Early adaptive radiations of Aplodontoidea (Rodentia, Mammalia) on the Holarctic region: systematics, and phylogenetic and paleobiogeographic implications

The Aplodontoidea, now restricted to only one North American species (Aplodontia rufa), have shown a wide Holarctic extension since the Upper Eocene. As their fossil record is poor, their phylogenetic relationships and the origins of their successive radiations remain unclear. We perform here phylogenetic analyses, primarily based on dental evidence (94 dental of 97 characters), restricted to Paleogene and early Miocene taxa (46 taxa) in order to avoid biases introduced by substantially derived (divergent) taxa. We confirm the inclusion of some problematic genera such as Cedromus or Douglassciurus within Sciuroidea rather than in Aplodontoidea. Ephemeromys and Lophallomys appear as the most basal members of the Aplodontoidea, and Epeiromys is the closest outgroup of the Sciuroidea-Aplodontoidea clade. The relationships among the “prosciurines” remain unclear, with paraphyletic genera such as Prosciurus and Haplomys. Their diagnoses are reevaluated and a new genus is described. The Aplodontidae, including the clade of the latter, and Haplomys liolophus display a dichotomy between Ansomyinae and Aplodontinae, the two crown groups. The first clade formed by the European species argoviensis and descendens (referred to a new genus) can be proposed as a sister group of the species of Ansomys. The second branch of the dichotomy includes the European Plesispermophilus and Sciurodon as basal groups. The species of Parallomys do not form a clade, and the genus appears paraphyletic. The last dichotomy separates the Allomys clade from the ‘meniscomyine’ clade. Comparisons of the selected species allow consideration of their patterns of dental evolution (e.g. enlargement of P4, development of a metaloph—protoloph disto-mesial connection, of crescentic shape in main cusps and ectoloph, of a buccal protruding compressed mesostyle, of a metastylid crest or an anterior spur of the hypoconid, etc.). The split between sciuroids and aplodontoids occurred in North America, and then aplodontoids dispersed rapidly throughout the whole Holarctic region. The first aplodontid adaptive radiation took place either in North America or in Asia. Periodic exchanges occurred between Europe, Asia and North America, and the last radiations (meniscomyines) were restricted in North America.     

Limited genetic divergence among Australian alpine Poa tussock grasses coupled with regional structuring points to ongoing gene flow and taxonomic challenges

Background and Aims

While molecular approaches can often accurately reconstruct species relationships, taxa that are incompletely differentiated pose a challenge even with extensive data. Such taxa are functionally differentiated, but may be genetically differentiated only at small and/or patchy regions of the genome. This issue is considered here in Poa tussock grass species that dominate grassland and herbfields in the Australian alpine zone.

Methods

Previously reported tetraploidy was confirmed in all species by sequencing seven nuclear regions and five microsatellite markers. A Bayesian approach was used to co-estimate nuclear and chloroplast gene trees with an overall dated species tree. The resulting species tree was used to examine species structure and recent hybridization, and intertaxon fertility was tested by experimental crosses.

Key Results

Species tree estimation revealed Poa gunnii, a Tasmanian endemic species, as sister to the rest of the Australian alpine Poa. The taxa have radiated in the last 0·5–1·2 million years and the non-gunnii taxa are not supported as genetically distinct. Recent hybridization following past species divergence was also not supported. Ongoing gene flow is suggested, with some broad-scale geographic structure within the group.

Conclusions

The Australian alpine Poa species are not genetically distinct despite being distinguishable phenotypically, suggesting recent adaptive divergence with ongoing intertaxon gene flow. This highlights challenges in using conventional molecular taxonomy to infer species relationships in recent, rapid radiations.     

