New species of ternstroemia (Theaceae) from the Guayana Highland

Three new species ofTernstroemia are described and illustrated. Two of these,Ternstroemia aracae andT., prancei, are high-elevation taxa endemic to Serra Aracá. Brazil. The third,T. campinicola, is a low-elevation, widespread species restricted to white-sand savannas in the Brazilian, Venezuelan, and Colombian Guayana.     

ANCIENT TEPUI SUMMITS HARBOR YOUNG RATHER THAN OLD LINEAGES OF ENDEMIC FROGS

The flattop mountains (tepuis) of South America are ancient remnants of the Precambrian Guiana Shield plateau. The tepui summits, isolated by their surrounding cliffs that can be up to 1000 m tall, are thought of as “islands in the sky,” harboring relict flora and fauna that underwent vicariant speciation due to plateau fragmentation. High endemicity atop tepui summits support the idea of an ancient “Lost World” biota. However, recent work suggests that dispersal between lowlands and summits has occurred long after tepui formation indicating that tepui summits may not be as isolated from the lowlands as researchers have long suggested. Neither view of the origin of the tepui biota (i.e., ancient vicariance vs. recent dispersal) has strong empirical support owing to a lack of studies. We test diversification hypotheses of the Guiana Shield highlands by estimating divergence times of an endemic group of treefrogs, Tepuihyla. We find that diversification of this group does not support an ancient origin for this taxon; instead, divergence times among the highland species are 2–5 Ma. Our data indicate that most highland speciation occurred during the Pliocene. Thus, this unparalleled landscape known as “The Lost World” is inhabited, in part, not by Early Tertiary relicts but neoendemics.     

Phylogenetic relationships in the Marcetia alliance (Melastomeae,Melastomataceae) and implications for generic circumscription

The Marcetia alliance of Melastomataceae is an exclusively Neotropical group that includes at least 12 genera of mostly herbs and subshrubs, occurring in the cerrado of central Brazil and savannas of the Amazon region and Guayana highlands. This study aimed to test the monophyly of genera in the Marcetia alliance, evaluate their phylogenetic relationships and generic boundaries, and investigate morphological characters as potential synapomorphies for delimiting clades or genera. We used nuclear (ITS, ETS) and plastid (accD‐psaI, atpH‐atpF, trnS‐trnG) DNA sequences of 107 terminals in 12 genera from the alliance. Aciotis, Fritzschia, Marcetia and Siphanthera were shown to be monophyletic and supported by molecular and morphological characters. Other genera with variable morphology and wider distributions, such as Acisanthera, Comolia, Ernestia and Macairea, were recovered as paraphyletic or polyphyletic. Most morphological characters analysed were found to be homoplastic, but when combined they are potentially useful for the diagnosis of genera and infrageneric groups. This study represents a major step in understanding internal relationships and provides the basis for a revision of the generic classification in the Marcetia alliance.     

Melastomeae come full circle: biogeographic reconstruction and molecular clock dating

Rhexia, with 11 species in the Coastal Plain province of North America, is the only temperate zone endemic of the tropical eudicot family Melastomataceae. It is a member of the only pantropical tribe of that family, Melastomeae. Based on the chloroplast gene ndhF, we use a fossil-calibrated molecular clock to address the question of the geographic origin and age of Rhexia. Sequences from 37 species in 21 genera representing the tribe's geographical range were analyzed together with five outgroups. To obtain better clade support, another chloroplast region, the rpl16 intron, was added for 24 of the species. Parsimony analysis of the combined data and maximum-likelihood analysis of ndhF alone indicate that the deepest split is between Rhexia plus its sister group, a small Central American genus, and all other Melastomeae. Old World Melastomeae are monophyletic and nested within New World Melastomeae. Although likelihood-ratio tests of clock and nonclock substitution models for the full or moderately pruned datasets rejected the clock, these models yielded identical topologies (for 30 taxa) with few significantly different branch lengths as assessed by a Student's t-test. Age estimates obtained were 22 million years ago (Mya) for the divergence of Rhexia from its sister group, 12 Mya for the dispersal of Melastomeae from the New World to West Africa, and 1 Mya for the diversification of Melastoma in Southeast Asia. The only other genus of Melastomeae to have reached Southeast Asia from Africa or Madagascar is Osbeckia. The age and geographic distribution of fossils, which come from Miocene sites throughout Eurasia, suggest that Melastomeae once ranged from Eurasia across Beringia to North America from whence they reached South America and subsequently Africa and Southeast Asia. Climate deterioration led to their extinction in the Northern Hemisphere, with Rhexia possibly surviving in Coastal Plain refugia.     

