New Glyptodont from the Codore Formation (Pliocene), Falcón State, Venezuela, its relationship with theAsterostemma problem, and the paleobiogeography of the Glyptodontinae

One of the basal Glyptodontidae groups is represented by the Propalaehoplophorinae (late Oligocene — middle Miocene), whose genera (Propalaehoplophorus, Eucinepeltus, Metopotoxus, Cochlops, andAsterostemma) were initially recognized in Argentinian Patagonia. Among these,Asterostemma was characterized by its wide latitudinal distribution, ranging from southernmost (Patagonia) to northernmost (Colombia, Venezuela) South America. However, the generic assignation of the Miocene species from Colombia and Venezuela (A.? acostae, A. gigantea, andA. venezolensis) was contested by some authors, who explicitly accepted the possibility that these species could correspond to a new genus, different from those recognized in southern areas. A new comparative study of taxa from Argentinian Patagonia, Colombia and Venezuela (together with the recognition of a new genus and species for the Pliocene of the latter country) indicates that the species in northern South America are not Propalaehoplophorinae, but represent the first stages in the cladogenesis of the Glyptodontinae glyptodontids, the history of which was heretofore restricted to the late Miocene — early Holocene of southernmost South America. Accordingly, we propose the recognition of the new genusBoreostemma for the species from northern South America and the restriction ofAsterostemma to the Miocene of Patagonia. Thus, the available data indicate that the Glyptodontinae would in fact have arisen in the northernmost regions of this continent. Their arrival to more southerly areas coincides with the acme of the “Age of Southern Plains”. The Propalaehoplophorinae are geographically restricted to Patagonia.     

Coral reefs as drivers of cladogenesis: expanding coral reefs, cryptic extinction events, and the development of biodiversity hotspots

Diversification rates within four conspicuous coral reef fish families (Labridae, Chaetodontidae, Pomacentridae and Apogonidae) were estimated using Bayesian inference. Lineage through time plots revealed a possible late Eocene/early Oligocene cryptic extinction event coinciding with the collapse of the ancestral Tethyan/Arabian hotspot. Rates of diversification analysis revealed elevated cladogenesis in all families in the Oligocene/Miocene. Throughout the Miocene, lineages with a high percentage of coral reef-associated taxa display significantly higher net diversification rates than expected. The development of a complex mosaic of reef habitats in the Indo-Australian Archipelago (IAA) during the Oligocene/Miocene appears to have been a significant driver of cladogenesis. Patterns of diversification suggest that coral reefs acted as a refuge from high extinction, as reef taxa are able to sustain diversification at high extinction rates. The IAA appears to support both cladogenesis and survival in associated lineages, laying the foundation for the recent IAA marine biodiversity hotspot.     

INVESTIGATING PROCESSES OF NEOTROPICAL RAIN FOREST TREE DIVERSIFICATION BY EXAMINING THE EVOLUTION AND HISTORICAL BIOGEOGRAPHY OF THE PROTIEAE (BURSERACEAE)

Andean uplift and the collision of North and South America are thought to have major implications for the diversification of the Neotropical biota. However, few studies have investigated how these geological events may have influenced diversification. We present a multilocus phylogeny of 102 Protieae taxa (73% of published species), sampled pantropically, to test hypotheses about the relative importance of dispersal, vicariance, habitat specialization, and biotic factors in the diversification of this ecologically dominant tribe of Neotropical trees. Bayesian fossil‐calibrated analyses date the Protieae stem at 55 Mya. Biogeographic analyses reconstruct an initial late Oligocene/early Miocene radiation in Amazonia for Neotropical Protieae, with several subsequent late Miocene dispersal events to Central America, the Caribbean, Brazil's Atlantic Forest, and the Chocó. Regional phylogenetic structure results indicate frequent dispersal among regions throughout the Miocene and many instances of more recent regional in situ speciation. Habitat specialization to white sand or flooded soils was common, especially in Amazonia. There was one significant increase in diversification rate coincident with colonization of the Neotropics, followed by a gradual decrease consistent with models of diversity‐dependent cladogenesis. Dispersal, biotic interactions, and habitat specialization are thus hypothesized to be the most important processes underlying the diversification of the Protieae.     

