The hypothesis of sympatric speciation as the dominant generator of endemism in a global hotspot of biodiversity

Allopatric or sympatric speciation influence the degree to which closely related species coexist in different manners, altering the patterns of phylogenetic structure and turnover among and between communities. The objective of this study was to examine whether phylogenetic community structure and turnover in the Brazilian Atlantic Forest permit conclusions about the dominant process for the formation of extant angiosperm richness of tree species. Therefore, we analyzed phylogenetic community structure (MPD, MNTD) as well as taxonomic (Jaccard similarity) and phylogenetic turnover (betaMPD, betaMNTD) among and between 49 tree communities distributed among three different habitat types. Mean annual precipitation and mean annual temperature in each survey area were estimated. Phylogenetic community structure does not differ between habitat types, although MPD reduces with mean annual temperature. Jaccard similarity decreases and betaMNTD increases with spatial distance and environmental differences between study sites. Spatial distance explains the largest portions of variance in the data, indicating dispersal limitation and the spatial aggregation of recently formed taxa, as betaMNTD is related to more recent evolutionary events. betaMPD, that is related to deep evolutionary splits, shows no spatial or environmental pattern, indicating that older clades are equally distributed across the Brazilian Atlantic Forest. While similarity pattern indicates dispersal limitations, the spatial turnover of betaMNTD is consistent with a high degree of sympatric speciation generating extant diversity and endemism in the Brazilian Atlantic Forest. More comprehensive approaches are necessary to reduce spatial sampling bias, uncertainties regarding angiosperm diversification patterns and confirm sympatric speciation as the dominant generator for the formation of extant species diversity in the Brazilian Atlantic Forest.     

Global dispersal and diversification of the genus Schoenus (Cyperaceae) from the Western Australian biodiversity hotspot

The predominantly austral genus Schoenus L. is the largest genus in tribe Schoeneae and one of the ten most species-rich Cyperaceae genera, with over 150 accepted species found mostly in Australia, New Zealand, southeast Asia, and southern Africa. Here, we use data based on two nuclear and three plastid DNA regions to present one of the most comprehensive phylogenetic reconstructions of a genus in Cyperaceae to date, covering over 70% of described species of Schoenus. After recent taxonomic realignments in the last 4 years have both added and removed species from the genus, we show that Schoenus is now monophyletic. In addition, our results indicate that Schoenus originated in Western Australia in the Paleocene and eventually dispersed to surrounding continents, but rarely back. The diversification rate of the genus appears to have slightly decreased over time, and there has not been an increase associated with the establishment of the Cape clade endemic to the sclerophyllous fynbos vegetation type, such as has been reported in other plant lineages endemic to the Cape region. These results will serve as a template to understanding the complex patterns of genome size evolution and to untangle drivers of diversification in this genus.     

Limited dispersal and local adaptation promote allopatric speciation in a biodiversity hotspot

Recently diverged or diverging populations can offer unobstructed insights into early barriers to gene flow during the initial stages of speciation. The current study utilised a novel insect system (order Mantophasmatodea) to shed light on the early drivers of speciation. The members of this group have limited dispersal abilities, small allopatric distributions and strong habitat associations in the Cape Floristic Region biodiversity hotspot in South Africa. Sister taxa from the diverse family Austrophasmatidae were chosen as focal species (Karoophasma biedouwense, K. botterkloofense). Population genetics and Generalized Dissimilarity Modelling (GDM) were used to characterise spatial patterns of genetic variation and evaluate the contribution of environmental factors to population divergence and speciation. Extensive sampling confirmed the suspected allopatry of these taxa. However, hybrids were identified in a narrow region occurring between the species' distributions. Strong population structure was found over short geographic distances; particularly in K. biedouwense in which geographic distance accounted for 32% of genetic variation over a scale of 50 km (r = .56, p < .001). GDM explained 42%–78% of the deviance in observed genetic dissimilarities. Geographic distance was consistently indicated to be important for between species and within population differentiation, suggesting that limited dispersal ability may be an important neutral driver of divergence. Temperature, altitude, precipitation and vegetation were also indicated as important factors, suggesting the possible role of adaptation to local environmental conditions for species divergence. The discovery of the hybrid-zone, and the multiple allopatric species pairs in Austrophasmatidae support the idea that this could be a promising group to further our understanding of speciation modes.     

