2. Here we identify divergent behavioural, morphological and physiological traits of bourgeois and parasitic male morphs in Lamprologus callipterus, a Lake Tanganyika cichlid with an extreme size dimorphism among males. We focus on limiting factors and compare these between large, nest-building males and dwarf males parasitizing their reproductive effort. 
 
3. Only nest males invest in courtship, and they exhibit much more aggression than dwarf males. In contrast, dwarf males spend 20% of their time feeding, whereas nest males hardly ever feed. 
 
4. Nest males accumulate reserves before breeding and use these up before taking a reproductive break, thereby performing a ‘capital breeder’ strategy. In contrast, dwarf males use assimilated energy immediately for reproduction, thus acting as ‘income breeders’. This is a requirement of their spawning tactic, which only works out with a small and slim body. 
 
5. A field experiment showed that nest males lose weight by their restricted feeding opportunities while holding a nest, which would allow them to hold a territory for 103 days on average. Due to their reproductive investment, however, they held territories only for a mean period of 33 days, which reveals the relative importance of opportunity costs and reproductive expenditure. 
 
6. Nest males are also limited by the requirement to fertilize each egg of a clutch with a separate ejaculate. Their ejaculation rate and the number of sperm released both decline sharply after 5 h, whereas undisturbed spawning lasts 2–4 h longer than that. 
 
7. There is a strong allometric relationship between body mass and gonad weight, with smaller males of both tactics investing disproportionately more in testes than large males. The major limitation of dwarf males is apparently access to spawning females, which is prevented by the monopolization of nest owners and becomes more difficult with increasing size of dwarf males. 
 
8. Our results show that different males in a population may act as capital or income breeders depending on tactic and may face very different limitations, which is a direct result of highly divergent spawning tactics and resulting body sizes. 
 
9. We argue that capital and income breeding are useful concepts to understand divergent life history decisions associated with alternative reproductive tactics, i.e. behavioural polymorphisms within a species and within one sex. It might turn out that in general, bourgeois tactics rather adopt a capital breeding strategy whereas parasitic tactics are inclined to perform as income breeders, due to the diverging constraints faced by these types of reproduction, although we discuss possible exceptions.

title: Why are reproductively parasitic fish males so small?—influence of tactic-specific selection
abstract: Despite the wide prevalence of alternative reproductive tactics, little attention has been paid to why reproductively parasitic males are so small. In this study, we tackled this issue in a shell-brooding fish Lamprologus callipterus. Sneaky ‘dwarf males’ of this fish remain much smaller than bourgeois conspecifics throughout their life and employ a unique parasitic tactic, i.e. entering into a gastropod shell where a female is spawning, passing through the space between the female and shell wall and staying behind her to ejaculate throughout the spawning event. Here, we tested the prediction that they remain small to get past her through the shell spaces by interpopulation comparison. We showed, across populations, a negative allometry for sexual size dimorphism, an exponential increase of female size with an increase in shell size and a negative correlation between the magnitude of sexual size dimorphism and shell size. These results suggest that the inner spaces strongly regulate dwarf male size. We conclude that the small bodies of dwarf males arise from adaptation to their unique reproductive behaviour.

title: Description of alternative male reproductive tactics in a shell-brooding cichlid, Telmatochromis vittatus, in Lake Tanganyika
abstract: Telmatochromis vittatus (Cichlidae) is a Tanganyikan substrate brooder which spawns in the gastropod-shell nests of a cichlid, Lamprologus callipterus. We describe male reproductive tactics of T. vittatus in and around the shell nests, where males of various sizes were found. Based on utilization patterns of the shell nests, interactions among males, and spawning behaviors, males could be categorized into four types based on reproductive tactics and in order of body size: sneaker males, satellite males, territorial males and piracy males. Size range of males in tactic groups rarely overlapped. Territorial males defended shell nests harboring multiple females, but during pair-spawning they were occasionally taken over by large piracy males that visited several nests repeatedly. Small sneaker males darted to pair-spawning territorial males and might ejaculate sperm. Satellite males did not perform parasitic spawning but pair-spawned in a single shell outside the nests. Spawning of satellite males was infrequently parasitized. The largest gonado-somatic index (GSI) was found in sneaker males followed by piracy males, territorial males and satellite males, suggesting that gonadal investment of males using the four tactics may be consistent with intensity or risk of sperm competition.

title: Reproductive-Tactic-Specific Variation in Sperm Swimming Speeds in a Shell-Brooding Cichlid1
abstract: Theory predicts that males experiencing elevated levels of sperm competition will invest more in gonads and produce faster-swimming sperm. Although there is ample evidence in support of the first prediction, few studies have examined sperm swimming speed in relation to sperm competition. In this study, we tested these predictions from sperm competition theory by examining sperm characteristics in Telmatochromis vittatus, a small shell-brooding cichlid fish endemic to Lake Tanganyika. Males exhibit four different reproductive tactics: pirate, territorial, satellite, and sneaker. Pirate males temporarily displace all other competing males from a shell nest, whereas sneaker males always release sperm in the presence of territorial and satellite males. Due to the fact that sneakers spawn in the presence of another male, sneakers face the highest levels of sperm competition and pirates the lowest, whereas satellites and territorials experience intermediate levels. In accordance with predictions, sperm from sneakers swam faster than sperm from males adopting the other reproductive tactics, whereas sperm from pirates was slowest. Interestingly, we were unable to detect any variation in sperm tail length among these reproductive tactics. Thus, sperm competition appears to have influenced sperm energetics in this species without having any influence on sperm size.