Pliocene forest dynamics as a primary driver of African bird speciation

Aim Montane tropics are areas of high endemism, and mechanisms driving this endemism have been receiving increasing attention at a global scale. A general trend is that climatic factors do not explain the species richness of species with small to medium‐sized geographic ranges, suggesting that geological and evolutionary processes must be considered. On the African continent, several hypotheses including both refugial and geographic uplift models have been advanced to explain avian speciation and diversity in the lowland forest and montane regions of central and eastern Africa; montane regions in particular are recognized as hotspots of vertebrate endemism. Here, we examine the possible role of these models in driving speciation in a clade of African forest robins. Location Africa. Methods We constructed the first robustly supported molecular phylogenetic hypothesis of forest robins. On this phylogeny, we reconstructed habitat‐based distributions and geographic distributions relative to the Albertine Rift. We also estimated the timing of lineage divergences via a molecular clock. Results Robust estimates of phylogenetic relationships and clock‐based divergences reject Miocene tectonic uplift and Pleistocene forest refugia as primary drivers of speciation in forest robins. Instead, our data suggest that most forest robin speciation took place in the Late Pliocene, from 3.2 to 2.2 Ma. Distributional patterns are complex, with the Albertine Rift region serving as a general east–west break across the group. Montane distributions are inferred to have evolved four times. Main conclusions Phylogenetic divergence dates coincide with a single period of lowland forest retraction in the late Pliocene, suggesting that most montane speciation resulted from the rapid isolation of populations in montane areas, rather than montane areas themselves being drivers of speciation. This conclusion provides additional evidence that Pliocene climate change was a major driver of speciation in broadly distributed African animal lineages. We further show that lowland forest robins are no older than their montane relatives, suggesting that lowland areas are not museums which house ‘ancient’ taxa; rather, for forest robins, montane areas should be viewed as living museums of a late Pliocene diversification event. A forest refugial pattern is operating in Africa, but it is not constrained to the Pleistocene.     

The combined role of dispersal and niche evolution in the diversification of Neotropical lizards

Ecological requirements and environmental conditions can influence diversification across temporal and spatial scales. Understanding the role of ecological niche evolution under phylogenetic contexts provides insights on speciation mechanisms and possible responses to future climatic change. Large‐scale phyloclimatic studies on the megadiverse Neotropics, where biomes with contrasting vegetation types occur in narrow contact, are rare. We integrate ecological and biogeographic data with phylogenetic comparative methods, to investigate the relative roles of biogeographic events and niche divergence and conservatism on the diversification of the lizard genus Kentropyx Spix, 1825 (Squamata: Teiidae), distributed in South American rainforests and savannas. Using five molecular markers, we estimated a dated species tree, which recovered three clades coincident with previously proposed species groups diverging during the mid‐Miocene. Biogeography reconstruction indicates a role of successive dispersal events from an ancestral range in the Brazilian Shield and western Amazonia. Ancestral reconstruction of climatic tolerances and niche overlap metrics indicates a trend of conservatism during the diversification of groups from the Amazon Basin and Guiana Shield, and a strong signal of niche divergence in the Brazilian Shield savannas. Our results suggest that climatic‐driven divergence at dynamic forest‐savanna borders might have resulted in adaptation to new environmental niches, promoting habitat shifts and shaping speciation patterns of Neotropical lizards. Dispersal and ecological divergence could have a more important role in Neotropical diversification than previously thought.     