Diversity, endemism and species turnover of millipedes within the south-western Australian global biodiversity hotspot

Aim To examine how current and historical environmental gradients affect patterns of millipede (Diplopoda) endemism and species turnover in a global hotspot of floristic diversity, and to identify regions of high endemism and taxonomic distinctness for conservation management. Location South‐western Australia. Methods Museum database records of millipedes (subclasses Pentazonia and Helminthomorpha), supplemented with extensive fieldwork, were used to map species richness, species turnover (β‐diversity), weighted endemism, average taxonomic distinctness and variation in taxonomic distinctness in half‐degree grid squares (c. 2500 km²). Generalized linear models were used to examine relationships between these parameters with rainfall (present day and historical), topography and human disturbance (clearing for agriculture and urbanization). Results Millipede species richness, particularly within the order Spirostreptida, and millipede endemism were positively associated with large within‐cell differences in elevation (mountainous regions). Large variation in taxonomic distinctness (unevenness in the taxonomic tree) in higher‐rainfall areas was mainly due to speciation within the Spirostreptida genus Atelomastix. Hotspots of millipede endemism and taxonomic distinctness were identified within three categories of importance: primary (Stirling Range East, Cape Le Grand, Cape Arid, Walpole, Porongurups), secondary (Mount Manypeaks, Bremer Bay, Stirling Range West, Duke of Orleans Bay, Ravensthorpe, Albany, Busselton) and tertiary (Nornalup). A species turnover boundary was positively associated with rainfall, broadly located in the transition zone of 300–600 mm year^?1. Main conclusions The current lack of knowledge on the endemism of invertebrates hampers their incorporation into conservation planning. With this knowledge we can identify global biodiversity hotspots and, at a smaller scale, significant conservation areas within a region. Here we have shown that weighted endemism and taxonomic distinctness are useful tools in identifying centres of high endemism and speciation for millipedes within the south‐west Australian hotspot. Moreover, it is unlikely that either vertebrates or vascular plants will be useful surrogates for identifying significant areas for invertebrate conservation. While other workers have shown that vascular plants, mammals and frogs have different centres of endemism within south‐west Australia, our results show that centres of endemism for millipedes encompass all of these plus other areas.     

Unexpected diversity in the catfish Pseudancistrus brevispinis reveals dispersal routes in a Neotropical center of endemism: the Guyanas Region

Neotropical freshwater fishes have reached an unrivalled diversity, organized into several areas of endemism, yet the underlying processes are still largely unknown. The topographical and ecological characteristics of the Guyanas Region make it an ideal area of endemism in which to investigate the forces that have shaped this great diversity. This region is thought to be inhabited by species descending from Amazonian ancestors, which would have used two documented routes that, however, hardly explain the entrance of species adapted to running waters. Here, we investigate the evolutionary history of Pseudancistrus brevispinis , a catfish endemic to this region and exclusively found in running waters, thus making it an ideal model for investigating colonization routes and dispersal in such habitats. Our analyses, based on mitochondrial and nuclear markers, revealed an unexpected diversity consisting of six monophyletic lineages within P. brevispinis , showing a disjoint distribution pattern. The lineages endemic to Guyanas coastal rivers form a monophyletic group that originated via an ancestral colonization event from the Amazon basin. Evidence given favours a colonization pathway through river capture between an Amazonian tributary and the Upper Maroni River. Population genetic analyses of the most widespread species indicate that subsequent dispersal among Guyanas coastal rivers occurred principally by temporary connections between adjacent rivers during periods of lower sea level, yet instances of dispersal via interbasin river captures are not excluded. During high sea level intervals, the isolated populations would have diverged leading to the observed allopatric species. This evolutionary process is named the sea level fluctuation (SLF) hypothesis of diversification.     

