The morphology and independent origin of ovoviviparity in Tiphobia and Lavigeria (Caenogastropoda: Cerithioidea: Paludomidae) from Lake Tanganyika

To better understand the diversification of the endemic thalassoid (i.e. marine-like) cerithioidean gastropods of Lake Tanganyika, as well as the origin and significance of brooding among lake species, we here redescribe the anatomy and ontogeny of the ovoviviparous Tiphobia horei from Lake Tanganyika and compare it to that of Lavigeria sp. A, representing another ovoviviparous lake clade that has acquired a uterine brood pouch independently. Within the phylogenetic framework provided by recent molecular analyses, the distant relation of these two taxa is corroborated by many external and internal anatomical differences. Comparison of the brood pouches demonstrates that they each bear unique features consistent with their independent modification for brooding. Despite representing functionally analogous structures, they also share several similarities in organization likely representing symplesiomorphies of the Lake Tanganyika species flock. The ontogeny is characterized by the presence of a velum and by delayed calcification producing a characteristically wrinkled embryonic cap. Comparison with other brooding cerithioideans reveals that T. horei and Lavigeria sp. A retain many more embryos than other freshwater cerithioideans of comparable size with a uterine brood pouch, possibly facilitated by the presence of longitudinal lamellae. Compartmentalization of the oviduct and delayed calcification is strongly linked to the brooding of embryos.     

Brood care among basommatophorans: a unique reproductive strategy in the freshwater limpet snail Protancylus (Heterobranchia: Protancylidae), endemic to ancient lakes on Sulawesi, Indonesia

In molluscan taxa inhabiting marine environments oviparity and reproduction via planktonic larvae is predominant while incubation and viviparity is most frequently found in taxa inhabiting brackish or freshwater aquatic habitats. Brooding has evolved repeatedly and independently in several limnic taxa among Bivalvia and Gastropoda. However, among basommatophoran gastropods no such cases were yet known. We here report on a unique reproductive strategy involving brood care in the lacustrine freshwater limpet genus Protancylus, endemic to the ancient lakes on central Sulawesi (former Celebes), Indonesia, namely the Lake Poso and the Malili lake system, because this constitutes the first known case of this behaviour among the Basommatophora. Protancylus live exclusively as epizoans on those pachychilid gastropods of the viviparous genus Tylomelania, also a Sulawesi endemic species, that inhabit mostly soft substrates. We found that the two known species Protancylus pileolus from Lake Poso and P. adhaerens from the Malili lake system both retain gelatinous egg strings underneath their outer mantle, where up to 15 (mostly eight or nine) shelled juveniles are brooded. Nourishment is provided within the egg capsule only. Thus, brood care in Protancylus resembles the reproductive strategy found recently among pachychilid gastropods Jagora from the Philippines, but differs from euviviparous (i.e. matrotrophic) incubation among thiarid gastropods possessing a brood pouch with juveniles being nourished via a ‘pseudoplacenta’ in several taxa.     

MORPHOLOGY AND BEHAVIOR OF CRABS AND GASTROPODS FROM LAKE TANGANYIKA,AFRICA: IMPLICATIONS FOR LACUSTRINE PREDATOR-PREY COEVOLUTION