A new Miocene vertebrate assemblage from the Río Yuca Formation (Venezuela) and the northernmost record of typical Miocene mammals of high latitude (Patagonian) affinities in South America

Geological explorations of the basal beds of the Río Yuca Formation (Tucupido region, Portuguesa State, western Venezuela) resulted in the recognition of a new vertebrate assemblage that includes eight taxa: the toxodont cf. Adinotherium, a Peltephilidae armadillo, the freshwaters fishes Platysilurus and Phractocephalus, the caiman Purussaurus, an indeterminate dolphin, turtles, and the previously recognized sloth Pseudoprepotherium venezuelanum. When compared with the higher latitudes faunas of Argentina and Chile, the presence of cf. Adinotherium and peltephilids in the Rio Yuca Formation is consistent, but not conclusive, with a Santacrucian to Frisian SALMA age. The associated fauna, as well recent apatite fission track analysis, indicates that the Río Yuca assemblage is more likely younger in age, specifically Middle to Late Miocene. So far, the Miocene localities of the northern part of South America have provided a less prolific fossil record compared to the southern part of the continent (e.g., Santacrucian and Friasian faunas of Patagonia), but the present work documents the surprising occurrence of two taxa (Peltephilidae and Nesodontinae) common in southern high latitude faunas of South America, implying the persistence of the Santacrucian-Friasian genus Adinotherium in younger strata from northern South America, and that peltephilids were much more widespread during the Miocene than previously recognized. The presence of these common Patagonian taxa (Peltephilidae and Nesodontinae) in Río Yuca also supports the hypothesis of prior researchers for the existence of biogeographical connections between the northern and southern portions of South America during the Late Oligocene or Early Miocene, which facilitated faunal interchange between the two regions. Finally, the biogeographical affinities of the freshwater fishes and the giant caiman (Purussaurus) indicate close relationships of the Tucupido region with the ancestral distribution of the Orinocoan–Amazonian drainage system.     

A Paleogeographic Overview of Tropical Fossil Sloths: Towards an Understanding of the Origin of Extant Suspensory Sloths?

Modern sloths are among the more characteristic mammals of South and Central American faunas. Recent discovery in four Paleogene, 22 Neogene, and dozens of Pleistocene fossiliferous localities in the tropics has revealed an unexpected paleobioversity constituted by some 81 fossil sloth species. Probably originating in southern South America near the Eocene/Oligocene transition, sloths were represented in the tropics during the late Oligocene by Pseudoglyptodon, Mylodontidae, and Megalonychidae. The latter occupied the West Indies between at least the late early Miocene and late Pleistocene, and two mylodontid clades, Octodontobradyinae and Urumacotheriinae, were characteristic of Amazonian localities from the Colhuehuapian and the Laventan periods, respectively, until the end of the Miocene. Megatheriinae and Nothrotheriidae appeared during the middle Miocene, colonizing the tropics and then North America, where Mylodontidae and Megalonychidae had already been present since the early late Miocene. Nothrotheriids are more abundant and diversified during the late Miocene in the tropics than in southern South America. Remains closely related to either of the modern sloths are absent from the fossil record, including those in the tropics. The characteristic suspensory posture of Bradypus and Choloepus appeared independently and likely after the Miocene epoch, and thus well after the hypothesized split suggested by molecular studies of the respective clades of these genera. Given their current widespread distribution in and reliance on the tropics, prospecting efforts for the direct fossil kin of suspensory sloths should concentrate on deposits in the Amazonian region, as this area has shown promise in producing fossil sloths.     