title: Sperm phenotypic plasticity in a cichlid: a territorial male's counterstrategy to spawning takeover
abstract: Studies of sperm competition in species with alternative reproductive tactics (ARTs) often pay attention to the differences in investments in sperm between sneakers facing a higher sperm competition risk and bourgeois males facing a lower risk. Here, we examined within-tactic as well as among-tactic variations in sperm investments in the Lake Tanganyika shell-brooding cichlid Telmatochromis vittatus, a species with 2 types of parasitic tactics. Territorial male T. vittatus suffer reproductive parasitism by both smaller sneaker males and larger pirate males ("pirates" take over the spawning event during which territorial males perform sneaking as a counterstrategy). We hypothesized that both territorial males living under the risk of pirates and sneakers face increased risk of sperm competition and therefore should produce high-quality sperm compared with both territorial males that experienced no piracy risk and pirates. As expected, field studies showed that the former 2 males produced longer lived sperm than the other males. Aquarium experiments demonstrated that a visual stimulus of a pirate was enough to induce an increase in sperm longevity in territorial males compared with when no such stimulus was given. These results indicate that territorial male T. vittatus can plastically adjust at least one sperm quality trait in response to piracy risk. Moreover, long-term monitoring of males in the field showed that small territorial males grow into large territorial males and finally into pirates, so ARTs are not fixed over life. Accordingly, male T. vittatus appear to ontogenetically change their sperm longevity in response to size-dependent sperm competition risks.

title: Mating craters of Cyathopharynx furcifer (Cichlidae) are individually specific, extended phenotypes
abstract: Sexually selected traits have been intensively studied in the recent past, but there is comparatively little information about the signalling value of the extended phenotype, despite its unique potential in mate choice and intersexual competition. Individuals manipulating their environment to create a nonbodily ornament may reveal their immediate quality more reliably than by using conventional signals. Nonbodily ornaments may also integrate a bearer’s behaviour over a recent interval, which would save the potential receiver repeated or prolonged sampling of the signaller’s behaviour. We investigated factors involved in the production of a nonbodily ornament and assessed its potential as a signal in the reproductive context. The crater-building behaviour of the Lake Tanganyika cichlid Cyathopharynx furcifer was observed and manipulated in the field and laboratory. Craters were maintained for prolonged periods, and crater size correlated with male body size, building behaviour and crater location in the lek. Larger craters had a higher ‘neatness index’, and this index correlated negatively with a male condition factor. Behaviour of cichlids after a storm and our manipulations of crater size showed that crater size is a repeatable, individual character; destroyed craters were rebuilt to their original sizes roughly within 1 day and both, experimentally enlarged and diminished craters were immediately reconstructed to their original sizes. We conclude that mating craters are an individually characteristic, extended phenotype of the building males and potentially a suitable signal for conspecifics, providing information about the owner’s size, capability and condition.

title: The Cichlid Fish Assemblages of Lake Tanganyika: Ecology, Behaviour and Evolution of its Species Flocks
abstract: Publisher Summary This chapter focuses on the cichlid fauna of Lake Tanganyika, the least studied of the African Great Lakes. It places particular emphasis on the faunas of the rocky shore habitats, and explores the great species diversity that may have arisen. The chapter describes the evolutionary scenarios within the lake and the biology of these fishes. Several of the more parsimonious explanations as to how the extant species diversity is maintained are considered, and examples of resource partitioning among coexisting species along a variety of resources axes, the most likely mechanism of coexistence, are given. It also illustrates several examples of the diversity of feeding and breeding habits seen both within groups and within species. Moreover, the cichlids of Lake Tanganyika offer splendid, perhaps unique, opportunities for studies of ecology, behavior and evolution, yet there is a paucity of quantitative information concerning the ecology and evolution of its species flocks, such that our present level of knowledge is insufficient on which to form comprehensive theories regarding factors structuring these communities.

title: Inter-population variation in the mating system of a substrate-breeding cichlid in lake tanganyika
abstract: Fishes of the tribe lamprologini in Lake Tanganyika are substrate brooders that guard young from predators until they become independent (Kuwamura, 1986, 1997; Yanagisawa, 1987; Barlow, 1991; Keenleyside, I991). In some species, the young are guarded by both parents, whereas in other species mainly or exclusively by the mother. Biparental guarding is prevalent in species of which young forage in assemblage in water and are vulnerable to predators. Removal of one of the two guardian parents results in quick loss of young by predation (Barlow, 1991). Biparental care keeps both parents from further mating, and promotes monogamy in these fishes (Kuwamura, 1986, 1997; Barlow, 1991; Keenleyside, 1991). By contrast, maternal care is seen in species of which young are benthic around nests and cryptic (Kuwamura 1986, 1997; Barlow, 1991; Keenleyside, 1991). Such maternal care allows the male to mate with other females, and promotes polygyny (Yanagisawa, 1987; Sato, 1994; Takemon & Nakanishi, 1998). Neolamprologus tetracanthus belonging to this tribe is a substrate brooder of which young ( f ry<l ,5 cm) are benthic and cryptically coloured (Kuwamura, 1986, 1997; Brichard, 1989). This benthophagous species commonly inhabits the sandy bottom, which is covered with pebbles or stones, and is found from most shores all around the lake (Brichard, 1989; Kohda et al., 1996). We found great differences in the mating system of this fish between the two populations at the northern and southern ends of this lake: monogamy in the north, whereas extreme-polygyny (a male encompassing up to 14 females at a time) in the south. To our knowledge, such wide variation between populations within a species has not been reported in cichlids, or in fishes. Here we report the mating systems and brood guarding patterns in the two populations of N. tetracanthus.

title: Unusual allometry for sexual size dimorphism in a cichlid where males are extremely larger than females
abstract: When males are the larger sex, a positive allometric relationship between male and female sizes is often found across populations of a single species (i.e. Rensch’s rule). This pattern is typically explained by a sexual selection pressure on males. Here, we report that the allometric relationship was negative across populations of a shell-brooding cichlid fish Lamprologus callipterus, although males are extremely larger than females. Male L. callipterus collect and defend empty snail shells in each of which a female breeds. We found that, across six populations, male and female sizes are positively correlated with not only sexual and fecundity selection indices, but also with shell sizes. Given their different reproductive behaviours, these correlations mean that males are required to be more powerful, and thus larger, to transport larger shells, while female bodies are reduced to the shell size to enable them to enter the shells. Among the three size selections (sexual selection, fecundity selection and shell size), shell size explained the allometry, suggesting that females are more strongly subject to size selection associated with shell size availability than males. However, the allometry was violated when considering an additional population where size-selection regimes of males differed from that of other populations. Therefore, sexual size allometry will be violated by body size divergence induced by multiple selection regimes.