Recent diversification in African greenbuls (Pycnonotidae: Andropadus) supports a montane speciation model

It is generally accepted that accentuated global climatic cycles since the Plio-Pleistocene (2.8 Ma ago) have caused the intermittent fragmentation of forest regions into isolated refugia thereby providing a mechanism for speciation of tropical forest biota contained within them. However, it has been assumed that this mechanism had its greatest effect in the species rich lowland regions. Contrary evidence from molecular studies of African and South American forest birds suggests that areas of recent intensive speciation, where mostly new lineages are clustered, occur in discrete tropical montane regions, while lowland regions contain mostly old species. Two predictions arise from this finding. First, a species phylogeny of an avian group, represented in both lowland and montane habitats, should be ordered such that montane forms are represented by the most derived characters. Second, montane speciation events should predominate within the past 2.8 Ma. In order to test this model I have investigated the evolutionary history of the recently radiated African greenbuls (genus Andropadus), using a molecular approach. This analysis finds that montane species are a derived monophyletic group when compared to lowland species of the same genus and recent speciation events (within the Plio-Pleistocene) have exclusively occurred in montane regions. These data support the view that montane regions have acted as centres of speciation during recent climatic instability.     

Over the hills and far away: New plant records for the Guayana Shield in Brazil

The Guayana Shield is one of the oldest geological formations in South America, ranging from southern Colombia, Venezuela, and the Guianas to the extreme north of the Brazilian states of Amazonas and Roraima. Because of its ancient origin and isolation from other mountain ranges in South America, it harbors a rich flora with high levels of endemism. Recent expeditions to remote areas in the Brazilian portion of the Guayana Shield generated many new herbarium collections. Through these efforts, we report the first records for Brazil of 57 species and infraspecific taxa of vascular plants. In most cases, the taxa are narrowly distributed and were previously known from nearby populations in other countries. In other cases, however, the new Brazilian records expand considerably the known geographical distributions of the taxa.     

Herbaceous ecosystems on the Guayana Shield, a regional overview

Aims Ecosystems dominated by herbaceous plant communities are amongst the most diversified landscape units in the Guayana (Guiana) Shield region. This paper aims to present a synthetic overview of the wide array of herbaceous ecosystems found in the region in an attempt to furnish a more concrete baseline for a better understanding of the pattern of variation, and to clarify some of the differences that occur in the vegetation of the area. Location The Guayana Shield region, and the area of north‐eastern South America extending between the Orinoco River to the North and the Amazon River to the South (c. 8° N to 1° S). Methods Floristic and ecological field data gathered from over 300 study sites located at different altitudinal levels in the Venezuelan Guayana and the northern Brazilian Amazon are evaluated and interpreted in the light of personal observations and existing literature. Results The diversification pattern includes physiognomic as well as floristic variation and shows two opposing tendencies in relation to their altitudinal location: grass‐dominated meadows (savannas) predominate in the macrothermic lowlands, whereas non‐gramineous, broadleaved herbaceous communities are dominant in the mesothermic highlands of the Guayanan mountains. In detail, the biogeographic region of Guayana (Guiana) consists of three easily recognized altitudinal levels, each of which contains a highly characteristic set of landscape types with their associated ecosystems and endemic plant communities. Between 0 and c. 500 m a.s.l. the extensive macrothermic (mean annual temperature, MAT > 24 °C) lowlands are found, where an enormous – still largely unexplored – diversity of forest types represents the main plant cover of the plains, peneplains, glacis and piedmont slopes. Sparsely distributed within this forest cover are numerous herbaceous ecosystems, ranging from true grass savannas to unusual and poorly understood meadows developed on extremely oligotrophic white sandy soils dominated by non‐gramineous genera of Rapateaceae and Xyridaceae. In the intermediate submesothermic (MAT 24–18 °C) Guayana uplands, extending roughly between 500 and 1500 m a.s.l., the grasslands of the Gran Sabana in south‐eastern Venezuela reach their upper altitudinal limit. At the same altitudinal level, however, several distinct herbaceous communities are found, in which other genera of Rapateaceae, together with Bromeliaceae and Xyridaceae, are predominant. Finally, at the uppermost altitudinal level, i.e. in Pantepui (which includes the characteristically flat topped mountain summits (tepuis) of the Guayana highlands) between 1500 and 3000 m a.s.l., encompassing a range of meso‐ to submicrothermic temperature regimes (MAT 18–8 °C), the extensive herbaceous ecosystems are developed either on deep organic soils (peat) or on open sandstone surfaces. These high‐tepui meadows present a considerable physiognomic diversification and are formed by a variety of endemic genera of the Rapateaceae, Bromeliaceae, Xyridaceae, Eriocaulaceae and Cyperaceae families, often dominated by locally endemic species on each of the larger tepui massifs. In contrast, grass dominated plant communities are very rare and restricted to only a few high‐tepui sites. Main conclusions A marked floristic and ecologic differentiation of herbaceous ecosystems in the Guayana Shield region can be recognized. The ecological differentiation results primarily from the wide spectrum of variations in the substrate found at the various altitudinal levels of the Guayana Shield region. A possible explanation for the present‐day pattern of herbaceous vegetation types may be the following: Non‐gramineous meadows representing ancient species pools of Guayana‐centred families had evolved successful colonization strategies in occupying extremely nutrient poor sites at all altitudinal levels. In contrast, the more modern grass savannas, which preferentially occupy the peripheral Guayana landscapes, are restricted to richer soil conditions with better internal drainage and water retention conditions.     

