Ancestral Distribution Research Articles

Multiple origins of circumboreal taxa in Pyrola (Ericaceae), a group with a Tertiary relict distribution.

In the Northern Hemisphere, Tertiary relict disjunctions involve older groups of warm affinity and wide disjunctions, whereas circumboreal distributions in Arctic-Alpine taxa tend to be younger. Arctic-Alpine species are occasionally derived from Tertiary relict groups, but Pyrola species, in particular, are exceptional and they might have occurred multiple times. The aim of this study was to reconstruct the biogeographic history of Pyrola based on a clear phylogenetic analysis and to explore how the genus attained its circumboreal distribution. Estimates of divergence times and ancestral geographical distributions based on neutrally evolving DNA sequence variation were used to develop a spatio-temporal model of colonization patterns for Pyrola. Pyrola originated and most diversification occurred in Asia; North America was reached first by series Scotophyllae in the late Miocene, then by sub-clades of series Pyrola and Ellipticae around the Pliocene. The three circumboreal taxa, P. minor, P. chlorantha and the P. rotundifolia complex, originated independently of one another, with the last two originating in Asia. Three circumboreal Pyrola lineages have arisen independently and at least two of these appear to have originated in Asia. The cool, high-altitude habitats of many Pyrola species and the fact that diversification in the genus coincided with global cooling from the late Miocene onwards fits a hypothesis of pre-adaptation to become circumboreal within this group.

Historical biogeography of the lacertid lizard Mesalina in North Africa and the Middle East

AbstractAimWe explored the phylogenetic relationships of species of Mesalina, using one nuclear and two mitochondrial loci. This genus of lacertid lizards is widely distributed in North Africa and the Middle East and our goal was to develop a scenario capable of explaining the current distribution and evolutionary patterns within the genus in the context of the wider historical biogeography of the region.LocationNorth Africa and the Middle East.MethodsThe assembled dataset consisted of 193 Mesalina individuals, representing 12 species distributed across the geographical range of the genus. Bayesian and maximum likelihood methods were used to support phylogenetic inferences on two mitochondrial (cytochrome b and 16S ribosomal RNA) and one nuclear (beta‐fibrinogen intron 7) markers. Palaeogeographical and palaeoclimatic data were used to support the inferred phylogeographical patterns.ResultsMesalina lizards exhibit high genetic diversity and complex phylogenetic patterns, leading to an unsatisfactory systematic hypothesis of one paraphyletic and three polyphyletic traditional species. The estimated divergence times place the origin of the genus in the early Miocene (c. 22 Ma) and the divergence of most currently recognized species in the middle to late Miocene. The inferred ancestral distribution suggests that the genus and most of its species originated somewhere in Arabia or the Middle East, with the exception of the Mesalina olivieri complex, which may be of African origin.Main conclusionsPhylogenetic reconstruction based on the three loci studied suggests a higher than expected cryptic diversity of Mesalina in North Africa and the Middle East. We suggest that the tectonic movements of the Arabian plate, coupled with the climatic changes occurring since the Miocene, may be responsible for the phylogeographical patterns of North African and Middle Eastern Mesalina.

Early diversification trend and Asian origin for extent bat lineages.

Bats are a unique mammalian group, which belong to one of the largest and most diverse mammalian radiations, but their early diversification is still poorly understood, and conflicting hypotheses have emerged regarding their biogeographic history. Understanding their diversification is crucial for untangling the enigmatic evolutionary history of bats. In this study, we elucidated the rate of diversification and the biogeographic history of extant bat lineages using genus-level chronograms. The results suggest that a rapid adaptive radiation persisted from the emergence of crown bats until the Early Eocene Climatic Optimum, whereas there was a major deceleration in diversification around 35-49 Ma. There was a positive association between changes in the palaeotemperature and the net diversification rate until 35 Ma, which suggests that the palaeotemperature may have played an important role in the regulation of ecological opportunities. By contrast, there were unexpectedly higher diversification rates around 25-35 Ma during a period characterized by intense and long-lasting global cooling, which implies that intrinsic innovations or adaptations may have released some lineages from the intense selective pressures associated with these severe conditions. Our reconstruction of the ancestral distribution suggests an Asian origin for bats, thereby indicating that the current panglobal but disjunct distribution pattern of extant bats may be related to events involving seriate cross-continental dispersal and local extinction, as well as the influence of geological events and the expansion and contraction of megathermal rainforests during the Tertiary.