title: The evolution of reproductive and genomic diversity in ray-finned fishes
abstract: Collectively, ray-finned fishes (Actinopterygii) display far more diversity in many reproductive and genomic features than any other major vertebrate group. Recent large-scale comparative phylogenetic analyses have begun to reveal the evolutionary patterns and putative causes for much of this diversity. Several such recent studies have offered clues to how different reproductive syndromes evolved in these fishes, as well as possible physiological and genomic triggers. In many cases, repeated independent origins of complex reproductive strategies have been uncovered, probably reflecting convergent selection operating on common suites of underlying genes and hormonal controls. For example, phylogenetic analyses have uncovered multiple origins and predominant transitional pathways in the evolution of alternative male reproductive tactics, modes of parental care and mechanisms of sex determination. They have also shown that sexual selection in these fishes is repeatedly associated with particular reproductive strategies. Collectively, studies on reproductive and genomic diversity across the Actinopterygii illustrate both the strengths and the limitations of comparative phylogenetic approaches on large taxonomic scales.

title: On the evolutionary pathway of parental care in mouth–brooding cichlid fishes
abstract: Evolutionary theory predicts that differences in parental care patterns among species arose from interspecific differences in the costs and benefits of care for each sex. In Galilee St Peter's fish, Sarotherodon galilaeus (Cichlidae), male care, female care and biparental care all occur in the same population. We exploit this unusual variability to isolate conditions favouring biparental versus uniparental mouth–brooding by males or females. We first review a game–theoretic model of parental care evolution, predictions of which we test experimentally in this paper. Manipulations of the operational sex ratio show that males and females desert their offspring more frequently when the costs of care are high (in terms of lost mating opportunities). Breeding trials with males of different sizes show that small fathers desert more frequently than large fathers. We attribute this to the associated difference in the fitness benefit of biparental care relative to female–only care. Our experimental results confirm that in St Peter's fish the probability of caring is determined facultatively according to current conditions at each spawn. The experiments and model together suggest that interspecific variation in remating opportunities and clutch size may be responsible for differences in care patterns within the sub–family Tilapiini. Our results support the hypothesis that biparental mouth–brooding was the ancestral state of both male and female uniparental mouth–brooding in cichlid fishes.

title: PHYLOGENETIC PERSPECTIVES IN THE EVOLUTION OF PARENTAL CARE IN RAY-FINNED FISHES
abstract: Among major vertebrate groups, ray-finned fishes (Actinopterygii) collectively display a nearly unrivaled diversity of parental care activities. This fact, coupled with a growing body of phylogenetic data for Actinopterygii, makes these fishes a logical model system for analyzing the evolutionary histories of alternative parental care modes and associated reproductive behaviors. From an extensive literature review, we constructed a supertree for ray-finned fishes and used its phylogenetic topology to investigate the evolution of several key reproductive states including type of parental care (maternal, paternal, or biparental), internal versus external fertilization, internal versus external gestation, nest construction behavior, and presence versus absence of sexual dichromatism (as an indicator of sexual selection). Using a comparative phylogenetic approach, we critically evaluate several hypotheses regarding evolutionary pathways toward parental care. Results from maximum parsimony reconstructions indicate that all forms of parental care, including paternal, biparental, and maternal (both external and internal to the female reproductive tract) have arisen repeatedly and independently during ray-finned fish evolution. The most common evolutionary transitions were from external fertilization directly to paternal care and from external fertilization to maternal care via the intermediate step of internal fertilization. We also used maximum likelihood phylogenetic methods to test for statistical correlations and contingencies in the evolution of pairs of reproductive traits. Sexual dichromatism and nest construction proved to be positively correlated with the evolution of male parental care in species with external fertilization. Sexual dichromatism was also positively correlated with female-internal fertilization and gestation. No clear indication emerged that female-only care or biparental care were evolutionary outgrowths of male-only care, or that biparental care has been a common evolutionary stepping stone between paternal and maternal care. Results are discussed in the context of prior thought about the evolution of alternative parental care modes in vertebrates.

title: Evolutionary transitions in parental care in cichlid fish
abstract: Cichlid fishes (Cichlidae) are well suited for testing theories of the evolution of vertebrate parental care. These freshwater teleost fish provide parental care for their offspring, display many different forms of care and have interspecific variation in which sex stays with the young. Here, we assemble the first family–wide composite phylogeny based on morphological and molecular studies, and trace two sets of character evolution: form of care (substrate guarding and mouthbrooding), and sex of care–giver (biparental, female–only, and male–only). Mouthbrooding has evolved from ancestral substrate guarding with 10 to 14 transitions and 0 to 3 reversals. The data support hypothesized transitions in the sex of care–giver, with uniparental female care having arisen from biparental care 21 to 30 times with 0 to 10 reversals. There is also evidence that male–only care evolved once from biparental care. These transitions in parental care characters are the most numerous reported for any family of vertebrates and, to our knowledge, provide the first quantitative support for models of parental care evolution in fish.