A rapid diversification of rainforest trees (Guatteria; Annonaceae) following dispersal from Central into South America

Several recent studies have suggested that a substantial portion of today's plant diversity in the Neotropics has resulted from the dispersal of taxa into that region rather than vicariance, but more data are needed to substantiate this claim. Guatteria (Annonaceae) is, with 265 species, the third largest genus of Neotropical trees after Inga (Fabaceae) and Ocotea (Lauraceae), and its widespread distribution and frequent occurrence makes the genus an excellent model taxon to study diversification patterns. This study reconstructed the phylogeny of Guatteria and inferred three major biogeographical events in the history of the genus: (1) a trans-oceanic Miocene migration from Central into South America before the closing of the Isthmus of Panama; (2) a major diversification of the lineage within South America; and (3) several migrations of South American lineages back into Central America via the closed Panamanian land bridge. Therefore, Guatteria is not an Amazonian centred-genus sensu Gentry but a major Miocene diversification that followed its dispersal into South America. This study provides further evidence that migration into the Neotropics was an important factor in the historical assembly of its biodiversity. Furthermore, it is shown that phylogenetic patterns are comparable to those found in Ocotea and Inga and that a closer comparison of these genera is desirable.     

Early Oligocene fruits and leaves of Burretiodendron (Malvaceae s.l.) from South China

The genus Burretiodendron Rehder is currently endemic to an area near the China–Vietnam border and the limestone mountains of Thailand and Myanmar. The fossil records of this genus were previously found only from the Miocene of Yunnan, Southwest China, and the Oligocene of Guangxi, South China. Here, we describe fossil fruits and associated leaves of Burretiodendron, which were discovered in the lower Oligocene Shangcun Formation of the Maoming Basin, Guangdong, South China. Morphological comparison with extant and fossil Burretiodendron taxa indicates that fruit fossils belong to the species Burretiodendron parvifructum J. Lebreton Anberrée & Z. K. Zhou. This is one of the earliest fossil records of the genus, providing additional evidence for the early biogeographic history of this genus and supporting the inference that the genus originated in South China. According to the habitat conditions of modern species, we speculate that there were limestone mountains around the Maoming Basin in the early Oligocene.     

From pampa to puna: Biogeography and diversification of a group of Neotropical obligate grassland birds (Anthus: Motacillidae)

The evolution of Neotropical birds of open landscapes remains largely unstudied. We investigate the diversification and biogeography of a group of Neotropical obligate grassland birds (Anthus: Motacillidae). We use a multilocus phylogeny of 22 taxa of Anthus to test the hypothesis that these birds radiated contemporaneously with the development of grasslands in South America. We employ the R package DDD to analyze the dynamics of Anthus diversification across time in Neotropical grasslands, explicitly testing for shifts in dynamics associated with the Miocene development of grasslands, the putative Pleistocene expansion of arid lowland biomes, and Pleistocene sundering of Andean highland grasslands. A lineage‐through‐time plot revealed increases in the number of lineages, and DDD detected shifts to a higher clade‐level carrying capacity during the late Miocene, indicating an early burst of diversification associated with grassland colonization. However, we could not corroborate the shift using power analysis, probably reflecting the small number of tips in our tree. We found evidence of a divergence at ~1 Mya between northern and southern Amazonian populations of Anthus lutescens, countering Haffer's idea of Pleistocene expansion of open biomes in the Amazon Basin. We used BioGeoBears to investigate ancestral areas and directionality of colonization of Neotropical grasslands. Members of the genus diversified into, out of, and within the Andes, within‐Andean diversification being mostly Pleistocene in origin.     

A molecular phylogeny of warbling-finches (Poospiza): paraphyly in a neotropical Emberizid genus

