Gene flow simulations demonstrate resistance of long-lived species to genetic erosion from habitat fragmentation

Habitat fragmentation restricts the movement of individuals across a landscape. In terrestrial and aquatic systems, barriers to movement can modify population and community dynamics at local or regional scales. This study contrasted life history traits related to lifespan with habitat fragmentation to determine impacts on species population genetic structure in the Neuse River Basin, USA. For this, we simulated gene flow among evenly-spaced populations in a river network and tracked individual and population genetics for 200 years. The modeled scenarios represent a full cross between five life history strategies and four riverscapes representing varying degrees of fragmentation. The five life history strategies include species (based on freshwater mussels) with average lifespans ranging from 10 to 50 years and age at maturity from 2 to 6 years. The movement landscapes included a (1) panmictic, (2) stepping-stone landscape allowing movement to only neighboring populations during each dispersal event, (3) partially-fragmented landscape divided by dams currently in the network, and (4) fully-fragmented landscape. Results suggest species with shorter lifespans have higher population genetic structure in fragmented landscapes than species with longer lifespans. Furthermore, species with shorter lifespans in highly fragmented landscapes may be harboring genetic degradation or decline as allele fixation and loss. Although anthropogenic fragmentation of many river systems is only 100–200 years old, the simulation indicates that species can respond genetically in that period of time. Additionally, the time frame of the simulation suggests that genetic impacts of habitat fragmentation in some species present in the Neuse River Basin may not yet be manifesting and restoration activities could be successful.     

Evolutionary history of the endemic Lake Tanganyika cichlid fish Tylochromis polylepis: a recent intruder to a mature adaptive radiation

With an age of 9–12 million years (Myr) Lake Tanganyika holds the oldest and most complex species flock of cichlid fishes. It is believed to be of polyphyletic origin and rooted in nine ancient African lineages, six of which underwent diversification, while three remained monotypic. Here, the evolutionary history, route and timing of colonization were analyzed, as well as intraspecific genetic diversity of Tylochromis polylepis , the single albeit endemic representative of the tribe Tylochromini in Lake Tanganyika. The role of Tylochromis in the radiation was uncertain and the species was suggested to be either an ancient colonizer or a recent addition to the fauna. With 2.8–4.2% sequence divergence to four congeners living in Lakes Bangweulu and Mweru (Upper Congo River) as well as from Luozi River and Lake Etsotso (Lower Congo River), the species appears to be a recent colonizer, which is likely to have diverged from its riverine allies less than 510 000 years ago. The ability to enter an adaptive radiation at a highly mature stage with its densely packed species community and fine-tuned niche segregation seems remarkable. So far it was assumed that all Tanganyikan endemics evolved within the lake ecosystem by intralacustrine speciation, and that complex competitive interactions among endemics tighten the system against intruders. The intruding species can be classified as a generalist living over muddy bottom in the lake and in river estuaries. Microgeographic substructuring is suggested, possibly because of geographic segregation of breeding sites. Concerning the route of colonization, a downstream movement via one of the inflowing rivers situated in the south of Lake Tanganyika is suggested.     

River barriers and cryptic biodiversity in an evolutionary museum

The Riverine Barriers Hypothesis (RBH) posits that tropical rivers can be effective barriers to gene flow, based on observations that range boundaries often coincide with river barriers. Over the last 160 years, the RBH has received attention from various perspectives, with a particular focus on vertebrates in the Amazon Basin. To our knowledge, no molecular assessment of the RBH has been conducted on birds in the Afrotropics, despite its rich avifauna and many Afrotropical bird species being widely distributed across numerous watersheds and basins. Here, we provide the first genetic evidence that an Afrotropical river has served as a barrier for birds and for their lice, based on four understory bird species collected from sites north and south of the Congo River. Our results indicate near‐contemporaneous, Pleistocene lineage diversification across the Congo River in these species. Our results further indicate differing levels of genetic variation in bird lice; the extent of this variation appears linked to the life‐history of both the host and the louse. Extensive cryptic diversity likely is being harbored in Afrotropical forests, in both understory birds and their lice. Therefore, these forests may not be “museums” of old lineages. Rather, substantial evolutionary diversification may have occurred in Afrotropical forests throughout the Pleistocene, supporting the Pleistocene Forest Refuge Hypothesis. Strong genetic variation in birds and their lice within a small part of the Congo Basin forest indicates that we may have grossly underestimated diversity in the Afrotropics, making these forests home of substantial biodiversity in need of conservation.     