title: What, if anything, is a Tilapia?-mitochondrial ND2 phylogeny of tilapiines and the evolution of parental care systems in the African cichlid fishes.
abstract: We estimated a novel phylogeny of tilapiine cichlid fish (an assemblage endemic to Africa and the Near East) within the African cichlid fishes on the basis of complete mitochondrial NADH dehydrogenase subunit 2 (ND2) gene sequences. The ND2 (1,047 bp) gene was sequenced in 39 tilapiine cichlids (38 species and 1 subspecies) and in an additional 14 nontilapiine cichlid species in order to evaluate the traditional morphologically based hypothesis of the respective monophyly of the tilapiine and haplochromine cichlid fish assemblages. The analyses included many additional cichlid lineages, not only the so-called tilapiines, but also lineages from Lake Tanganyika, east Africa, the Neotropics and an out-group from Madagascar with a wide range of parental care and mating systems. Our results suggest, in contrast to the historical morphology-based hypotheses from Regan (1920, 1922 ), Trewavas (1983), and Stiassny (1991), that the tilapiines do not form a monophyletic group because there is strong evidence that the genus Tilapia is not monophyletic but divided into at least five distinct groups. In contrast to this finding, an allozyme analysis of Pouyaud and Agnèse (1995), largely based on the same samples as used here, found a clustering of the Tilapia species into only two groups. This discrepancy is likely caused by the difference in resolution power of the two marker systems used. Our data suggest that only type species Tilapia sparrmanii Smith (1840) should retain the genus name TILAPIA: One particular group of tilapiines (composed of genera Sarotherodon, Oreochromis, Iranocichla, and Tristramella) is more closely related to an evolutionarily highly successful lineage, the haplochromine cichlids that compose the adaptive radiations of cichlid species flocks of east Africa. It appears that the highly adaptable biology of tilapiines is the ancestral state for all African cichlids and that the more stenotypic lifestyle of the haplochromine cichlids is derived from this condition. We reconstructed the evolution of the highly variable parental care systems on the basis of the most inclusive composite phylogeny to date of the African, Neotropical, and Madagascan cichlids with special emphasis on a group of tilapiines comprising the substrate-spawning genus Tilapia, and the mouthbrooding genera Sarotherodon and OREOCHROMIS: We demonstrate several independent origins of derived mouthbrooding behaviors in the family Cichlidae.

title: Trade up polygyny and breeding synchrony in avian populations.
abstract: The advent of the molecular techniques used to assign paternity has focused attention on the differences between the social and the genetic mating systems of sexual species. In particular, the interrelations between breeding synchrony-the degree to which the fertility periods of individual females in a population overlap and the degree of extra pair paternity (EPP) in that population, has became a subject of a lively debate. Investigation of the subject can be facilitated by examining the criteria that females use in choosing extra pair partners. These preferences constitute a continuum ranging between two extremes. At one end, there are situations wherein all the females in a population exhibit a preference for males with particular phenotypic markers, and females mated to males lacking such "quality" markers seek extra pair fertilizations from males that do -trade up polygyny. At the other extreme, there are situations wherein females seek to maximize the total number of male partners, rather than secure fertilization by males of particular type -indiscriminate polygyny. Previously, we used game theoretical methods to model the interrelations between breeding synchrony and EPP in the context of indiscriminate polygyny. Here we present an analogous investigation in the context of trade up polygyny. Our results for the two cases, which delimit the range of the possible behavior, are similar. That is, we see that it is the pursuit of extra pair fertilizations opportunities that determines breeding synchrony of populations, rather than the vice versa as has been previously suggested.

title: The environmental potential for polygyny and sexual selection in the sand goby, Pomatoschistus minutus
abstract: How nest size and temporal availability of females affects sexual selection was tested experimentally in the sand goby, Pomatoschistus minutus, a small fish with paternal brood care. The distribution of matings depended on both nest size and female arrival pattern, supporting the model for mating system evolution presented by Emlen & Oring (1977). Polygyny and the opportunity for sexual selection was greatest when nests were large and females arrived asynchronously. In these replicates a single male frequently spawned with all four females. The total number of females spawning was the same for each treatment. Male size was important for mating success in the large nest treatment, whereas female arrival pattern did not affect the size difference between mated and unmated males. This study demonstrates that sexual selection can depend on resource distribution even if all individuals can obtain similar quality resources, a situation where resource monopolization per se should not differentially affect access to mates.

title: Synchronous Female Spawning and Male Mating Behavior in a Land-Locked Population of Japanese Charr, Salvelinus leucomaenis japonicus
abstract: In resource-based promiscuous mating systems, synchronous spawning of females affects competition among males and variation in the reproductive success of males. We documented the mating behavior of Japanese charr (Salvelinus leucomaenis japonicus) through an annual breeding season to examine the relationship between female spawning synchrony and male mating behavior. Females spawned highly synchronously in the population studied, i.e., approximately half the spawning was finished within the first three days of the entire spawning season (11 days). The daily operational sex ratio (OSR) was nearly 1:1 through the spawning period. The number of males around a spawning female was very small (1.21±0.49 males per female) over the spawning ground and period, suggesting that a competitive male could effectively chase subordinate males away from a spawning female. A few males attempted to sneak near the oviposition site of females (16%; 9 of 57 breeding groups), while some males adopted sneaking tactics in the initial phase of females’ spawning (24%). We did not observe any males to succeed in sneak fertilizations. We conclude that in this Japanese charr population, the synchronous spawning of females was related to the unbiased daily OSR, male aggregation around females, and consequently whether and how efficiently males engaged in sneak mating behavior.

title: Complete mitochondrial DNA replacement in a Lake Tanganyika cichlid fish
abstract: We used nuclear and mitochondrial DNA (mtDNA) sequences from specimens collected throughout Lake Tanganyika to clarify the evolutionary relationship between Lamprologus callipterus and Neolamprologus fasciatus. The nuclear data support the reciprocal monophyly of these two shell‐breeding lamprologine cichlids. However, mtDNA sequences show that (i) L. callipterus includes two divergent and geographically disjunct (North–South) mtDNA lineages; and that (ii) N. fasciatus individuals cluster in a lineage sister group to the northern lineage of L. callipterus. The two mtDNA lineages of L. callipterus diverged c. 684 kya to 1.2 Ma, coinciding with a major water level low stand in Lake Tanganyika, which divided the lake into isolated sub‐lakes. This suggests that the two mtDNA lineages originated as the result of the separation of L. callipterus populations in different sub‐basins. The incongruent phylogenetic position of N. fasciatus can best be explained by an ancient unidirectional introgression from L. callipterus into N. fasciatus. Remarkably, our data indicate that this event resulted in the complete mtDNA replacement in N. fasciatus. Our data suggest that hybridization occurred soon after the divergence of the two L. callipterus mtDNA lineages, probably still during the water level low stand, and that subsequently the invading mtDNA lineage spread throughout the lake.