Toad egg-jelly as a source of divalent cations essential for fertilization

Dejellied uterine eggs of the toad Bufo bufo japonicus are not fertilizable in 1/20 De Boer's solution (1/20 DB), but are fertilized when inseminated in a uv-solubilized jelly (UVJ) or the dialyzate of UVJ (UVJD). The present study was carried out to define this fertilization-supporting activity of egg-jelly. Dejellied eggs were fertilized in a high frequency when inseminated in a medium containing the ashes obtained by heating UVJD at 600 degrees C for 16 hr. Similarly, a reconstituted salt solution (RSS), which mimics the ionic composition of UVJD, supported a high rate of fertilization. To be effective in fertilization, however, RSS had to be present at the time of insemination. Analyses of individual salts revealed that dejellied eggs are successfully fertilized in CaCl2 and/or MgCl2 at 1-5 mM, only slightly in KCl at 10 mM, but not at all in NaCl at any of the concentrations tested. The activity of UVJD was lost reversibly when divalent cations were chelated by EDTA. The fertilization of dejellied eggs is therefore possible in a medium without any organic components of egg-jelly, provided that 2-5 mM Ca2+ or Mg2+ is present. Sperm were motile in media containing cations below 20-25 mM, regardless of the ionic composition. The egg-jelly possessed cations in a concentration of about 130 mM, but most ions were lost from intact jelly on immersion of eggs in water for 2-3 min, accompanied by the acquisition of fertilizability by sperm. Examination of the behavior of salts on dialysis or gel-filtration of jelly molecules revealed that the jelly retains Ca2+ and Mg2+, and possibly K+ as well, but not Na+ and Cl-. We propose that toad egg-jelly plays a function in fertilization by retaining Ca2+ and/or Mg2+ around each egg at the level necessary for successful sperm entrance into the egg.     

Marine caves of the Mediterranean Sea: a sponge biodiversity reservoir within a biodiversity hotspot

Marine caves are widely acknowledged for their unique biodiversity and constitute a typical feature of the Mediterranean coastline. Herein an attempt was made to evaluate the ecological significance of this particular ecosystem in the Mediterranean Sea, which is considered a biodiversity hotspot. This was accomplished by using Porifera, which dominate the rocky sublittoral substrata, as a reference group in a meta-analytical approach, combining primary research data from the Aegean Sea (eastern Mediterranean) with data derived from the literature. In total 311 species from all poriferan classes were recorded, representing 45.7% of the Mediterranean Porifera. Demospongiae and Homoscleromorpha are highly represented in marine caves at the family (88%), generic (70%), and species level (47.5%), the latter being the most favored group along with Dictyoceratida and Lithistida. Several rare and cave-exclusive species were reported from only one or few caves, indicating the fragmentation and peculiarity of this unique ecosystem. Species richness and phylogenetic diversity varied among Mediterranean areas; the former was positively correlated with research effort, being higher in the northern Mediterranean, while the latter was generally higher in caves than in the overall sponge assemblages of each area. Resemblance analysis among areas revealed that cavernicolous sponge assemblages followed a pattern quite similar to that of the overall Mediterranean assemblages. The same pattern was exhibited by the zoogeographic affinities of cave sponges: species with Atlanto-Mediterranean distribution and Mediterranean endemics prevailed (more than 40% each), 70% of them having warm-water affinities, since most caves were studied in shallow waters. According to our findings, Mediterranean marine caves appear to be important sponge biodiversity reservoirs of high representativeness and great scientific interest, deserving further detailed study and protection.     

The building of a biodiversity hotspot across a land-bridge in the Mediterranean

Many of the macroevolutionary processes that have shaped present-day phylogenetic patterns were caused by geological events such as plate tectonics and temporary land-bridges. The study of spatial patterns of phylogenetic diversity can provide insights into these past events. Here we focus on a western Mediterranean biodiversity hotspot located in the southern Iberian Peninsula and northwest Africa, two regions that are separated by the Strait of Gibraltar. We explore the spatial structure of the phylogenetic relationships within and across large-scale plant assemblages. Significant turnover in terminal lineages tends to occur between landmasses, whereas turnover in deep lineages tends to occur within landmasses. Plant assemblages in the western ecoregions of this hotspot tend to be phylogenetically overdispersed but are phylogenetically clustered on its eastern margins. We discuss our results in the light of potential scenarios of niche evolution (or conservatism) and lineage diversification. The significant turnover between landmasses suggests a common scenario of allopatric speciation that could have been facilitated by the intermittent joining of the two continents. This may have constituted an important stimulus for diversification and the emergence of this western Mediterranean biodiversity hotspot.     

