Inferring the biogeographic origins of inter‐continental disjunct endemics using a Bayes‐DIVA approach

The arcto‐Tertiary relictual flora is comprised of many genera that occur non‐contiguously in the temperate zones of eastern Asia, Europe, eastern North America, and western North America. Within each distributional area, species are typically endemic and may thus be widely separated from closely related species within the other areas. It is widely accepted that this common pattern of distribution resulted from of the fragmentation of a once more‐continuous arcto‐Tertiary forest. The historical biogeographic events leading to the present‐day disjunction have often been investigated using a phylogenetic approach. Limitations to these previous studies have included phylogenetic uncertainty and uncertainty in ancestral range reconstructions. However, the recently described Bayes‐DIVA method handles both types of uncertainty. Thus, we used Bayes‐DIVA analysis to reconstruct the stem lineage distributions for 185 endemic lineages from 23 disjunct genera representing 17 vascular plant families. In particular, we asked whether endemic lineages within each of the four distributional areas more often evolved from (1) widespread ancestors, (2) ancestors dispersed from other areas, or (3) endemic ancestors. We also considered which of these three biogeographic mechanisms may best explain the origins of arcto‐Tertiary disjunct endemics in the neotropics. Our results show that eastern Asian endemics more often evolved from endemic ancestors compared to endemics in Europe and eastern and western North America. Present‐day endemic lineages in the latter areas more often arose from widespread ancestors. Our results also provide anecdotal evidence for the importance of dispersal in the biogeographic origins of arcto‐Tertiary species endemic in the neotropics.     

Patterns of endemism in riverine fish of the Northern Hemisphere

Loss of endemic species represents a symptom of general degrading ecosystem conditions that is the indirect result of biodiversity alteration. Here, we developed a predictive model relating species richness of endemic riverine fishes to measured biological, climatic, and historical variables using data from 118 rivers distributed all over the Northern Hemisphere. In a minimally adequate multiple general least square model, total riverine fish species richness, historical biogeography (Pleistocene glaciations), and comtemporary climate accounted for 63% of the variability in endemic species richness; the strongest correlate being riverine fish species richness. Our findings suggest that (i) endemism and richness patterns are generally similar (fish diversity "hot-spots" areas sustain higher endemic species richness); (ii) glaciation in the Pleistocene have had a significant negative influence on endemic species richness in the more septentrional areas; and (iii) certain basins situated in desertic areas (subtropical dry-zone of deserts) have unusually high numbers of endemics. These last areas should not be overshadowed when setting conservation priorities.     

Inferences of biogeographical histories within subfamily Hyacinthoideae using S-DIVA and Bayesian binary MCMC analysis implemented in RASP (Reconstruct Ancestral State in Phylogenies)

Background and Aims

Subfamily Hyacinthoideae (Hyacinthaceae) comprises more than 400 species. Members are distributed in sub-Saharan Africa, Madagascar, India, eastern Asia, the Mediterranean region and Eurasia. Hyacinthoideae, like many other plant lineages, show disjunct distribution patterns. The aim of this study was to reconstruct the biogeographical history of Hyacinthoideae based on phylogenetic analyses, to find the possible ancestral range of Hyacinthoideae and to identify factors responsible for the current disjunct distribution pattern.

Methods

Parsimony and Bayesian approaches were applied to obtain phylogenetic trees, based on sequences of the trnL-F region. Biogeographical inferences were obtained by applying statistical dispersal-vicariance analysis (S-DIVA) and Bayesian binary MCMC (BBM) analysis implemented in RASP (Reconstruct Ancestral State in Phylogenies).

Key Results

S-DIVA and BBM analyses suggest that the Hyacinthoideae clade seem to have originated in sub-Saharan Africa. Dispersal and vicariance played vital roles in creating the disjunct distribution pattern. Results also suggest an early dispersal to the Mediterranean region, and thus the northward route (from sub-Saharan Africa to Mediterranean) of dispersal is plausible for members of subfamily Hyacinthoideae.

