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.     

Floristic patterns of epiphytes in the Brazilian Atlantic Forest,a biodiversity hotspot

To assess the main factors driving epiphytic angiosperm distribution throughout the Brazilian Atlantic Forest, we compiled 57 floristic surveys and analysed species composition under the influence of environmental variables, space and vegetation type using canonical correspondence analysis (CCA), similarity (Sorensen) and Mantel's tests. The indicator value index (IndVal) was used to find indicator species of each Brazilian Atlantic Forest vegetation type. Group sharpness analysis was performed in order to determine the appropriate group partition level. CCA showed a separation of the epiphytic flora reflecting temperature and rainfall gradients. Mantel's test showed that environment and space were highly correlated with floristic similarity. Cluster analysis, indicating floristic similarity, resolved five groups, mainly grouped by region. Clear differentiation of the Brazilian Atlantic Forest epiphytic flora on a north–south axis with a strong correlation with temperature and rainfall gradients was found. The role of space and environment on species composition varied according to distinct epiphytic species groups. In particular, for Bromeliaceae and Orchidaceae, the main factor associated with floristic similarity was space. Indicator species were found for all vegetation types apart from the Seasonal Semideciduous Forest that seemed to represent a subset of a more humid forest type. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 587–601.     

Plant and small mammal richness correlate positively in a biodiversity hotspot

Hotspots of biodiversity are important areas in facilitating an understanding of species richness and its maintenance. Herbivores can increase plant richness by reducing dominant plant species thus providing space for subdominant species. As small mammals are abundant in the Succulent Karoo and therefore might affect plant richness by means of herbivory, we tested if this mechanism might exist in the Succulent Karoo in southern Africa, a biodiversity hotspot due to its extraordinary plant richness. At ten ecologically different study sites we measured plant and small mammal richness and diversity and determined 11 abiotic factors including soil composition, altitude and rainfall. We found positive correlations between plant richness and the number of small mammal species. A general linear model revealed that the number of small mammal species was more important than abiotic factors in explaining variation in plant richness. To test whether small mammals might directly influence plant richness, we studied the influence of the bush-Karoo rat Otomys unisulcatus, a central place forager, on the plant community. The immediate surroundings of occupied O. unisulcatus nests showed significantly higher plant richness than control areas. We conclude that small mammals can have a positive effect on plant richness in the Succulent Karoo. While experimental data are needed to support these correlative results, the results of our study indicate that areas of high small mammal richness should be included in conservation programs of the Succulent Karoo.     

Biodiversity responses to land‐use and restoration in a global biodiversity hotspot: Ant communities in Brazilian Cerrado

Given that land‐use change is the main cause of global biodiversity decline, there is widespread interest in adopting land‐use practices that maintain high levels of biodiversity, and in restoring degraded land that previously had high biodiversity value. In this study, we use ant taxonomic and functional diversity to examine the effects of different land uses (agriculture, pastoralism, silviculture and conservation) and restoration practices on Cerrado (Brazilian savanna) biodiversity. We also examine the extent to which ant diversity and composition can be explained by vegetation attributes that apply across the full land management spectrum. We surveyed vegetation attributes and ant communities in five replicate plots of each of 13 land‐use and restoration treatments, including two types of native vegetation as reference sites: cerrado sensu stricto and cerradão. Several land‐use and restoration treatments had comparable plot richness to that of the native reference habitats. Ant species and functional composition varied systematically among land‐use treatments following a gradient from open habitats such as agricultural fields to forested sites. Tree basal area and grass cover were the strongest predictors of ant species richness. Losses in ant diversity were higher in land‐use systems that transform vegetation structure. Among productive systems, therefore, uncleared pastures and old pine plantations had similar species composition to that occurring in cerrado sensu stricto. Restoration techniques currently applied to sites that were previously Cerrado have focused on returning tree cover, and have failed to restore ant communities typical of savanna. To improve restoration outcomes for Cerrado biodiversity, greater attention needs to be paid to the re‐establishment and maintenance of the grass layer, which requires frequent fire. At the broader scale, conservation planning in agricultural landscapes, should recognize the value of land‐use mosaics and the risks of homogenization.     

Searching for heat in a marine biodiversity hotspot

Coral reefs exhibit highly congruent patterns of biodiversity, with a prominent hotspot in the Indo-Australian Archipelago (IAA). Unlike many terrestrial systems, the IAA hotspot exhibits extensive latitudinal and longitudinal biodiversity gradients. Conflicting hypotheses have highlighted the importance of the area as a centre of origin, overlap or accumulation, with the location of endemics being used as the primary criterion for testing these hypotheses, by identifying the presumed geographical origins of species. We evaluate the utility of marine endemics for resolving these hypotheses, and examine recent molecular phylogenetic evidence for coral reef species that has revealed the antiquity of the endemics and the other species that make up this hotspot. These analyses emphasize the importance of the IAA in the survival rather than the origins of species.     