Cooperative Breeding and Long-Distance Dispersal: A Test Using Vagrant Records

Cooperative breeding is generally associated with increased philopatry and sedentariness, presumably because short-distance dispersal facilitates the maintenance of kin groups. There are, however, few data on long-distance dispersal in cooperative breeders—the variable likely to be important for genetic diversification and speciation. We tested the hypothesis that cooperative breeders are less likely to engage in long-distance dispersal events by comparing records of vagrants outside their normal geographic range for matched pairs (cooperatively vs. non-cooperatively breeding) of North American species of birds. Results failed to support the hypothesis of reduced long-distance dispersal among cooperative breeders. Thus, our results counter the conclusion that the lower rate of speciation among cooperative breeding taxa found in recent analyses is a consequence of reduced vagility.     

Polemoniaceae phylogeny and classification: implications of sequence data from the chloroplast gene ndhF

The chloroplast gene ndhF was used to study phylogenetic relationships of the Polemoniaceae at two levels: among members of the Ericales and among genera of the family. Sequence data for interfamilial analyses consisted of 2266 bp for 14 members of the Ericales, including four species of the Polemoniaceae, plus three outgroup taxa. The Polemoniaceae were found to be related to Diospyros, Fouquieria, the Primulales, Rhododendron, and Impatiens, but relationships among taxa were generally not well supported. The precise position of the Polemoniaceae within the Ericales remains obscure. Data for intrafamilial analyses consisted of 1031 bp for 27 species of the Polemoniaceae, including at least one species from most genera of the family, plus five outgroup taxa. A single most parsimonious tree was identified. The analyses suggested that subfamily Cobaeoideae, excluding Loeselia, is monophyletic and that Huthia is sister to Cantua. Acanthogilia was sister to the remainder of subfamily Cobaeoideae. Subfamily Polemonioideae plus Loeselia formed four subclades that were strongly supported as monophyletic and represent the major lineages of the subfamily.     

New Zealand's pteridophyte flora—Plants of ancient lineage but recent arrival?

A hypothesis is presented that most pteridophytes arrived in New Zealand relatively recently, by long-distance dispersal. The flora comprises 194 native species, of which 89 (46%) are endemic and 105 (54%) are widespread. Of the latter, 90% are shared with temperate Australasia, 53% with tropical regions, 14% with temperate southern Africa and 13% with the circum-Antarctic islands and South America. New Zealand has undergone such dramatic changes in location, land area, and topography since initial separation from Gondwana 85 Ma that it seems improbable that the 95 species shared with temperate Australasia could have remained conspecific throughout that time. Modern fossil and molecular evidence strongly suggest that many families of ferns had not even evolved prior to separation, and palynological evidence from New Zealand indicates that 78% of pteridophyte genera first appeared there only after separation from Gondwana. Present-day distributions in New Zealand suggest that ferns have greater dispersal potential than flowering plants, and that pteridophyte distributions are more heavily influenced by temperature, rainfall, and geothermal activity than by geological history. Most endemic pteridophyte species have a predominantly southern distribution pattern and are characteristic of cool, lowland to montane forest. Pteridophytes in the northern part of New Zealand show a lower level of endemism than elsewhere and tend to be widespread species that have arrived from temperate Australasian and tropical regions. There is also evidence that at least some pteridophytes have migrated from New Zealand to Australia. It is suggested that the hypothesis of long-distance dispersal of pteridophytes across the Tasman Sea could be tested by molecular techniques.     