title: Evolutionary history of Lake Tanganyika's scale-eating cichlid fishes.
abstract: Although Lake Tanganyika is not the most species-rich of the Great East African Lakes it comprises by far the greatest diversity of cichlid fishes in terms of morphology, ecology, and breeding styles. Our study focuses on the Tanganyikan cichlid tribe Perissodini, which exhibits one of the most peculiar feeding strategies found in cichlids-scale-eating. Their evolutionary history was reconstructed from 1416 bp DNA sequence of two mitochondrial genes (ND2 and partial control region) and from 612 AFLP markers. We confirm the inclusion of the zooplanktivorous genus Haplotaxodon in the tribe Perissodini, and species status of Haplotaxodon trifasciatus. Within the Perissodini, the major lineages emerged within a short period roughly 1.5-2 MYA, which makes their radiation slightly younger than that of other Tanganyikan cichlid tribes. Most scale-eaters evolved in deep-water habitat, perhaps associated with the previously documented radiations of other deep-water dwelling cichlid lineages, and colonized the shallow habitat only recently.

title: Rapid radiation, ancient incomplete lineage sorting and ancient hybridization in the endemic Lake Tanganyika cichlid tribe Tropheini.
abstract: The evolutionary history of the endemic Lake Tanganyika cichlid tribe Tropheini, the sister group of the species flocks of Lake Malawi and the Lake Victoria region, was reconstructed from 2009 bp DNA sequence of two mitochondrial genes (ND2 and control region) and from 1293 AFLP markers. A period of rapid cladogenesis at the onset of the diversification of the Tropheini produced a multitude of specialized, predominantly rock-dwelling aufwuchs-feeders that now dominate in Lake Tanganyika's shallow habitat. Nested within the stenotopic rock-dwellers is a monophyletic group of species, which also utilize more sediment-rich habitat. Most of the extant species date back to at least 0.7 million years ago. Several instances of disagreement between AFLP and mtDNA tree topology are attributed to ancient incomplete lineage sorting, introgression and hybridization. A large degree of correspondence between AFLP clustering and trophic types indicated fewer cases of parallel evolution of trophic ecomorphology than previously inferred from mitochondrial data.

title: Pinniped phylogenetic relationships inferred using AFLP markers
abstract: Amplified fragment length polymorphisms (AFLPs) are widely used for phylogenetic reconstruction in plants but their use in animal taxa has been limited, and in mammals rare. In addition, their use has been largely limited to shallow relationships amongst species or subspecies. Here, we genotype 23 pinniped species for 310 AFLP markers and find a strong phylogenetic signal, with individuals coclustering within species, and overall a good agreement between our phylogeny and those constructed using mitochondrial DNA and nuclear sequences even at nodes ∼15 million years old. Although supporting the existing ideas about pinniped relationships, our data shed light on relationships within the hitherto relatively unresolved Phocine species group, and provide further supporting evidence for raising two subspecies of Zalophus californianus, Z. c. californianus and Z. c. wollebaeki, to species level. Plotting AFLP divergence time estimates against those based on both mtDNA and nuclear sequences we find strong linear relationships, suggesting that the different markers are evolving in a clocklike fashion. These data further emphasize the utility of AFLP markers as general tools for phylogenetic reconstruction.

title: AFLP phylogeny of the snubnose darters and allies (Percidae: Etheostoma) provides resolution across multiple levels of divergence.
abstract: The snubnose darters (Percidae: subgenus Ulocentra) are a group of small, brightly colored North American freshwater fish belonging to the genus Etheostoma. Phylogenetic relationships among snubnose species have been a challenge to resolve at all levels of divergence, from the monophyly of species to deeper relationships among subgenera. Here, we used amplified fragment length polymorphisms (AFLPs) to estimate phylogenetic relationships among species from three closely related subgenera: Ulocentra, Etheostoma, and Nanostoma. With nearly complete sampling of recognized species, our analysis yielded a robust tree with statistical support at all nodes. Support was strongest for shallower relationships; support for internal nodes was either comparable to or greater than that of previous studies based on mitochondrial sequence data. Most recovered relationships were consistent with earlier hypotheses based on morphology or mtDNA sequences, with the exception of Etheostoma histrio, which was recovered as sister to Ulocentra. Our analysis indicates that careful use of AFLPs can yield statistically robust estimates of evolutionary relationships across multiple levels of divergence.

title: The influence of sexual selection and ecological constraints on an extreme sexual size dimorphism in a cichlid
abstract: The degree of sexual size dimorphism (SSD) in the cichlid Lamprologus callipterus is the greatest recorded among species in which males exceed females in body size. Males collect and defend empty snail shells in which females breed. We report on the potential importance of different selection mechanisms and constraints on male and female body sizes. To test for the importance of sexual selection, we measured the reproductive behaviour of differently sized males and their reproductive success in the field. Larger males did not have larger nests than smaller males. In the laboratory, we performed male–male competition and female choice experiments and found an indirect effect of large body size on competitive ability and male reproductive success. Neither in the field nor in the laboratory did females choose particular males. To test for the importance of ecological constraints, we studied the significance of shell size for male and female body sizes. Shell-carrying experiments revealed a minimum male threshold size for the ability to carry Neothauma shells. Females chose larger shells than available on average in males' nests, and their reproductive success increased with shell size. However, their size is limited by the availability of large shells. We conclude that the divergent influence of an ecological constraint on male and female body sizes is primarily responsible for this extreme SSD, in combination with size-dependent mechanisms of intrasexual selection.