Causes of endemic radiation in the Caribbean: evidence from the historical biogeography and diversification of the butterfly genus Calisto (Nymphalidae: Satyrinae: Satyrini).

BackgroundCalisto is the largest butterfly genus in the West Indies but its systematics, historical biogeography and the causes of its diversification have not been previously rigorously evaluated. Several studies attempting to explain the wide-ranging diversity of Calisto gave different weights to vicariance, dispersal and adaptive radiation. We utilized molecular phylogenetic approaches and secondary calibrations points to estimate lineage ages. In addition, we used the dispersal-extinction-cladogenesis model and Caribbean paleogeographical information to reconstruct ancestral geographical distributions. We also evaluated different models of diversification to estimate the dynamics of lineage radiation within Calisto. By understanding the evolution of Calisto butterflies, we attempt to identify the main processes acting on insular insect diversity and the causes of its origin and its maintenance.ResultsThe crown age of Calisto was estimated to the early Oligocene (31 ± 5 Ma), and a single shift in diversification rate following a diversity-dependent speciation process was the best explanation for the present-day diversity found within the genus. A major increase in diversification rate was recovered at 14 Ma, following geological arrangements that favoured the availability of empty niches. Inferred ancestral distributional ranges suggested that the origin of extant Calisto is in agreement with a vicariant model and the origin of the Cuban lineage was likely the result of vicariance caused by the Cuba-Hispaniola split. A long-distance dispersal was the best explanation for the colonization of Jamaica and the Bahamas.ConclusionsThe ancestral geographical distribution of Calisto is in line with the paleogeographical model of Caribbean colonization, which favours island-to-island vicariance. Because the sister lineage of Calisto remains ambiguous, its arrival to the West Indies remains to be explained, although, given its age and historical biogeography, the hypothesized GAARlandia land bridge might have been a plausible introduction route from continental America. Intra-island radiation caused by ecological innovation and the abiotic creation of niche spaces was found to be the main force shaping Calisto diversity and island endemism in Hispaniola and Cuba.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-014-0199-7) contains supplementary material, which is available to authorized users.

Extension of historical range of Betylobraconinae (Hymenoptera: Braconidae) into Palaearctic Region based on a Baltic amber fossil, and description of a new species of Mesocentrus Szépligeti from Papua New Guinea.

Two new species of the parasitic wasp genus Mesocentrus Szépligeti (Betylobraconinae) are described. One based on a new species from Papua New Guinea, the other the first Palaearctic member of the subfamily based on a 30+ mya, species from Baltic amber. The second species is illustrated using synchrotron X-ray microtomography. Whereas the extant Betylobraconinae are restricted to Australia, New Guinea and New Caledonia, their ancestral distribution is now known to have extended considerably further. A key to the four species of Mesocentrus known from Papua New Guinea is provided. Both species possess some putatively plesiomorphic characters absent in other extant Mesocentrus spp. The new extant species differs in having a considerably larger number of antennal segments and a less laterally depressed frons, while the extinct one has the clypeus separated from the face dorsally and strongly developed hypoclypeal depression. Availability of sequence data for this species enabled further analysis of the relationships of the subfamily, which we present in a phylogenetic analysis additionally including the release of a number of new sequences of related taxa.

Tracing the roots of syntax with Bayesian phylogenetics.

