The marine East Indies: diversity and speciation

Aim To discuss the impact of new diversity information and to utilize recent findings on modes of speciation in order to clarify the evolutionary significance of the East Indies Triangle. Location The Indo‐Pacific Ocean. Methods Analysis of information on species diversity, distribution patterns and speciation for comparative purposes. Results Information from a broad‐scale survey of Indo‐Pacific fishes has provided strong support for the theory that the East Indies Triangle has been operating as a centre of origin. It has become apparent that more than two‐thirds of the reef fishes inhabiting the Indo‐Pacific are represented in the Triangle. An astounding total of 1111 species, more than are known from the entire tropical Atlantic, were reported from one locality on the small Indonesian island of Flores. New information on speciation modes indicates that the several unique characteristics of the East Indian fauna are probably due to the predominance of competitive (sympatric) speciation. Main conclusions It is proposed that, within the East Indies, the high species diversity, the production of dominant species, and the presence of newly formed species, are due to natural selection being involved in reproductive isolation, the first step in the sympatric speciation process. In contrast, speciation in the peripheral areas is predominately allopatric. Species formed by allopatry are the direct result of barriers to gene flow. In this case, reproductive isolation may be seen as a physical process that does not involve natural selection. Allopatric species formation often takes millions of years, while the sympatric process is generally much faster. Following species formation, dispersal from the East Indies appears to take place according to the centrifugal hypothesis.     

COINCIDENT BIOGEOGRAPHIC PATTERNS: INDO-WEST PACIFIC OCEAN

The majority of tropical marine families demonstrate their greatest concentration of species within the relatively small East Indies Triangle. In every direction, the species diversity decreases with distance from the East Indies. Other patterns suggest that the East Indies is where the average generic age is youngest, where some historical routes of dispersal originate, where the most apomorphic species occur, where genetic diversity is the greatest, and where extinctions are likely to originate. These coincident patterns provide support for the hypothesis that the East Indies has been operating as a center of evolutionary radiation. The driving force for this dynamic system is apparently the predominance of successful speciation involving relatively large populations with higher genetic diversity. This mechanism fits the centrifugal speciation model that was proposed more than 50 years ago.     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

A phylogenetic test of multiple proposals for the origins of the East Indies coral reef biota

Aim  We analysed data on Indo-Pacific coral reef taxa to test four mechanisms proposed for the origins of the biodiversity centre in the East Indies Triangle: (1) the centre of origin hypothesis, (2) the centre of accumulation hypothesis, (3) the centre of overlap hypothesis, and (4) the bioaccumulation hypothesis.
Location  The Indian and western Pacific oceans.
Methods  The data set consisted of eight clades of fishes, four of corals and one of molluscs, consisting of a total of 95 species, with 72 informative nodes, in 29 areas. We used phylogenetic analysis for comparing trees (PACT) to reconstruct reticulate area relationships.
Results  An analysis using PACT produced very little congruence between area relationships among the histories of the taxa examined. Only two sets of two taxa each showed substantial congruence; several other taxa showed partial congruence.
Main conclusions  Despite numerous 'explanatory' claims for the incredible biodiversity of the marine macrofauna of the East Indies Triangle, the patterns obtained here did not accord with any of the hypotheses proposed. It is possible that, with the addition of more taxa, additional patterns would emerge. Much more systematic work within the East Indies is required to resolve this problem.     

Antitropical distribution and evolution in the Indo-West Pacific Ocean

Antitropical distributions of continental shelf, Indo-West Pacific species are probably not due to transgression of the tropics during the glacial periods, isothermic submergence, island integration, rising Neogene temperatures, or the Mesozoic dispersal of fragments from a Pacific continental mass. Characteristics of common antitropical patterns, plus information from systematic works on a variety on a variety of animal and plant groups, indicate that the long discarded "relict theory" of Theel (1885) appears to best fit the evidence, for it provides a mechanism whereby antitropical distribution may be brought about. The relict theory is compatible with the concept that the East Indies part of the Indo-West Pacific has been functioning as a center of evolutionary origin. It suggests that antitropical and associated disjunct patterns are produced as an older species, that has spread out to occupy a broad range, loses ground and gradually becomes supplanted by a younger species that had subsequently evolved in the East Indies. As this process goes on, the older species becomes restricted to a few isolated localities on the fringe of its original range. These isolates are often found to the north and south of the equatorial region but may include relict populations at the western edge of the Indian Ocean.     