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.     

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.     

Landscape heterogeneity and the confluence of regional faunas promote richness and structure community assemblage in a tropical biodiversity hotspot

Crane flies (Limoniidae; Limoniinae) were sampled at national parks and protected areas across central to northern Thailand to observe patterns of species richness and faunal turnover in the Indo-Burma biodiversity hotspot. Prior to sampling, the crane fly fauna of this region was poorly known and no taxonomic keys existed for specimen identification. Utilizing a multi-access taxonomic key to the Limoniinae genera of the Oriental Region designed for this project, identification of collected specimens revealed a crane fly fauna displaying higher richness than inventories from temperate regions. Sixty-six morphospecies from 29 genera/subgenera were collected using a combination of light trapping and Malaise trapping. Richness estimators projected that a total of 70–81 species are to be collected with future sampling, with mountainous northern Thailand projected to have the highest richness. The faunas of Central and Northern Thailand were different, with the north generally composed of more temperate genera and the south composed of more tropical genera. The increased diversity in northern Thailand was significantly influenced by landscape topology. Sampling that spread across two mountain ranges displayed faunas that were divided into both high elevation (>1,000 m) and lower elevation (<1,000 m) faunas. This change in community assemblage across elevation illustrates faunas that were more alike at similar elevations between mountain ranges than they were within national parks.     

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.     

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.     

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.     

Assessing the vulnerability of species richness to anthropogenic climate change in a biodiversity hotspot

Aim To compare theoretical approaches towards estimating risks of plant species loss to anthropogenic climate change impacts in a biodiversity hotspot, and to develop a practical method to detect signs of climate change impacts on natural populations. Location The Fynbos biome of South Africa, within the Cape Floristic Kingdom. Methods Bioclimatic modelling was used to identify environmental limits for vegetation at both biome and species scale. For the biome as a whole, and for 330 species of the endemic family Proteaceae, tolerance limits were determined for five temperature and water availability‐related parameters assumed critical for plant survival. Climate scenarios for 2050 generated by the general circulation models HadCM2 and CSM were interpolated for the region. Geographic Information Systems‐based methods were used to map current and future modelled ranges of the biome and 330 selected species. In the biome‐based approach, predictions of biome areal loss were overlayed with species richness data for the family Proteaceae to estimate extinction risk. In the species‐based approach, predictions of range dislocation (no overlap between current range and future projected range) were used as an indicator of extinction risk. A method of identifying local populations imminently threatened by climate change‐induced mortality is also described. Results A loss of Fynbos biome area of between 51% and 65% is projected by 2050 (depending on the climate scenario used), and roughly 10% of the endemic Proteaceae have ranges restricted to the area lost. Species range projections suggest that a third could suffer complete range dislocation by 2050, and only 5% could retain more than two thirds of their range. Projected changes to individual species ranges could be sufficient to detect climate change impacts within ten years. Main conclusions The biome‐level approach appears to underestimate the risk of species diversity loss from climate change impacts in the Fynbos Biome because many narrow range endemics suffer range dislocation throughout the biome, and not only in areas identified as biome contractions. We suggest that targeted vulnerable species could be monitored both for early warning signs of climate change and as empirical tests of predictions.     

Fine‐scale patterns of species and phylogenetic turnover in a global biodiversity hotspot: Implications for climate change vulnerability

Question: What is the relative importance of environmental and spatial factors for species compositional and phylogenetic turnover? Location: High‐rainfall zone of the Southwest Australian Floristic Region (SWAFR). Methods: Correlates of species compositional turnover were assessed using quadrat‐based floristic data, and establishing relationships with environmental and spatial factors using canonical correspondence analyses and Mantel tests. Between‐quadrat phylogenetic distance measures were computed and examined for correlations with environmental and spatial attributes. Processes structuring pa2t2terns of beta diversity were also evaluated within four broad floristic assemblages defined a priori. Results: Floristic diversity was strongly related to environmental attributes. A low significance of spatial variables on assemblage patterns suggested no evident effect of dispersal limitations. Species compositional turnover was especially high within the swamp and outcrop assemblage. Phylogenetic turnover was closely coupled to species compositional turnover, implying the occurrence of many locally endemic and phylogenetically relict taxa. Beta diversity patterns within assemblages were also significantly correlated with the local environment, and relevant correlates differed between floristic assemblage types. Conclusion: Phylogenetic diversity in the SWAFR high‐rainfall zone is clustered within edaphic microhabitats in a generally subdued landscape. A clustered rather than dispersed distribution of phylogenetic diversity increases the probability of significant plant diversity loss during periods of climate change. Climate change susceptibility of the region's flora is accordingly estimated to be high. We highlight the conservation significance of swamp and outcrops that are characterized by distinct hydrological properties and may provide refugial habitat for plant diversity during periods of moderate climate change.     

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.     

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.