title: An optimality approach to male and female body sizes in an extremely size-dimorphic cichlid fish
abstract: Can intrasexual selection on male size and natural selection on female size in the shell-brooding cichlid Lamprologus callipterus explain the greatest male/female sexual size dimorphism (SSD) reported to date among animals?  (1) Mortality model to predict male and female body sizes. (2) Gain rate maximization and (3) combined mortality/gain rate maximization model to predict female size. We compared observed body sizes in the field with predictions from these models.  (1) Body size at sexual maturity is determined by a trade-off between survival to sexual maturity and (a) number of mates in males and (b) fecundity in females. (2) and (3) Female body size is constrained by increasing search costs, because shells large enough for breeding become limited as females become larger.  (1) Male body size predicted by the mortality model matches size as found in the field closely, but the model strongly overestimated female size. (2) The female body size predicted by the gain rate maximization model fitted well within the range of sizes observed in the field. (3) Including the effect of mortality into this model hardly changes the prediction.  (1) Intersexual selection can explain male body size, but fecundity selection alone cannot explain female size. (2) and (3) The breeding substrate strongly constrains female size. The extreme male-biased SSD is due to strong selection on the body sizes of both sexes in opposite directions.

title: Variance in reproductive success and the opportunity for selection in a serially monogamous species: simulations of the mating system of Tropheus (Teleostei: Cichlidae)
abstract: Sexual selection is believed to play a major role in speciation processes of cichlid fishes in the East African Great Lakes, in particular Lakes Malawi and Victoria, by driving diversification in male nuptial coloration. In Lake Tanganyika, a high rate of color pattern diversification is found in the genus Tropheus, where more than 100, mostly allopatric, color morphs have been described. Whether color pattern differentiation in Tropheus could follow from sexual selection as well, has not yet been conclusively shown. Unlike typical sexually selected species, Tropheus are sexually monomorphic, establish temporary pair bonds and spawn monogamously. Variance in mating success among individuals would still allow for sexual selection, but due to their high population census sizes, parentage data from natural populations are difficult to obtain. Simulations designed according to existing data on mating behavior in Tropheus suggested that variance in male reproductive success can be substantial and lead to levels of opportunity for selection, which are similar to values estimated, for example, from natural populations of sexually dimorphic songbirds. Variance in male success was mostly affected by the way females perceived and reacted to differences between males, and to a lesser extent by the duration of components of the reproductive cycles of males and females. These results indicate that future work on the importance of sexual selection in Tropheus should concentrate on mate choice cues and female mate choice behavior, but does not depend critically on the acquisition of more detailed life history data.

title: Concordant female mate preferences in the cichlid fish Tropheus moorii
abstract: Discriminating female mate preferences enhance the variance in reproductive success among males of a population and create a potential for sexual selection, which can account for trait evolution and diversification. Fish color patterns are among the prime targets of mate choice-driven sexual selection. Populations of the cichlid Tropheus from Lake Tanganyika display remarkable geographic color pattern variation, but the role of female choice in their rapid and rich phenotypic diversification is unclear. Males and females establish a pair bond prior to spawning monogamously, but as brood care is strictly maternal, female investment in reproduction is high and the operational sex ratio is male-biased. Therefore, variance in male reproductive success can accrue if individual males succeed repeatedly in securing a mate. To test this prediction in the red colored Tropheus moorii “Chimba”, four pairs of males were presented to a series of females and female mate preferences were inferred from pairwise interactions. There was a significant difference in mating success between the males of each pair (P < 0.001 over all trials), as—with one exception—females shared preferences for the same males. Male courtship activity was strongly correlated with female choice. Our experiment suggests that female choice contributes to the variance in male reproductive success in the tested population.

title: The mismeasurement of sexual selection
abstract: Sexual selection can explain major micro‐ and macro‐evolutionary patterns. Much of current theory predicts that the strength of sexual selection (i) is driven by the relative abundance of males and females prepared to mate (i.e. the operational sex ratio, OSR) and (ii) can be generally estimated by calculating intra‐sexual variation in mating success (e.g. the opportunity for sexual selection, Is). Here, we demonstrate the problematic nature of these predictions. The OSR and Is only accurately predict sexual selection under a limited set of circumstances, and more specifically, only when mate monopolization is extremely strong. If mate monopolization is not strong, using OSR or Is as proxies or measures of sexual selection is expected to produce spurious results that lead to the false conclusion that sexual selection is strong when it is actually weak. These findings call into question the validity of empirical conclusions based on these measures of sexual selection.

title: On the Opportunity for Sexual Selection, the Bateman Gradient and the Maximum Intensity of Sexual Selection
abstract: Bateman's classic paper on fly mating systems inspired quantitative study of sexual selection but also resulted in much debate and confusion. Here, I consider the meaning of Bateman's principles in the context of selection theory. Success in precopulatory sexual selection can be quantified as a “mating differential,” which is the covariance between trait values and relative mating success. The mating differential is converted into a selection differential by the Bateman gradient, which is the least squares regression of relative reproductive success on relative mating success. Hence, a complete understanding of precopulatory sexual selection requires knowledge of two equally important aspects of mating patterns: the mating differential, which requires a focus on mechanisms generating covariance between trait values and mating success, and the Bateman gradient, which requires knowledge of the genetic mating system. An upper limit on the magnitude of the selection differential on any sexually selected trait is given by the product of the standard deviation in relative mating success and the Bateman gradient. This latter view of the maximum selection differential provides a clearer focus on the important aspects of precopulatory sexual selection than other methods and therefore should be an important part of future studies of sexual selection.