Biogeography in the Death Valley region: evidence from springsnails (Hydrobiidae: Tryonia)

Allozyme and mitochondrial DNA variation were analysed to examine evolution of the nine species of springsnails (genus Tryonia) living in the Death Valley system (Owens and Amargosa basins) of southeastern California and southwestern Nevada. Both allozyme and mtDNA evidence indicate that this highly endemic fauna is non-monophyletic. Species from the upper Amargosa basin comprise a clade most closely related to snails living in the Colorado basin. Snails from the lower Amargosa basin (Death Valley trough) reflect a complex evolutionary history and two of these species are more closely related to an estuarine species from western California than to other snails of the region. These results indicate a commonality of pattern with the well-studied Death Valley pupfishes (Cyprinodon), which also are non-monophyletic and include species that are most closely related to Colorado basin congeners. These biogeographic patterns are interpreted within the context of a recently proposed model for the early history of the lower Colorado River.     

Subterranean phylogeography of freshwater crayfishes shows extensive gene flow and surprisingly large population sizes

Subterranean animals are currently viewed as highly imperiled, precariously avoiding extinction in an extreme environment of darkness. This assumption is based on a hypothesis that the reduction in visual systems and morphology common in cave faunas reflects a genetic inability to adapt and persist coupled with the perception of a habitat that is limited, disconnected, and fragile. Accordingly, 95% of cave fauna in the United States are presumed endangered due to surface environmental degradation and limited geographic distributions. Our study explores the subterranean phylogeography of stygobitic crayfishes in the southeastern United States, a global hotspot of groundwater biodiversity, using extensive geographic sampling and molecular data. Despite their endangered status, our results show that subterranean crayfish species have attained moderate to high levels of genetic diversity over their evolutionary histories with large population sizes and extensive gene flow among karst systems. We then compare the subterranean population histories to those of common surface stream-dwelling crayfishes. Our results show recent drastic declines in genetic variability in the surface crayfish and suggest that these species also warrant conservation attention.     

塔里木盆地鱼类入侵及区系演变趋势

鱼类入侵是塔里木盆地土著鱼类濒危的关键性影响因素之一,结合历史文献数据及近年调查数据,系统综述了塔里木盆地鱼类入侵现状,并就鱼类入侵引起的鱼类区系均一化问题进行了分析。结果显示,该地区记录鱼类63种,其中土著种19种,入侵种44种。尽管入侵鱼类提升了塔里木盆地鱼类群落属级及科级的多样性水平,但相应的G-F多样性指数却明显下降,表明该区鱼类群落从物种组成简单、分化明显的特点演变为物种成分复杂、分化贫乏的特点。入侵种中源于长江和额尔齐斯河分别有30种(68.2%)和7种(15.9%),源于美欧和亚洲其他区域的有7种(15.9%)。入侵鱼类的进入引起塔里木盆地与其相关区域鱼类区系相似性显著上升(P0.01),并导致该区鱼类区系从单一的中亚高原山区复合体演变为多区系混合体。进一步利用鱼类个体生态矩阵分析方法研究显示,外来种在塔里木盆地适应性广泛,能够占据水域各种栖息生境,显著压缩了土著种的栖息空间,进而通过捕食、食物竞争及其他作用途径严重危及土著鱼类种群的续存,提示须采取有力措施限制入侵种的进一步扩散及对濒危土著物种迅速开展人工保育工作。     

