Ecophenotypic plasticity leads to extraordinary gastropod shells found on the “Roof of the World”

The often extraordinary shell forms and shapes of gastropods found in palaeolakes, such as the highly diverse Gyraulus fauna of the famous Steinheim Basin, have been puzzling evolutionary biologists for centuries, and there is an ongoing debate whether these aberrant shell forms are indicative of true species (or subspecies) or ecophenotypic morphs. Interestingly, one of the Steinheim Gyraulus morphs – a corkscrew-like open-coiled shell – has a recent analogue in the Lake Bangong drainage system on the western Tibetan Plateau. Therefore, a combination of morphological, molecular, palaeolimnological, and ecological analyses was used in this study to assess whether the extraordinary shell shape in Gyraulus sp. from this drainage system represents a (young) ecophenotypic phenomenon or if it has been genetically fixed over an extended period of time. Our morphological, ecological, and palaeolimnological data suggest that the corkscrew-like specimens remain restricted to a small pond near Lake Bangong with an elevated pH value and that the colonization may have occurred recently. The phylogenetic reconstruction based on two gene fragments shows that these nonplanispiral specimens cluster within the previous described Tibetan Plateau Gyraulus clade N2. A network analysis indicates that some haplotypes are even shared by planispiral and nonplanispiral specimens. Given the ephemerality of the phenomenon, the compact network patterns inferred, the likely young phylogenetic age of the aberrant Gyraulus shells studied, and the ecological peculiarities of the study site, we suggest that the evolution of the aberrant shell forms on the Tibetan Plateau could likely be considered as a rapid ecophenotypic response, possibly induced by ecological stress. This finding may thus have implications for the ongoing debate about the processes that have caused the extraordinary shell diversity in palaeolakes such as the Steinheim Basin.     

Predation on freshwater snails in Miocene Lake Steinheim: a trigger for intralacustrine evolution?

Shells of the freshwater gastropods Gyraulus trochiformis (Planorbidae or ramshorn snails) and Bania pseudoglobula (formerly Pseudamnicola, Hydrobiidae) from Miocene Steinheim Basin in SW Germany contain small holes with a mean diameter of 0.8 mm. Analyses of comparable holes are so far unknown from fossil or extant assemblages of freshwater shells. This analysis of the perforated shells suggests that the holes were not formed by post‐depositional or pure taphonomic processes. Instead, they were most likely produced by predators. This analysis widens the means for identification of predation on freshwater snail shells that can be used in other palaeolimnological studies. The co‐occurrence of fish teeth and perforated shells in the studied samples as well as the perforation features suggest that the predator was either barbel or tench fish. The correlation between shell sizes and hole diameters suggests a size relationship between predator and prey that may generally be related to gape‐limited fish predators. The co‐occurrence of perforated shells in these lake sediments with a dominance of large and armoured shells suggest that these larger shells with protuberances and knobs could not be crushed by the gape‐limited fish predators. This analysis is the basis for a hypothesis that the endemic evolution of Gyraulus in Lake Steinheim, with some varied forms of shell thickness and morphology, was triggered by a predator–prey relationship based on adaptations to avoid shell‐breaking predators.     

The Steinheim snails (Miocene; Schwäbische Alb) from a neo-darwinian point of view: A discussion

The meteorite crater lake of Steinheim was invaded byGyraulus kleini which is widely distributed in the Upper-Freshwater Molasse Formation of Southern Germany. This species is the founder-species of an endemic lineage which speciated repeatedly. The reason for the endemism of the lineage may have been the exceptionally high ion-content of the lake water to which the animals were adapted. The change in morphology within the lineage is only indirectly connected with the geochemical regime.     

Endemism and diversification in freshwater insects of Madagascar revealed by coalescent and phylogenetic analysis of museum and field collections