The shells of most lacustrine gastropods are typically small, weakly calcified, and modestly ornamented to unornamented. Similarly, most lacustrine crabs are usually small detritivores with weak chelae. A number of invertebrate taxa in Lake Tanganyika, however, deviate from these generalities. This study explores a predator-prey coevolution model as an explanation for the large, heavily calcified, and ornate gastropods and the robust, durophagous crabs of Lake Tanganyika. The endemic thiarid and viviparid gastropods from Lake Tanganyika have significantly thicker shells and higher frequencies of terminal apertural lip thickening than closely related cosmopolitan taxa from outside the lake. Tanganyikan gastropods also display considerably higher incidence of shell repair, following nonlethal shell damage, than cosmopolitan taxa of the same families. There is a strong positive correlation between gastropod apertural lip thickness and shell repair frequency among all the gastropod species analyzed. The endemic Tanganyikan potamonautid crab Platytelphusa armata (a molluscivore) possesses larger, more robust crushing chelae than other African potamonautid or potamonid crabs. In contrast with the cosmopolitan African crabs, the Tanganyikan crabs display molariform, rather than serrate dentition on their crushing chelipeds. In shell-crushing experiments, the Tanganyikan gastropod shells were an order of magnitude stronger than typical lacustrine gastropod shells, many well within the range of tropical marine gastropod shell strengths. Predation experiments with the endemic gastropods Spekia, Neothauma, Lavigeria spp., Paramelania spp. and the crab Platytelphusa armata showed that increased size, apertural lip thickness or shell sculpture reduced the successful predation rate of P. armata. Crabs with large chelae have a greater ratio of successful: unsuccessful attacks than crabs with small chelae. Among cases of successful predation, crabs with large chelae employed predation methods that required less time and energy (such as crushing the shell in the cheliped) than the methods employed by crabs with small chelae (such as peeling the shell from the aperture or the spire). The morphological, shell-crushing, and aquarium experiment data, considered in concert, provide strong support for the idea that the endemic gastropods and crabs of Lake Tanganyika have coevolved over the past 7 million years.     

Four new species of Gomphocythere (Crustacea, Ostracoda) from Lake Tanganyika, East Africa

Four new species of the ostracod genus Gomphocythere are described from Lake Tanganyika (East Africa): Gomphocythere downingi n. sp. G. coheni n. sp., G. wilsoni n. sp., and G. woutersi n. sp. All species are endemic to the lake and are found within a variety of substrates and depths. The addition of these four new species brings the total number of endemic Gomphocythere species in Lake Tanganyika to nine. Other Gomphocythere species are known from water bodies throughout East and South Africa and in the Levant. Brooding is an important, but not a unique, preadaptation for the persistence and taxic prolific speciation of this lineage in Lake Tanganyika.     

Ancient lakes as evolutionary reservoirs: evidence from the thalassoid gastropods of Lake Tanganyika

Ancient lakes are often collectively viewed as evolutionary hot spots of diversification. East Africa's Lake Tanganyika has long been the subject of scientific interest owing to dramatic levels of endemism in species as diverse as cichlid fishes, paludomid gastropods, decapod and ostracod crustaceans and poriferans. It is the largest and deepest of the African rift lakes, and its endemic fauna has been presented with a stable inland environment for over 10 Myr, offering unique opportunities for within-lake diversification. Although astonishing diversification has been documented in the endemic cichlid fauna of the lake, similar patterns of rapid diversification have long been assumed for other groups. In contrast to this hypothesis of rapid speciation, we show here that there has been no acceleration in the rate of speciation in the thalassoid gastropods of the lake following lake colonization. While limited within-lake speciation has occurred, the dramatic conchological diversity of gastropods presently found within the lake has evolved from at least four major lineages that pre-date its formation by as much as 40 Myr. At the same time, a widespread group of African gastropods appears to have evolved from taxa presently found in the lake. While Lake Tanganyika has been a cradle of speciation for cichlid fishes, it has also been an important evolutionary reservoir of gastropod lineages that have been extirpated outside the basin.     

Spermatophores of thalassoid gastropods (Paludomidae) in Lake Tanganyika, East Africa, with a survey of their occurrence in Cerithioidea: functional and phylogenetic implications