Conclusions

Biogeographical analyses reveal that subfamily Hyacinthoideae has originated in sub-Saharan Africa. S-DIVA indicates an early dispersal event to the Mediterranean region followed by a vicariance event, which resulted in Hyacintheae and Massonieae tribes. By contrast, BBM analysis favours dispersal to the Mediterranean region, eastern Asia and Europe. Biogeographical analysis suggests that sub-Saharan Africa and the Mediterranean region have played vital roles as centres of diversification and radiation within subfamily Hyacinthoideae. In this bimodal distribution pattern, sub-Saharan Africa is the primary centre of diversity and the Mediterranean region is the secondary centre of diversity. Sub-Saharan Africa was the source area for radiation toward Madagascar, the Mediterranean region and India. Radiations occurred from the Mediterranean region to eastern Asia, Europe, western Asia and India.     

Extinct genus Lagokarpos reveals a biogeographic connection between Tibet and other regions in the Northern Hemisphere during the Paleogene

The biodiversity history on the Qinghai–Tibetan Plateau (QTP) has remained unclear for a long time. Recent paleobotanical investigations provide opportunities for revealing the phytogeographic history on the QTP in deep time and for testing phylogeographic hypotheses. Lagokarpos is an extinct genus with unknown modern affinities. It is easily distinguished by its fruits, which have two long wings and an elliptical fruit body. Previously, Lagokarpos fossils have only been found from North America and Germany, ranging from the latest Paleogene to early middle Eocene. Recently, we found fossil fruits of this genus from the Niubao Formation, near Bangoin County in the central QTP. A new species, namely Lagokarpos tibetensis H. Tang, T. Su & Z. K. Zhou sp. nov., is described. This is the first fossil record of the genus in Asia. The occurrence of L. tibetensis indicates a close floristic linkage between the QTP and other floras in the Northern Hemisphere during the Paleogene. According to the floristic assemblages, we suggest the central QTP experienced a tropical or subtropical humid climate during that period.     

Bat metacommunity structure on Caribbean islands and the role of endemics

Aim We evaluate characteristics of species ranges (i.e. coherence, species turnover and range boundary clumping) to determine the structure of bat metacommunities and metaensembles from Caribbean islands. We evaluate the effects of endemic species on that structure, and quantify associations between island characteristics and latent environmental gradients that structure these metacommunities and metaensembles. Location Sixty‐five Caribbean islands throughout the Bahamas, Greater Antilles and Lesser Antilles. Methods Metacommunity structure is an emergent property of a set of ecological communities at different sites defined by species distributions across geographic or environmental gradients. We analysed elements of metacommunity structure (coherence, range turnover and range boundary clumping) to determine the best‐fit pattern for metacommunities from all Caribbean islands, as well as from the Bahamas, the Greater Antilles and the Lesser Antilles separately. For each island group, analyses were conducted for all bats and for each of two broadly defined guilds (i.e. carnivores and herbivores). In addition, analyses were conducted for all species and for a subset in which endemic species were removed from the fauna. Spearman rank correlations identified island characteristics (area, elevation, latitude, longitude) that were associated significantly with island scores for ordination axes based on reciprocal averaging. Results Metacommunity structure for all bats and for carnivores was similar for each island group, with Clementsian distributions (i.e. discrete communities with groups of species replacing other groups of species along the gradient) for all islands, the Bahamas and the Lesser Antilles, but with nested distributions for the Greater Antilles. Herbivore distributions were random for the Bahamas, but were Clementsian for all other island groups. Removal of endemic species affected the best‐fit model of metacommunity structure in only 3 of 12 cases. In general, ordination scores for islands were correlated with longitude or latitude, but not with island area or elevation. Main conclusions Characteristics of bat species ranges and associated metacommunity structure were primarily dependent on the number and geographic arrangement of primary sources of colonization, and not on interspecific interactions, species‐specific levels of environmental tolerance, or the physical characteristics of islands. Endemic species did not greatly affect metacommunity structure in Caribbean bats.     