The biodiversity and endemism of Madagascar are among the most extraordinary and endangered in the world. This includes the island’s freshwater biodiversity, although detailed knowledge of the diversity, endemism, and biogeographic origin of freshwater invertebrates is lacking. The aquatic immature stages of mayflies (Ephemeroptera) are widely used as bio-indicators and form an important component of Malagasy freshwater biodiversity. Many species are thought to be microendemics, restricted to single river basins in forested areas, making them particularly sensitive to habitat reduction and degradation. The Heptageniidae are a globally diverse family of mayflies (>500 species) but remain practically unknown in Madagascar except for two species described in 1996. The standard approach to understanding their diversity, endemism, and origin would require extensive field sampling on several continents and years of taxonomic work followed by phylogenetic analysis. Here we circumvent this using museum collections and freshly collected individuals in a combined approach of DNA taxonomy and phylogeny. The coalescent-based GMYC analysis of DNA barcode data (mitochondrial COI) revealed 14 putative species on Madagascar, 70% of which were microendemics. A phylogenetic analysis that included African and Asian species and data from two mitochondrial and four nuclear loci indicated the Malagasy Heptageniidae are monophyletic and sister to African species. The genus Compsoneuria is shown to be paraphyletic and the genus Notonurus is reinstalled for African and Malagasy species previously placed in Compsoneuria. A molecular clock excluded a Gondwanan vicariance origin and instead favoured a more recent overseas colonization of Madagascar. The observed monophyly and high microendemism highlight their conservation importance and suggest the DNA-based approach can rapidly provide information on the diversity, endemism, and origin of freshwater biodiversity. Our results underline the important role that museum collections can play in molecular studies, especially in critically endangered biodiversity hotspots like Madagascar where entire species or populations may go extinct very quickly.     

Reappraisal of the Genus Tapirus (Perissodactyla, Tapiridae): Systematics and Phylogenetic Affinities of the South American Tapirs

The record of the genus Tapirus in South America is associated with the faunistic events of the Great American Biotic Interchange (GABI). The taxon is considered an immigrant of Holarctic origin. Although remains are scarce and incomplete during the Pleistocene, an analysis of these materials allowed us to consider valid seven fossil species : Tapirus tarijensis, T. cristatellus, T. greslebini, T. rioplatensis, T. oliverasi, T. mesopotamicus, and T. rondoniensis. A phylogenetic analysis was carried out in order to elucidate the relationships of the American fossil and extant species. Our result is consistent with a paraphyletic hypothesis for South American tapirs and suggests that a second dispersal event would have occurred from South America to North America, of a form closely related to T. cristatellus, resulting in the derived forms of North America.     

Species richness and endemism in the Western Australian flora

Aim Estimates of endemic and non‐endemic native vascular plant species in each of the three Western Australian Botanical Provinces were made by East in 1912 and Beard in 1969. The present paper contains an updated assessment of species endemism in the State. Location Western Australia comprises one third of the continental Australian land mass. It extends from 13° to 35° S and 113° to 129° W. Methods Western Australia is recognized as having three Botanical Provinces (Northern, Eremaean and South‐West) each divided into a number of Botanical Districts. Updated statistics for number of species and species endemism in each Province are based on the Census of Western Australian Plants data base at the Western Australian Herbarium ( Western Australian Herbarium, 1998 onwards). Results The number of known species in Western Australia has risen steadily over the years but reputed endemism has declined in the Northern and Eremaean Provinces where cross‐continental floras are common. Only the isolated South‐West Province retains high rates of endemism (79%). Main conclusions With 5710 native species, the South‐West Province contains about the same number as the California Floristic Province which has a similar area. The Italian mediterranean zone also contains about this number but in a smaller area, while the much smaller Cape Floristic Region has almost twice as many native species. The percentage of endemic species is highest at the Cape, somewhat less in south‐western Australia and less again in California. Italy, at 12.5%, has the lowest value. Apart from Italy, it is usual for endemism to reach high values in the largest plant families. In Western Australia, these mainly include woody sclerophyll shrubs and herbaceous perennials with special adaptations to environmental conditions. While those life forms are prominent in the Cape, that region differs in the great importance of herbaceous families and succulents, both of which are virtually absent from Western Australia. In California and Italy, most endemics are in families of annual, herbaceous perennial and soft shrub plants. It is suggested that the dominant factor shaping the South‐West Province flora is the extreme poverty of the area’s soils, a feature that emphasizes sclerophylly, favours habitat specialization and ensures relatively many local endemic species.     

Diversification,Biogeographic Pattern,and Demographic History of Taiwanese Scutellaria Species Inferred from Nuclear and Chloroplast DNA