Argentine ants displace floral arthropods in a biodiversity hotspot

Argentine ant (Linepithema humile (Mayr)) invasions are often associated with the displacement of ground‐dwelling arthropods. Argentine ant invasions can also exert other effects on the community through interactions with plants and their associated arthropods. For example, carbohydrate resources (e.g. floral or extrafloral nectar) may influence foraging behaviour and interactions among ants and other arthropods. In South Africa's Cape Floristic Region, Argentine ants and some native ant species are attracted to the floral nectar of Leucospermum conocarpodendron Rourke (Proteaceae), a native tree that also has extrafloral nectaries (EFNs). Despite having relatively low abundance in pitfall traps, Argentine ants visited inflorescences more frequently and in higher abundance than the most frequently observed native ants, Camponotus spp., though neither native nor Argentine ant floral foraging was influenced by the EFNs. Non‐metric multidimensional scaling revealed significant dissimilarity in arthropod communities on inflorescences with Argentine ants compared to inflorescences with native or no ants, with Coleoptera, Diptera, Hymenoptera, Arachnida, Orthoptera, and Blattaria all being underrepresented in inflorescences with Argentine ants compared to ant‐excluded inflorescences. Native honeybees (Apis mellifera capensis Eschscholtz) spent 75% less time foraging on inflorescences with Argentine ants than on inflorescences without ants. Neither Argentine ant nor native ant visits to inflorescences had a detectable effect on seed set of Le. conocarpodendron. However, a pollen supplementation experiment revealed that like many other proteas, Le. conocarpodendron is not pollen‐limited. Flower predation was negatively associated with increased ant visit frequency to the inflorescences, but did not differ among inflorescences visited by native and Argentine ants. Displacement of arthropods appears to be a consistent consequence of Argentine ant invasions. The displacement of floral arthropods by Argentine ants may have far‐reaching consequences for this biodiversity hotspot and other regions that are rich in insect‐pollinated plants.     

Beta diversity of orchid bees in a tropical biodiversity hotspot

A controversial issue in ecology and conservation is whether community composition is controlled by niche or dispersal assembly. We assessed the importance of climatic factors and geographic gradients on the distribution of orchid bees in a severely-fragmented and species-rich tropical forest region in Brazil. Orchid-bee males were attracted to 17 different scent baits and collected in 15 forest sites. In total we captured 11,081 bees from 40 species. Climatic variables explained twice as much of the observed variation in the bee species data set as did pure spatial variation. However, most of the climatic explained variation was spatially structured, indicating that the species and the climatic data have a similar spatial arrangement. In fact, part of the observed latitudinal changes in community composition appears to be explained by a concomitant gradient in precipitation seasonality. Similarly, reduced temperatures and a more seasonal precipitation may help to explain the relative distinctiveness of the fauna from some of the westernmost sites. The level of similarity among the sampled sites, although highly variable, decayed both as function of the climatic and geographic distances among these sites. The greatest pairwise dissimilarities in the composition of the orchid-bee fauna were observed among sites 200–300 km from each other, since in many case those sites were more dissimilar in terms of climate than those further apart. It is suggested that global warming and consequent altered climatic regimes will influence the distribution patterns of orchid bees in a region already threatened by deforestation and forest fragmentation.     

Probing into the diversity of trypanosomatid flagellates parasitizing insect hosts in South-West China reveals both endemism and global dispersal

Flagellates of the class Kinetoplastea are known to frequently parasitize insects. We have collected 67 isolates from 407 Heteroptera hosts captured in several locations of South-West China. Their splice leader (SL) RNA gene repeats and small subunit (SSU) rRNA genes were PCR amplified from the infected tissue samples. In most cases, parasites were found in the midgut, rarely the infection was confined to the Malpighian tubes. Phylogenetic analysis of the obtained sequences has significantly expanded the known diversity of these monoxenous parasites. Fifteen typing units were found among these isolates including 11 potentially new species. Four typing units matched the previously known typing units from the Neotropics indicating a global distribution of the respective parasite species. At the same time, new clades appeared, testifying for a certain level of endemism. The host record of the parasites found indicated a variable specificity level of the host–parasite association including several cases of a very broad host range. Our results disprove the “one host – one parasite” paradigm and show that although the global diversity of monoxenous parasites is high, it is not as enormous as suggested earlier. Moreover, phylogenetic analysis revealed the presence, among the isolated strains, of a new Phytomonas species, which is the first documentation of this potentially pathogenic dixenous parasite of plants in China.     

Pervasive defaunation of forest remnants in a tropical biodiversity hotspot

Tropical deforestation and forest fragmentation are among the most important biodiversity conservation issues worldwide, yet local extinctions of millions of animal and plant populations stranded in unprotected forest remnants remain poorly explained. Here, we report unprecedented rates of local extinctions of medium to large-bodied mammals in one of the world's most important tropical biodiversity hotspots. We scrutinized 8,846 person-years of local knowledge to derive patch occupancy data for 18 mammal species within 196 forest patches across a 252,669-km(2) study region of the Brazilian Atlantic Forest. We uncovered a staggering rate of local extinctions in the mammal fauna, with only 767 from a possible 3,528 populations still persisting. On average, forest patches retained 3.9 out of 18 potential species occupancies, and geographic ranges had contracted to 0-14.4% of their former distributions, including five large-bodied species that had been extirpated at a regional scale. Forest fragments were highly accessible to hunters and exposed to edge effects and fires, thereby severely diminishing the predictive power of species-area relationships, with the power model explaining only ～9% of the variation in species richness per patch. Hence, conventional species-area curves provided over-optimistic estimates of species persistence in that most forest fragments had lost species at a much faster rate than predicted by habitat loss alone.     