Abstract. With few exceptions, spermatozoa-encapsulating packages in molluscs are known mostly from cephalopods and pulmonate gastropods. Among non-stylommatophoran gastropods, the marine Cerithioidea are second only to the Neritimorpha in the number of species that posess a spermatophore, but they have only rarely been found in freshwater taxa of this superfamily. We describe and compare here the sperm packages of 11 paludomid cerithioideans as part of an ongoing study on the evolution and systematics of the thalassoid (i.e., "marine-like") endemic species flock from Lake Tanganyika. Stereomicroscopic and SEM examination revealed unexpected complexity in shape and structure of spermatophores within paludomids. In addition, we present a survey of other Cerithioidea, which revealed that spermatophores are in general structurally simple and confirmed their presence in 12 marine species (five families) and 15 limnic species (four families), including those of 10 thalassoid species for which spermatophores are described herein. Based on histological sections of the male genital tract, we hypothesize that spermatophores are formed anteriorly, wholly or in part, by the so-called spermatophore-forming organ, and which is considered a synapomorphy of the Paludomidae. In addition, we briefly discuss functional aspects inferred from morphological study of the spermatophore-forming organ, with possible implications for spermatozoan transfer and fertilization. Finally, we place the features of the spermatophores and the spermatophore-forming organ in a phylogenetic framework of cerithioidean and paludomid systematics, which suggests that the spermatophore-forming organ is a synapomorphy of Paludomidae, that a bifurcate spermatophore structure is plesiomorphic, and that the evolution of structurally complex, spiny spermatophores has occurred independently in disparate lineages within the thalassoid species flock.     

Brood parasitism of an open-water spawning cichlid by the cuckoo catfish

The cuckoo catfish Synodontis multipunctatus and S. grandiops are endemic to Lake Tanganyika and the only known nonavian vertebrates that exhibit obligate interspecific brood parasitism. Seven maternal mouth-brooding cichlid fish species are reported to be natural hosts of the parasitic catfish and share a common reproductive behaviour that the catfish exploits: cichlid females spawn eggs on the bottom, allowing the catfish female to place her eggs near the cichlid eggs, and the cichlid females collect the catfish eggs by mouth together with their own eggs. However, so far it has not been reported that the cuckoo catfish exploit different spawning behaviours. The genus Cyprichromis consists of five maternal mouth-brooding species endemic to Lake Tanganyika, most of which spawn and collect eggs in open water. This study reports that the cuckoo catfish also parasitizes the open-water spawning Cyprichromis coloratus, although it may not be a regular host.     

On the Cyprideis species flock (Crustacea, Ostracoda) in Lake Tanganyika, with the description of four new species

Four new species, belonging to three genera, of ostracods from Lake Tanganyika are described: Mesocyprideis nitida sp.nov., Romecytheridea belone sp.nov., R. plegma sp. nov. and Cyprideis loricata sp. nov. The Cyprideis species flock of Lake Tanganyika now consists of 17 formally described species (16 endemic) in 6 genera (4 endemic). The geographical and ecological distribution of the four species is briefly discussed and some supplementary remarks on the radiation of the Cyprideis species flock in Lake Tanganyika are offered.     

Anatomy and systematic affinity of Stanleya neritinoides (Smith, 1880), an enigmatic member of the thalassoid gastropod species flock in Lake Tanganyika, East Africa (Cerithioidea, Paludomidae)

The radiation of gastropods in Lake Tanganyika is an ideal system for testing competing hypotheses of species flock formation. Yet, much of the basic biology of these species remains unknown. In an ongoing effort to understand the evolution of Tanganyikan gastropods, we here describe Stanleya neritinoides. Alcohol‐preserved material of the soft parts is rare, consequently, the systematic position of the species, and a repeated suggested affinity to Tanganyicia rufofilosa, have been based primarily on features of the shell. However, features of the radula and operculum are unique and do not suggest an affinity to any other Tanganyikan species. Thus, S. neritinoides has remained a particularly poorly known and enigmatic member of the species flock. This investigation confirmed that several aspects of internal and external anatomy are shared between S. neritinoides, T. rufofilosa, and other Tanganyikan gastropods, but that S. neritinoides is unique in features of the radula and seminal receptacle. Moreover, S. neritinoides differs from T. rufofilosa in features of the foregut, midgut, hindgut, kidney, nervous system, reproductive system and reproductive strategy. These new data are inconsistent with an interpretation of identity of Stanleya and Tanganyicia. In addition, given the pervasive differences between the two, a sister‐group relationship between the two is unlikely. More precise systematic placement of S. neritinoides awaits the establishment of a phylogenetic framework for all Tanganyikan gastropods.     