title: Assortative mating preferences between colour morphs of the endemic Lake Tanganyika cichlid genus Tropheus
abstract: Female mate preferences effectuate reproductive isolation among and sexual selection within species. Both mechanisms have been associated with the diversification and speciation of cichlid species flocks of the East African Great Lakes. In Lake Tanganyika, the endemic genus Tropheus has diversified into >100 geographic colour morphs. Although distributed allopatrically at present, water level fluctuations have repeatedly displaced and merged the benthic, rock-dwelling populations. Tests for assortative mating were performed to explore the potential for reproductive isolation between morphs in secondary contact, and to assess the importance of sexual selection for the diversification of this group. In contrast to other haplochromine cichlids, Tropheus is a sexually monochromatic, territorial and maternally mouthbrooding fish, which establishes temporary pair bonds prior to spawning. Female mate preference trials involved two-way choices between a homotypic and a heterotypic male and were conducted on allopatric populations of red and blue morphs from the southern part of Lake Tanganyika. Female affiliation time near each male’s compartment did not predict the mate preferences subsequently expressed in unrestrained interactions after removal of the compartment separators (spawning, pseudospawning and courtship). Consequently, mate preferences were inferred from unrestrained interactions with one test male at a time in replicate observation sessions. Of the 23 females tested, 13 courted, pseudospawned or spawned with the homotypic male, one blue female courted a red male, and nine females expressed no sexual motivation. The assortative mate preferences in the experiments (P < 0.01) suggest that colour differentiation between Tropheus populations can effectuate reproductive isolation, and is consistent with the notion that sexual selection contributed to the diversification of the genus.

title: Variable discrimination and asymmetric preferences in laboratory tests of reproductive isolation between cichlid colour morphs
abstract: Mating behaviour affects reproductive isolation and phenotypic differentiation. In Lake Tanganyika, the cichlid fish Tropheus moorii diversified into numerous, currently allopatric colour variants. Allopatric isolation is periodically interrupted by dispersal and secondary contact during lake level fluctuations, making long‐term differentiation partly dependent on assortative mating. Laboratory experiments with two moderately distinct morphs revealed assortative female preferences in one (Nakaku), but random mate choice in the other morph (Mbita). No discrimination was apparent between two subtly differentiated morphs (Chimba and Moliro). Tested against each other in a previous study, the highly distinct Moliro and Nakaku exhibited strong assortative preferences. The correlation between colour pattern similarity and mate discrimination suggests that allopatry and philopatric behaviour are less crucial for the maintenance of differentiation between highly distinct morphs than for more similar morphs. Interestingly, the asymmetric isolation in one pair of morphs is congruent with a pattern of unidirectional mitochondrial introgression between populations.

title: Size-assortative mating in the absence of mate choice
abstract: Size-assortative mating is one of the most common mating patterns in nature. Nevertheless, the underlying behavioural mechanisms have received little attention. Assortment is typically assumed to result from mate choice, which can be coupled with differences in competitive potential. We investigated the behavioural mechanisms underlying size-assortative mating in a monogamous, biparental goby cichlid, where mutual mate choice should be expected. We performed three field experiments with females and males of Eretmodus cyanostictus to test for the existence of mate preferences in general and with regard to size: (1) a sequential presentation of differently sized potential partners; (2) a removal of partners combined with surveillance until re-pairing with a new partner; and (3) the simultaneous release of new and original partners on the experimental territories. In the removal experiment, we found evidence for weak preferences for large partners relative to own size and to the original partner's size, but pairs were formed irrespective of these preferences. The ecological importance of being paired appears to reduce choosiness and to override mate preferences. Territory ownership was quickly decided by aggressive interactions between original and new partners, and in both sexes the larger contestant won and was immediately accepted as partner by the resident. Our results suggest that strong intrasexual competition can be a powerful promoter of size-assortative mating even in the absence of active mate choice.

title: Who to include in measures of sexual selection is no trivial matter.
abstract: In many animals acquiring limited reproductive opportunities involves competition for resources, mates and opposite-sex gametes. There is ambiguity in which competitive steps are included in measures of sexual selection: individuals who fail to obtain resources necessary for reproduction are often excluded. We illustrate the implications of variation in who is included in measures of selection. We quantified selection on male length and the opportunity for selection associated with nest acquisition, mate acquisition, and fertility of mates at two levels of density and two levels of nest availability in the sand goby. Both measures varied significantly across the three episodes of selection. Nest and mate acquisition contributed substantially to the overall opportunity for selection and selection on male size. Focusing only on males with nests led to lower estimates of selection. The effects of density and nest availability depended on the selective episodes considered. While there is nothing wrong with focusing on particular episodes of interest, inconsistency in who is included in measures of sexual selection across studies will make it difficult to answer broad research questions.

title: How cuckoldry can decrease the opportunity for sexual selection: Data and theory from a genetic parentage analysis of the sand goby, Pomatoschistus minutus
abstract: Alternative mating strategies are common in nature and are generally thought to increase the intensity of sexual selection. However, cuckoldry can theoretically decrease the opportunity for sexual selection, particularly in highly polygamous species. We address here the influence of sneaking (fertilization thievery) on the opportunity for sexual selection in the sand goby Pomatoschistus minutus, a marine fish species in which males build and defend nests. Our microsatellite-based analysis of the mating system in a natural sand goby population shows high rates of sneaking and multiple mating by males. Sneaker males had fertilized eggs in ≈50% of the assayed nests, and multiple sneakers sometimes fertilized eggs from a single female. Successful males had received eggs from 2 to 6 females per nest (mean = 3.4). We developed a simple mathematical model showing that sneaking in this polygynous sand goby population almost certainly decreases the opportunity for sexual selection, an outcome that contrasts with the usual effects of cuckoldry in socially monogamous animals. These results highlight a more complex and interesting relationship between cuckoldry rates and the intensity of sexual selection than previously assumed in much of the literature on animal mating systems.