Application of the unified species concept reveals distinct lineages for disjunct endemics of the Brassica repanda (Brassicaceae) complex

Species delimitation is an important issue in terms of conservation priorities, especially for narrow endemics under threat of extinction. The Alpine endemics Brassica repanda subsp. glabrescens and subsp. baldensis belong to a highly polymorphic species complex, although their disjunct distribution suggests favourable conditions for independent evolution. In the present study, we applied the unified species concept to test whether the endemics form distinct evolutionary lineages, both from one another and also from the remainder of the complex. Compliance with the criteria of monophyly, diagnosability, and genotypic clustering was examined, primarily by making use of amplified fragment length polymorphism data. Both endemics were indicated as monophyletic by phylogenetic analyses, and diagnostic characters were found for both taxa. Population structure analyses showed clear genetic discontinuity for each of the endemics, with little admixture among the clusters. This evidence indicates that the endemics have acquired multiple properties that satisfy each of the species criteria considered. Hence, we suggest the taxonomic recognition of B. baldensis and B. glabrescens as separate species. Comparative population genetics analyses show the lack of marked genetic structuring within either taxon, as well as low levels of heterozygosity. Conclusions on the status of threat and on recommended conservation actions are drawn. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 482–497.     

Phylogenetic analysis of Toxicodendron （Anacardiaceae） and its biogeographic implications on the evolution of north temperate and tropical intercontinental disjunctions

Toxicodendron is a genus in the Rhus complex of Anacardiaceae with a disjunct distribution between eastern Asia and North America, extending to southeastern Asia and the neotropics. Nuclear （internal transcribed spacer, external transcribed spacer, and NIA-i3） and chloroplast （ndhF and trnL-F） sequences were used to construct phylogenetic relationships of Toxicodendron. Phylogenetic analysis of these data strongly support Toxieodendron as a monophyletic group distinct from other genera of the Rhus complex, and the phylogeny does not fully corroborate classification at the sectional level. Two temperate disjunct lineages were detected, one from section Toxicodendron and the other between the eastern North American Toxicodendron vernix and the eastern Asian Toxicodendron vernieifluum. Their divergence times were estimated to be 13.46 （7.95-19.42） and 7.53 （2.76-12.86） mya, respectively. The disjunction between section Griffithii （taxa from warm temperate to tropical Asia） and Toxieodendron striatum （from the neotropics） was supported and their divergence time was estimated to be 20.84 （11.1 6-30.52） mya in the early Miocene. Our biogeographic results and the paleontological data support the Bering land bridge as the most likely route to explain the temperate disjunctions, yet the tropical disjunction in Toxicodendron seems to be best explained by the North Atlantic land bridge hypothesis.     

Intercontinental disjunctions between eastern Asia and western North America in vascular plants highlight the biogeographic importance of the Bering land bridge from late Cretaceous to Neogene

This review shows a close biogeographic connection between eastern Asia and western North America from the late Cretaceous to the late Neogene in major lineages of vascular plants (flowering plants, gymnosperms, ferns and lycophytes). Of the eastern Asian–North American disjuncts, conifers exhibit a high proportion of disjuncts between eastern Asia and western North America. Several lineages of ferns also show a recent disjunct pattern in the two areas. In flowering plants, the pattern is commonly shown in temperate elements between northeastern Asia and northwestern North America, as well as elements of the relict boreotropical and Neogene mesophytic and coniferous floras. The many cases of intercontinental biogeographic disjunctions between eastern Asia and western North America in plants supported by recent phylogenetic analyses highlight the importance of the Bering land bridge and/or the plant migrations across the Beringian region from the late Cretaceous to the late Neogene, especially during the Miocene. The Beringian region has permitted the filtering and migration of certain plant taxa since the Pliocene after the opening of the Bering Strait, as many conspecific taxa or closely related species occur on both sides of Beringia.     

Crossing the (Wallace) line: local abundance and distribution of mammals across biogeographic barriers

Past and ongoing vertebrate introductions threaten to rearrange ecological communities in the Indo‐Malay Archipelago, one of Earth's most biodiverse regions. But the consequences of these translocations are difficult to predict. We compared local abundance and distributions in four tropical mammal lineages that have crossed from Asia to Wallacea or New Guinea. The local abundance of macaques (Macaca spp.), which naturally crossed Wallace's Line, was higher in Sulawesi (east of the line; mean = 3.7 individuals per camera station, 95% CI = 2.2: 5.1) than in Borneo (west of the line; mean = 1.1, CI = 0.8: 1.4), but the local abundance of Malay civets (Viverra tangalunga), Rusa deer, and Sus pigs was similar in their native ranges and where they had been introduced by humans east of Wallace's Line. Proximity to rivers increased Malay Civet local abundance and decreased the local abundance of pigs in parts of their introduced ranges (Maluku and New Guinea, respectively), while having no effect on local abundance in their native ranges (Borneo) or other areas where they have been introduced (Sulawesi). That local abundance was higher east of Wallace's Line in just one of four mammal lineages is consistent with findings from plant invasions, where most species have similar abundance in their native and introduced ranges. However, species’ ecology may change as they enter new communities, for example, their patterns of abundance at local scales. This could make it difficult to predict community structure in the face of ongoing species introductions.     