Mastacembelid eels support Lake Tanganyika as an evolutionary hotspot of diversification

Background  

Lake Tanganyika (LT) is the oldest of the African Rift Lakes and is one of the richest freshwater ecosystems on Earth, with high levels of faunal diversity and endemism. The endemic species flocks that occur in this lake, such as cichlid fishes, gastropods, catfish and crabs, provide unique comparative systems for the study of patterns and processes of speciation. Mastacembelid eels (Teleostei: Mastacembelidae) are a predominately riverine family of freshwater fish, occurring across Africa and Asia, but which also form a small species flock in LT.     

SHELL MICROSTRUCTURE OF GASTROPODS FROM LAKE TANGANYIKA,AFRICA: ADAPTATION,CONVERGENT EVOLUTION,AND ESCALATION

Gastropod shells from Lake Tanganyika, with their heavy calcification, coarse noded ribbing, spines, apertural lip thickening and repair scars, resemble marine shells more closely than they resemble other lacustrine shells. This convergence between Tanganyikan and marine gastropod shells, however, is not just superficial. Scanning electron microscope (SEM) studies reveal that the Tanganyikan shells are primarily layers of crossed-lamellar crystal architecture (that is, needle-like aragonite crystals arranged into laths that are packed into sheets such that the aragonite needles of adjacent laths are never parallel). The number of crossed-lamellar layers can vary from one to four between different Tanganyikan gastropod species. In species with two or more crossed-lamellar layers, the orientation of the lamellae is offset by approximately 90° between the different layers. The number of crossed-lamellar layers in the shell wall is positively correlated with shell strength and with predation resistance. Three and four crossed-lamellar layers in the shell wall evolved several times independently within the endemic thiarid gastropod radiation in Lake Tanganyika. Repeated origins of three and four crossed-lamellar layers suggest that they may be specific adaptations by Tanganyikan gastropods to strengthen their shells as a defense against shell-crushing predators.     

Mitochondrial phylogeny of the Lamprologini, the major substrate spawning lineage of cichild fishes from Lake Tanganyika in eastern Africa

Lake Tanganyika harbors the oldest, morphologically and behaviorally most diverse flock of cichlid species. While the cichlids in Lakes Malawi and Victoria breed their eggs exclusively by buccal incubation (termed "mouthbrooding"), the Tanganyikan cichlid fauna comprise mouthbrooding and substrate-spawning lineages (fish spawn on rocks, and never orally incubate eggs or wrigglers). The substrate-spawning tribe Lamprologini appears to occupy a key position that might allow one to elucidate the origin of the Tanganyika flock, because five riverine (therefore nonendemic) species from the Zaire River system have been assigned to this tribe, in addition to the lake's endemic species, which make up almost 50% of all 171 species known from this lake (Poll 1986). From 16 species (18 individuals) of the tribe Lamprologini, a 402-bp segment of the mitochondrial cytochrome b gene was sequenced, and, from 25 lamprologine species (35 individuals), sequences from the mitochondrial control region were obtained. To place the Lamprologini into a larger phylogenetic framework, orthologous sequences were obtained from eight nonlamprologine Tanganyikan cichlid species (13 individuals). The Lamprologini are monophyletic, and a clade of six Tanganyikan lineages of mouthbrooders, representing five tribes (Poll 1986), appears to be their sister group. Comparisons of sequence divergences of the control region indicate that the Lamprologini may be older than the endemic Tanganyikan tribe Ectodini, and short basal branches might suggest a rapid formation of lineages at an early stage of the Tanganyika radiation. It is interesting that three analyzed riverine members of the tribe form a monophyletic group; however, they are not the most ancestral branch of the Lamprologini. This might indicate that they are derived from an endemic lamprologine ancestor that left Lake Tanganyika by entering the Zaire River system. These riverine species may not have seeded the Tanganyikan radiation, as currently thought, but may have recently recolonized the river after a long period of isolation, as soon as the lake was connected to the Zaire River again about 2 Mya. Neolamprologus moorii, endemic to Lake Tanganyika, appears to represent the most basal clade of the Lamprologini. Complex breeding behavior, involving the usage of gastropod shells and associated with dwarfism, is likely to have evolved in parallel in several lineages among the Lamprologini. The tribe Lamprologini may be in need of revision, since several genera appear to be polyphyletic.      