High diversity and local endemism in Aotearoa New Zealand's groundwater crustacean fauna

We used DNA barcoding to assess the diversity and distribution of New Zealand''s groundwater amphipods and isopods (Crustacea) and to determine whether biodiversity and endemism within tectonically active New Zealand are similar to those of more tectonically stable continents. Sixty‐five wells were sampled in seven aquifers across four regions within the North and South islands of New Zealand, and resident invertebrates were morphologically identified and then assessed using sequencing of the mitochondrial DNA cytochrome c oxidase subunit one (COI) gene. Invertebrates were found in 54 wells. Of the 228 individual amphipods and isopods found in 36 of the wells, 154 individuals were successfully sequenced for COI (68% success rate) from 25 wells, with at least one well in each aquifer containing sequenced individuals. Of the 45 putative species identified using Barcode Index Numbers (BINs), 30 BINs (78% of all taxa and 83% of amphipods) were previously unrecorded. Substantial morphologically cryptic, species‐level diversity was revealed, particularly within the amphipod Family Paraleptamphopidae. Similarly, one isopod taxon morphologically identified as Cruregens fontanus was assigned to five well‐separated BINs based on COI sequences. Endemism appeared high, with all taxa regionally endemic; 87% of species were restricted to one aquifer and more than 50% restricted to one well. Non‐saturated species accumulation curves indicated that, while additional sampling may increase the range of some currently identified taxa, additional range‐restricted taxa are also likely to be discovered. Patterns of diversity and short‐range endemism were similar to those found elsewhere, including locations which are more tectonically stable. The predominance of local endemism within New Zealand''s groundwater fauna suggests that land‐use activities and groundwater extraction require careful evaluation to minimize threats to groundwater biodiversity.     

Explosive speciation in the New World Dendroica warblers

The 27 species of Dendroica wood-warblers represent North America''s most spectacular avian adaptive radiation. Dendroica species exhibit high levels of local sympatry and differ in plumage and song, but the group contrasts with other well-known avian adaptive radiations such as the Hawaiian honeycreepers and Galapagos finches in that Dendroica species have differentiated modestly in morphometric traits related to foraging. Instead, sympatric Dendroica tend to partition resources behaviourally and they have become a widely cited example of competitive exclusion. We explored the temporal structure of Dendroica diversification via a phylogeny based on 3639 nucleotides of protein-coding mitochondrial DNA (mtDNA). The taxa sampled included 60 individuals representing 24 Dendroica species and a variety of other paruline warbler and outgroup species. Mitochondrial divergences among Dendroica species were generally large (mean pairwise interspecific distances, 10.0%) and many species were rooted in a basal polytomy. The prevalence of long terminal branches indicates that these species have evolved efficient isolating mechanisms that have prevented mtDNA introgression despite the many opportunities for hybridization resulting from local sympatry. Comparisons with a null model of random bifurcation–extinction demonstrate that cladogenesis in Dendroica has been clustered non-randomly with respect to time, with a significant burst of speciation occurring early in the history of the genus, possibly as long ago as the Late Miocene or Early Pliocene periods. Although this non-random clustering of speciation is consistent with the pattern expected of an adaptive radiation, the age of the Dendroica radiation suggests it is an ''ancient species flock'' in which most extant species represent lineages that have long been evolutionarily independent.     

Molecular phylogeny of Anaphalis (Asteraceae, Gnaphalieae) with biogeographic implications in the Northern Hemisphere

Anaphalis is the largest Asian genus in the tribe Gnaphalieae (Asteraceae) and has its greatest species diversity in the eastern Himalayas. The nuclear internal and external transcribed spacers were sequenced for Anaphalis species, with an emphasis on the eastern Himalayan taxa to examine the monophyly and construct the phylogenetic relationships of and within the genus. The results suggest that all species of Anaphalis are nested with Helichrysum, showing a close relationship with a Mediterranean–Asian group of Helichrysum. Although the monophyly of Anaphalis is only weakly supported, two clades within the genus are well recognized, each consisting of two subgroups. The inferred phylogenetic relationships within Anaphalis correspond to the shape of leaf base, rather than the morphology of the capitula and phyllaries that are usually used for species delimitation and classification in the genus. All four subgroups of Anaphalis are common and diversified in the eastern Himalayas with multiple dispersals out of this region. The sole North American species of Anaphalis is best hypothesized to be the result of long-distance dispersal or overland migration via Bering land bridge from Asia. Our analyses suggest that the extant distribution of Anaphalis has most likely resulted one radiation into the eastern Himalayas followed by repeated independent dispersals and/or radiations mostly into eastern Asia but also into the western Himalayas, North America, and southeast Asia.     