Tree species diversity and community composition in a human-dominated tropical forest of Western Ghats biodiversity hotspot,India

The structure, function, and ecosystem services of tropical forest depend on its species richness, diversity, dominance, and the patterns of changes in the assemblages of tree populations over time. Long-term data from permanent vegetation plots have yielded a wealth of data on the species diversity and dynamics of tree populations, but such studies have only rarely been undertaken in tropical forest landscapes that support large human populations. Thus, anthropogenic drivers and their impacts on species diversity and community structure of tropical forests are not well understood. Here we present data on species diversity, community composition, and regeneration status of tropical forests in a human-dominated landscape in the Western Ghats of southern India. Enumeration of 40 plots (50 m × 20 m) results a total of 106 species of trees, 76 species of saplings and 79 species of seedlings. Detrended Correspondence Analysis ordination of the tree populations yielded five dominant groups, along disturbance and altitudinal gradients on the first and second axes respectively. Abundant species of the area such as Albizia amara, Nothopegia racemosa and Pleiospermum alatum had relatively few individuals in recruiting size classes. Our data indicate probable replacement of rare, localized, and old-growth ‘specialists’ by disturbance-adapted generalists, if the degradation is continuing at the present scale.     

Conservation of invertebrate biodiversity on a mountain in a global biodiversity hotspot,Cape Floral Region

Mountains present particular challenges for biodiversity conservation. Table Mountain is a significant mountain in a global biodiversity hotspot, the Cape Floristic Region. It has outstanding angiosperm diversity and endemism. Yet, aerial and foliage invertebrates in the area have been poorly studied, despite their importance as pollinators and predators. These plant and invertebrate assemblages are under great pressure from human disturbance. Aerial and foliage invertebrates were sampled with a range of techniques. Sites were chosen to make comparisons between vegetation structure and type, elevation and aspect. In total, 216 species from 63 families and 14 orders were recorded. Vegetation structure (fynbos or forest) and elevation were the most important environmental variables for both aerial and foliage invertebrates. Peak time for aerial invertebrate abundance was spring and summer in the fynbos and spring in the forests, while the foliage invertebrates showed very little seasonal variation. There was no correlation between the diversity of aerial and foliage invertebrates. When these results were compared with others on epigaeic invertebrates, it became clear that epigaeic and aerial invertebrates are not correlated, while epigaeic and foliage invertebrates were only partially correlated, but not sufficiently so to consider one as a reliable estimator of the other. The management pointer from this study is that sites at all elevations are vital for the conservation of biodiversity on Table Mountain. Both the aerial and epigaeic/foliage invertebrate assemblages will need to be monitored separately to maintain the mountain’s conservation status.     

Phylogeographic evidence for two mesic refugia in a biodiversity hotspot

Phylogeographic studies of flora in species-rich south-western Australia point to complex evolutionary histories, reflecting patterns of persistence and resilience to climatic changes during the Pleistocene. We asked whether coastal areas of the mid-west and south, as well as granite outcrops and inland ranges, have acted as major refugia within this region during Pleistocene climatic fluctuations by analysing phylogeographic patterns in the shrub Calothamnus quadrifidus R.Br. (Myrtaceae). We determined variation in chloroplast DNA data for 41 populations across the geographic range. Relationships and major clades were resolved using parsimony and Bayesian analyses. We tested for demographic and spatial expansion of the major clades and estimated clade divergence dates using an uncorrelated, lognormal relaxed clock based on two conservative chloroplast mutation rates. Two distinct phylogeographic clades were identified showing divergence during the Pleistocene, consistent with other phylogeographic studies of south-west Australian flora, emphasising the impact of climatic oscillations in driving divergence in this landscape. The southern clade was more diverse, having higher haplotype diversity and greater genetic structure, while the northern clade showed evidence of fluctuation in population size. Regions of high haplotype diversity with adjacent areas of low diversity observed in each clade indicated the locations of two coastal refugia: one on the south coast and another along the mid-west coast. This is the first evidence for major Pleistocene refugia using chloroplast genetic data in a common, widespread species from this region.