DOES NICHE CONSERVATISM PROMOTE SPECIATION? A CASE STUDY IN NORTH AMERICAN SALAMANDERS

Recent speciation research has generally focused on how lineages that originate in allopatry evolve intrinsic reproductive isolation, or how ecological divergence promotes nonallopatric speciation. However, the ecological basis of allopatric isolation, which underlies the most common geographic mode of speciation, remains poorly understood and largely unstudied. Here, we explore the ecological and evolutionary factors that promote speciation in Desmognathus and Plethodon salamanders from temperate eastern North America. Based on published molecular phylogenetic estimates and the degree of geographic range overlap among extant species, we find strong evidence for a role for geographic isolation in speciation. We then examine the relationship between climatic variation and speciation in 16 sister-taxon pairs using geographic information system maps of climatic variables, new methods for modeling species' potential geographic distributions, and data on geographic patterns of genetic variation. In contrast to recent studies in tropical montane regions, we found no evidence for parapatric speciation along climatic gradients. Instead, many montane sister taxa in the Appalachian Highlands inhabit similar climatic niches and seemingly are allopatric because they are unable to tolerate the climatic conditions in the intervening lowlands. This temporal and spatial-ecological pattern suggests that niche conservatism, rather than niche divergence, plays the primary role in promoting allopatric speciation and montane endemism in this species-rich group of vertebrates. Our results demonstrate that even the relatively subtle climatic differences between montane and lowland habitats in eastern North America may play a key role in the origin of new species.     

Generic delimitation and relationships in Ebenaceae sensu lato: evidence from six plastid DNA regions

Phylogenetic relationships of the pantropical family Ebenaceae s.l. were investigated using plastid DNA sequence data from six regions: atpB, matK, ndhF, trnK intron, trnL intron, and trnL-trnF spacer. Sampling included representatives of all currently recognized genera of Ebenaceae, Diospyros, Euclea, and Lissocarpa, and nearly all taxa that were previously recognized at the generic level, e.g., Cargillia, Gunisanthus, Maba, Macreightia, Royena, and Tetraclis. Our results strongly support monophyly of Ebenaceae s.l. and demonstrate that the previous infrafamilar classifications of the family do not circumscribe monophyletic groups. A new infrafamilial classification based on a phylogenetic approach is proposed here and consists of two subfamilies, Lissocarpoideae and Ebenoideae, and four genera, Lissocarpa, Euclea, Royena, and Diospyros. Relationships and potential synapomorphic characters are discussed and summarized. This study supports a western Gondwanan origin of family and indicates that both vicariant and long-distance dispersal events played an important role in attaining current distributions.     

苦苣苔科镜像花的多样性及演化

泛热带分布的苦苣苔科(Gesneriaceae)在我国南方具有极高的物种丰富度与特有率,花部特征变化丰富,是研究物种形成与适应演化的代表类群。镜像花(mirror-image flowers)是极为特化的传粉系统,在苦苣苔科中出现了较多的不同类型,可能与苦苣苔科物种多样性形成与维持有关。该研究总结与分析了苦苣苔科镜像花的类型多样性以及系统分布与适应演化等,讨论了镜像花对苦苣苔科物种形成与维持的积极意义。结果表明:镜像花仅分布在亚洲和非洲的苦苣苔亚科(Didymocarpoideae)的7个属,在历史上就至少发生了5次独立起源。长冠苣苔属(Rhabdothamnopsis)、南洋苣苔属(Henckelia)及长蒴苣苔属(Didymocarpus)镜像花的花柱与可育雄蕊分别向左、右两侧偏转,形成互补镜像花;蛛毛苣苔属(Paraboea)、喜鹊苣苔属(Ornithoboea)、非洲堇属(Saintpaulia)镜像花缺乏与花柱对应侧偏的可育雄蕊(非互补镜像花);而海角苣苔属(Streptocarpus)直立堇兰亚属(subg.Streptocarpella Engler)则同时出现了互补、非互补镜像花。不同于其他被子植物(离瓣花、缺乏花冠筒),苦苣苔科中的镜像花大多伴随着明显的花冠筒、内藏的雄蕊、合生的花药,以非互补镜像花为主;传粉者以小型的无垫蜂(Amegilla spp.)和熊蜂(Bombus spp.)为主。这些特殊的花部综合征与特化的传粉机制,提高了传粉精确性,可能促进了传粉隔离与物种适应辐射。今后的一个研究重点应通过分子系统发育方法,进一步揭示苦苣苔亚科互补与非互补镜像花的进化顺序及其在物种分化与长距离扩散过程中的可能作用。     