We investigated the phylogenetic relationships of 12 species within a single genus of neotropical passerine (Poospiza) using 849 bp (283 codons) of the cytochrome b mitochondrial gene. We further explored evolutionary affinities of these taxa using sequence from an additional 47 thraupine (tanagers) and 7 emberizine (sparrows and buntings) genera, members of the predominantly New World family Emberizidae. Poospiza have traditionally been considered part of the emberizine radiation. However, our analyses suggest that members of this genus are more closely related to some thraupine lineages than they are to the other neotropical emberizine genera included in our study (Atlapetes, Embernagra, Melopyrrha, Phrygilus, Saltatricula, Tiaris). Although member taxa are closely related, the genus Poospiza appears to be paraphyletic with representatives of 6 thraupine genera (Cnemoscopus, Cypsnagra, Hemispingus, Nephelornis, Pyrrhocoma, Thylpopsis) interspersed among four well-supported Poospiza clades. The majority of species within this Poospiza-thraupine clade have geographic ranges that are exclusive to, or partially overlap with, the Andes Mountains. It is probable that these mountains have played an important role in driving cladogenesis within this group. Sequence divergence (transversions only; mean 4.7+/-1.3%) within the clade suggests that much of this diversification occurred within the late Miocene and Pliocene, a period coincident with major orogenic activity in central-western South America.     

Major Radiations in the Evolution of Caviid Rodents: Reconciling Fossils,Ghost Lineages,and Relaxed Molecular Clocks

Background

Caviidae is a diverse group of caviomorph rodents that is broadly distributed in South America and is divided into three highly divergent extant lineages: Caviinae (cavies), Dolichotinae (maras), and Hydrochoerinae (capybaras). The fossil record of Caviidae is only abundant and diverse since the late Miocene. Caviids belongs to Cavioidea sensu stricto (Cavioidea s.s.) that also includes a diverse assemblage of extinct taxa recorded from the late Oligocene to the middle Miocene of South America (“eocardiids”).

Results

A phylogenetic analysis combining morphological and molecular data is presented here, evaluating the time of diversification of selected nodes based on the calibration of phylogenetic trees with fossil taxa and the use of relaxed molecular clocks. This analysis reveals three major phases of diversification in the evolutionary history of Cavioidea s.s. The first two phases involve two successive radiations of extinct lineages that occurred during the late Oligocene and the early Miocene. The third phase consists of the diversification of Caviidae. The initial split of caviids is dated as middle Miocene by the fossil record. This date falls within the 95% higher probability distribution estimated by the relaxed Bayesian molecular clock, although the mean age estimate ages are 3.5 to 7 Myr older. The initial split of caviids is followed by an obscure period of poor fossil record (refered here as the Mayoan gap) and then by the appearance of highly differentiated modern lineages of caviids, which evidentially occurred at the late Miocene as indicated by both the fossil record and molecular clock estimates.

Conclusions

The integrated approach used here allowed us identifying the agreements and discrepancies of the fossil record and molecular clock estimates on the timing of the major events in cavioid evolution, revealing evolutionary patterns that would not have been possible to gather using only molecular or paleontological data alone.     

A new species of Ameripodius (Aves: Galliformes: Quercymegapodiidae) from the Lower Miocene of France

The Quercymegapodiidae, primitive galliforms resembling recent megapodes, have been described from the Upper Eocene of Quercy, France. They have also been identified in the Upper Oligocene–Lower Miocene of Brazil, where they are represented by the genus Ameripodius Alvarenga. A new species of this genus, Ameripodius alexis sp. nov., from the Lower Miocene of France, is described here. The occurrence of the same genus on both sides of the Atlantic Ocean emphasizes the similarities between South American and European avifaunas during the early Tertiary. New discoveries indicate that a similar avifauna was also present in North America, and that a characteristic association of taxa can be defined for the group that includes South America, North America and Eurasia. However, so far as is known, the same avifauna does not occur in contemporaneous African avifaunas.     

Arrival and diversification of caviomorph rodents and platyrrhine primates in South America