Anatomy of an adaptive radiation: a unique reproductive strategy in the endemic freshwater gastropod Tylomelania (Cerithioidea: Pachychilidae) on Sulawesi, Indonesia and its biogeographical implications

The patterns of adaptive radiations in ancient lakes provide valuable clues to mechanisms of speciation and adaptation. In contrast to vertebrate radiations, for instance in fishes or finches, invertebrate species flocks have been largely neglected. While the increase in molecular data narrows this gap, the anatomical basis for interpreting these data against the background of evolutionary hypotheses is still widely lacking. Here we evaluate anatomical findings in the live‐bearing pachychilid freshwater gastropod Tylomelania, which has radiated extensively in ancient lakes in the Indonesian island, Sulawesi; we have aimed at reconciling these data with recently obtained molecular phylogenetic evidence. Discovered more than a century ago, the speciose and phenotypically diverse species flock with 34 currently described taxa was only occasionally cited as an example of adaptive radiation in ancient lakes, while anatomical data were entirely lacking. Our study of anatomical characters reveals very low qualitative variation at the species level. Thus, contrary to earlier views we suggest the existence of a single monophyletic lineage endemic to this island. The most conspicuous feature of Tylomelania is its uterine brooding strategy, i.e. retaining eggs and embryos in the pallial oviduct. This is unique among South‐East Asian pachychilids. Within the uterine brood pouch the offspring is surrounded by considerable amounts of nutritive material produced by a very large albumin gland, and the embryos are produced continuously. The shelled juveniles of some species are the largest known so far in viviparous gastropods, measuring almost 2 cm in length when hatching. This combination of reproductive features in Tylomelania, characterized by a high amount of maternal investment, is considered to be ovoviviparous, rendering its brooding strategy unique also among other gastropods. In addition, our data reject a previously assumed close relationship to other South‐East Asian pachychilids and instead suggest the North Australian Pseudopotamis as sister group to Tylomelania. These findings have significant consequences for the phylogenetic interpretation of morphological characters of Tylomelania in an evolutionary and biogeographical context, leading to the hypothesis that the common ancestor of both genera originated somewhere on the northern Australian continental margin. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 513–542.     

Colour-assortative mating among populations of Tropheus moorii, a cichlid fish from Lake Tanganyika, East Africa

The species flocks of cichlid fishes in the East African Lakes Tanganyika, Malawi and Victoria are prime examples of adaptive radiation and explosive speciation. Several hundreds of endemic species have evolved in each of the lakes over the past several thousands to a few millions years. Sexual selection via colour-assortative mating has often been proposed as a probable causal factor for initiating and maintaining reproductive isolation. Here, we report the consequences of human-mediated admixis among differentially coloured populations of the endemic cichlid fish Tropheus moorii from several localities that have accidentally been put in sympatry in a small harbour bay in the very south of Lake Tanganyika. We analysed the phenotypes (coloration) and genotypes (mitochondrial control region and five microsatellite loci) of almost 500 individuals, sampled over 3 consecutive years. Maximum-likelihood-based parenthood analyses and Bayesian inference of population structure revealed that significantly more juveniles are the product of within-colour-morph matings than could be expected under the assumption of random mating. Our results clearly indicate a marked degree of assortative mating with respect to the different colour morphs. Therefore, we postulate that sexual selection based on social interactions and female mate choice has played an important role in the formation and maintenance of the different colour morphs in Tropheus, and is probably common in other maternally mouthbrooding cichlids as well.     

Repeated parallel evolution of parental care strategies within Xenotilapia, a genus of cichlid fishes from Lake Tanganyika

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.     