Phylogeny of slug species of the genus Arion: evidence of monophyly of Iberian endemics and of the existence of relict species in Pyrenean refuges

The Iberian Peninsula contains the majority of the Paleartic land slug species of the genus Arion, which exhibits diverse taxonomic problems. The present study investigated Arion taxonomy on the basis of analyses of the mitochondrial ND1 gene and nuclear internal transcribed spacer 1 (ITS1) sequences. The Iberian endemic species were monophyletically clustered in two divergent sister clades. The topotype specimens of Arion lusitanicus and the closely related species Arion nobrei and Arion fuligineus, as well as Arion hispanicus and Arion flagellus, were grouped into an ‘Atlantic’ clade, whereas Arion baeticus, Arion gilvus, Arion anguloi, Arion wiktori and Arion paularensis were included in a ‘Continental–Mediterranean’ clade. Calibration of mutation rate in the ND1 gene suggested that the divergence of these two clades occurred around the Pliocene–Pleistocene boundary, with subsequent speciation events during the Pleistocene. A group of ancestral and divergent endemic species with distribution centred in the Pyrenean mountain range (Arion molinae, Arion lizarrusti, Arion antrhacius and Arion iratii) arose in the Pliocene and survived through the Pleistocene in geographically confined small populations. Arion lusitanicus showed up to be polyphyletic: specimens, sampled outside the geographic range of the topotype in the north‐western Iberian Peninsula, were included in a non‐monophyletic clade together with the widely distributed species Arion ater and Arion rufus. The divergent species with a wide European distribution (Arion subfuscus, Arion hortensis, Arion fagophilus and Arion intermedius) were located in basal positions in all topologies. The evolutionary history of these slug species (highly sensitive to climatic factors, with capacity for both outcrossing and selfing, and with low dispersal ability) appears to have been moulded by Pliocene–Pleistocene climate events and by the rugged topography of southern Europe, giving rise to repeated cycles of population isolation during periods of glaciation alternating with interglacial expansions limited by geographic barriers.     

The Branchiopoda (Crustacea: Anomopoda, Ctenopoda and Cyclestherida) of the rain forests of Cameroon, West Africa: low abundances, few endemics and a boreal–tropical disjunction

Aim We provide the first in‐depth study of the Branchiopoda of the rain forests of Cameroon and also of the African continent. Location Surface water environments, Cameroon. Methods Qualitative plankton samples were collected in all types of surface water environments present, ranging from big lakes to water collected in rock crevices or fallen fruit cavities. A tow or hand‐held plankton net of mesh size 100 μm was used, and water volumes filtered were at least several m³ in large water bodies, or half to whole water volume in small water bodies. Results We recorded 61 species (53 first records for the country), based on 700+ samples collected between September 1998 and March 2002. Anomopoda (92%) was the dominant order, followed by Ctenopoda (6.5%) and Cyclestherida (1.5%). Chydoridae (67%) was the most speciose family followed by Macrothricidae (6.5%) and Daphniidae (5%). Alona (11%) was the dominant genus followed by Chydorus (10%) and Pleuroxus (8%). Several species of Chydorinae, especially of the genus Pleuroxus, are shared with continental Eurasia–North America, but are absent from the Mediterranean and desert–steppe–savanna zones of Africa (boreal–tropical disjunction). Daphnia was absent, as in most tropical lowlands. No single species was really abundant, and a majority were rare to very rare, and of restricted occurrence within the rain forest patches. Comparing Africa, South America and Southeast Asia, we found a current total of 196 species for the combined rain forest areas, out of a world total of 500+ species. Systematic trends in richness at three taxonomic levels were the same for all continents: Anomopoda–Ctenopoda–Cyclestherida at ordinal level, Chydoridae–Daphniidae–Macrothricidae–Sididae at family level and Alona–Chydorus–Macrothrix–Diaphanosoma at genus level. Southeast Asia was richest (111 species, 14 endemics) with South America a close second (110 species, 27 endemics). Africa was the most species‐poor (95 species, of which only 5 are endemics). Main conclusions We hypothesize that the post‐Miocene cooling and aridization of the world climate hit the freshwater biota of Africa particularly hard, with more extinction here than elsewhere, and little recolonization. Most extinction occurred in the savanna‐desert belt, and eight disjunct boreal species (four Pleuroxus, Picripleuroxus laevis, Kurzia latissima, Alonella exigua, and Monospilus dispar) survive morphologically unchanged since pre‐Pleistocene times in the Cameroon rain forest. Slow evolution thus appears typical of these cyclic parthenogenetic branchiopods in which sexual recombination occurs only at intervals. Illustrative of the same slow evolution is the fact that the two endemic cladocerans of Cameroon (Nicsmirnovius camerounensis and Bryospilus africanus) belong to tropicopolitan genera of Gondwanan age.     