Comparative Phylogeography in Rainforest Trees from Lower Guinea,Africa

Comparative phylogeography is an effective approach to assess the evolutionary history of biological communities. We used comparative phylogeography in fourteen tree taxa from Lower Guinea (Atlantic Equatorial Africa) to test for congruence with two simple evolutionary scenarios based on physio-climatic features 1) the W-E environmental gradient and 2) the N-S seasonal inversion, which determine climatic and seasonality differences in the region. We sequenced the trnC-ycf6 plastid DNA region using a dual sampling strategy: fourteen taxa with small sample sizes (dataset 1, mean n = 16/taxon), to assess whether a strong general pattern of allele endemism and genetic differentiation emerged; and four taxonomically well-studied species with larger sample sizes (dataset 2, mean n = 109/species) to detect the presence of particular shared phylogeographic patterns. When grouping the samples into two alternative sets of two populations, W and E, vs. N and S, neither dataset exhibited a strong pattern of allelic endemism, suggesting that none of the considered regions consistently harboured older populations. Differentiation in dataset 1 was similarly strong between W and E as between N and S, with 3–5 significant F _ST tests out of 14 tests in each scenario. Coalescent simulations indicated that, given the power of the data, this result probably reflects idiosyncratic histories of the taxa, or a weak common differentiation pattern (possibly with population substructure) undetectable across taxa in dataset 1. Dataset 2 identified a common genetic break separating the northern and southern populations of Greenwayodendron suaveolens subsp. suaveolens var. suaveolens, Milicia excelsa, Symphonia globulifera and Trichoscypha acuminata in Lower Guinea, in agreement with differentiation across the N–S seasonal inversion. Our work suggests that currently recognized tree taxa or suspected species complexes can contain strongly differentiated genetic lineages, which could lead to misinterpretation of phylogeographic patterns. Therefore the evolutionary processes of such taxa require further study in African tropical rainforests.     

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.     

Italian neo-endemism in a widespread group of leafhoppers insects: A revision of the Eupteryx aurata group (Auchenorrhyncha: Cicadellidae: Typhlocybinae) using morphology,ecology and genetics

The Eupteryx aurata group is characterised by several features, primarily by colouration and aedeagus morphology. It is distributed mainly in Europe, with some species also in Asia and one species perhaps in North Africa. At least one species is introduced in North America. Until now 12 taxa were known. Two new species, Eupteryx divulsa from central Italy and Eupteryx oscorum from central and southern Italy, are described in this paper. Eupteryx petasitidis Ferrari, 1882, described as Eupteryx carpini petasitidis and later synonymised with Eupteryx atropunctata (Goeze, 1778), is reestablished on species level and redescribed including genital morphology. Ten taxa of the species group are studied in relation to their morphological and molecular characters. The colouration, aedeagus, and pygofer appendage are figured for all of these taxa. Molecular study showed that the E. aurata group is monophyletic. The little or moderate distances in sequences (below 6.0% for mtDNA and up to 2.5% for nuclear ITS2) between the different taxa of the E. aurata group and incongruence of genetic signals from mtDNA and nuclear DNA could be the result of incomplete lineage sorting in young species and/or hybridisation and introgression of DNA among taxa. Of particular importance in this study is material from Italy, a peripheral region in relation to the distribution of the species group and due to its geomorphological features favouring isolation and speciation. A distribution map for the E. aurata species group in Italy and an identification key for all 15 taxa of the aurata group are given. Phylogenetic relationships within the aurata group are discussed in relation to morphological, molecular, ecological, and zoogeographic aspects.     