The ordering of subject, verb, and object is one of the fundamental components of the syntax of natural languages. The distribution of basic word orders across the world's languages is highly nonuniform, with the majority of languages being either subject-object-verb (SOV) or subject-verb-object (SVO). Explaining this fact using psychological accounts of language acquisition or processing requires understanding how the present distribution has resulted from ancestral distributions and the rates of change between orders. We show that Bayesian phylogenetics can provide quantitative answers to three important questions: how word orders are likely to change over time, which word orders were dominant historically, and whether strong inferences about the origins of syntax can be drawn from modern languages. We find that SOV to SVO change is more common than the reverse and VSO to SVO change is more common than VSO to SOV, and that if the seven language families we consider share a common ancestor then that common ancestor likely had SOV word order, but also that there are limits on how confidently we can make inferences about ancestral word order based on modern-day observations. These results shed new light on old questions from historical linguistics and provide clear targets for psychological explanations of word-order distributions.

Pleistocene diversification of the Pomacentrus coelestis species complex (Pisces: Pomacentridae): historical biogeography and species boundaries

Pleistocene eustatic changes in sea level are often invoked to explain genetic divergence among marine organisms. However, molecular phylogenies have revealed relatively few examples of speciation events dating to the Pleistocene. We present a species-level hypothesis of the timing of evolution for the Pomacentrus coelestis species complex (Pomacentridae), based on the nuclear S7 intron and the mitochondrial Cytb gene, and reconstruct ancestral range distributions across the timetree. Ancestral range reconstruction suggests the complex originated in the Coral Triangle and East Indian Ocean, with subsequent range expansion outward from this region. We suggest that land barriers to dispersal (e.g., Indo-Pacific barrier) may be responsible for the divergence between Indian (P. alleni, P. similis, P. caeruleopunctatus, and P. caeruleus) and Pacific (P. micronesicus, P. auriventris, and P. coelestis) species groups, and subsequent isolation by Pleistocene sea-level fluctuations in certain areas of the Coral Triangle (glacial refugia) may play an important role in the diversification of this species complex. Additionally, our analyses show cryptic lineages within P. micronesicus and highlight the need for comprehensive sampling within and among species to reveal recent speciation events.

In the wake of invasion: tracing the historical biogeography of the South American cricetid radiation (Rodentia, Sigmodontinae).

The Great American Biotic Interchange (GABI) was greatly influenced by the completion of the Isthmus of Panama and impacted the composition of modern faunal assemblages in the Americas. However, the contribution of preceding events has been comparatively less explored, even though early immigrants in the fossil records are evidence for waif dispersals. The cricetid rodents of the subfamily Sigmodontinae are a classic example of a species-rich South American radiation resulting from an early episode of North American invasion. Here, we provide a temporal and spatial framework to address key aspects of the historical biogeography and diversification of this diverse mammal group by using mitochondrial and nuclear DNA datasets coupled with methods of divergence time estimation, ancestral area reconstruction and comparative phylogenetics. Relaxed-clock time estimates indicate that divergence of the Sigmodontinae began in the middle–late Miocene (ca. 12–9 Ma). Dispersal-vicariance analyses point to the arrival of a single lineage of northern invaders with a widespread ancestral distribution and imply that the initial differentiation between Central and South America gave rise to the most basal groups within the subfamily. These two major clades diversified in the late Miocene followed by the radiation of main tribes until the early Pliocene. Within the Oryzomyalia, tribes diverged initially in eastern South America whereas multiple dispersals into the Andes promoted further diversification of the majority of modern genera. A comparatively uniform background tempo of diversification explains the species richness of sigmodontines across most nodes, except for two akodontine genera with recent increases in diversification rates. The bridging of the Central American seaway and episodes of low sea levels likely facilitated the invasion of South America long before the onset of the post-Isthmian phase of the GABI.

Himalayan origin and evolution of Myricaria (Tamaricaeae) in the neogene.

