Cementation in the freshwater bivalve family Corbiculidae (Mollusca: Bivalvia): a new genus and species from Lake Poso,Indonesia

A species from Lake Poso, Indonesia represents the first instance of cementation in the freshwater bivalve family Corbiculidae. Posostrea anomioides gen. nov., sp. nov., which is cemented indifferently by its left or right valve to calcareous rocks, reaches a size of 16 mm. It has typical corbiculoid dentition, with serrated lateral teeth, and juveniles are brooded primarily in the inner ctenidia. Lake Poso occupies a tectonic fault in central Sulawesi. Posostrea adds to an interesting radiation of endemic fishes and molluscs, including the gastropod genera Miratesta and Tylomelania. This revised version was published online in July 2006 with corrections to the Cover Date.     

Molecular phylogeny and diversification of freshwater shrimps (Decapoda, Atyidae, Caridina) from ancient Lake Poso (Sulawesi, Indonesia)--the importance of being colourful

Ancient Lake Poso on the Indonesian island Sulawesi hosts a highly diverse endemic fauna, including a small species flock of atyid Caridina shrimps, which are characterized by conspicuous colour patterns. We used a mtDNA based molecular phylogeny to test the assumption of a monophyletic origin and intralacustrine radiation of the species flock and to assess the species specificity of some colour morphs. Our data reveal a rapid radiation of Caridina in the entire Poso drainage system, but provide no strong evidence for a monophyletic radiation of the lake species. Nevertheless each lacustrine species shows a varying degree of substrate or trophic specialization, usually considered a hallmark of adaptive radiation. Two distinct colour forms previously attributed to a single species, C. ensifera, lack distinguishing qualitative morphological characters, but are shown to be two different species. In contrast, morphologically rather distinct lake species lacking specific colour patterns may be hybridizing with riverine taxa. These results suggest that colour may play a similar role in species recognition and possibly speciation in ancient lake Caridina as hypothesized, e.g. for some African cichlids.     

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.     

Escalation and trophic specialization drive adaptive radiation of freshwater gastropods in ancient lakes on Sulawesi, Indonesia

Species flocks in ancient lakes have long been appreciated as ideal model systems for the study of speciation and adaptive processes. We here present data from a new invertebrate model system with intrinsic parameters distinct from those of other documented radiations. The ancient lakes on Sulawesi harbour an endemic species flock of at least 33 species of viviparous snails. Molecular data reveal multiple independent colonizations of the lakes by riverine ancestors. In each colonizing clade, parallel evolution of conspicuous shell morphologies, followed by a differentiation of trophic morphology and the development of habitat specificity can be observed. Extensive shell crushing experiments and strong dentition of the chelae observed in some lacustrine crab species suggest that coevolution with crabs, i.e. escalation, is the most likely cause of initial shell divergence. By contrast, repeated parallel evolution in radula morphology indicates that speciation within lineages is driven by divergent adaptation to different resources among sympatric taxa. The inclusion of coevolutionary processes is unique in this system compared with diversification models developed for vertebrate radiations.     

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.     

Molecular diversity of freshwater crabs from Sulawesi and the sequential colonization of ancient lakes

Ancient lakes represent one of the most stable freshwater environments on Earth, with a species richness clearly exceeding that of younger and more short-lived limnic habitats. In most cases, the biological colonization of old lake systems must have occurred via the surrounding rivers. Two ancient lake systems of Sulawesi (Malili lake system and Lake Poso) have been studied in terms of the taxonomy and phylogeny of freshwater crabs (Decapoda: Brachyura: Gecarcinucidae: Parathelphusinae). Both systems have been colonized twice independently, and in both systems we can find three trophic niches which are always occupied by different crab species: molluskivores, omnivores, and detritivores. In the present study, we reconstruct phylogenies of freshwater crabs from more than 20 river systems of Sulawesi. We thereby confirm two independent colonization events for both ancient lake systems, with subsequent radiations. The phylogenies imply that the lineages which evolved into the molluskivore forms were the first ones to colonize, whereas omnivores and detritivores are derived from later colonization events and, based on their monophyletic relationship, resulted from minor lacustrine radiations. Most of the diversity of freshwater crabs from Sulawesi remains taxonomically undescribed. This study uncovers several undocumented phylogenetic units, with long independent evolutionary histories according to patristic distances.     