Platyrrhine primates and caviomorph rodents are clades of mammals that colonized South America during its period of isolation from the other continents, between 100 and 3 million years ago (Mya). Until now, no molecular study investigated the timing of the South American colonization by these two lineages with the same molecular data set. Using sequences from three nuclear genes (ADRA2B, vWF, and IRBP, both separate and combined) from 60 species, and eight fossil calibration constraints, we estimated the times of origin and diversification of platyrrhines and caviomorphs via a Bayesian relaxed molecular clock approach. To account for the possible effect of an accelerated rate of evolution of the IRBP gene along the branch leading to the anthropoids, we performed the datings with and without IRBP (3768 sites and 2469 sites, respectively). The time window for the colonization of South America by primates and by rodents is demarcated by the dates of origin (upper bound) and radiation (lower bound) of platyrrhines and caviomorphs. According to this approach, platyrrhine primates colonized South America between 37.0 +/- 3.0 Mya (or 38.9 +/- 4.0 Mya without IRBP) and 16.8 +/- 2.3 (or 20.1 +/- 3.3) Mya, and caviomorph rodents between 45.4 +/- 4.1 (or 43.7 +/- 4.8) Mya and 36.7 +/- 3.7 (or 35.8 +/- 4.3) Mya. Considering both the fossil record and these molecular datings, the favored scenarios are a trans-Atlantic migration of primates from Africa at the end of the Eocene or beginning of the Oligocene, and a colonization of South America by rodents during the Middle or Late Eocene. Based on our nuclear DNA data, we cannot rule out the possibility of a concomitant arrival of primates and rodents in South America. The caviomorphs radiated soon after their arrival, before the Oligocene glaciations, and these early caviomorph lineages persisted until the present. By contrast, few platyrrhine fossils are known in the Oligocene, and the present-day taxa are the result of a quite recent, Early Miocene diversification.     

Systematics and evolution of the Australian knob-tail geckos (Nephrurus, Carphodactylidae, Gekkota): plesiomorphic grades and biome shifts through the Miocene

Clades that predate the origin of biomes that they inhabit provide unique opportunities to examine both when major environmental transitions occurred, and how lineages adapted to these changes. The isolated island continent Australia has undergone a profound environmental transition through the Miocene, from relatively mesic to predominantly arid; however, we have much to learn about both the timing of this change, and how organisms may have responded to it. The family Carphodactylidae is an ancient Gondwanan group of geckos that occurs across all major Australian biomes. A multilocus (ND2, Rag-1, c-mos) phylogenetic and dating analysis of the most ecologically diverse clade within this group, the genus Nephrurus (sensuBauer, 1990) reveals that two of three morphological taxa historically recognized (the 'spiny knob-tails' and 'Underwoodisaurus') are relatively species depauperate, pleisomorphic basal grades that diversified through the late Oligocene and early Miocene, and are now absent from most of the arid biome. Based on their deep divergence and morphological distinctiveness we recognize two lineages (milii and sphyrurus) as monotypic genera, the later of which is named herein (Uvidicolus nov. gen). In contrast, a third morphological group, the 'smooth knob-tails,' is a monophyletic group of five exclusively arid zone burrowing species that has radiated relatively recently (mid-Miocene). Our phylogeny indicates that successful colonization of this novel and challenging biome by Nephrurus correlates with an initial shift to terrestriality and adaptation to at least seasonally arid conditions around the early Miocene, and the eventual evolution and subsequent mid-Miocene radiation of a lineage specialized for burrowing.     

<Emphasis Type="Italic">Gryposuchus</Emphasis> (Crocodylia,Gavialoidea) from the early Miocene of Venezuela

Here, a fragment of a mandible recently discovered in the Cerro Zamuro site (Castillo Formation, Lara State, northwestern Venezuela) is assigned to the giant gavialoid Gryposuchus. This specimen, recovered from putative brackish environments of the early Miocene (~18 Ma) age, is unequivocally the earliest record of the genus in South America. Gryposuchus, together with the other gryposuchine previously recognized from the Castillo Formation, Siquisiquesuchus venezuelensis, increases the early Miocene taxonomic diversity of the group in the northern Neotropics. This new information from the Castillo Formation supports the conclusion that early gryposuchine evolutionary stages were in coastal, shallow marine or brackish environments, while the presence of some genera, such as Gryposuchus, in middle to late Miocene freshwater environments, is secondary habitat colonization late in the evolution of the clade. Freshwater colonization is probably the result of the gradual adaptation of early marine-adapted gryposuchines to the extensive estuarine-like environments of northern South America lowlands associated with marine transgressions that systematically occurred during the middle Eocene to early Oligocene. This new record is evidence of the wide chronological distribution of Gryposuchus in northern South America, highlighting the importance of this area as the center of origin and radiation of this successful Miocene gavialoid.     