Time and origin of cichlid colonization of the lower Congo rapids

Most freshwater diversity is arguably located in networks of rivers and streams, but, in contrast to lacustrine systems riverine radiations, are largely understudied. The extensive rapids of the lower Congo River is one of the few river stretches inhabited by a locally endemic cichlid species flock as well as several species pairs, for which we provide evidence that they have radiated in situ. We use more that 2,000 AFLP markers as well as multilocus sequence datasets to reconstruct their origin, phylogenetic history, as well as the timing of colonization and speciation of two Lower Congo cichlid genera, Steatocranus and Nanochromis. Based on a representative taxon sampling and well resolved phylogenetic hypotheses we demonstrate that a high level of riverine diversity originated in the lower Congo within about 5 mya, which is concordant with age estimates for the hydrological origin of the modern lower Congo River. A spatial genetic structure is present in all widely distributed lineages corresponding to a trisection of the lower Congo River into major biogeographic areas, each with locally endemic species assemblages. With the present study, we provide a phylogenetic framework for a complex system that may serve as a link between African riverine cichlid diversity and the megadiverse cichlid radiations of the East African lakes. Beyond this we give for the first time a biologically estimated age for the origin of the lower Congo River rapids, one of the most extreme freshwater habitats on earth.     

Factors affecting the persistence of endangered Ganges River dolphins (Platanista gangetica gangetica)

The Ganges–Brahmaputra–Meghna and Karnaphuli (GBMK) River Basin in Nepal, India, and Bangladesh is among the world's most biodiverse river basins. However, human‐induced habitat modification processes threaten the ecological structure of this river basin. Among the GBMK’s diverse flora and fauna of this freshwater ecosystem, the endemic Ganges River dolphin (Platanista gangetica gangetica; GRD) is one of the most charismatic species in this freshwater ecosystem. Though a >50% population size reduction has occurred since 1957, researchers and decision‐makers often overlook the persistence (or evolutionary potential) of this species in the highly fragmented GBMK. We define the evolutionary potential as the ability of species/populations to adapt in a changing environment by maintaining their genetic diversity. Here, we review how evolutionary trap mechanisms affect the dynamics and viability of the GRD (hereafter Ganges dolphin) populations after rapid declines in their population size and distribution. We detected six potential trap mechanisms that might affect the Ganges dolphin populations discretely or in combination: (a) habitat modification; (b) occurrence of finite and geographically restricted local populations; (c) ratio of effective to estimate population size; (d) increasing risk of inbreeding depression in genetically isolated groups; (e) at‐risk behavioral attributes; and (f) direct fisheries–dolphin interactions. Because evolutionary traps appear most significant during low water season, they adversely affect demographic parameters, which reduce evolutionary potential. These traps have already caused local extirpation events; therefore, we recommend translocation among populations, including restoring and preserving essential habitats as immediate conservation strategies. Integrative evolutionary potential information based on demographic, genetic, and environmental data is still lacking. Thus, we identify gaps in the knowledge and suggest integrative approaches to understand the future of Ganges dolphins in South Asian waterways.     

Phylogeography and conservation genetics of Eld's deer (Cervus eldi)

Eld's deer (Cervus eldi) is a highly endangered cervid, distributed historically throughout much of South Asia and Indochina. We analysed variation in the mitochondrial DNA (mtDNA) control region for representatives of all three Eld's deer subspecies to gain a better understanding of the genetic population structure and evolutionary history of this species. A phylogeny of mtDNA haplotypes indicates that the critically endangered and ecologically divergent C. eldi eldi is related more closely to C. e. thamin than to C. e. siamensis, a result that is consistent with biogeographic considerations. The results also suggest a strong degree of phylogeographic structure both between subspecies and among populations within subspecies, suggesting that dispersal of individuals between populations has been very limited historically. Haplotype diversity was relatively high for two of the three subspecies (thamin and siamensis), indicating that recent population declines have not yet substantially eroded genetic diversity. In contrast, we found no haplotype variation within C. eldi eldi or the Hainan Island population of C. eldi siamensis, two populations which are known to have suffered severe population bottlenecks. We also compared levels of haplotype and nucleotide diversity in an unmanaged captive population, a managed captive population and a relatively healthy wild population. Diversity indices were higher in the latter two, suggesting the efficacy of well-designed breeding programmes for maintaining genetic diversity in captivity. Based on significant genetic differentiation among Eld's deer subspecies, we recommend the continued management of this species in three distinct evolutionarily significant units (ESUs). Where possible, it may be advisable to translocate individuals between isolated populations within a subspecies to maintain levels of genetic variation in remaining Eld's deer populations.     