Phylogeography of a Morphologically Cryptic Golden Mole Assemblage from South-Eastern Africa

The Greater Maputaland-Pondoland-Albany (GMPA) region of southern Africa was recently designated as a centre of vertebrate endemism. The phylogeography of the vertebrate taxa occupying this region may provide insights into the evolution of faunal endemism in south-eastern Africa. Here we investigate the phylogeographic patterns of an understudied small mammal species assemblage (Amblysomus) endemic to the GMPA, to test for cryptic diversity within the genus, and to better understand diversification across the region. We sampled specimens from 50 sites across the distributional range of Amblysomus, with emphasis on the widespread A. hottentotus, to analyse geographic patterns of genetic diversity using mitochondrial DNA (mtDNA) and nuclear intron data. Molecular dating was used to elucidate the evolutionary and phylogeographic history of Amblysomus. Our phylogenetic reconstructions show that A. hottentotus comprises several distinct lineages, or evolutionarily significant units (ESUs), some with restricted geographic ranges and thus worthy of conservation attention. Divergence of the major lineages dated to the early Pliocene, with later radiations in the GMPA during the late-Pliocene to early-Pleistocene. Evolutionary diversification within Amblysomus may have been driven by uplift of the Great Escarpment c. 5–3 million years ago (Ma), habitat changes associated with intensification of the east-west rainfall gradient across South Africa and the influence of subsequent global climatic cycles. These drivers possibly facilitated geographic spread of ancestral lineages, local adaptation and vicariant isolation. Our study adds to growing empirical evidence identifying East and southern Africa as cradles of vertebrate diversity.     

Disturbance,seeds, restoration,and the importance of experiments and long‐term observations: the Editors' Award for 2005

Question: Can the geographic patterning of endemic plant species inform reserve selection in a region of high endemism? Location: The Southeastern Coastal Plain of North America, focusing primarily on the imperiled longleaf pine (Pinus palustris P. Miller) ecosystem. Methods: We documented the high level of plant endemism in the region, and characterized the endemic taxa into distributional subregions. Results: A total of 1630 plant taxa are endemic to the Coastal Plain, a large proportion of which are endemic to phytogeographical subregions within the Coastal Plain, with particularly large numbers of narrow endemics occurring in the East Gulf Coastal Plain and Florida Peninsula. Conclusions: This pattern of local endemism presents challenges in conserving the full biota of the region: a reserve system focusing on few and large conservation areas has theoretical benefits for long‐term management and viability, but will fail to capture many local endemics. We propose that the dispersed distribution of endemic species will require a mixture of large core reserves and smaller satellite reserves.     

Diversity of North American map and sawback turtles (Testudines: Emydidae: Graptemys)

Map turtles of the genus Graptemys are native to North America, where a high degree of drainage endemism is believed to have shaped current diversity. With 14 species and one additional subspecies, Graptemys represents the most diverse genus in the family Emydidae. While some Graptemys species are characterized by pronounced morphological differences, previous phylogenetic analyses have failed yet to confirm significant levels of genetic divergence for many taxa. As a consequence, it has been debated whether Graptemys is taxonomically inflated or whether the low genetic divergence observed reflects recent radiations or ancient hybridization. In this study, we analysed three mtDNA blocks (3228 bp) as well as 12 nuclear loci (7844 bp) of 89 specimens covering all species and subspecies of Graptemys. Our analyses of the concatenated mtDNA sequences reveal that the widespread G. geographica constitutes the sister taxon of all other Graptemys species. These correspond to two clades, one comprised of all broad‐headed Graptemys species and another clade containing the narrow‐headed species. Most species of the broad‐headed clade are reciprocally monophyletic, except for G. gibbonsi and G. pearlensis, which are not differentiated. By contrast, in the narrow‐headed clade, many currently recognized species are not monophyletic and divergence is significantly less pronounced. Haplotype networks of phased nuclear loci show low genetic divergence among taxa and many shared haplotypes. Principal component analyses using coded phased nuclear DNA sequences revealed eight distinct clusters within Graptemys that partially conflict with the terminal mtDNA clades. This might be explained by male‐mediated gene flow across drainage basins and female philopatry within drainage basins. Our results support that Graptemys is taxonomically oversplit and needs to be revised.     