Background Myricaria consists of about twelve-thirteen species and occurs in Eurasian North Temperate zone, most species in the Qinghai-Tibet Plateau (QTP) and adjacent areas.Methodology/Principal FindingsTwelve species of Myricaria plus two other genera Tamarix and Reaumuria in Tamaricaceae, were sampled, and four markers, ITS, rps16, psbB-psbH, and trnL-trnF were sequenced. The relaxed Bayesian molecular clock BEAST method was used to perform phylogenetic analysis and molecular dating, and Diva, S-Diva, and maximum likelihood Lagrange were used to estimate the ancestral area. The results indicated that Myricaria could be divided into four phylogenetic clades, which correspond to four sections within the genus, of them two are newly described in this paper. The crown age of Myricaria was dated to early Miocene ca. 20 Ma, at the probable early uplifting time of the Himalayas. The Himalayas were also shown as the center of origin for Myricaria from the optimization of ancestral distribution. Migration and dispersal of Myricaria were indicated to have taken place along the Asian Mountains, including the Himalayas, Kunlun, Altun, Hendukosh, Tianshan, Altai, and Caucasus etc., westward to Europe, eastward to Central China, and northward to the Mongolian Plateau.Conclusions/Significance Myricaria spatiotemporal evolution presented here, especially the Himalayan origin at early Miocene ca. 20 Ma, and then migrated westward and eastward along the Asian mountains, offers a significant evolutionary case for QTP and Central Asian biogeography.

Phylogeny, Historical Biogeography and the Evolution of Migration in Accipitrid Birds of Prey (Aves: Accipitriformes)

Abstract Migration plays a fundamental part in the life of most temperate bird species. The regular, large-scale seasonal movements that characterize temperate migration systems appear to have originated in parallel with the postglacial northern expansion of tropical species. Migratoriness is also influenced by a number of ecological factors, such as the ability to survive harsh winters. Hence, understanding the origins and evolution of migration requires integration of the biogeographic history and ecology of birds in a phylogenetic context. We used molecular dating and ancestral state reconstruction to infer the origins and evolutionary changes in migratory behavior and ancestral area reconstruction to investigate historical patterns of range evolution in accipitrid birds of prey (Accipitriformes). Migration evolved multiple times in birds of prey, the earliest of which occurred in true hawks (Accipitrinae), during the middle Miocene period, according to our analyses. In most cases, a tropical ancestral distribution was inferred for the non-migratory ancestors of migratory lineages. Results from directional evolutionary tests indicate that migration evolved in the tropics and then increased the rate of colonization of temperate habitats, suggesting that temperate species might be descendants of tropical ones that dispersed into these seasonal habitats. Finally, we found that diet generalization predicts migratoriness in this group.

Late Miocene origin of an Ibero‐Maghrebian clade of ground beetles with multiple colonizations of the subterranean environment

AbstractAimTo test different biogeographical scenarios for the evolution of the ground beetles of the Trechus fulvus group, a lineage with many narrowly distributed flightless subterranean species, highly suitable for tracing their biogeographical history.LocationThe Western Palaearctic, focusing on the Betic–Rifean area between south‐east Iberia and north Morocco.MethodsWe sequenced 3.3 kb of four mitochondrial and two nuclear genes of 30 individuals of 15 species of the T. fulvus group, plus 29 outgroups. We reconstructed their phylogeny and estimated divergence times using Bayesian probabilities and a priori evolutionary rates, and their ancestral distribution using maximum likelihood.ResultsThe phylogenetic reconstruction uncovered multiple independent colonizations of the subterranean environment within the T. fulvus group, a scenario also supported by variation in troglomorphic characters. Most of the Moroccan and south‐east Iberian species form a clade with strong geographical structure, including the former genus Antoinella. The biogeographical model best fitting the current distribution and phylogeny of the group was a late Miocene palaeogeographical scenario with isolated populations on the Betic and Rifean areas and a south‐eastern Iberian origin of the north Moroccan species. The widespread T. fulvus was sister to a central Moroccan species, suggesting a second Iberian–North African vicariance event within the group. One of the species, T. lallemantii, expanded its range to the east (Algeria and Tunisia) and the north (extreme south of the Iberian Peninsula) during the Pleistocene.Main conclusionsThe T. fulvus group originated in the early Miocene, and the south‐eastern Iberian lineage in the Tortonian. The lineage dispersed to Morocco during the Messinian, diversifying in rapid succession in the Atlas and the Rif and colonizing the subterranean environment multiple times. The geography of the Betic–Rifean region at the end of the Miocene can still be traced from the distribution of the extant species of the group.