Distribution and vegetation dynamics of humid savannas in Africa and Asia

Abstract. A review is presented on the literature about the distribution of savannas in humid climates in Africa and Asia and their vegetation dynamics. Sections are devoted to African lowland and montane savannas (the latter divided into southern, eastern, western and northern African), Madagascar, Indian subcontinent, SE Asia and New Guinea. It is concluded that the extension of savannas under humid climatic conditions and the relation to the distribution of forests is a function of cultivation, grazing by domestic and wild animals, present and previous climate, geomorphology and soil characteristics. Once established, savannas are often maintained by fires, both natural and man-made. Montane savannas are generally brought about by man's clearing, cultivation and burning. Fire is a stochastic variable; it creates an ecotone sensu stricto (an environmentally stochastic stress zone) at the forest/savanna border. On the other hand, if geomorphology and soil are the determinants, the transition between forest and savanna would have the character of an ecocline (a gradient zone) with fundamentally different conditions. In humid African lowland climates forests expand into savannas if the latter are not maintained by man. Whether forests also expand in less humid climates is disputed. In montane areas forest expansion may be delayed on degraded soils and when diaspores are lacking.     

Phylogeny of Acridocarpus-Brachylophon (Malpighiaceae): implications for tertiary tropical floras and Afroasian biogeography

Abstract.— A major tenet of African Tertiary biogeography posits that lowland rainforest dominated much of Africa in the late Cretaceous and was replaced by xeric vegetation as a response to continental uplift and consequent widespread aridification beginning in the late Paleogene. The aridification of Africa is thought to have been a major factor in the extinction of many African humid-tropical lineages, and in the present-day disparity of species diversity between Africa and other tropical regions. This primarily geologically based model can be tested with independent phylogenetic evidence from widespread African plant groups containing both humid- and xeric-adapted species. We estimated the phylogeny and lineage divergence times within one such angiosperm group, the acridocarpoid clade (Malpighiaceae), with combined ITS, ndhF , and trnL-F data from 15 species that encompass the range of morphological and geographic variation within the group. Dispersal-vicariance analysis and divergence-time estimates suggest that the basal acridocarpoid divergence occurred between African and Southeast Asian lineages approximately 50 million years ago (mya), perhaps after a southward ancestral retreat from high-latitude tropical forests in response to intermittent Eocene cooling. Dispersion of Acridocarpus from Africa to Madagascar is inferred between approximately 50 and 35 mya, when lowland humid tropical forest was nearly continuous between these landmasses. A single dispersal event within Acridocarpus is inferred from western Africa to eastern Africa between approximately 23 and 17 mya, coincident with the widespread replacement of humid forests by savannas in eastern Africa. Although the spread of xeric environments resulted in the extinction of many African plant groups, our data suggest that for others it provided an opportunity for further diversification.     

Vertical transmission as the key to the colonization of Madagascar by fungus-growing termites?

The mutualism between fungus-growing termites (Macrotermitinae) and their mutualistic fungi (Termitomyces) began in Africa. The fungus-growing termites have secondarily colonized Madagascar and only a subset of the genera found in Africa is found on this isolated island. Successful long-distance colonization may have been severely constrained by the obligate interaction of the termites with fungal symbionts and the need to acquire these symbionts secondarily from the environment for most species (horizontal symbiont transmission). Consistent with this hypothesis, we show that all extant species of fungus-growing termites of Madagascar are the result of a single colonization event of termites belonging to one of the only two groups with vertical symbiont transmission, and we date this event at approximately 13 Mya (Middle/Upper Miocene). Vertical symbiont transmission may therefore have facilitated long-distance dispersal since both partners disperse together. In contrast to their termite hosts, the fungal symbionts have colonized Madagascar multiple times, suggesting that the presence of fungus-growing termites may have facilitated secondary colonizations of the symbiont. Our findings indicate that the absence of the right symbionts in a new environment can prevent long-distance dispersal of symbioses relying on horizontal symbiont acquisition.     