Speciation and phylogeography of Hawaiian terrestrial arthropods

The Hawaiian archipelago is arguably the world's finest natural laboratory for the study of evolution and patterns of speciation. Arthropods comprise over 75% of the endemic biota of the Hawaiian Islands and a large proportion belongs to species radiations. We classify patterns of speciation within Hawaiian arthropod lineages into three categories: (i) single representatives of a lineage throughout the islands; (ii) species radiations with either (a) single endemic species on different volcanoes or islands, or (b) multiple species on each volcano or island; and (iii) single widespread species within a radiation of species that exhibits local endemism. A common pattern of phylogeography is that of repeated colonization of new island groups, such that lineages progress down the island chain, with the most ancestral groups (populations or species) on the oldest islands. While great dispersal ability and its subsequent loss are features of many of these taxa, there are a number of mechanisms that underlie diversification. These mechanisms may be genetic, including repeated founder events, hybridization, and sexual selection, or ecological, including shifts in habitat and/or host affiliation. The majority of studies reviewed suggest that natural selection is a primary force of change during the initial diversification of taxa.     

Diverse pollination systems of the twin-spurred orchid genus Satyrium in African grasslands

The large terrestrial orchid genus Satyrium underwent evolutionary radiations in the Cape floral region and the grasslands of southern and eastern Africa. These radiations were accompanied by tremendous diversification of the unusual twin-spurred flowers that characterize the genus, but pollination data required to interpret these patterns of floral evolution have been lacking for grassland species in the genus. Here we document pollinators, nectar properties, and levels of pollination success for 11 grassland Satyrium species in southern and south-central Africa. Pollinators of these species include bees, beetles, butterflies, hawkmoths, noctuid moths, long-proboscid flies, and sunbirds. Most species appear to be specialized for pollination by one functional pollinator group. Long-proboscid fly pollination systems are reported for the first time in Satyrium (in S. macrophyllum and a high-altitude form of S. neglectum). Floral morphology, especially spur length and rostellum structure, differs markedly among plants with different pollinators, while nectar volume, concentration, and sugar composition are fairly uniform across species. Most taxa exhibited high levels of pollination success (>50% of flowers pollinated), a trend that can be attributed to the presence of nectar in the twin spurs.     

Plant diversity of Hyrcanian relict forests: An annotated checklist,chorology and threat categories of endemic and near endemic vascular plant speciesOA

In this paper a critical annotated checklist of 256 endemic and near endemic species belonging to 152 genera and 50 families of flowering plants known from Hyrcanian relict forests is presented. Distribution maps of taxa, elevational range, number of known records, chorotypes, life forms, IUCN threat categories and habitat types are also provided. The chorotypes are categorized into eight main patterns: 1) the Omni-Hyrcanian pattern(OH), 2) West Hyrcanian pattern(WH), 3) Manjil-Rudbar pattern(MR...     

Recent origin,active speciation and dispersal for the lichen genus Nephroma (Peltigerales) in Macaronesia

Aim We reconstructed the phylogeny of the lichen genus Nephroma (Peltigerales) to assess the relationships of species endemic to Macaronesia. We estimated dates of divergences to test the hypothesis that the species arose in Macaronesia (neo‐endemism) versus the oceanic archipelagos serving as refugia for formerly widespread taxa (palaeo‐endemism). Location Cosmopolitan with a special focus on the archipelagos of the Azores, Madeira and the Canary Islands. Methods DNA sequences were obtained from 18 species for three loci and analysed using maximum parsimony, maximum likelihood and Bayesian inferences. Divergence dates were estimated for the internal transcribed spacer (ITS)‐based phylogeny using a relaxed molecular clock. Reconstruction of the ancestral geographical range was conducted using the Bayesian 50% majority rule consensus tree under a parsimony method. Results The backbone phylogenetic tree was fully supported, with Nephroma plumbeum as sister to all other species. Four strongly supported clades were detected: the Nephroma helveticum, the N. bellum, the N. laevigatum and the N. parile clades. The latter two share a common ancestor and each includes a widespread Holarctic species (N. laevigatum and N. parile, respectively) and all species endemic to Macaronesia. The data suggest a neo‐endemic origin of Macaronesian taxa, a recent range expansion from Macaronesia of both widespread species, a range expansion limited to the Mediteranean Basin and south‐western Europe for another taxon, and a long dispersal event that resulted in a speciation event in the western parts of North America. Main conclusions The Macaronesian endemic species belong to two sister clades and originated from a most recent common ancestor (MRCA) shared with one widely distributed taxon, either N. parile or N. laevigatum. Estimates of the mean divergence dates suggest that the endemics originated in the archipelagos after the rise of the volcanic islands, along with the ancestor to the now widespread species, which probably expanded their range beyond Macaronesia via long‐distance dispersal. This study provides the first phylogenetic evidence of Macaronesian neo‐endemism in lichenized fungi and provides support for the hypothesis that oceanic islands may serve as a source for the colonization of continents. However, further data are needed to properly assess the alternative hypothesis, namely colonization from western North America.     