Using Comparative Biogeography to Retrace the Origins of an Ecosystem: The Case of Four Plants Endemic to the Central Florida Scrub

Premise of research. Plant communities assemble through historical, adaptive, and stochastic processes, with their relative contributions varying among communities. This study traces the origin of the highly threatened Florida scrub ecosystem, a series of relic sand dunes with exceptionally high endemism. We used dated phylogenies with four endemic angiosperms to test two hypotheses on the origins of the Florida scrub: the western hypothesis, which emphasizes the role of mid-Pliocene fragmentation of the xeric belt between southwestern North America and Florida, and the eastern hypothesis, which emphasizes the glacial cycles of the Pleistocene. Methodology. Augmenting existing phylogenies, we sequenced the nuclear internal transcribed spacer region and several plastid loci of species of Prunus (Rosaceae), Polygala (Polygalaceae), Persea (Lauraceae), and Ilex (Aquifoliaceae) to phylogenetically place the Florida scrub endemics and identify their putative sister taxa. We used topology tests, ancestral area reconstruction analyses, and time-calibrated trees to evaluate the western and eastern hypotheses both geographically and temporally. Pivotal results. Results for Polygala and Ilex support the eastern hypothesis for the origin of the central Florida endemics, while data for Persea and Prunus are ambiguous. However, all species show an eastern ancestral distribution to some degree. Molecular dating analyses suggest that extant Ilex opaca populations diverged in the Pleistocene, with older Pliocene origins for the North American Persea clade, Polygala lewtonii, and Prunus geniculata. Conclusions. Taken together, our results support eastern North America as the dominant origin of the plant species of the central Florida scrub. However, contrary to the current eastern hypothesis, molecular dating suggests that the origins of the four endemics predate the last glacial cycles. These results have implications for the age of the Florida scrub itself as well as its component species, suggesting that the Florida scrub or its precursor existed before Pleistocene glaciation. Furthermore, the more ancient age of this assemblage of species than envisioned by most accounts suggests that interspecific interactions in this community are not recently formed (i.e., since the end of the Pleistocene) and may have been established over much longer periods of geological history. These fundamental associations argue for enhanced conservation efforts of both the individual species and the persistent remnants of the scrub community.

Beyond Moa's Ark and Wallace's Line: extralimital distribution of new species of Austronothrus (Acari, Oribatida, Crotoniidae) and the endemicity of the New Zealand oribatid mite fauna.

The genus Austronothrus was previously known from three species recorded only from New Zealand. Austronothrus kinabalu sp. nov. is described from Sabah, Borneo and A. rostralis sp. nov. from Norfolk Island, south-west Pacific. A key to Austronothrus is included. These new species extend the distribution of Austronothrus beyond New Zealand and confirms that the subfamily Crotoniinae is not confined to former Gondwanan landmasses. The distribution pattern of Austronothrus spp., combining Oriental and Gondwanan localities, is indicative of a curved, linear track; consistent with the accretion of island arcs and volcanic terranes around the plate margins of the Pacific Ocean, with older taxa persisting on younger island though localised dispersal within island arc metapopulations. Phylogenetic analysis and an area cladogram are consistent with a broad ancestral distribution of Austronothrus in the Oriental region and on Gondwanan terranes, with subsequent divergence and distribution southward from the Sunda region to New Zealand. This pattern is more complex than might be expected if the New Zealand oribatid fauna was derived from dispersal following re-emergence of land after inundation during the Oligocene (25 mya), as well as if the fauna emanated from endemic, relictual taxa following separation of New Zealand from Gondwana during the Cretaceous (80 mya).