Origins of social parasitism: the importance of divergence ages in phylogenetic studies

Phylogenetic studies on insect social parasites have found very close host-parasite relationships, and these have often been interpreted as providing evidence for sympatric speciation. However, such phylogenetic inferences are problematic because events occurring after the origin of parasitism, such as extinction, host switching and subsequent speciation, or an incomplete sampling of taxa, could all confound the interpretation of phylogenetic relationships. Using a tribe of bees where social parasitism has repeatedly evolved over a wide time-scale, we show the problems associated with phylogenetic inference of sympatric speciation. Host-parasite relationships of more ancient species appear to support sympatric speciation, whereas in a case where parasitism has evolved very recently, sympatric speciation can be ruled out. However, in this latter case, a single extinction event would have lead to relationships that support sympatric speciation, indicating the importance of considering divergence ages when analysing the modes of social parasite evolution.     

Marine longitudinal biodiversity: causes and conservation

The horizontal temperature zones of the earth tend to restrict the latitudinal ranges of species but allow the possibility of exceedingly broad longitudinal dispersals. In the Tropical Zone, biodiversity on the continental shelves is not homogeneous but is concentrated in two conspicuous peaks, one in the Indo‐Pacific Ocean and the other in the Atlantic. The Indo‐Pacific biodiversity peak is located within a relatively small area called the East Indies Triangle. The Atlantic peak is located in the southern Caribbean Sea. Evidence that has been accumulated over the years indicates that each area functions as a centre of origin and evolutionary radiation. What are the causes of these concentrations and their present functions? A newly published model indicates a positive relationship between environmental temperature and the rate of speciation. While this helps to explain the generally high tropical diversity, and the negative relationship between diversity and latitude, it does not provide a reason for the longitudinal concentrations. But, other new research serves to substantiate previous indications of a positive relationship between speciation rate and species diversity. The existence of this positive feedback, together with some contributory factors, provides the reason why concentrations occur. The evolutionary radiation probably begins when the build‐up of species diversity reaches a critical level. The warm‐temperate biotas are derived from the tropics. Their northern longitudinal relationships tend to be minor but, in the southern hemisphere, the West Wind Drift is an important dispersal mechanism for both warm‐temperate and cold‐temperate species. The cold‐temperate biotas peaked in two areas, the North Pacific and the Antarctic; each has developed into a centre of origin. The continuous dispersal of well‐adapted species from the centres helps peripheral communities maintain diversity.     

The West Indies as a laboratory of biogeography and evolution

Islands have long provided material and inspiration for the study of evolution and ecology. The West Indies are complex historically and geographically, providing a rich backdrop for the analysis of colonization, diversification and extinction of species. They are sufficiently isolated to sustain endemic forms and close enough to sources of colonists to develop a dynamic interaction with surrounding continental regions. The Greater Antilles comprise old fragments of continental crust, some very large; the Lesser Antilles are a more recent volcanic island arc, and the low-lying Bahama Islands are scattered on a shallow oceanic platform. Dating of island lineages using molecular methods indicates over-water dispersal of most inhabitants of the West Indies, although direct connections with what is now southern Mexico in the Early Tertiary, and subsequent land bridges or stepping stone islands linking to Central and South America might also have facilitated colonization. Species-area relationships within the West Indies suggest a strong role for endemic radiations and extinction in shaping patterns of diversity. Diversification is promoted by opportunities for allopatric divergence between islands, or within the large islands of the Greater Antilles, with a classic example provided by the Anolis lizards. The timing of colonization events using molecular clocks permits analysis of colonization-extinction dynamics by means of species accumulation curves. These indicate low rates of colonization and extinction for reptiles and amphibians in the Greater Antilles, with estimated average persistence times of lineages in the West Indies exceeding 30Myr. Even though individual island populations of birds might persist an average of 2Myr on larger islands in the Lesser Antilles, recolonization from within the archipelago appears to maintain avian lineages within the island chain indefinitely. Birds of the Lesser Antilles also provide evidence of a mass extinction event within the past million years, emphasizing the time-heterogeneity of historical processes. Geographical dynamics are matched by ecological changes in the distribution of species within islands over time resulting from adaptive radiation and shifts in habitat, often following repeatable patterns. Although extinction is relatively infrequent under natural conditions, changes in island environments as a result of human activities have exterminated many populations and others--especially old, endemic species--remain vulnerable. Conservation efforts are strengthened by recognition of aesthetic, cultural and scientific values of the unique flora and fauna of the West Indies.     