The Radiation of Hydrobioid Gastropods (Caenogastropoda, Rissooidea) in Ancient Lake Poso, Sulawesi

A survey of the fauna of hydrobioid gastropods living in ancient Lake Poso in Sulawesi revealed a total of 16 species, 14 of them new, belonging to two genera, Sulawesidrobia and Keindahan gen. nov. Most species occurred on hard substrates, water plants or rootlets of trees. Since only the upper 0.5 m of the lake have been sampled, many more species probably remain to be discovered. Already, Lake Posoȁ9s fauna ranks among the four most diverse hydrobioid lake faunas worldwide. The Sulawesi lakes including Lake Poso and the Malili Lakes are the only lakes where sizeable radiations of hydrobioid and cerithioid gastropods coexist. Outside Lake Poso, hydrobioid gastropods have not been investigated so far apart from a single species reported from Lake Lindoe similar to or identical with S. bonnei [Abbott, 1945. Occasional Papers on Mollusks 1: 1–4], which has its type locality in Lake Poso and may in fact be a complex of species. Therefore endemicity in Lake Poso cannot be estimated nor is it possible to say, whether the radiation is of lacustrine origin. The introduction of alien fish has had a severe impact on the native fish fauna. The impact on the invertebrate fauna is not known but chances are that this survey based on collections from 1991 no longer reflects the original assemblage in the lake.     

Ancient lakes as hotspots of diversity: a morphological review of an endemic species flock of <Emphasis Type="Italic">Tylomelania</Emphasis> (Gastropoda: Cerithioidea: Pachychilidae) in the Malili lake system on Sulawesi,Indonesia

The viviparous freshwater gastropod Tylomelania (Caenogastropoda: Cerithioidea: Pachychilidae) endemic to the Indonesian island Sulawesi has radiated extensively in two ancient lake systems. We here present the first systematic species-level review of taxa in the five lakes of the Malili lake system, which contains the most diverse and best studied freshwater fauna on Sulawesi. Our results indicate a significantly higher diversity of Tylomelania in these lakes than previously perceived based on morphological evidence for delimiting the taxa. We describe nine new species, thus increasing the number of taxa known from the Malili lakes to 25. Tylomelania species are inhabiting all available substrates in the lakes, and the diversity of habitats is reflected in an unparalleled range of radula types in this closely related group. Several species show a high intraspecific variability in some characters, and their closer investigation will probably lead to the discovery of more cryptic species. As it is, this species flock on Sulawesi is among the largest freshwater mollusc radiations known. Since the Malili lake system also contains other large endemic species flocks of e.g. crustaceans and fishes, it is a major hotspot of freshwater biodiversity in Asia to become a conservation priority. Handling editor: K. Martens     

Rapid evolution of an ancient lake species flock: Freshwater limpets (Gastropoda: Ancylidae) in the Balkan Lake Ohrid

Ancient lakes have long been recognized as evolutionary theatres and hot spots of endemism; the evolution of their morphologically often highly diverse species flocks has received much attention. However, as each ancient lake has its own geological and evolutionary history, modes of speciation may differ from system to system. Ancient lakes can act as evolutionary reservoirs that assure the survival of relict species, but at the same time extant species may evolve through intralacustrine speciation. Other aspects of interest are the actual rates of immigration, diversification or extinction as well as the temporal framework of morphological change. Many of these questions have been addressed in the African (e.g. Lake Tanganyika) and Asian (e.g. Lake Baikal) ancient lakes. For an European ancient lakes (e.g. Lakes Ohrid and Prespa), such studies are largely missing. In the present paper, extraordinarily shaped endemic freshwater limpets of the genus Ancylus from the Balkan Lake Ohrid are used in a phylogeographic and phylogenetic context to test whether they represent an ancient lake species flock, to study the mode of speciation, and to assess the timing of morphological change. Based on DNA data from two mitochondrial genes (COI, LSU rDNA), it has been found that the Lake Ohrid Ancylus species form an endemic monophyletic group. In addition, the lake's feeder springs are inhabited by another, undescribed Ancylus species. All other studied waterbodies within the watershed do not support their own Ancylus lineages but are inhabited by a widespread Mediterranean taxon. The split between the species endemic to the lake and its sister taxon is dated to 1.4±0.6 million years ago. The study presents the first genetic confirmation for the existence of a species flock in a European ancient lake. Contrary to the prevailing opinion it shows that, concerning Ancylus, Lake Ohrid represents a site of intralacustrine speciation rather than an evolutionary reservoir. Moreover, it provides the first evidence for rapid morphological change in an European ancient lake species flock. See also Electronic Supplement at: http://www.senckenberg.de/odes/06-12.htm.     