The ragged topography created by orogenesis generates diversified habitats for plants in Taiwan. In addition to colonization from nearby mainland China, high species diversity and endemism of plants is also present in Taiwan. Five of the seven Scutellaria species (Lamiaceae) in Taiwan, for example, are endemic to the island. Hypotheses of multiple sources or in situ radiation have arisen to explain the high endemism of Taiwanese species. In this study, phylogenetic analyses using both nuclear and chloroplast markers revealed the multiple sources of Taiwanese Scutellaria species and confirmed the rapid and recent speciation of endemic species, especially those of the “indica group” composed of S. indica, S. austrotaiwanensis, S. tashiroi, and S. playfairii. The common ancestors of the indica group colonized first in northern Taiwan and dispersed regionally southward and eastward. Climate changes during glacial/interglacial cycles led to gradual colonization and variance events in the ancestors of these species, resulting in the present distribution and genetic differentiation of extant populations. Population decline was also detected in S. indica, which might reflect a bottleneck effect from the glacials. In contrast, the recently speciated endemic members of the indica group have not had enough time to accumulate much genetic variation and are thus genetically insensitive to demographic fluctuations, but the extant lineages were spatially expanded in the coalescent process. This study integrated phylogenetic and population genetic analyses to illustrate the evolutionary history of Taiwanese Scutellaria of high endemism and may be indicative of the diversification mechanism of plants on continental islands.     

Comparative Anatomy of the Petrosal Bone of Dichobunoids,Early Members of Artiodactylamorpha (Mammalia)

Among Artiodactylamorpha, dichobunoids are some of the oldest fossil species that have been associated with Artiodactyla, the crown clade that includes hippopotamids, camelids, suoids, ruminants, and cetaceans. These important fossil species are known from early Eocene rocks of North America, Europe, and Asia, but their phylogenetic position has yet to be well resolved. Before generating such a phylogeny, it is first critical to document all of the anatomy of known dichobunoid fossils. Here we use CT scans to describe previously undescribed anatomy of the petrosal bone, a complex part of the mammalian skull that contains many variable and phylogenetically informative features. Results show that these extinct species share a number of features that are not documented in modern species including a lateral process of the epitympanic wing constituting the medial border of the piriform fenestra, and a tegmen tympani foramen that may have given passage to the ramus superior of the stapedial artery. Future comprehensive phylogenetic studies may show that many of these characters are plesiomophic for Artiodactylamopha. Some species (Diacodexis, Homacodon and ?Helohyus) exhibit a dorsolateral exposure of the mastoid region of the petrosal on the temporal part of the cranium. This uncommon feature has, to our knowledge, not been reported in another euungulate group.     

Amplified fragment length polymorphisms,the evolution of the land snail genus Theba (Stylommatophora: Helicidae), and an objective approach for relating fossils to internal nodes of a phylogenetic tree using geometric morphometrics

Time‐trees are commonly calibrated based on fossils attributed to internal nodes, thus defining the minimum age of these nodes. However, in the absence of synapomorphies, the phylogenetic position of a fossil can only be inferred based on similarity. In this study, we objectively allocated fossil shells to internal nodes based on the reconstruction of ancestral shapes and sizes in a geometric morphometric framework. Our phylogenetic analysis of 24 (putative sub)species was based on 2524 amplified fragment length polymorphism loci. In this well‐supported tree the taxa occurring in north‐west Africa and on the Iberian peninsula were paraphyletic with respect to the (sub)species from the Canary and Selvagen Islands, indicating a continental origin of the genus in contrast to our earlier sequence‐based account. Ancestral shell shapes and sizes were inferred based on landmark data using squared‐change parsimony. In a subsequent principal component analysis, only three of 20 fossil shells could be unequivocally allocated to internal nodes. However, these fossils were all Quaternary, and thus too young to infer meaningfully narrow confidence intervals for divergence estimates that probably reach back into the Miocene or even Oligocene. The apparent failure of allocating older fossils to internal nodes and achieving a reliable tree calibration was caused by the absence of phylogenetic signal in the shells of Theba due to extensive, sometimes rapid, convergent evolution, including reversals. © 2014 The Linnean Society of London     

Biodiversity,habitat and range-size aspects of a flora from a winter-rainfall desert in north-western Namaqualand,South Africa

We examined patterns of richness and endemism in relation to phylogeny (family membership), growth form and habitat in the flora of an arid (<60 mm annual rainfall), sandy coastal belt between Port Nolloth and Alexander Bay on the north-west coast of Namaqualand, South Africa. This region forms part of the species-rich Succulent Karoo biome. In an area of 750 km₂, 300 plant species were recorded with 24% endemnicity. This level of diversity and endemism is exceptionally high for a desert region. Plant distribution is strongly influenced by habitat type. Diversity, especially of endemic species, was concentrated on rocky or unusual (lag-gravel pavements) habitats. Endemics were not random phylogenetic nor biological assemblages. Dwarf leaf-succulent shrubs, especially Mesembryanthemaceae, were significantly over-represented amongst the endemic component. The results are discussed in terms of a model for plant diversification in an arid environment with relatively predictable sources of moisture.     