Spatial patterns of disparity and diversity of the Recent cuttlefishes (Cephalopoda) across the Old World

Aim Diversity and disparity metrics of all Recent cuttlefishes are studied at the macroevolutionary scale (1) to establish the geographical biodiversity patterns of these cephalopods at the species level and (2) to explore the relationships between these two metrics. Location Sampling uses what is known about these tropical, subtropical and warm temperate cephalopods of the Old World based on a literature review and on measurements of museum specimens. Some 111 species spread across seventeen biogeographical areas serve as basic units for exploring diversity and disparity metrics in space. Methods Landmarks describe the shape of the cuttlebone (the inner shell of the sepiids) and differences between shapes are quantified using relative warp analyses. Relative warps are thus used as the morphological axis for constructing morphospaces whose characteristics are described by disparity indices: total variance, range, and minimum and maximum of relative warps. These are analysed and then compared with the diversity (species richness) metric. Results Results show no significant latitudinal or longitudinal gradients either for diversity or for disparity. Around the coast of southern Africa, disparity is high regardless of whether diversity (species richness) is high or low. In the ‘East Indies’ area disparity is low despite the high diversity. Main conclusions The relationship between diversity and disparity is clearly not linear and no simple adjustment models seem to fit. The number of species in a given area does not predict its disparity level. The particular pattern of southern Africa may be the result of paleogeographical changes since the Eocene, whereas that of the ‘East Indies’ may indicate that this area could act as a centre of origin. However, the lack of any clear phylogenetical hypothesis precludes the study from providing any explanation of the observed patterns.     

Faster Speciation and Reduced Extinction in the Tropics Contribute to the Mammalian Latitudinal Diversity Gradient

The increase in species richness from the poles to the tropics, referred to as the latitudinal diversity gradient, is one of the most ubiquitous biodiversity patterns in the natural world. Although understanding how rates of speciation and extinction vary with latitude is central to explaining this pattern, such analyses have been impeded by the difficulty of estimating diversification rates associated with specific geographic locations. Here, we use a powerful phylogenetic approach and a nearly complete phylogeny of mammals to estimate speciation, extinction, and dispersal rates associated with the tropical and temperate biomes. Overall, speciation rates are higher, and extinction rates lower, in the tropics than in temperate regions. The diversity of the eight most species-rich mammalian orders (covering 92% of all mammals) peaks in the tropics, except that of the Lagomorpha (hares, rabbits, and pikas) reaching a maxima in northern-temperate regions. Latitudinal patterns in diversification rates are strikingly consistent with these diversity patterns, with peaks in species richness associated with low extinction rates (Primates and Lagomorpha), high speciation rates (Diprotodontia, Artiodactyla, and Soricomorpha), or both (Chiroptera and Rodentia). Rates of range expansion were typically higher from the tropics to the temperate regions than in the other direction, supporting the “out of the tropics” hypothesis whereby species originate in the tropics and disperse into higher latitudes. Overall, these results suggest that differences in diversification rates have played a major role in shaping the modern latitudinal diversity gradient in mammals, and illustrate the usefulness of recently developed phylogenetic approaches for understanding this famous yet mysterious pattern.     

The phytogeographical studies of Thermopsis (Fabaceae)

The present article is the first comprehensive treatment of phytogeography of Thermopsis (Fabaceae) in the world. Thermopsis is one of the few genera within Fabaceae with the distribution pattern of the East Asia-North American disjunction. The distribution patterns of 5 recognized sections (including a new one) covering 21 species in Thermopsis are analyzed, and the results show four centres of frequency of the genus: the Eastern Asiatic Region (9 spp. / 3 sects., including 4 endemic species), the Irano-Turanian Region (7 spp./3 sects., including 3 endemic species), the Rocky Mountain Region (7 spp./2 sects., all endemic), and the Atlantic North American Region (3 spp. / 1 sect., all endemic). In the light of the fact that most species and sections, a number of phylogenetic series of the genus, and the most primitive sections and most advanced sections in Thermopsis occur in the East Asia, the Eastern Asiatic Region might be the centre of diversity of the genus. As the Irano-Turanian Region and the Rocky Mountain Region were just second to that of Eastern Asiatic Region in number of sections and species, and many polyploids appeared in these regions, they were considered as the secondary centres of distribution and speciation of the genus. The speciation looks to be frequent and complex in these regions, and many new taxa have been described from there while many new reduced or incorporated taxa have happened over there. However, recent molecular data has shown that two reduced taxa of Thermopsis are distinct in these regions. Based on the modern distribution patterns and evolutionary trends in morphological characters of the genus, and available fossil record of the genus and the historical geology, we speculate that Thermopsis had already existed on Eurasia and North America before the Late Miocene, and probably originated from an ancestral form of Sophora-like taxa with lupine alkaloids somewhere in the Laurasia in the Early Tertiary or Late Cretaceous. After the separation of the two continents, species on different continents developed distinctly under influences of different evolutionary factors. In Asia, the late Tertiary orogeny, disappearing of the Tethys and aridity and freezing caused by the Quaternary glaciation were the main forces to promote the speciation and evolutionary processes, whereas in North America it was the Quaternary glaciation and the orogeny of partial area to promote evolution of the genus. According to the evolutionary trends in Thermopsis and the distribution pattern of the primitive taxa, Sino-Japanese Subregion of Eastern Asiatic Region may be considered asthe centre of primitive forms of Thermopsis.     

Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity

Many groups show higher species richness in tropical regions but the underlying causes remain unclear. Despite many competing hypotheses to explain latitudinal diversity gradients, only three processes can directly change species richness across regions: speciation, extinction and dispersal. These processes can be addressed most powerfully using large-scale phylogenetic approaches, but most previous studies have focused on small groups and recent time scales, or did not separate speciation and extinction rates. We investigate the origins of high tropical diversity in amphibians, applying new phylogenetic comparative methods to a tree of 2871 species. Our results show that high tropical diversity is explained by higher speciation in the tropics, higher extinction in temperate regions and limited dispersal out of the tropics compared with colonization of the tropics from temperate regions. These patterns are strongly associated with climate-related variables such as temperature, precipitation and ecosystem energy. Results from models of diversity dependence in speciation rate suggest that temperate clades may have lower carrying capacities and may be more saturated (closer to carrying capacity) than tropical clades. Furthermore, we estimate strikingly low tropical extinction rates over geological time scales, in stark contrast to the dramatic losses of diversity occurring in tropical regions presently.     

Can sympatric speciation by disruptive sexual selection explain rapid evolution of cichlid diversity in Lake Victoria?

Rapid speciation can occur on ecological time scales and interfere with ecological processes, resulting in species distribution patterns that are difficult to reconcile with ecological theory. The haplochromine cichlids in East African lakes are an extreme example of rapid speciation. We analyse the causes of their high speciation rates. Various studies have identified disruptive sexual selection acting on colour polymorphisms that might cause sympatric speciation. Using data on geographical distribution, colouration and relatedness from 41 species endemic to Lake Victoria, we test predictions from this hypothesis. Plotting numbers of pairs of closely related species against the amount of distributional overlap between the species reveals a bimodal distribution with modes on allopatric and sympatric. The proportion of sister species pairs that are heteromorphic for the traits under disruptive selection is higher in sympatry than in allopatry. These data support the hypothesis that disruptive sexual selection on colour polymorphisms has caused sympatric speciation and help to explain the rapid radiation of haplochromine species flocks.     

Two decades of interaction between the MacArthur-Wilson model and the complexities of mammalian distributions

More than two decades after its publication, MacArthur and Wilson's equilibrium model of insular biogeography continues to provide the conceptual foundation for investigating the distribution of species on islands and the composition of insular biotas. During this period, studies of the distributions of mammals among insular habitats have tested, modified, and extended MacArthur and Wilson's simple formalism to enhance greatly our understanding of the complexities of biogeographic patterns and processes. The papers in this symposium summarize many of the past contributions of mammalian biogeographers and introduce important new data and ideas. The diversity of biological characteristics and associated distributional patterns exhibited by mammals has facilitated this endeavour. Some insular mammalian faunas appear to represent approximate equilibria between opposing rates of contemporary colonization and extinction. Other faunas are currently decreasing in diversity because of extinctions, owing either to natural habitat fragmentation that has occurred since the Pleistocene or to human activities within the last few centuries. Still other faunas have been increasing in diversity (at least until recent human impacts) because limiting rates of origination, both colonization and speciation, have been extremely low. The questions and analyses of island biogeography can also be applied to continents with comparable overall results: the distributions of continental faunas reflect the consequences of similar processes of colonization, speciation and extinction. Analyses of insular distributions show unequivocally that probabilities of extinction, colonization and speciation are highly deterministic and vary in predictable ways among different taxa and archipelagos. These findings have important implications for applying the theory and data of insular biogeography to the pressing practical problems of designing natural reserves to preserve native species.     