Testing two contrasting evolutionary patterns in ancient lakes: species flock versus species scatter in valvatid gastropods of Lake Ohrid

Ancient lakes have long been recognized as “hot spots of evolution” and “evolutionary theatres” and they have significantly contributed to a better understanding of speciation and radiation processes in space and time. Yet, phylogenetic relationships of many ancient lake taxa, particularly invertebrate groups, are still unresolved. Also, the lack of robust morphological, anatomical, and phylogeographical data has largely prevented a rigorous testing of evolutionary hypotheses. For the freshwater gastropod genus Valvata—a group with a high degree of endemism in several ancient lakes—different evolutionary scenarios are suggested for different ancient lakes. Lake Baikal, for example, is inhabited by several endemic Valvata taxa that presumably do not form a monophyletic group. For such an evolutionary pattern, the term ancient lake species scatter is introduced here. In contrast, for the Balkan Lake Ohrid, workers previously suggested the presence of a monophyletic group of endemic Valvata species, that is, an ancient lake species flock. Sequence data of the mitochondrial cytochrome oxidase c subunit I gene (COI) from worldwide taxa, with a strong emphasis on Balkan species, are here used to test whether the putative Ohrid Valvata endemics represent an ancient lake species flock and to study patterns of speciation both on the Ohrid and the Balkan scale. The study reveals three distinct clades of endemic Valvata in Lake Ohrid. Monophyly of these taxa, however, is rejected, and they therefore do not represent an ancient lake species flock, but rather an ancient lake species scatter. Also, in contrast to many other gastropod groups in Lake Ohrid, the valvatids apparently did not radiate. Many Valvata taxa in ancient lakes are characterized by enhanced levels of shell complexity. However, it remains unclear whether these patterns are associated with ancient lake environments per se. It is here suggested that similarities in shell structure between North American and Balkan taxa might simply be due to convergent evolution.     

Biodiversity and distribution of epibiontic communities on Caridina ensifera (Crustacea,Decapoda, Atyidae) from Lake Poso: comparison with another ancient lake system of Sulawesi (Indonesia)

Fernandez‐Leborans, G. and von Rintelen, K. 2010. Biodiversity and distribution of epibiontic communities on Caridina ensifera (Crustacea, Decapoda, Atyidae) from Lake Poso: comparison with another ancient lake system of Sulawesi (Indonesia). — Acta Zoologica (Stockholm) 91 : 163–175 The epibiont communities of the shrimp Caridina ensifera, endemic to Lake Poso (Sulawesi, Indonesia), were analysed. Most of the epibiont species were ciliated protozoa belonging to three suctorian genera (Acineta, Podophrya and Spelaeophrya), three peritrich genera (Zoothamnium, Vorticella and Cothurnia), and a haptorid genus (Amphileptus). There was also a rotifer epibiont of the genus Embata. Epibionts were identified to species level. There were 14 to 1114 epibionts per shrimp. The distribution of the epibiont species on the surface of the basibiont was recorded, calculating the number on the different colonized individuals of C. ensifera. The most abundant species, Zoothamnium intermedium and Acineta sulawesiensis, were also the most widely distributed. There was a significant difference between the spatial distributions of the different epibiont species. The analysis of the number of the epibiont species throughout the anteroposterior axis of the shrimp showed a gradient from the anterior to the posterior end of the body. Data from Lake Poso were compared with those of the Malili lake system (Sulawesi), obtained from its endemic shrimp, Caridina lanceolata. Lake Poso had the highest mean diversity, while Lake Mahalona showed the highest maximum diversity. All lakes were correlated with respect to the mean number of epibionts on the anatomical units of the shrimp, which showed a similar general distribution. The distributions of the different epibiont species were compared between the lakes. The possible adaptations of the epibionts as well as the colonization patterns were discussed. From the statistical results and the analysis of the distributions, we propose that in these communities epibiont species have a pattern of colonization in which they follow a behaviour as a whole; each species has a differential distribution, with the species occupying the available substratum with the particular requirements of each functional group, but there is a trend towards maintaining an equilibrium among species and groups, compensating for diversity and number of individuals. In all lakes there was an epibiont distribution model comprising the maintenance of an anteroposterior axis gradient, which was supported by the fluctuation in diversity and number of individuals of the different functional groups of epibiont species. The functional role of the different groups of species seems to tend towards sustainability with little global variation among the lakes.     