Multiple facets of biodiversity are threatened by mining-induced land-use change in the Brazilian Amazon

Aim

Mining is increasingly pressuring areas of critical importance for biodiversity conservation, such as the Brazilian Amazon. Biodiversity data are limited in the tropics, restricting the scope for risks to be appropriately estimated before mineral licensing decisions are made. As the distributions and range sizes of other taxa differ markedly from those of vertebrates—the common proxy for analysis of risk to biodiversity from mining—whether mining threatens lesser-studied taxonomic groups differentially at a regional scale is unclear.

Location

Brazilian Amazon.

Methods

We assess risks to several facets of biodiversity from industrial mining by comparing mining areas (within 70 km of an active mining lease) and areas unaffected by mining, employing species richness, species endemism, phylogenetic diversity and phylogenetic endemism metrics calculated for angiosperms, arthropods and vertebrates.

Results

Mining areas contained higher densities of species occurrence records than the unaffected landscape, and we accounted for this sampling bias in our analyses. None of the four biodiversity metrics differed between mining and nonmining areas for vertebrates. For arthropods, species endemism was greater in mined areas. Mined areas also had greater angiosperm species richness, phylogenetic diversity and phylogenetic endemism, although less species endemism than unmined areas.

Main Conclusions

Unlike for vertebrates, facets of angiosperm and arthropod diversity are relatively higher in areas of mining activity, underscoring the need to consider multiple taxonomic groups and biodiversity facets when assessing risk and evaluating management options for mining threats. Particularly concerning is the proximity of mining to areas supporting deep evolutionary history, which may be impossible to recover or replace. As pressures to expand mining in the Amazon grow, impact assessments with broader taxonomic reach and metric focus will be vital to conserving biodiversity in mining regions.     

Diversification in the tropical pacific: comparisons between marine and terrestrial systems and the importance of founder speciation

Patterns of distribution and processes of differentiation haveoften been contrasted between terrestrial and marine biotas.The islands of Oceania offer an excellent setting to explorethis contrast, because the geographic setting for terrestrialand shallow-water, benthic, marine organisms are the same: themyriad islands strewn across the vast Pacific. The size of speciesranges and the geographic distribution of endemism are two biogeographicattributes that are thought to differ markedly between terrestrialand marine biotas in the Pacific. While terrestrial speciesare frequently confined to single islands or archipelagoes throughoutOceania, marine species tend to have wide to very wide distributions,and are rarely restricted to single island groups except forthe most isolated archipelagoes. We explore the conditions underwhich species can reach an island by dispersal and differentiate.Genetic differentiation can occur either through founder speciationor vicariance; these processes are requisite ends of a continuum.We show that founder speciation is most likely when few propagulesenter the dispersal medium and survive well while they travelfar. We argue that conditions favorable to founder speciationare common in marine as well as terrestrial systems, and thatterrestrial-type, archipelagic-level endemism is likely commonin marine taxa. We give examples of marine groups that showarchipelagic level endemism on most Pacific island groups aswell as of terrestrial species that are widespread. Thus boththe patterns and processes of insular diversification are variable,and overlap more between land and sea than previously considered.     

Phylogenetic endemism of the orchids of Megamexico reveals complementary areas for conservation

Orchid diversity provides a unique opportunity to further our understanding of biotic and abiotic factors linked to patterns of richness, endemism, and phylogenetic endemism in many regions. However, orchid diversity is consistently threatened by illegal trade and habitat transformation. Here, we identified areas critical for orchid conservation in the biogeographic province of Megamexico. For this purpose, we evaluated orchid endemism, phylogenetic diversity, and phylogenetic endemism within Megamexico and characterized orchid life forms. Our results indicate that the majority of the regions with the highest estimates of endemism and phylogenetic endemism are in southern Mexico and northern Central America, mostly located on the Pacific side of Megamexico. Among the most important orchid lineages, several belong to epiphytic lineages such as Pleurothallidinae, Laeliinae and Oncidiinae. We also found that species from diverse and distantly related lineages converge in montane forests where suitable substrates for epiphytes abound. Furthermore, the southernmost areas of phylogenetic diversity and endemism of Megamexico are in unprotected areas. Thus, we conclude that the most critical areas for orchid conservation in Megamexico are located in southern Mexico and northern Central America. We recommend that these areas should be given priority by the Mexican system of natural protected areas as complementary conservation areas.     