Evolution in caves: Darwin’s ‘wrecks of ancient life’ in the molecular era

Cave animals have historically attracted the attention of evolutionary biologists because of their bizarre ‘regressive’ characters and convergent evolution. However, understanding of their biogeographic and evolutionary history, including mechanisms of speciation, has remained elusive. In the last decade, molecular data have been obtained for subterranean taxa and their surface relatives, which have allowed some of the classical debates on the evolution of cave fauna to be revisited. Here, we review some of the major studies, focusing on the contribution of phylogeography in the following areas: biogeographic history and the relative roles of dispersal and vicariance, colonization history, cryptic species diversity and modes of speciation of cave animals. We further consider the limitations of current research and prospects for the future. Phylogeographic studies have confirmed that cave species are often cryptic, with highly restricted distributions, but have also shown that their divergence and potential speciation may occur despite the presence of gene flow from surface populations. Significantly, phylogeographic studies have provided evidence for speciation and adaptive evolution within the confines of cave environments, questioning the assumption that cave species evolved directly from surface ancestors. Recent technical developments involving ‘next generation’ DNA sequencing and theoretical developments in coalescent and population modelling are likely to revolutionize the field further, particularly in the study of speciation and the genetic basis of adaptation and convergent evolution within subterranean habitats. In summary, phylogeographic studies have provided an unprecedented insight into the evolution of these unique fauna, and the future of the field should be inspiring and data rich.     

Phylogenetic effects on functional traits and life history strategies of Australian freshwater fish

Understanding the biogeographic and phylogenetic basis to interspecific differences in species’ functional traits is a central goal of evolutionary biology and community ecology. We quantify the extent of phylogenetic influence on functional traits and life‐history strategies of Australian freshwater fish to highlight intercontinental differences as a result of Australia's unique biogeographic and evolutionary history. We assembled data on life history, morphological and ecological traits from published sources for 194 Australian freshwater species. Interspecific variation among species could be described by a specialist–generalist gradient of variation in life‐history strategies associated with spawning frequency, fecundity and spawning migration. In general, Australian fish showed an affinity for life‐history strategies that maximise fitness in hydrologically unpredictable environments. We also observed differences in trait lability between and within life history, morphological and ecological traits where in general morphological and ecological traits were more labile. Our results showed that life‐history strategies are relatively evolutionarily labile and species have potentially evolved or colonised in freshwaters frequently and independently allowing them to maximise population performance in a range of environments. In addition, reproductive guild membership showed strong phylogenetic constraint indicating that evolutionary history is an important component influencing the range and distribution of reproductive strategies in extant species assemblages. For Australian freshwater fish, biogeographic and phylogenetic history contribute to broad taxonomic differences in species functional traits, while finer scale ecological processes contribute to interspecific differences in smaller taxonomic units. These results suggest that the lability or phylogenetic relatedness of different functional traits affects their suitability for testing hypothesis surrounding community level responses to environmental change.     

Landscape connectivity of Cercidiphyllum japonicum,an endangered species and its implications for conservation

Cercidiphyllum japonicum, a Tertiary relict, recolonized areas north of the Yangtze River after the last glacial; however, little is known about its specific colonization corridors. Together with distribution models, the least cost path (LCP) analysis has been used to reveal the landscape connectivity of species. In this study, we utilized the categorical LCP method, combining the species distribution with genetic data from cpDNA and nuclear markers, to identify the possible dispersal routes of C. japonicum after the LGM. Across time periods and genetic markers, the results revealed that the species generally spread from the western edge of the Sichuan Basin, while the highest degree of dispersal potential corresponds with the year 2080 and the cpDNA haplotype. Furthermore, shifts in the species' range and the indication of an area of low genetic divergence further support the existence of a dispersal corridor. Overall, we believe that a dispersal route from the western edge of the Sichuan Basin through the Qinling Mountains and further to the northeast could exist, and therefore, the results are an important supplement to the evolutionary history of C. japonicum. In the future, we believe species distribution models (SDM) and connectivity assessment in relation to climate change will provide increasingly useful information and new implications for prioritizing the conservation of the endangered species.     