Phylogeny of the Lake Tanganyika cichlid species flock and its relationship to the Central and East African haplochromine cichlid fish faunas

Lake Tanganyika, the oldest of the East African Great Lakes, harbors the ecologically, morphologically, and behaviorally most complex of all assemblages of cichlid fishes, consisting of about 200 described species. The evolutionary old age of the cichlid assemblage, its extreme degree of morphological differentiation, the lack of species with intermediate morphologies, and the rapidity of lineage formation have made evolutionary reconstruction difficult. The number and origin of seeding lineages, particularly the possible contribution of riverine haplochromine cichlids to endemic lacustrine lineages, remains unclear. Our phylogenetic analyses, based on mitochondrial DNA sequences of three gene segments of 49 species (25% of all described species, up to 2,400 bp each), yield robust phylogenies that provide new insights into the Lake Tanganyika adaptive radiation as well as into the origin of the Central- and East-African haplochromine faunas. Our data suggest that eight ancient African lineages may have seeded the Tanganyikan cichlid radiation. One of these seeding lineages, probably comprising substrate spawning Lamprologus-like species, diversified into six lineages that evolved mouthbrooding during the initial stage of the radiation. All analyzed haplochromines from surrounding rivers and lakes seem to have evolved within the radiating Tanganyikan lineages. Thus, our findings contradict the current hypothesis that ancestral riverine haplochromines colonized Lake Tanganyika to give rise to at least part of its spectacular endemic cichlid species assemblage. Instead, the early phases of the Tanganyikan radiation affected Central and East African rivers and lakes. The haplochromines may have evolved in the Tanganyikan basin before the lake became a hydrologically and ecologically closed system and then secondarily colonized surrounding rivers. Apparently, therefore, the current diversity of Central and East African haplochromines represents a relatively young and polyphyletic fauna that evolved from or in parallel to lineages now endemic to Lake Tanganyika.     

Molecular evidence reveals a polyphyletic origin and chromosomal speciation of Lake Baikal's endemic asellid isopods

The six endemic isopod species of Lake Baikal have been regarded as a small species flock with uncertain affinities to related asellids. We provide evidence from 16S rRNA sequences for polyphyletic origins of Baikalian Asellidae. One clade of two species is related to the Eurasian genus Asellus. The other clade, Baicalasellus, shows affinities to North American asellids and may have a long evolutionary history within the lake basin. Some speciation events within Baicalasellus clearly have a chromosomal basis. In contrast with numerous taxa exhibiting monophyletic radiations in ancient lakes, the endemic Baikalian isopods arose by multiple invasions and chromosomal mechanisms.     

ADAPTIVE RADIATION AND ECOLOGICAL DIVERSIFICATION OF SULAWESI'S ANCIENT LAKE SHRIMPS

Ancient lakes are natural laboratories for the study of adaptive radiation. Recently, two lake systems on the Indonesian island Sulawesi have emerged as promising new model systems. A species flock of atyid freshwater shrimps in the Malili lake system comprises 15 colorful endemic taxa. Mitochondrial DNA data suggest two independent colonizations by riverine ancestors. Only one colonization event led to subsequent radiation into 14 species, while the second clade comprises just one species. The vast majority of species (n= 12) are habitat specialists, which are confined to the larger Malili clade and include all taxa with species‐specific color patterns and a restricted distribution within the five connected Malili lakes. Morphological, genetic, and ecological data are consistent with the existence of an adaptive radiation in the Malili lakes, involving the habitat‐specific diversification of trophic morphology. In addition to testing criteria for the recognition of an adaptive radiation, an ancestral state reconstruction reveals an equal probability for either a riverine generalist or a lacustrine specialist as ancestor of the large Malili clade, which is interpreted as indicative of an early stage of habitat specialization within this radiation. Finally, our results suggest that species diversification may have been primarily driven by ecological specialization and allopatric speciation.     