Spatial patterns of phylogenetic diversity and endemism in the Western Ghats,India: A case study using ancient predatory arthropods

The Western Ghats (WG) mountain chain in peninsular India is a global biodiversity hotspot, one in which patterns of phylogenetic diversity and endemism remain to be documented across taxa. We used a well‐characterized community of ancient soil predatory arthropods from the WG to understand diversity gradients, identify hotspots of endemism and conservation importance, and highlight poorly studied areas with unique biodiversity. We compiled an occurrence dataset for 19 species of scolopendrid centipedes, which was used to predict areas of habitat suitability using bioclimatic and geomorphological variables in Maxent. We used predicted distributions and a time‐calibrated species phylogeny to calculate taxonomic and phylogenetic indices of diversity, endemism, and turnover. We observed a decreasing latitudinal gradient in taxonomic and phylogenetic diversity in the WG, which supports expectations from the latitudinal diversity gradient. The southern WG had the highest phylogenetic diversity and endemism, and was represented by lineages with long branch lengths as observed from relative phylogenetic diversity/endemism. These results indicate the persistence of lineages over evolutionary time in the southern WG and are consistent with predictions from the southern WG refuge hypothesis. The northern WG, despite having low phylogenetic diversity, had high values of phylogenetic endemism represented by distinct lineages as inferred from relative phylogenetic endemism. The distinct endemic lineages in this subregion might be adapted to life in lateritic plateaus characterized by poor soil conditions and high seasonality. Sites across an important biogeographic break, the Palghat Gap, broadly grouped separately in comparisons of species turnover along the WG. The southern WG and Nilgiris, adjoining the Palghat Gap, harbor unique centipede communities, where the causal role of climate or dispersal barriers in shaping diversity remains to be investigated. Our results highlight the need to use phylogeny and distribution data while assessing diversity and endemism patterns in the WG.     

BOTANICAL AFFINITIES OF SOME TRIPROJECTACITES FOSSIL POLLEN

During Late Cretaceous to Oligocene times, fossil pollen of the Triprojectacites group (also known as Aquilapolles or triprojectates), comprised a temporally and environmentally distinctive element of palynofloras in eastern Asia and western North America. Several species of this group serve as biostratigraphic index fossils for this interval. Using electron microscope and numerical analyses, primarily of North American triprojectate pollen, it is possible to recognize the presence of three distinct subgroups. One group, corresponding to the fossil genus Mancicorpus, has no morphologically close modern representative. Characters resembling those in Santalaceae (Santalales, Rosidae) occur in the second group, which is represented by a previously underscribed fossil triprojectate genus. The third group contains retipilate, isopolar pollen and strioreticulate, isopolar pollen. Forms exhibiting the latter morphology are commonly assigned to the genus Integricorpus, while the retipilate morphology characterizes another underscribed genus. This third group may have some phylogenetic connection to Apiaceae (Apiales, Asteridae). Pollen of some other extant families exhibits triprojectate features, although no close fossil representative can be presently identified. The triprojectate morphology is thus interpreted in modern and fossil forms as resulting from convergence rather than close phylogenetic relationships at the group level.     

Biogeography of the Lizard Genus Tropidurus Wied-Neuwied, 1825 (Squamata: Tropiduridae): Distribution,Endemism, and Area Relationships in South America

Based on comprehensive distributional records of the 23 species currently assigned to the lizard genus Tropidurus, we investigated patterns of endemism and area relationships in South America. Two biogeographic methods were applied, Parsimony Analysis of Endemicity (PAE) and Brooks Parsimony Analysis (BPA). Two areas of endemism were detected by PAE: the first within the domains of the semiarid Brazilian Caatinga, which includes seven endemic species, and the second in the region of the Serranía de Huanchaca, eastern Bolivia, in which three endemic species are present. The area cladograms recovered a close relationship between the Atlantic Forest and areas of the South American open corridor. The results revealed a close relationship among the provinces Caatinga (Cerrado, Parana Forest (Pantanal+Chaco)). The uplift of the Brazilian Central Plateau in the Late Pliocene-Early Pleistocene (4-2 Myr BP) has been interpreted as a major event responsible for isolation and differentiation of biotas along these areas. However, we emphasize that without the establishment of a temporal framework concerning the diversification history of Tropidurus it is premature to correlate cladogenetic events with specific time periods or putative vicariant scenarios. The limiting factors hampering the understanding of the biogeographic history of this genus include (1) the absence of temporal references in relation to the diversification of distinct clades within Tropidurus; (2) the lack of an appropriate taxonomic resolution of the species complexes currently represented by widely distributed forms; and (3) the need for a comprehensive phylogenetic hypothesis. We suggest that these three important aspects should be prioritized in future investigations.     