Resilient forest faunal communities in South Africa: a legacy of palaeoclimatic change and extinction filtering?

Aim To examine the influence of climatic extinction filtering during the last glacial maximum (LGM; c. 18,000 yr bp ) and of the subsequent recolonization of forest faunas on contemporary assemblage composition in southern African forests. Location South Africa, Mozambique, Swaziland, Zimbabwe. Methods Data comprised presence/absence by quarter‐degree grid cell for forest‐dependent and forest‐associated birds, non‐volant mammals and frogs. Twenty‐one forest subregions were assigned to one of three previously identified forest types: Afrotemperate, scarp, and Indian Ocean coastal belt. Differences among forest types were examined through patterns and gradients of species richness and endemism, assemblage similarity, species turnover, and coefficients of species dispersal direction. The influence of contemporary environment on assemblage composition was investigated using partial canonical correspondence analysis. Several alternative biogeographical hypotheses for the recolonization of forest faunas were tested. Results Afrotemperate faunas are relatively species‐poor, have low species turnover, and are unsaturated and infiltrated by generalist species. In northern and central regions, communities are supplemented by recolonization from scarp forest refugia, and among frogs by autochthanous speciation in localized refugia. Scarp faunas are relatively species‐rich, contain many forest‐dependent species, have high species turnover, and overlap with coastal and Afrotemperate faunas. Coastal forests are relatively species‐rich with high species turnover. Main conclusions Afrotemperate communities were affected most by climatic extinction filtering events. Scarp forests were Afrotemperate refugia during the LGM and are a contemporary overlap zone between Afrotemperate and coastal forest. Coastal faunas derive from post‐LGM colonization along the eastern seaboard from tropical East African refugia. The greatest diversity is achieved in scarp and coastal forest faunas in northern KwaZulu–Natal province. This historical centre of diversity has influenced the faunal diversity of nearly all other forests in South Africa. The response of vertebrate taxa to large‐scale, historical processes is dependent on their relative mobility: forest birds best illustrate patterns resulting from post‐glacial faunal dispersal, while among mammals and frogs the legacy of climatic extinction filtering remains stronger.     

Heterogeneity, speciation/extinction history and climate: explaining regional plant diversity patterns in the Cape Floristic Region

This paper investigates the role of heterogeneity and speciation/extinction history in explaining variation in regional scale (c. 0.1–3000 km²) plant diversity in the Cape Floristic Region of south‐western Africa, a species‐ and endemic‐rich biogeographical region. We used species‐area analysis and analysis of covariance to investigate geographical (east vs. west) and topographic (lowland vs. montane) patterns of diversity. We used community diversity as a surrogate for biological heterogeneity, and the diversity of naturally rare species in quarter degree squares as an indicator of differences in speciation/extinction histories across the study region. We then used standard statistical methods to analyse geographical and topographic patterns of these two measures. There was a clear geographical diversity pattern (richer in the west), while a topographic pattern (richer in mountains) was evident only in the west. The geographical boundary coincided with a transition from the reliable winter‐rainfall zone (west) to the less reliable non‐seasonal rainfall zone (east). Community diversity, or biological heterogeneity, showed no significant variation in relation to geography and topography. Diversity patterns of rare species mirrored the diversity pattern for all species. We hypothesize that regional diversity patterns are the product of different speciation and extinction histories, leading to different steady‐state diversities. Greater Pleistocene climatic stability in the west would have resulted in higher rates of speciation and lower rates of extinction than in the east, where for the most, Pleistocene climates would not have favoured Cape lineages. A more parsimonious hypothesis is that the more predictable seasonal rainfall of the west would have favoured non‐sprouting plants and that this, in turn, resulted in higher speciation and lower extinction rates. Both hypotheses are consistent with the higher incidence of rare species in the west, and higher levels of beta and gamma diversity there, associated with the turnover of species along environmental and geographical gradients, respectively. These rare species do not contribute to community patterns; hence, biological heterogeneity is uniform across the region. The weak topography pattern of diversity in the west arises from higher speciation rates and lower extinction rates in the topographically complex mountains, rather than from the influence of environmental heterogeneity on diversity.     