ORIGIN AND RECENT ENDEMIC DIVERGENCE OF A CASPIAN MYSIS SPECIES FLOCK WITH AFFINITIES TO THE “GLACIAL RELICT” CRUSTACEANS IN BOREAL LAKES

Aspects of the evolution of intralacustrine species flocks and of the origin of the Arctic or “glacial-relict” zoogeographical element in Eurasian inland waters were elucidated in an allozyme study of the crustacean genus Mysis. This element, of supposedly northern marine ancestry, is represented by vicarious taxa in the deeper parts of the Caspian Sea (an enclosed ancient basin) and in young boreal lakes. The three endemic Caspian Mysis species studied are very close genetically (Nei's D = 0.06), which suggests a recent intrabasin radiation and rapid morphological divergence. This is in contrast to the pattern in postglacial Holarctic boreal lakes, where the Mysis relicta group is represented by a set of morphologically uniform but probably much older sibling species (D = 0.3–0.6). The results provide a parallel to those on the recent diversification of some fish species flocks in ancient freshwater lakes. The situation is, however, unusual in that the Caspian sympatric Mysis flock is pelagic, and conditions promoting speciation through allopatric isolation or spatial segregation by trophic substrate specialization seem implausible. The monophyletic Caspian Mysis clade shows a relatively strong divergence from both the northern lacustrine and the Arctic marine congeners (D = 0.6–1.0); the phylogenetic branching order of these three zoogeographical groups is not conclusively resolved. The results contradict the prevailing hypothesis of a recent Pleistocene origin of the Caspian Arctic element by invasion from Eastern European continental proglacial lakes that drained south to the Caspian basin during the glacial maxima and served as refugia for the boreal lacustrine taxa.     

Origin and radiation of rissooidean gastropods (Caenogastropoda) in ancient lakes of Sulawesi

Zielske, S., Glaubrecht, M. & Haase, M. (2010). Origin and radiation of rissooidean gastropods (Caenogastropoda) in ancient lakes of Sulawesi. —Zoologica Scripta, 40, 221–237. We investigated diversity and origin of as well as relationships among rissooidean gastropods from the ancient lakes of Sulawesi. Analyses of morphological, anatomical and genetic data (16S rRNA, COI and 18S rRNA) revealed ten new species: Sulawesidrobia perempuan sp. n., which is probably parthenogenetic, Sulawesidrobia soedjatmokoi sp. n., Sulawesidrobia mahalonaensis sp. n., Sulawesidrobia anceps sp. n., Sulawesidrobia bicolor sp. n., Sulawesidrobia megalodon sp. n., Sulawesidrobia abreui sp. n., Sulawesidrobia datar sp. n., Sulawesidrobia yunusi sp. n. and Sulawesidrobia towutiensis sp. n. In two cases morphological and anatomical differentiation between pairs of species were not paralleled by genetic differentiation suggesting young speciation events and incomplete lineage sorting as a likely explanation. A Bayesian phylogenetic analysis including representatives of most rissooidean families placed Sulawesidrobia Ponder & Haase 2005 among Tateidae and thus indicated an Australasian origin of the genus. Relationships among the Sulawesian species suggested that isolated Lake Poso was colonised only once whereas the lakes of the Malili system experienced repeated colonizations.     

Novel reproductive modes in freshwater clams: brooding and larval morphology in Southeast Asian taxa of Corbicula (Mollusca, Bivalvia, Corbiculidae)