Local endemism and within‐island diversification of shrews illustrate the importance of speciation in building Sundaland mammal diversity

Island systems are important models for evolutionary biology because they provide convenient, discrete biogeographic units of study. Continental islands with a history of intermittent dry land connections confound the discrete definitions of islands and have led zoologists to predict (i) little differentiation of terrestrial organisms among continental shelf islands and (ii) extinction, rather than speciation, to be the main cause of differences in community composition among islands. However, few continental island systems have been subjected to well‐sampled phylogeographic studies, leaving these biogeographic assumptions of connectivity largely untested. We analysed nine unlinked loci from shrews of the genus Crocidura from seven mountains and two lowland localities on the Sundaic continental shelf islands of Sumatra and Java. Coalescent species delimitation strongly supported all currently recognized Crocidura species from Sumatra (six species) and Java (five species), as well as one undescribed species endemic to each island. We find that nearly all species of Crocidura in the region are endemic to a single island and several of these have their closest relative(s) on the same island. Intra‐island genetic divergence among allopatric, conspecific populations is often substantial, perhaps indicating species‐level diversity remains underestimated. One recent (Pleistocene) speciation event generated two morphologically distinct, syntopic species on Java, further highlighting the prevalence of within‐island diversification. Our results suggest that both between‐ and within‐island speciation processes generated local endemism in Sundaland, supplementing the traditional view that the region's fauna is relictual and primarily governed by extinction.     

The earthworm fauna of the Carpathian Basin with new records and description of three new species (Oligochaeta: Lumbricidae)

A review of the earthworm fauna of the Carpathian Basin recorded a total of 97 earthworm species of which three: Dendrobaena vladeasa, Octodriloides izanus and Octodrilus parvivesiculatus spp. nov. are new to science. A biogeographic analysis of the earthworm fauna revealed an extraordinarily high rate of endemism (40.21%) which is explained by the special Quaternary and tectonic history of the region.     

Comparative phylogenetic histories of two louse genera found on Catharus thrushes and other birds

The louse genera Brueelia (Ischnocera) and Myrsidea (Amblycera) are broadly codistributed on songbirds (Passeriformes), but differ in a variety of life history characteristics. We used mitochondrial and nuclear DNA sequences to assess levels of genetic divergence and reconstruct phylogenies of these 2 genera, focusing especially on Catharus thrushes in North America. We then qualitatively compared the phylogenies and levels of divergence within these 2 genera of codistributed parasites. Neither Brueelia nor Myrsidea appears to cospeciate with Catharus thrushes or passerine birds in general. The Myrsidea phylogeny exhibits significant levels of biogeographic structure, whereas the Brueelia phylogeny does not. Myrsidea and Brueelia also differ in their levels of intra-generic genetic divergence, with Myrsidea showing higher levels of genetic divergence and host specificity than Brueelia. Our genetic data support traditional morphology-based taxonomy in several instances in which the same species of Brueelia has been reported on multiple host taxa, e.g., all migrant Catharus spp. carry B. antiqua, with little haplotype divergence. Myrsidea found on each Catharus sp. are in general genetically distinct, except for M. incerta, which parasitizes both Catharus ustulatus and Catharus minimus. The strong biogeographic signal in the Myrsidea phylogeny and higher relative levels of host specificity of Myrsidea spp. suggest that infrequent host-switching, followed by speciation, is shaping the evolutionary history of this group. In contrast, the relatively lower host specificity of Brueelia spp. suggests that host-switching, combined with more frequent ongoing dispersal, has been more important in the evolutionary history of Brueelia.     