Origin and diversification of the Milla Clade (Brodiaeoideae, Asparagaceae): A Neotropical group of six geophytic genera

The Milla clade currently comprises six genera of geophytic plants distributed from Arizona to Guatemala. Three genera (Behria, Jaimehintonia and Petronymphe) are monotypic while the remaining genera (Bessera, Dandya and Milla) contain from two to ten (Milla) species. Parsimony, Maximum Likelihood and Bayesian Inference analyses were conducted with plastid and nuclear DNA sequences from a total of 181 plants belonging to 15 species in all six genera. Molecular dating was performed under a relaxed clock model. We examined the phylogenetic relationships of the genera and species, estimated origin-divergence times for the clade and genera and determined the ancestral distribution area of the clade by optimizing ancestral areas given current biogeographic distributions. The phylogenetic results suggest that final decisions on limits of the six genera in the Milla clade will have to be established until further taxonomic work is completed for Milla, in particular for the group of populations included under the name M. biflora. The later genus is rendered polyphyletic by other genera of the family. The origin of the Milla clade is estimated at 15.8Ma. Ancestral area of the clade most likely was located in the California Floristic Province and dispersal occurred most likely to the Chihuahuan-Coahuila Plateaus and the Trans-Mexican Volcanic Belt and from there to Baja California and the Sierra Madre del Sur. Two hypotheses that need further testing are proposed to explain complex relationships of genera and polyphyly of Milla, one in relation to fragmentation of populations and pollinator shifts and another suggesting that populations remained in refugia in the Trans-Mexican Volcanic Belt.

Historical biogeography of Aganope, Brachypterum and Derris (Fabaceae, tribe Millettieae): insights into the origins of Palaeotropical intercontinental disjunctions and general biogeographical patterns in Southeast Asia

AbstractAimThe historical biogeography of three Palaeotropical legume genera, Aganope, Brachypterum and Derris, was investigated with the aim of (1) evaluating competing hypotheses on the origins of Palaeotropical intercontinental disjunctions (PIDs), and (2) inferring spatio‐temporal diversification patterns in tropical Southeast Asia.LocationPalaeotropics.MethodsPlastid (trnL–F IGS, psbA–trnH IGS and trnK–matK ORF) and nuclear ribosomal (ITS/5.8S) DNA sequence data, covering the geographical distribution of all three genera, were analysed using an uncorrelated‐rates relaxed molecular clock model. Ancestral areas were reconstructed using a likelihood approach implementing the dispersal–extinction–cladogenesis model (Lagrange) and a Bayesian approach to dispersal–vicariance analysis (S‐DIVA).ResultsA wide ancestral distribution in Africa and Asia was inferred for the Aganope stem and crown groups, with a vicariance event between Africa and Asia in the early Miocene. The Southeast Asian mainland was inferred as the ancestral area for both the Brachypterum and the Derris crown groups. The reconstructions indicated numerous dispersal events westwards to India, and eastwards across Wallace's Line to New Guinea from the middle Miocene onwards. Two dispersal events from Asia to Africa, in the Miocene–Pliocene in Brachypterum and in the Pliocene–Pleistocene in Derris, were deduced.Main conclusionsThe PID in Aganope is likely to be the result of vicariance, caused by climatic deterioration subsequent to the Middle Miocene Climatic Optimum. The inferred PIDs in Brachypterum and Derris in the middle Miocene to Pliocene–Pleistocene are consistent with long‐distance dispersal. The biogeographical patterns of Brachypterum and Derris are similar to patterns identified in other Southeast Asian plant taxa, and highly congruent with geological events in Southeast Asia facilitating dispersal from the early Miocene onwards. Preadaptation to several environmental conditions and habitats including mangrove swamps, and high dispersal capabilities by hydrochory may explain the wide distributions of some species and frequent dispersal across oceanic water bodies separating western and eastern Malesia.