The species flocks of lacustrine gastropods: <Emphasis Type="Italic">Tylomelania</Emphasis> on Sulawesi as models in speciation and adaptive radiation

Endemic radiations provide splendid opportunities for studies in evolutionary biology. Species flocks in ancient lakes, such as in Tanganyika, Malawi or Baikal, have featured prominently in evolutionary biology, viewing these “evolutionary theatres” as hotspots of diversification. However, following a century of neglect, the endemic evolution of limnic cerithioidean gastropods in the two central lake systems on the Indonesian island of Sulawesi (i.e. Lake Poso and the lakes of the Malili system, e.g. Danau Matano, Mahalona and Towuti) also provide instructive model cases for the study of speciation mechanisms, adaptive radiation and annidation (i.e. niche exploitation). We here discuss the evolutionary and taxonomic implications of the lacustrine species flocks in Tylomelania from these lakes in Sulawesi as an exceptional endemic assemblage of morphologically distinct viviparous pachychilid gastropods. This first comprehensive compilation of data on both ancient lake systems, Poso and Malili, offers a new perspective on ecological differentiation in this radiation. Presented here within the framework of the theory of evolutionary ecology it provides a research program for acquiring a synthetical perspective that includes morphology, molecular genetics, ecology and biogeography. In this context, it will be possible to compare the species flocks of these truly “Darwinian snails” on Sulawesi with the long enigmatic, so-called thalassoid (i.e. marine-like) gastropod radiation in East African’s Lake Tanganyika.     

Brooding season, sex ratio, and brood pouch development in the seaweed pipefish, Syngnathus schlegeli, in Otsuchi Bay, Japan

Males of the seaweed pipefish, Syngnathus schlegeli, take care of their eggs in the brood pouch. These pipefish were periodically collected from the shallow seagrass beds in Otsuchi Bay on the Pacific coast of northern Honshu, Japan, from spring to autumn to investigate the basic reproductive ecology. Appearance of the pipefish in the coastal seagrass beds coincided with the initiation of reproduction. The reproductive season was from May to at least October, with its peak in July. A rearing experiment revealed that the brooding period of the male had a negative correlation with water temperature, and it was estimated to last about 1 month in the bay. Almost all males were brooding during the peak of the reproductive season. Although, the brood pouch of most males was either full or devoid of eggs, 6.2% of the males had a partially filled (20%–90%) brood pouch, and multiple clutches were identified in the brood pouch of some males, indicating that the mating system of the pipefish is polygamous, perhaps polygynous. Sex ratio fluctuated among months, and the overall sex ratio tended to be biased to male. Body size of males with an immature brood pouch had a wide range, from 133 to 215 mm standard length (SL). The smallest brooding male was 134 mm SL. Mean SL of brooding males was significantly larger than that of nonbrooding mature males. The number of males with an immature brood pouch was greater at the beginning than later in the reproductive season. The results seem to collectively indicate that the occurrence of a larger proportion of immature males at the onset of the reproductive season may be ascribed to both new recruitment and larger body size at maturation, resulting from the males trading the reproductive effort to somatic growth, perhaps to increase future reproductive success. Received: April 4, 2000 / Revised: September 21, 2000 / Accepted: January 16, 2001     

Mitochondrial phylogeny of the endemic mouthbrooding lineages of cichlid fishes from Lake Tanganyika in eastern Africa

Of the three cichlid species flocks in eastern Africa, Lake Tanganyika harbors the oldest species assemblage, which is also the most diverse morphologically and behaviorally. For 12 species (20 individuals) of 12 genera of the tribe Ectodini, 852 bp from two segments (cytochrome b and control region) of the mitochondrial genome were sequenced. In addition, orthologous sequences were obtained from eight species (11 individuals) representing other mouthbrooding lineages from Lake Tanganyika. Comparisons of sequence divergences revealed that the single Tanganyikan tribe Ectodini appears to be approximately five times older than the whole Lake Malawi cichlid species flock, suggesting that the radiation of the Tanganyikan mouthbrooding lineages took place long before the species flocks of Lakes Malawi and Victoria evolved. Seven of nine surveyed tribes of Tanganyikan cichlids appear to be approximately equally divergent, and this seems to corroborate the hypothesis of a rapid simultaneous formation of lineages at an early stage in the history of the Lake Tanganyika species flock. The close genetic relationship between the endemic Tropheus lineage and a nonendemic "Haplochromine," Astatotilapia burtoni, indicates that members of the tribe Tropheini may be the sister group of the cichlid flocks of Lakes Malawi and Victoria. The phylogenetic analyses demonstrate the monophyly of the Ectodini and identify the Cyprichromini as their sister group among the Tanganyikan cichlids. Within the tribe Ectodini the molecular data suggest both a branching pattern different than that previously proposed and a subdivision of the Ectodini into four clades, instead of the two originally described. The previously postulated model of morphological transformations believed to be responsible for the drastically different types of ecological specialization found among the Ectodini might therefore be in need of reinterpretation. Characters immediately related to foraging and nutrition seem to be particularly prone to homoplasy, even among members of a single lineage of cichlid fishes.      