While the majority of marine bivalves are oviparous, the two freshwater families among the order Veneroida, i.e. Corbiculidae and Sphaeriidae, comprise species with ovoviviparous and viviparous reproduction. Within the Corbiculidae, the genus Corbicula, which is well‐known for its invasive and, thus, ecologically important representatives, is characterized by (i) a wide range of limnic habitats, inhabiting both brackish water and freshwater environments, and (ii) contrasting modes of reproduction, including ‘planktonic’ development via a free‐swimming larva vs. intrabranchial incubation (brooding) of shelled juveniles. The present investigation of five species of Corbicula from the Indonesian islands Sumatra and Sulawesi, which were hitherto not studied anatomically, adds to the diversity in reproductive patterns in this genus. As a unique feature among Corbicula we here report on two newly observed modes of brooding in species endemic to Sulawesi, (i) tetragenous brooding (i.e. in both demibranchs) in Corbicula possoensis Sarasin & Sarasin, 1898 from Lake Poso, and (ii) prolonged incubation in the maternal gills, with juvenile shells reaching up to 1.3 mm in length and with a well‐developed hinge in C. linduensis Bollinger, 1914 from the Lindu River system. In contrast, a third method is seen in the following taxa that incubate their young in their inner demibranchs only until the stage of juveniles with straight‐hinged shells (D‐shaped): C. matannensis Sarasin & Sarasin, 1898 from Lake Matano and Lake Mahalona, C. loehensis Kruimel, 1913 from Lake Masapi (all on Sulawesi) and C. moltkiana Prime, 1878 from Lakes Maninjau and Singkarak (on Sumatra). Details of the anatomical and histological features of ctenidia are described for each type of brooding, and some trends in the evolution of reproductive strategies within the Corbiculidae are discussed, comparing them with those known from other limnic molluscs.     

The endemic gastropod fauna of Lake Titicaca: correlation between molecular evolution and hydrographic history

Lake Titicaca, situated in the Altiplano high plateau, is the only ancient lake in South America. This 2- to 3-My-old (where My is million years) water body has had a complex history that included at least five major hydrological phases during the Pleistocene. It is generally assumed that these physical events helped shape the evolutionary history of the lake's biota. Herein, we study an endemic species assemblage in Lake Titicaca, composed of members of the microgastropod genus Heleobia, to determine whether the lake has functioned as a reservoir of relic species or the site of local diversification, to evaluate congruence of the regional paleohydrology and the evolutionary history of this assemblage, and to assess whether the geographic distributions of endemic lineages are hierarchical. Our phylogenetic analyses indicate that the Titicaca/Altiplano Heleobia fauna (together with few extralimital taxa) forms a species flock. A molecular clock analysis suggests that the most recent common ancestor (MRCAs) of the Altiplano taxa evolved 0.53 (0.28-0.80) My ago and the MRCAs of the Altiplano taxa and their extralimital sister group 0.92 (0.46-1.52) My ago. The endemic species of Lake Titicaca are younger than the lake itself, implying primarily intralacustrine speciation. Moreover, the timing of evolutionary branching events and the ages of two precursors of Lake Titicaca, lakes Cabana and Ballivián, is congruent. Although Lake Titicaca appears to have been the principal site of speciation for the regional Heleobia fauna, the contemporary spatial patterns of endemism have been masked by immigration and/or emigration events of local riverine taxa, which we attribute to the unstable hydrographic history of the Altiplano. Thus, a hierarchical distribution of endemism is not evident, but instead there is a single genetic break between two regional clades. We also discuss our findings in relation to studies of other regional biota and suggest that salinity tolerance was the most likely limiting factor in the evolution of Altiplano species flocks.     

The number of endemic species of freshwater sponges (Malawispongiidae; Spongillina; Porifera) from Lake Kinneret is overestimated

It is well accepted that the freshwater sponges (Porifera; Haplosclerida; Spongillina) currently comprise six extant families: Spongillidae, Lubomirskiidae, Malawispongiidae, Metaniidae, Metschnikowiidae and Potamolepidae, but the phylogeny of this group is poorly understood. Family Malawispongiidae includes five genera: Malawispongia, Spinospongilla, Cortispongilla, Ochridaspongia, Pachydictyum, which inhabit ancient lakes: Malawi and Tanganyika (African Rift Valley), Kinneret (Middle East), Ohrid (Europe) and Poso (Central Sulawesi). We show via nuclear and mitochondrial markers (cox 1, 28S rRNA and ribosomal ITS regions) that both endemic species Cortispongilla barroisi and Ephydatia syriaca from Lake Kinneret are synonymous with the cosmopolitian species Ephydatia fluviatilis, which also supports suggestions that the family Malawispongiidae is polyphyletic. Our findings also suggest that Nudospongilla is a synthetic taxon and that the number of endemic freshwater sponge species is overestimated.