Climatic Niche Dynamics and Its Role in the Insular Endemism of <Emphasis Type="Italic">Anolis</Emphasis> Lizards

Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.     

Phylogenetic analysis of the endemic New Caledonian cockroach Lauraesilpha. Testing competing hypotheses of diversification

New Caledonia is a tropical hotspot of biodiversity with high rates of regional and local endemism. Despite offering an ideal setting to study the evolution of endemism, New Caledonia has received little attention compared with the other nearby hotspots, particularly New Zealand. Most studies of the Neocaledonian endemism have been carried out at the regional level, comparing the various groups and species present in New Caledonia but absent in neighboring territories. In addition, remarkably high short‐range endemism has been documented among plants, lizard and invertebrates, although these have usually been done, lacking a phylogenetic perspective. Most studies of Neocaledonian endemism have referred to the geological Gondwanan antiquity of the island and its metalliferous soils derived from ultramafic rocks. Very old clades are thought to have been maintained in refugia and diversified on the metalliferous soils. The present study documents the pattern of diversification and establishment of short‐range endemism in a phylogenetic context using the Neocaledonian cockroach genus Lauraesilpha. Mitochondrial and nuclear genes were sequenced to reconstruct phylogenetic relationships among the species of this genus. These relationships, in the light of the species distribution, do not support the hypothesis that species diversified via an adaptive radiation on metalliferous soils and are not consistent with areas of highest rainfall. Species of Lauraesilpha have similar altitudinal ranges and ecological habits and are short‐range endemics on mountains. What our analysis did reveal was that closely related species are found on nearby or contiguous mountains, and thus these formations probably played the key role establishing short‐range endemism (in association with recent climatic changes). © The Willi Hennig Society 2008.     

Phylogenetic endemism: a new approach for identifying geographical concentrations of evolutionary history

We present a new, broadly applicable measure of the spatial restriction of phylogenetic diversity, termed phylogenetic endemism (PE). PE combines the widely used phylogenetic diversity and weighted endemism measures to identify areas where substantial components of phylogenetic diversity are restricted. Such areas are likely to be of considerable importance for conservation. PE has a number of desirable properties not combined in previous approaches. It assesses endemism consistently, independent of taxonomic status or level, and independent of previously defined political or biological regions. The results can be directly compared between areas because they are based on equivalent spatial units. PE builds on previous phylogenetic analyses of endemism, but provides a more general solution for mapping endemism of lineages. We illustrate the broad applicability of PE using examples of Australian organisms having contrasting life histories: pea-flowered shrubs of the genus Daviesia (Fabaceae) and the Australian species of the Australo-Papuan tree frog radiation within the family Hylidae.     

Biogeography and genome size evolution of the oldest extant vascular plant genus,Equisetum (Equisetaceae)

Background and AimsExtant plant groups with a long fossil history are key elements in understanding vascular plant evolution. Horsetails (Equisetum, Equisetaceae) have a nearly continuous fossil record dating back to the Carboniferous, but their phylogenetic and biogeographic patterns are still poorly understood. We use here the most extensive phylogenetic analysis to date as a framework to evaluate their age, biogeography and genome size evolution.MethodsDNA sequences of four plastid loci were used to estimate divergence times and investigate the biogeographic history of all extant species of Equisetum. Flow cytometry was used to study genome size evolution against the framework of phylogenetic relationships in Equisetum.Key ResultsOn a well-supported phylogenetic tree including all extant Equisetum species, a molecular clock calibrated with multiple fossils places the node at which the outgroup and Equisetum diverged at 343 Mya (Early Carboniferous), with the first major split among extant species occurring 170 Mya (Middle Jurassic). These dates are older than those reported in some other recent molecular clock studies but are largely in agreement with a timeline established by fossil appearance in the geological record. Representatives of evergreen subgenus Hippochaete have much larger genome sizes than those of deciduous subgenus Equisetum, despite their shared conserved chromosome number. Subgenus Paramochaete has an intermediate genome size and maintains the same number of chromosomes.ConclusionsThe first divergences among extant members of the genus coincided with the break-up of Pangaea and the resulting more humid, warmer climate. Subsequent tectonic activity most likely involved vicariance events that led to species divergences combined with some more recent, long-distance dispersal events. We hypothesize that differences in genome size between subgenera may be related to the number of sperm flagellae.