Phylogenetic relationships and historical biogeography of neotropical parrots (Psittaciformes: Psittacidae: Arini) inferred from mitochondrial and nuclear DNA sequences

Previous hypotheses of phylogenetic relationships among Neotropical parrots were based on limited taxon sampling and lacked support for most internal nodes. In this study we increased the number of taxa (29 species belonging to 25 of the 30 genera) and gene sequences (6388 base pairs of RAG-1, cyt b, NADH2, ATPase 6, ATPase 8, COIII, 12S rDNA, and 16S rDNA) to obtain a stronger molecular phylogenetic hypothesis for this group of birds. Analyses of the combined gene sequences using maximum likelihood and Bayesian methods resulted in a well-supported phylogeny and indicated that amazons and allies are a sister clade to macaws, conures, and relatives, and these two clades are in turn a sister group to parrotlets. Key morphological and behavioral characters used in previous classifications were mapped on the molecular tree and were phylogenetically uninformative. We estimated divergence times of taxa using the molecular tree and Bayesian and penalized likelihood methods that allow for rate variation in DNA substitutions among sites and taxa. Our estimates suggest that the Neotropical parrots shared a common ancestor with Australian parrots 59 Mya (million of years ago; 95% credibility interval (CrI) 66, 51 Mya), well before Australia separated from Antarctica and South America, implying that ancestral parrots were widespread in Gondwanaland. Thus, the divergence of Australian and Neotropical parrots could be attributed to vicariance. The three major clades of Neotropical parrots originated about 50 Mya (95% CrI 57, 41 Mya), coinciding with periods of higher sea level when both Antarctica and South America were fragmented with transcontinental seaways, and likely isolated the ancestors of modern Neotropical parrots in different regions in these continents. The correspondence between major paleoenvironmental changes in South America and the diversification of genera in the clade of amazons and allies between 46 and 16 Mya suggests they diversified exclusively in South America. Conversely, ancestors of parrotlets and of macaws, conures, and allies may have been isolated in Antarctica and/or the southern cone of South America, and only dispersed out of these southern regions when climate cooled and Antarctica became ice-encrusted about 35 Mya. The subsequent radiation of macaws and their allies in South America beginning about 28 Mya (95% CrI 22, 35 Mya) coincides with the uplift of the Andes and the subsequent formation of dry, open grassland habitats that would have facilitated ecological speciation via niche expansion from forested habitats.     

Phylogeography of the bushmaster (Lachesis tnuta: Viperidae): implications for neotropical biogeography, systematics, and conservation

We used mitochondrial gene sequences to reconstruct phylogenetic relationships among subspecies of the bushmaster, Lachesis muta. These large vipers are widely distributed in lowland tropical forests in Central and South America, where three of four allopatric subspecies are separated by montane barriers. Our phylogeny indicates that the four subspecies belong to two clades, the Central American and South American lineages. We use published molecular studies of other taxa to estimate a 'reptilian mtDNA rate' and thus temporal boundaries for major lineage divergences in Lachesis. We estimate that the Central and South American forms diverged 18-6 Mya, perhaps due to the uplifting of the Andes, whereas the two Central American subspecies may have diverged 11-4 Mya with the uprising of the Cordillera de Talamanca that separates them today. South American bushmasters from the Amazon Basin and the Atlantic Forest are not strongly differentiated, perhaps due to episodic gene flow during the Pleistocene, when suitable habitat for this species was at times more continuous. Our results agree with previous evidence that genetic divergence among some neotropical vertebrates pre-dated Pleistocene forest fragmentation cycles and the appearance of the Panamanian Isthmus. Based on morphological, behavioral, and molecular evidence, we recognize three species of Lachesis. In addition to L. muta, the widespread South American form, the Central American forms are treated as distinct species (L. meknocephak and L. stenophrys), each deserving of special conservation status due to restricted distribution and habitat destruction.