Multilocus phylogenetic and geospatial analyses illuminate diversification patterns and the biogeographic history of Malagasy endemic plated lizards (Gerrhosauridae: Zonosaurinae)

Although numerous studies have attempted to find single unifying mechanisms for generating Madagascar's unique flora and fauna, little consensus has been reached regarding the relative importance of climatic, geologic and ecological processes as catalysts of diversification of the region's unique biota. Rather, recent work has shown that both biological and physical drivers of diversification are best analysed in a case‐by‐case setting with attention focused on the ecological and life‐history requirements of the specific phylogenetic lineage under investigation. Here, we utilize a comprehensive analytical approach to examine evolutionary drivers and elucidate the biogeographic history of Malagasy plated lizards (Zonosaurinae). Data from three genes are combined with fossil information to construct time‐calibrated species trees for zonosaurines and their African relatives, which are used to test alternative diversification hypotheses. Methods are utilized for explicitly incorporating phylogenetic uncertainty into downstream analyses. Species distribution models are created for 14 of 19 currently recognized species, which are then used to estimate spatial patterns of species richness and endemicity. Spatially explicit analyses are employed to correlate patterns of diversity with both topographic heterogeneity and climatic stability through geologic time. We then use inferred geographic ranges to estimate the biogeographic history of zonosaurines within each of Madagascar's major biomes. Results suggest constant Neogene and Quaternary speciation with divergence from the African most recent common ancestor ~30 million years ago when oceanic currents and African rivers facilitated dispersal. Spatial patterns of diversity appear concentrated along coastal regions of northern and southern Madagascar. We find no relationship between either topographic heterogeneity or climatic stability and patterns of diversity. Ancestral state reconstructions suggest that western dry forests were important centres of origin with recent invasion into spiny and rain forest. These data highlight the power of combining multilocus phylogenetic and spatially explicit analyses for testing alternative diversification hypotheses within Madagascar's unique biota and more generally, particularly as applied to phylogenetically and biologically constrained systems.     

Biogeography of Eocene bryozoans from the St Vincent Basin, South Australia

The first extensive and stratigraphically detailed taxonomic study of the Middle to Late Eocene Bryozoa of the St Vincent Basin has identified more than 200 species of Cheilostomata and 50 species of Cyclostomata. There are three biogeographic groups: basin endemic, Australian and global. Two-thirds (116) of the cheilostome species and seven genera are currently considered endemic to this basin. Most species are endemic to Australia and similar to those found in the Oligo-Miocene of Victoria. The Cellariidae are a common component of most Australian Cainozoic deposits, but the species are highly dissimilar, with 13 of the 17 species here being new. The global component indicates that biogeographic links with regions outside Australia still existed in the Eocene. The cyclostome genus Reticrescis is only known from the Australian and Antarctic Eocene. Ten genera have their first occurrence in the Eocene St Vincent Basin. The Phidoloporidae and Smittinidae represent the most diverse and ubiquitous groups at a geological time close to their time of origination. Contemporaneous sediments in Antarctica, eastern Europe and North America also have a diverse fauna of this family, pointing to a strong Tethyan link. Rhamphosmittina lateralis (MacGillivray) is still extant in New Zealand, having an exceptionally long time range of 40 million years. Overall, the fauna has a distinct Late Cretaceous character. A new genus of Onychocellidae appears similar to genera that were common in Cretaceous Tethyan faunas but rare during the Cainozoic. This similarity ends in the Oligocene, after which the Australian bryozoan became endemic     

Genetic differentiation of the African dwarf crocodile Osteolaemus tetraspis Cope, 1861 (Crocodylia: Crocodylidae) and consequences for European zoos