Floristic division of the Arctic

Abstract: The progress in the floristic study of the circumpolar Arctic since the 1940s is summarized and a new floristic division of this region is presented. The treeless areas of the North Atlantic and North Pacific with an oceanic climate, absence of permafrost and a very high proportion of boreal taxa are excluded from the Arctic proper. It is argued that the Arctic deserves the status of a floristic region. The tundra zone and some oceanic areas are divided into subzones according to their flora and vegetation. Two groups of subzones are recognized: the Arctic group (including the Arctic tundras proper and the High Arctic) and the Hypoarctic group. The Arctic phytochorion is floristically divided into sectors: 6 provinces and 20 subprovinces reflecting the regional features of each sector in connection with flora history, physiography and continentality-oceanity of the climate. Each sector is described and differentiated by a set of differential and co-differential species. The peculiarities of the Arctic flora are manifest in different ways in the various sectors, and endemism is not the universal criterion for subdivision.     

Biodiversity in the Gulf of Guinea: an overview

Patterns of species-richness and endemism in the Gulf of Guinea reflect the region's biogeographic history. Bioko is a continental-shelf island that was recently connected to the African mainland, whereas Príncipe, São Tomé and Annobón are truly oceanic and have never been connected with each other or with the mainland. As a result, Bioko supports a much more diverse flora and fauna but with relatively low levels of endemism at the species level, whereas the oceanic islands are relatively depauperate because of their isolation but rich in endemic taxa. Species endemism is 0–3% on Bioko for angiosperms, bats, birds, reptiles and amphibians, compared with much higher values on Principe for these same taxa of 8% (plants) to 100% (amphibians), on São Tomé between 14% (plants) and 100% (amphibians), and on Annobón 0% (bats) to 71% (reptiles). On a global scale, for their size both Príncipe and São Tomé support unusually high numbers of single-island endemic species of birds, reptiles and amphibia. For its tiny size, Annobón is also notable for its endemic birds and reptiles. Among terrestrial molluscs the rates of endemism are in general higher than for plants and vertebrates, from ca 50% on Bioko to ca 80% on the oceanic islands. In contrast and as might be expected, only Bioko supports a rich freshwater fish fauna and it contains many endemic taxa, whereas the oceanic islands support only a few salt-tolerant species. The Gulf of Guinea islands are also important for their marine organisms, amongst which coral reef fish and marginellid molluscs show high levels of endemism, though they are not especially species-rich. The Gulf of Guinea islands are of great interest to conservationists and evolutionary biologists. Each island, of greatly differing size and degree of isolation, has acquired its unique sub-set of plants and animals separately from the neighbouring mainland, followed by adaptive radiations in situ. For this reason the conservation value of the archipelago as a whole is greater than the sum of the biodiversity contained in its individual islands. Conservation initiatives in the Gulf of Guinea should therefore ensure that representative terrestrial, freshwater and marine habitats and groups of organisms are targeted in a co-ordinated manner among the islands.     

From ratites to rats: the size of fleshy fruits shapes species' distributions and continental rainforest assembly

Seed dispersal is a key process in plant spatial dynamics. However, consistently applicable generalizations about dispersal across scales are mostly absent because of the constraints on measuring propagule dispersal distances for many species. Here, we focus on fleshy-fruited taxa, specifically taxa with large fleshy fruits and their dispersers across an entire continental rainforest biome. We compare species-level results of whole-chloroplast DNA analyses in sister taxa with large and small fruits, to regional plot-based samples (310 plots), and whole-continent patterns for the distribution of woody species with either large (more than 30 mm) or smaller fleshy fruits (1093 taxa). The pairwise genomic comparison found higher genetic distances between populations and between regions in the large-fruited species (Endiandra globosa), but higher overall diversity within the small-fruited species (Endiandra discolor). Floristic comparisons among plots confirmed lower numbers of large-fruited species in areas where more extreme rainforest contraction occurred, and re-colonization by small-fruited species readily dispersed by the available fauna. Species'' distribution patterns showed that larger-fruited species had smaller geographical ranges than smaller-fruited species and locations with stable refugia (and high endemism) aligned with concentrations of large fleshy-fruited taxa, making them a potentially valuable conservation-planning indicator.     