Evolutionary genetics and biogeographic structure of Rhizobium gallicum sensu lato, a widely distributed bacterial symbiont of diverse legumes

We used phylogenetic and population genetics approaches to evaluate the importance of the evolutionary forces on shaping the genetic structure of Rhizobium gallicum and related species. We analysed 54 strains from several populations distributed in the Northern Hemisphere, using nucleotide sequences of three 'core' chromosomal genes (rrs, glnII and atpD) and two 'auxiliary' symbiotic genes (nifH and nodB) to elucidate the biogeographic history of the species and symbiotic ecotypes (biovarieties) within species. The analyses revealed that strains classified as Rhizobium mongolense and Rhizobium yanglingense belong to the chromosomal evolutionary lineage of R. gallicum and harbour symbiotic genes corresponding to a new biovar; we propose their reclassification as R. gallicum bv. orientale. The comparison of the chromosomal and symbiotic genes revealed evidence of lateral transfer of symbiotic information within and across species. Genetic differentiation analyses based on the chromosomal protein-coding genes revealed a biogeographic pattern with three main populations, whereas the 16S rDNA sequences did not resolve that biogeographic pattern. Both the phylogenetic and population genetic analyses showed evidence of recombination at the rrs locus. We discuss our results in the light of the contrasting views of bacterial species expressed by microbial taxonomist and evolutionary biologists.     

Protecting a single endangered species and meeting multiple conservation goals: an approach with Guaiacum sanctum in Yucatan Peninsula,Mexico

Aim New protected areas should consider safeguarding high conservation value sites based on multiple criteria and not just the presence of a single endangered or charismatic species. However, the extent to which complementary criteria coincide is usually unknown. We use the case of Guaiacum sanctum (Zygopyllaceae), an endangered timber tree species, to explore whether the protection of forests where this species is most abundant would meet other complementary conservation goals, such as capturing regional plant biodiversity, protecting other threatened/endemic species or safeguarding ecosystem services. Location Yucatan Peninsula, southern Mexico. Methods We conducted an analysis of the structure, composition and diversity of tree communities (including stems ≥5 cm dbh) at eight G. sanctum forest sites. We identified endemic and threatened tree species and quantified above‐ground tree biomass and carbon storage in these G. sanctum forests. Results Guaiacum sanctum forests contain 35–59 tree species on plots as small as 1000 m². The species composition of tree communities changed rapidly (high β‐diversity) across soil boundaries and rainfall regimes. Twenty‐one endemic and eight threatened tree species were recorded in our inventories. Individuals of G. sanctum represented up to 55% of the above‐ground carbon for trees ≥5 cm dbh. The high basal area of G. sanctum forests plus the high wood density, abundance, large size and longevity (more than 500 years) of G. sanctum and other tree species enhance the potential importance of these forests for carbon storage. Main conclusions A conservation strategy focused on protecting important populations of G. sanctum in the Yucatan Peninsula would have significant co‐benefits for conservation of regional tree species biodiversity and provision of critical ecosystem services. Our study illustrates a multiple criteria approach useful for the selection of areas with high conservation value on the basis of endemic, threatened species, species richness and ecosystem services.