The endangered African dwarf crocodile Osteolaemus tetraspis is distributed in Central and Western Africa. Conventionally, two subspecies were distinguished: Osteolaemus tetraspis tetraspis and Osteolaemus tetraspis osborni. The taxonomic significance of diagnostic morphological characters is still being discussed and the existence of additional species in the Osteolaemus group remains unclear. Recent molecular studies suggest the existence of three allopatric species in the genus Osteolaemus. These results supported a division of the dwarf crocodile into a Congo Basin form (O. osborni), an Ogooué Basin form (O. tetraspis), and a third separate evolutionary lineage from Western Africa. Several European zoos host African dwarf crocodiles. For reasons of conservation and possible reintroduction, it is important to clarify provenance of these zoo animals. Therefore, we conducted molecular and phylogenetic analyses of three mitochondrial and two nuclear gene sequences with all available samples from European zoos and museums. We also estimated the origin of the zoo animals by comparing sequences of wild animals and museum samples of known provenance. Our study strongly supports three distinct lineages of Osteolaemus as recently postulated, but also reveals a fourth evolutionary lineage. We demonstrate that, of the European zoo animals sampled, only one dwarf crocodile corresponds to the Congo Basin form (O. osborni) whereas the majority of individuals correspond to the three other forms. Four zoo animals belong to the new fourth group; but their provenance is still unresolved. The origin of these animals is probably located in an African region from which no wild animal samples are currently available. Further investigations and sampling of other regions should be completed to clarify the identity of this fourth lineage. We found potential hybrids from European zoological gardens using nuclear DNA sequences. The European Studbook will use these results for further breeding programmes to keep genetically suitable ex-situ populations as reassurance colonies for prospective reintroduction into African countries.     

Evolutionary history and biogeographic affinities of the serranochromine cichlids in Zambian rivers

Zambian rivers belong to two major drainages: the Luapula-Congo and the Zambezi River system. Tectonic activity repeatedly altered drainage systems in Africa, so that current fish faunas can only be understood in the context of historic drainage capture events. We use phylogenetic relationships of one widespread lineage of haplochromine cichlids, the serranochromines, to trace their biogeographic expansion and diversification in Zambia. The mitochondrial DNA phylogeny suggests five ancient clades, and their common ancestor was likely to have invaded from the Lower Congo River. The branching intervals in the linearized tree analysis suggest three major cladogenesis events and two periods of faunal exchange. The five clades originating in the Congo River drainage diversified further; one stayed in the Congo drainage, one diversified in the Zambezi system only, while the three clades of mixed distribution underwent diversification in the Zambezi system, to re-enter the Congo drainage very recently, as indicated by sometimes zero mutation differences among taxa from different drainages. Our hypothesis is consistent with the suggested radiation in the extinct Lake palaeo-Makgadikgadi, so that we propose that the Zambian serranochromine fauna in part represents survivors of the extinct lacustrine flock plus several novel species that originated in situ.     

Phylogeography and speciation in the <Emphasis Type="Italic">Pseudocrenilabrus philander</Emphasis> species complex in Zambian Rivers

Haplochromine cichlids form the most species-rich lineage of cichlid fishes that both colonized almost all river systems in Africa and radiated to species flocks in several East African lakes. The enormous diversity of lakes is contrasted by a relatively poor albeit biogeographically clearly structured species diversity in rivers. The present study analyzed the genetic structure and phylogeographic history of species and populations of the genus Pseudocrenilabrus in Zambian rivers that span two major African drainage systems, the Congo- and the Zambezi-system. The mtDNA phylogeny identifies four major lineages, three of which occur in the Congo-system and one in the Zambezi system. Two of the Congo-clades (Lake Mweru and Lunzua River) comprise distinct albeit yet undescribed species, while the fish of the third Congo-drainage clade (Chambeshi River and Bangweulu swamps), together with the fish of the Zambezi clade (Zambezi and Kafue River) are assigned to Pseudocrenilabrus philander. Concerning the intraspecific genetic diversity observed in the sampled rivers, most populations are highly uniform in comparison to lacustrine haplochromines, suggesting severe founder effects and/or bottlenecking during their history. Two bursts of diversification are reflected in the structure of the linearized tree. The first locates at about 3.9% mean sequence divergence and points to an almost simultaneous colonization of the sampled river systems. Subsequent regional diversification (with about 1% mean sequence divergence) occurred contemporaneously within the Kafue River and the Zambezi River. The clear-cut genetic biogeographic structure points to the dominance of geographic speciation in this lineage of riverine cichlid fishes, contrasting the importance of in situ diversification observed in lake cichlids.