Niche width impacts vertebrate diversification

Aim The size of the climatic niche of a species is a major factor determining its distribution and evolution. In particular, it has been proposed that niche width should be associated with the rate of species diversification. Here, we test whether species niche width affects the speciation and extinction rates of three main clades of vertebrates: amphibians, mammals and birds. Location Global. Methods We obtained the time‐calibrated phylogenies, IUCN conservation status, species distribution maps and climatic data for 2340 species of amphibians, 4563 species of mammals and 9823 species of birds. We computed the niche width for each species as the mean annual temperature across the species range. We estimated speciation, extinction and transition rates associated with lineages with either narrow (specialist) or wide (generalist) niches using phylogeny‐based birth–death models. We also tested if current conservation status was correlated with the niche width of species. Results We found higher net diversification rates in specialist species than in generalist species. This result was explained by both higher speciation rates (for the three taxonomic groups) and lower extinction rates (for mammals and birds only) in specialist than in generalist species. In contrast, current specialist species tended to be more threatened than generalist species. Main conclusions Our diversification analysis shows that the width of the climatic niche is strongly associated with diversification rates and may thus be a crucial factor for understanding the emergence of diversity patterns in vertebrates. The striking difference between our diversification results and current conservation status suggests that the current extinction process may be different from extinction rates estimated from the whole history of the group.     

Testing for equal rates of cladogenesis in diverse taxa

Taxa differ widely in numbers of species, which may be due either to chance alone or to factors that cause differences in speciation and extinction rates between taxa. To test whether an observed distribution of species over taxa differs from the distribution expected from chance alone, one must take into account that neither speciation nor extinction rates are known. This paper introduces a way to estimate speciation and extinction probabilities from the distribution of extant species over families and to test whether the observed distribution is different from expected. Application of this procedure to the distributions of bird, hexapod, primate, and angiosperm species over taxa provides statistical evidence of differences in rates of cladogenesis between taxa.     

粉条儿菜属(肺筋草科)的地理分布及其物种多样性和分化中心

粉条儿菜属(AletrisL.)隶属于肺筋草科,全世界有23种1变种,东亚有18种1变种,北美东南部有5种,为典型的东亚-北美间断分布的属.本文在种(变种)的水平上,研究了粉条儿菜属的地理分布及其分布中心和多样化中心,并对其起源和分化以及现代洲际间断分布格局的成因进行了分析.结果表明,(1)中国共分布有粉条儿菜属植物15种1变种,而广义的横断山地区集中分布有13种1变种,是东亚粉条儿菜属植物分布最为集中的地区,而且包含该属植物各个进化阶段的代表.因此,广义的横断山地区是粉条儿菜属在东亚的分布中心和多样化中心.(2)根据粉条儿菜属及其近缘属的分布格局推测,该属可能在不晚于第三纪早期,起源于古北大陆.东亚和北美的粉条儿菜属植物形态区别明显,应该是隔离分化的结果.(3)该属植物可能曾经广布于北半球,后来地质、气候以及冰川等因素的变化,导致该属在一些地区灭绝,而仅存于东亚和北美东南部.(4)尽管横断山及其周边地区是东亚粉条儿菜属的多样化中心,但该地区很可能并不是粉条儿菜属最早的分化中心,因横断山地区周边的一些特有种可能是在晚近的时期形成的新特有种;另外,东亚粉条儿菜属一些原始的种类主要分布于我国中东部到日本一带.所以,中国中东部到日本一带可能是粉条儿菜属早期的分化中心.     

Primate diversification inferred from phylogenies and fossils

Biodiversity arises from the balance between speciation and extinction. Fossils record the origins and disappearance of organisms, and the branching patterns of molecular phylogenies allow estimation of speciation and extinction rates, but the patterns of diversification are frequently incongruent between these two data sources. I tested two hypotheses about the diversification of primates based on ～600 fossil species and 90% complete phylogenies of living species: (1) diversification rates increased through time; (2) a significant extinction event occurred in the Oligocene. Consistent with the first hypothesis, analyses of phylogenies supported increasing speciation rates and negligible extinction rates. In contrast, fossils showed that while speciation rates increased, speciation and extinction rates tended to be nearly equal, resulting in zero net diversification. Partially supporting the second hypothesis, the fossil data recorded a clear pattern of diversity decline in the Oligocene, although diversification rates were near zero. The phylogeny supported increased extinction ～34 Ma, but also elevated extinction ～10 Ma, coinciding with diversity declines in some fossil clades. The results demonstrated that estimates of speciation and extinction ignoring fossils are insufficient to infer diversification and information on extinct lineages should be incorporated into phylogenetic analyses.