A Molecular Phylogeny of Warbling-Finches (Poospiza): Paraphyly in a Neotropical Emberizid Genus

We investigated the phylogenetic relationships of 12 species within a single genus of neotropical passerine (Poospiza) using 849 bp (283 codons) of the cytochrome b mitochondrial gene. We further explored evolutionary affinities of these taxa using sequence from an additional 47 thraupine (tanagers) and 7 emberizine (sparrows and buntings) genera, members of the predominantly New World family Emberizidae. Poospiza have traditionally been considered part of the emberizine radiation. However, our analyses suggest that members of this genus are more closely related to some thraupine lineages than they are to the other neotropical emberizine genera included in our study (Atlapetes, Embernagra, Melopyrrha, Phrygilus, Saltatricula, Tiaris). Although member taxa are closely related, the genus Poospiza appears to be paraphyletic with representatives of 6 thraupine genera (Cnemoscopus, Cypsnagra, Hemispingus, Nephelornis, Pyrrhocoma, Thylpopsis) interspersed among four well-supported Poospiza clades. The majority of species within this Poospiza–thraupine clade have geographic ranges that are exclusive to, or partially overlap with, the Andes Mountains. It is probable that these mountains have played an important role in driving cladogenesis within this group. Sequence divergence (transversions only; mean 4.7 ± 1.3%) within the clade suggests that much of this diversification occurred within the late Miocene and Pliocene, a period coincident with major orogenic activity in central-western South America.     

The Gondwanan and South American Episodes: Two Major and Unrelated Moments in the History of the South American Mammals

The first steps in the history of South American mammals took place ca. 130 Ma., when the South American plate, still connected to the Antarctic Peninsula, began to drift away from the African-Indian plate. Most of the Mesozoic history of South American mammals is still unknown, and we only have a few enigmatic taxa (i.e., a Jurassic Australosphenida and an Early Cretaceous Prototribosphenida) that pose more evolutionary and biogeographic questions than answers. The best-known Mesozoic, South American land-mammal fossils are from Late Cretaceous Patagonian beds. These fossils represent the last survivors of non- and pre-tribosphenic Pangaean lineages, all of them with varying endemic features: some with few advanced features (e.g., ?Eutriconodonta and “Symmetrodonta”), some very diversified as endemic groups (e.g., ?Docodonta Reigitheriidae), and others representing vicariant types of well known Laurasian Mesozoic lineages (e.g., Gondwanatheria as vicariant of Multituberculata). These endemic mammals lived as relicts (although advanced) of pangeic lineages when a primordial South American continent was still connected to the Antarctic Peninsula and, at the northern extreme, near the North American Plate. By the beginning of the Late Cretaceous, the volcanic and diastrophic processes that finally led to the differentiation of the Caribbean region and Central America built up transient geographic connections that permitted the initiation of an overland inter-American exchange that included, for example, dinosaurian titanosaurs from South America and hadrosaurs from North America. The immigration of other vertebrates followed the same route, for example, polydolopimorphian marsupials. These marsupials were assumed to have differentiated in South America prior to new discoveries from the North American Late Cretaceous. The complete extinction of endemic South American Mesozoic mammals by the Late Cretaceous-Early Paleocene, and the subsequent and in part coetaneous immigration of North American therians, respectively, represent two major moments in the history of South American mammals: a Gondwanan Episode and a South American Episode. The Gondwanan Episode was characterized by non- and pre-tribosphenic mammal lineages that descended from the Pangeic South American stage (but already with a pronounced Gondwanan accent, and wholly extinguished during the Late Cretaceous-Early Paleocene span). The South American Episode, in turn, was characterized only by therian mammals, mostly emigrated from the North American continent and already with a South American accent obtained through isolation. The southernmost extreme of South America (Patagonia) remained connected to the present Antarctic Peninsula at least up until about 30 Ma., and both provided the substratum where the primordial cladogenesis of “South American” mammals occurred. The resulting cladogenesis of South American therian mammals followed Gould's motto: early experimentation, later standardization. That is to say, early cladogenesis engendered a great variety of taxa with scarce morphological differentiation. After this early cladogenesis (Late Eocene-Early Oligocene), the variety of taxa became reduced, but each lineage became clearly recognizable distinctive by a constant morphologic pattern. At the same time, those mammals that underwent the “early experimentation” were part of communities dominated by archaic lineages (e.g., brachydont types among the native “ungulates”), whereas the subsequent communities were dominated by mammals of markedly “modern” stamp (e.g., protohypsodont types among the native “ungulates”). The Gondwanan and South American Episodes were separated by a critical latest Cretaceous-earliest Paleocene hiatus, it is as unknown as it is important in which South American land-mammal communities must have experienced extinction of the Gondwanan mammals and the arrival and radiation of the North American marsupials and placentals (with the probable exception of the xenarthrans, whose biogeographic origin is still unclear).     