In or Out-of-Madagascar?—Colonization Patterns for Large-Bodied Diving Beetles (Coleoptera: Dytiscidae)

High species diversity and endemism within Madagascar is mainly the result of species radiations following colonization from nearby continents or islands. Most of the endemic taxa are thought to be descendants of a single or small number of colonizers that arrived from Africa sometime during the Cenozoic and gave rise to highly diverse groups. This pattern is largely based on vertebrates and a small number of invertebrate groups. Knowledge of the evolutionary history of aquatic beetles on Madagascar is lacking, even though this species-rich group is often a dominant part of invertebrate freshwater communities in both standing and running water. Here we focus on large bodied diving beetles of the tribes Hydaticini and Cybistrini. Our aims with this study were to answer the following questions 1) How many colonization events does the present Malagasy fauna originate from? 2) Did any colonization event lead to a species radiation? 3) Where did the colonizers come from—Africa or Asia—and has there been any out-of-Madagascar event? 4) When did these events occur and were they concentrated to any particular time interval? Our results suggest that neither in Hydaticini nor in Cybistrini was there a single case of two or more endemic species forming a monophyletic group. The biogeographical analysis indicated different colonization histories for the two tribes. Cybistrini required at least eight separate colonization events, including the non-endemic species, all comparatively recent except the only lotic (running water) living Cybister operosus with an inferred colonization at 29 Ma. In Hydaticini the Madagascan endemics were spread out across the tree, often occupying basal positions in different species groups. The biogeographical analyses therefore postulated the very bold hypothesis of a Madagascan origin at a very deep basal node within Hydaticus and multiple out-of-Madagascar dispersal events. This hypothesis needs to be tested with equally intense taxon sampling of mainland Africa as for Madagascar.     

Historical Biogeography and Diversification of Truffles in the Tuberaceae and Their Newly Identified Southern Hemisphere Sister Lineage

Truffles have evolved from epigeous (aboveground) ancestors in nearly every major lineage of fleshy fungi. Because accelerated rates of morphological evolution accompany the transition to the truffle form, closely related epigeous ancestors remain unknown for most truffle lineages. This is the case for the quintessential truffle genus Tuber, which includes species with socio-economic importance and esteemed culinary attributes. Ecologically, Tuber spp. form obligate mycorrhizal symbioses with diverse species of plant hosts including pines, oaks, poplars, orchids, and commercially important trees such as hazelnut and pecan. Unfortunately, limited geographic sampling and inconclusive phylogenetic relationships have obscured our understanding of their origin, biogeography, and diversification. To address this problem, we present a global sampling of Tuberaceae based on DNA sequence data from four loci for phylogenetic inference and molecular dating. Our well-resolved Tuberaceae phylogeny shows high levels of regional and continental endemism. We also identify a previously unknown epigeous member of the Tuberaceae – the South American cup-fungus Nothojafnea thaxteri (E.K. Cash) Gamundí. Phylogenetic resolution was further improved through the inclusion of a previously unrecognized Southern hemisphere sister group of the Tuberaceae. This morphologically diverse assemblage of species includes truffle (e.g. Gymnohydnotrya spp.) and non-truffle forms that are endemic to Australia and South America. Southern hemisphere taxa appear to have diverged more recently than the Northern hemisphere lineages. Our analysis of the Tuberaceae suggests that Tuber evolved from an epigeous ancestor. Molecular dating estimates Tuberaceae divergence in the late Jurassic (∼156 million years ago), with subsequent radiations in the Cretaceous and Paleogene. Intra-continental diversification, limited long-distance dispersal, and ecological adaptations help to explain patterns of truffle evolution and biodiversity.