Mitogenomic perspectives into sciaenid fishes' phylogeny and evolution origin in the New World

Sciaenid fishes are widely distributed throughout the coastal waters and estuaries of the world. A total of 23 genera of this family are endemic to the Old World. However, evolutionary relationships among Old World sciaenid fishes and their origin have remained unresolved despite their diversity and importance. Besides, hypotheses that explain the origin and biogeographical distribution of sciaenid fishes are controversial. In this study, the complete mitochondrial genome sequences of seven representative sciaenid species were determined and a well-resolved tree was recovered. This new timescale demonstrated that the sciaenid originated during the late Jurassic to early Cretaceous Period. The estimated origin time of sciaenid fish is 208Mya, and the origin of Old World sciaenid is estimated at 126Mya. Reconstruction of ancestral distributions indicated a plesiomorphic distribution and center of origin in the New World, with at least one lineage subsequently dispersed to the Old World. Moreover, we conclude that the common ancestors of Old World sciaenid fishes were derived from species of New World.

Species delimitation, patterns of diversification and historical biogeography of the Neotropical frog genus Adenomera (Anura, Leptodactylidae)

AbstractAimFor many taxa, inaccuracy of species boundaries and distributions hampers inferences about diversity and evolution. This is particularly true in the Neotropics where prevalence of cryptic species has often been demonstrated. The frog genus Adenomera is suspected to harbour many more species than the 16 currently recognized. These small terrestrial species occur in Amazonia, Atlantic Forest (AF), and in the open formations of the Dry Diagonal (DD: Chaco, Cerrado and Caatinga). This widespread and taxonomically complex taxon provides a good opportunity to (1) test species boundaries, and (2) investigate historical connectivity between Amazonia and the AF and associated patterns of diversification.LocationTropical South America east of the Andes.MethodsWe used molecular data (four loci) to estimate phylogenetic relationships among 320 Adenomera samples. These results were integrated with other lines of evidence to propose a conservative species delineation. We subsequently used an extended dataset (seven loci) and investigated ancestral area distributions, dispersal–vicariance events, and the temporal pattern of diversification within Adenomera.ResultsOur conservative delineation identified 31 Confirmed Candidate Species (four remaining unconfirmed) representing a 94% increase in species richness. The biogeographical analysis suggested an Amazonian origin of Adenomera with as many as three dispersals to the DD and one to the AF during the Miocene. These dispersals were associated with habitat shifts from forest towards open habitats.Main conclusionsThe DD played a major role in the history of Adenomera in limiting dispersal and favouring diversification of open‐habitat lineages. Moreover, a forest bridge during the Miocene Climatic Optimum may have permitted dispersal from Amazonia towards the AF and subsequent diversification. Uncovering species boundaries and distributions might drastically change inferences based on currently perceived distribution patterns.

Global biogeography of scaly tree ferns (Cyatheaceae): evidence for Gondwanan vicariance and limited transoceanic dispersal.

AimScaly tree ferns, Cyatheaceae, are a well-supported group of mostly tree-forming ferns found throughout the tropics, the subtropics and the south-temperate zone. Fossil evidence shows that the lineage originated in the Late Jurassic period. We reconstructed large-scale historical biogeographical patterns of Cyatheaceae and tested the hypothesis that some of the observed distribution patterns are in fact compatible, in time and space, with a vicariance scenario related to the break-up of Gondwana.LocationTropics, subtropics and south-temperate areas of the world.MethodsThe historical biogeography of Cyatheaceae was analysed in a maximum likelihood framework using Lagrange. The 78 ingroup taxa are representative of the geographical distribution of the entire family. The phylogenies that served as a basis for the analyses were obtained by Bayesian inference analyses of mainly previously published DNA sequence data using MrBayes. Lineage divergence dates were estimated in a Bayesian Markov chain Monte Carlo framework using beast.ResultsCyatheaceae originated in the Late Jurassic in either South America or Australasia. Following a range expansion, the ancestral distribution of the marginate-scaled clade included both these areas, whereas Sphaeropteris is reconstructed as having its origin only in Australasia. Within the marginate-scaled clade, reconstructions of early divergences are hampered by the unresolved relationships among the Alsophila, Cyathea and Gymnosphaera lineages. Nevertheless, it is clear that the occurrence of the Cyathea and Sphaeropteris lineages in South America may be related to vicariance, whereas transoceanic dispersal needs to be inferred for the range shifts seen in Alsophila and Gymnosphaera.Main conclusionsThe evolutionary history of Cyatheaceae involves both Gondwanan vicariance scenarios as well as long-distance dispersal events. The number of transoceanic dispersals reconstructed for the family is rather few when compared with other fern lineages. We suggest that a causal relationship between reproductive mode (outcrossing) and dispersal limitations is the most plausible explanation for the pattern observed.