title: Genetic mating patterns studied in pools with manipulated nest site availability in two populations of Pomatoschistus minutus
abstract: Pomatoschistus minutus show paternal care in a resource defence mating system. We investigated the effect of nest‐site availability on parasitic spawning. Each experimental pool contained four potentially nest‐building males, four females and nests‐sites in shortage (2) or excess (6). Both treatments were conducted in two populations; one with natural nest‐sites in excess, one with a nest‐site shortage. Microsatellite‐DNA revealed that all nest‐holders had fertilized most of the eggs they tended. Yet, 35% of the nests contained eggs fertilized by another male and 14.4% of the males had performed parasitic spawning. There was no site or treatment effect. Several females spawned in two nests, which coincided with parasitic spawnings, suggesting a cost to the nest‐holder in terms of lost mating success. Nest‐holders with and without eggs and non‐nesting males all spawned parasitically, generating a significantly lower opportunity for sexual selection compared to if there had been no parasitic spawning.

title: Effects of extra-pair and within-pair reproductive success on the opportunity for selection in birds
abstract: The number of mates, their fecundity, and the number of extra-pair fertilizations can all affect male reproductive success in biparental species. Extra-pair mating in birds has been of particular interest, because it could generate strong levels of sexual selection even when a species is socially monogamous. We examined how extra-pair fertilizations affect the opportunity for selection in the sexually dimorphic common yellowthroat (Geothlypis trichas) and the sexually monomorphic house wren (Troglodytes aedon). We were able to identify sires for at least 95% of all nestlings, and, thus, we were able to make a nearly complete accounting of male reproductive success. Although extra-pair fertilizations were common in yellowthroats (26% of young) it contributed little (21%) to the total variance in male reproductive success. Most of the variance in reproductive success (58%) was attributable to the male’s within-pair success, which was influenced primarily by the number of young produced by each mate and the proportion of within-pair young sired. Despite a moderate level of extra-pair fertilizations (10% of young) in house wrens, almost all of the variance in male reproductive success (97%) was attributable to within-pair success, particularly the number of social mates. Although extra-pair fertilizations generally increase the variance in male reproductive success, withinpair reproductive success may be the major source of variation in male reproductive success. Thus, sexual dimorphism in monogamous birds may be influenced more by the number of mates and their fecundity than by extra-pair matings.  extra-pair paternity, sexual selection, common yellowthroat, house wren.

title: The strength of postcopulatory sexual selection within natural populations of field crickets
abstract: Sperm competition and cryptic female choice are likely to exert strong postcopulatory sexual selection and may amplify or ameliorate selection acting via male mating success. However, relatively few studies have estimated the potential strength of postcopulatory sexual selection acting within natural populations. Field crickets in the genus Teleogryllus have been used extensively as laboratory models to study the evolution of polyandry, and the mechanisms of sperm competition and cryptic female choice. However, we have no knowledge of the extent of multiple mating or patterns of sperm utilization in natural populations of these crickets. We used microsatellite loci to examine the strength of postcopulatory sexual selection in Teleogryllus oceanicus and T. commodus. Genotyping of sperm within sperm storage organs revealed high levels of multiple mating by females in both species, but females utilized sperm from just 75% and 60% of their mates, respectively. Paternity was skewed toward a subset of fathers; skew was strongest in T. oceanicus and declined with the number of fathers represented in both species. Thus, postcopulatory sexual selection is strong in natural populations of both species and must be taken into account when estimating total sexual selection acting on these species. More generally, we argue that more effort should be made to measure the strength of postcopulatory sexual selection in other species, and to integrate episodes of pre- and postcopulatory selection, if we are to gain an accurate appreciation of the evolutionary consequences of sexual selection.

title: Methods of parentage analysis in natural populations
abstract: The recent proliferation of hypervariable molecular markers has ushered in a surge of techniques for the analysis of parentage in natural and experimental populations. Consequently, the potential for meaningful studies of paternity and maternity is at an all‐time high. However, the details and implementation of the multifarious techniques often differ in subtle ways that can influence the results of parentage analyses. Now is a good time to reflect on the available techniques and to consider their strengths and weaknesses. Here, we review the leading techniques in parentage analysis, with a particular emphasis on those that have been implemented in readily useable software packages. Our survey leads to some important insights with respect to the utility of the different approaches. This review should serve as a useful guide to anyone who wishes to embark on the study of parentage.

title: Pedigree reconstruction in wild cichlid fish populations
abstract: It is common practice to use microsatellites to detect parents and their offspring in wild and captive populations, in order to reconstruct a pedigree. However, correct inference is often constrained by a number of factors, including the absence of demographic data and ignorance regarding the completeness of parental sampling. Here we present a new Bayesian estimator that simultaneously estimates the pedigree and the size of the unsampled population. The method is robust to genotyping error, and can estimate pedigrees in the absence of demographic data. Using a large‐scale microsatellite assay in four wild cichlid fish populations of Lake Tanganyika (1000 individuals in total), we assess the performance of the Bayesian estimator against the most popular assignment program, Cervus. We found small but significant pedigrees in each of the tested populations using the Bayesian procedure, but Cervus had very high type I error rates when the size of the unsampled population was assumed to be lower than what it was. The need of pedigree relationships to infer adaptive processes in natural populations places strong constraints on sampling design and identification of multigenerational pedigrees in natural populations.

title: Sexual Selection: Harem Size and the Variance in Male Reproductive Success
abstract: Sexual selection is potentially stronger than natural selection when the variance in male reproductive fitness exceeds all other components of fitness variance combined. However, measuring the variance in male reproductive fitness is difficult when nonmating males are absent, inconspicuous, or otherwise difficult to find. Omitting the nonmating males inflates estimates of average male reproductive success and diminishes the variance, leading to underestimates of the potential strength of sexual selection. We show that, in theory, the proportion of the total variance in male fitness owing to sexual selection is approximately equal to H, the mean harem size, as long as H is large and females are randomly distributed across mating males (i.e., 
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$$V_{\mathrm{harem}\,}=H$$
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$$H+( k-1) $$
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$$V_{\mathrm{harem}\,}/ H$$
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