First fossil fruits of Elaeocarpus (Elaeocarpaceae) in East Asia: Implications for phytogeography and paleoecology

The genus Elaeocarpus contains approximately 360 species and occurs in mesic forest communities from India, through to China, Southeast Asia, New Guinea, Australia, and New Caledonia. Elaeocarpus fossils are best known from the Eocene to the Miocene of Australia and the late Pliocene–early Pleistocene of India, but have not been documented from East Asia before. Here we describe six new species of Elaeocarpus, E. nanningensis sp. nov. from the late Oligocene Yongning Formation of the Nanning Basin, E. presikkimensis sp. nov. from the Miocene Erzitang Formation of the Guiping Basin, E. prerugosus sp. nov., E. prelacunosus sp. nov., E. preserratus sp. nov., and E. preprunifolioides sp. nov. from the late Miocene Foluo Formation of the Nankang Basin in Guangxi, South China. This is the first reliable report for the genus occurring in East Asia, and the fossils indicate that Elaeocarpus had colonized this region by the late Oligocene and represented by a morphologically diverse group of species by the late Miocene. This sheds new insights into the timing and migration patterns of the genus in East Asia. Elaeocarpus is typically a rainforest genus occurring in mesic forests. Based on the habitat of their morphologically similar modern relatives we propose that these three sedimentary basins were warm and wet adjacent to mountainous regions with the evergreen or rain forests during the late Oligocene to Miocene.     

Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot

Aim East Africa is one of the most biologically diverse regions, especially in terms of endemism and species richness. Hypotheses put forward to explain this high diversity invoke a role for forest refugia through: (1) accumulation of new species due to radiation within refugial habitats, or (2) retention of older palaeoendemic species in stable refugia. We tested these alternative hypotheses using data for a diverse genus of East African forest chameleons, Kinyongia. Location East Africa. Methods We constructed a dated phylogeny for Kinyongia using one nuclear and two mitochondrial markers. We identified areas of high phylogenetic diversity (PD) and evolutionary diversity (ED), and mapped ancestral areas to ascertain whether lineage diversification could best be explained by vicariance or dispersal. Results Vicariance best explains the present biogeographic patterns, with divergence between three major Kinyongia clades (Albertine Rift, southern Eastern Arc, northern Eastern Arc) in the early Miocene/Oligocene (> 20 Ma). Lineage diversification within these clades pre‐dates the Pliocene (> 6 Ma). These dates are much older than the Plio‐Pleistocene climatic shifts associated with cladogenesis in other East African taxa (e.g. birds), and instead point to a scenario whereby palaeoendemics are retained in refugia, rather than more recent radiations within refugia. Estimates of PD show that diversity was highest in the Uluguru, Nguru and East Usambara Mountains and several lineages (from Mount Kenya, South Pare and the Uluguru Mountains) stand out as being evolutionarily distinct as a result of isolation in forest refugia. PD was lower than expected by chance, suggesting that the phylogenetic signal is influenced by an unusually low number of extant lineages with long branch lengths, which is probably due to the retention of palaeoendemic lineages. Main conclusions The biogeographic patterns associated with Kinyongia are the result of long evolutionary histories in isolation. The phylogeny is dominated by ancient lineages whose origins date back to the early Miocene/Oligocene as a result of continental wide forest fragmentation and contraction due to long term climatic changes in Africa. The maintenance of palaeoendemic lineages in refugia has contributed substantially to the remarkably high biodiversity of East Africa.     