Neotropical diversification seen through glassfrogs

AbstractAimWe used frogs of the clade Allocentroleniae (Centrolenidae + Allophrynidae; c. 170 species endemic to Neotropical rain forests) as a model system to address the historical biogeography and diversification of Neotropical rain forest biotas.LocationNeotropical rain forests.MethodsWe used an extensive taxon (109 species) and gene (seven nuclear and three mitochondrial genes) sampling to estimate phylogenetic relationships, divergence times, ancestral area distributions, dispersal–vicariance events, and the temporal pattern of diversification rate.ResultsThe Allocentroleniae started to diversify in the Eocene in South America and by the early Miocene were present in all major Neotropical rain forests except in Central America, which was colonized through 11 late range expansions. The initial uplifts of the Andes during the Oligocene and early Miocene, as well as marine incursions in the lowlands, are coincidental with our estimates of the divergence times of most clades of Allocentroleniae. Clades with broad elevational distributions occupy more biogeographical areas. Most dispersals involve the Andes as a source area but the majority were between the Central and the Northern Andes, suggesting that the Andes did not play a major role as a species pump for the lowlands. The diversification of glassfrogs does not follow a south‐to‐north pattern of speciation for Andean clades, and the establishment of a transcontinental Amazon drainage system is coincidental in time with the isolation of the Atlantic Forest glassfrogs. Diversification analyses indicated that a model of constantly increasing diversity best fits the data, compatible with the ‘evolutionary museum’ hypothesis or ‘ancient cradle’ hypothesis.Main conclusionsOur work illustrates how the different geological and climatic historical events of the Neotropics shaped, at different levels of the phylogeny, the diversity of a species‐rich clade, highlighting the importance of studying large evolutionary radiations at a continental scale.

Adaptive radiation, correlated and contingent evolution, and net species diversification in Bromeliaceae

We present an integrative model predicting associations among epiphytism, the tank habit, entangling seeds, C3 vs. CAM photosynthesis, avian pollinators, life in fertile, moist montane habitats, and net rates of species diversification in the monocot family Bromeliaceae. We test these predictions by relating evolutionary shifts in form, physiology, and ecology to time and ancestral distributions, quantifying patterns of correlated and contingent evolution among pairs of traits and analyzing the apparent impact of individual traits on rates of net species diversification and geographic expansion beyond the ancestral Guayana Shield. All predicted patterns of correlated evolution were significant, and the temporal and spatial associations of phenotypic shifts with orogenies generally accorded with predictions. Net rates of species diversification were most closely coupled to life in fertile, moist, geographically extensive cordilleras, with additional significant ties to epiphytism, avian pollination, and the tank habit. The highest rates of net diversification were seen in the bromelioid tank-epiphytic clade (Dcrown=1.05My−1), associated primarily with the Serra do Mar and nearby ranges of coastal Brazil, and in the core tillandsioids (Dcrown=0.67My−1), associated primarily with the Andes and Central America. Six large-scale adaptive radiations and accompanying pulses of speciation account for 86% of total species richness in the family. This study is among the first to test a priori hypotheses about the relationships among phylogeny, phenotypic evolution, geographic spread, and net species diversification, and to argue for causality to flow from functional diversity to spatial expansion to species diversity.