Morphology, reproductive biology and molecular genetics of ovoviviparous freshwater gastropods (Cerithioidea, Pachychilidae) from the Philippines, with description of a new genus Jagora

The new genus Jagora of the family Pachychilidae Troschel, 1857 is described for the type species Melania asperata Lamarck, 1822 from the Philippines. In addition to J. asperata, a second endemic species of the new genus, J. dactylus (I. Lea & H.C. Lea, 1850), is recognized on the basis of shell morphometry and molecular genetic data (cytochrome C oxidase I and 16S rRNA). The taxonomic history of J. asperata and J. dactylus is revised and the recent distribution documented on basis of available museum material and the authors’ own field collections. J. asperata occurs on Luzon and its satellite islands Leyte and Samar, while J. dactylus is restricted to the Visayan islands Bohol, Cebu, and Guimeras. The morphology of the two species is presented and illustrated in detail, and compared to taxa of the closely related genus Brotia H. Adams, 1866 to which they were previously assigned. Among the South‐east Asian freshwater Cerithioidea of the family Pachychilidae, which were previously subsumed under Brotia for their more or less similar shell morphology and operculum, three distinct lineages can be distinguished, in particular by means of distinct reproductive anatomy: (1) the species of Brotia sensu stricto from mainland South‐east Asia, Sumatra, Borneo and Java, which all exhibit a subhaemocoelic brood pouch; (2) the pachychilid species endemic to Sulawesi, currently assigned to the genera Brotia and Tylomelania, as well as both species of Pseudopotamis endemic to the Torres Strait Islands, which possess a uterine brood pouch; (3) the females of the Philippine Jagora, which carry egg capsules, embryos and advanced juvenile stages within the mantle cavity — a unique reproductive feature. Associated with this mode of ovoviviparity, Jagora is characterized by additional unique properties of the reproductive system including a deeply incised and long sperm gutter in the medial lamina, a very short and posteriorly positioned spermatophore bursa formed by the medial lamina, and a prominent lateral ridge functioning as a seminal receptacle. These characteristics are exclusive to Jagora and are consequently considered to represent autapomorphies of this clade which is endemic only to the Philippines. The zoogeographical implications are discussed in connection with a recently developed palaeogeographical reconstruction.     

Genetic and morphological population differentiation in the rock-dwelling and specialized shrimp-feeding cichlid fish species <Emphasis Type="Italic">Altolamprologus compressiceps</Emphasis> from Lake Tanganyika,East Africa

With about 250 endemic species, Lake Tanganyika contains an extraordinarily diverse cichlid fish fauna, and thus represents an ideal model system for the study of pathways and processes of speciation. The Lamprologini form the most species-rich tribe in Lake Tanganyika comprising about 100 species in seven genera, most of which are endemic to the lake. They are territorial substrate-breeders and represent a monophyletic tribe. By combined analysis of population genetics and geometric morphometric markers, we assessed gene flow among three populations of the highly specialized shrimp-feeding rock-dweller Altolamprologus compressiceps, separated by geographic distance and ecological barriers. Five highly polymorphic microsatellite markers were analyzed in conjunction with 17 landmarks in order to compare genetic differences to body shape differences among populations. Both genetic and morphological analyses revealed significant differentiation among the three studied populations. A significant, but overall relatively low degree of genetic differentiation supports a very recent divergence. Phenotypic differentiation was primarily found in the head region of A. compressiceps. In agreement with findings in other cichlid species, similar adaptations to specialized feeding mechanisms can consequently lead to marginal shape changes in the trophic apparatus.