Large-scale patterns of diversification in the widespread legume genus Senna and the evolutionary role of extrafloral nectaries

Unraveling the diversification history of old, species-rich and widespread clades is difficult because of extinction, undersampling, and taxonomic uncertainty. In the context of these challenges, we investigated the timing and mode of lineage diversification in Senna (Leguminosae) to gain insights into the evolutionary role of extrafloral nectaries (EFNs). EFNs secrete nectar, attracting ants and forming ecologically important ant-plant mutualisms. In Senna, EFNs characterize one large clade (EFN clade), including 80% of its 350 species. Taxonomic accounts make Senna the largest caesalpinioid genus, but quantitative comparisons to other taxa require inferences about rates. Molecular dating analyses suggest that Senna originated in the early Eocene, and its major lineages appeared during early/mid Eocene to early Oligocene. EFNs evolved in the late Eocene, after the main radiation of ants. The EFN clade diversified faster, becoming significantly more species-rich than non-EFN clades. The shift in diversification rates associated with EFN evolution supports the hypothesis that EFNs represent a (relatively old) key innovation in Senna. EFNs may have promoted the colonization of new habitats appearing with the early uplift of the Andes. This would explain the distinctive geographic concentration of the EFN clade in South America.     

Marine incursions,cryptic species and ecological diversification in Amazonia: the biogeographic history of the croaker genus Plagioscion (Sciaenidae)

Aim We propose a phylogenetic hypothesis for the marine‐derived sciaenid genus Plagioscion in the context of geomorphology and adaptation to freshwaters of South America, and assess the extent to which contemporary freshwater hydrochemical gradients influence diversification within a widely distributed Plagioscion species, Plagioscion squamosissimus. Location Amazon Basin and South America. Methods Using nuclear and mitochondrial DNA sequence data, phylogenetic analyses were conducted on the five nominal Plagioscion species, together with representatives from Pachyurus and Pachypops, using character and model‐based methods. Genealogical relationships and population genetic structure of 152 P. squamosissimus specimens sampled from the five major rivers and three hydrochemical settings/‘colours’ (i.e. white, black and clear water) of the Amazon Basin were assessed. Results Phylogenetic analyses support the monophyly of Plagioscion in South America and identify two putative cryptic species of Plagioscion. Divergence estimates suggest that the Plagioscion ancestor invaded South America via a northern route during the late Oligocene to early Miocene. Within P. squamosissimus a strong association of haplotype and water colour was observed, together with significant population structure detected between water colours. Main conclusions Our analyses of Plagioscion are consistent with a biogeographic scenario of early Miocene marine incursions into South America. Based on our phylogenetic results, the fossil record, geomorphological history and distributional data of extant Plagioscion species, we propose that marine incursions into western Venezuela between the late Oligocene and early Miocene were responsible for the adaptation to freshwaters in Plagioscion species. Following the termination of the marine incursions during the late Miocene and the establishment of the modern Amazon River, Plagioscion experienced a rapid diversification. Plagioscion squamosissimus arose during that time. The formation of the Amazon River probably facilitated population and range expansions for this species. Further, the large‐scale hydrochemical gradients within the Amazon Basin appear to be acting as ecological barriers maintaining population discontinuities in P. squamosissimus even in the face of gene flow. Our results highlight the importance of divergent natural selection through time in the generation and maintenance of sciaenid diversity in Amazonia.