Tempo Of Diversification Research Articles

Diversification and evolutionary history of the African laminated-toothed rats (Rodentia, Otomyini)

The African continent was subjected to periodic climatic shifts during the Pliocene and Pleistocene. These habitat changes greatly affected the evolutionary processes and tempo of diversification in numerous, widely distributed mammals. The Otomyini (Family Muridae) comprises three African rodent genera, Parotomys, Otomys and Myotomys, characterized by unique laminated-shaped molars. Species within this tribe generally prefer open-habitat and show low dispersal capabilities, with previous studies suggesting that their diversification was closely associated with climatic oscillations over the last four million years. Our phylogenetic reconstructions, based on three mitochondrial (mtDNA) genes (Cytb, COI and 12S) and four nuclear introns (EF, SPTBN, MGF and THY), identified eight major genetic clades that are distributed across southern, eastern and western Africa. Our data permit the re-examination of the taxonomic status of the three genera as well as the previously proposed mesic-arid dichotomy of the 10 South African species. Moreover, multiple mtDNA species delimitation methods incorporating 168 specimens estimated the number of Otomyini species to be substantially higher than the ∼ 30 recognized, suggesting that the current taxonomy will necessitate an integrative approach to delimit extant species diversity within the Otomyini. The data suggests that the origin of the tribe can be dated back to ∼ 5.7 million years ago (Ma) in southern Africa. The distribution and phylogenetic associations among the eight major otomyine evolutionary lineages can best be explained by several waves of northward colonization from southern Africa, complemented by independent reversed dispersals from eastern back to southern Africa at different time periods. There is strong support for the hypothesis that the radiation, dispersion, and diversification of the otomyine rodents is closely linked to recent Plio-Pleistocene climatic oscillations.

How the African house gecko (Hemidactylus mabouia) conquered the world.

Alien species are among the greatest threats to biodiversity, but the evolutionary origins of invasiveness remain obscure. We conducted the first range-wide sampling of Hemidactylus mabouia from more than 120 localities across Africa, Madagascar and the Neotropics to understand the evolutionary history of one of the most widely distributed, invasive vertebrates in the world. We used a multi-locus phylogeny, species delimitation, fossil-calibrated timetree, ancestral area reconstruction and species distribution models (SDMs) to determine how many putative species-level lineages are contained within H. mabouia, the timing and tempo of diversification, and the origins of commensality—providing insights into the evolutionary origins of invasiveness. Our analyses suggest ‘H. mabouia’ originated in the Miocene in the Zambezian biogeographic region and includes as many as 20 putative species-level lineages, of which only Hemidactylus mabouia sensu stricto is invasive and widely distributed, including all Neotropical records. Zambezia is the hotspot for diversity within the group with 14 species in southeastern Zambezia. SDMs suggest that H. mabouia was able to establish in the Neotropics due to habitat suitability, and globalization and the slave trade probably allowed it to cross the Atlantic. Distribution models for the H. mabouia complex overpredict the range of the invasive H. mabouia sensu stricto—highlighting the importance of taxonomy in invasive species management.

Niche specificity influences gene flow across fine-scale habitat mosaics in Succulent Karoo plants.

While the tempo of diversification in biodiversity hotspots has received much attention, the spatial scale of diversification has often been overlooked. Addressing this deficiency requires understanding the drivers of population divergence and the spatial scales at which they operate in species-rich clades and ecosystems. South Africa's Succulent Karoo (SK) hotspot provides an excellent system for such research, being both compact (ca. 110,000km2 ) and home to spectacular in-situ radiations, such as the ruschioid Aizoaceae. Here we use GBS to document genetic structure in two co-occurring ruschioid species, at both coarse (>10km) and fine (<500m) spatial scales. Where Ruschia burtoniae shows strong between-population genetic differentiation and no gene flow, Conophytum calculus shows weak differentiation, with high levels of admixture suggesting recent or ongoing gene flow. Community analysis and transplant experiments reveal that R.burtoniae occupies a narrow, low-pH edaphic niche, and at scales of a few hundred metres, areas of elevated genetic turnover correspond to patches of edaphically unsuitable habitat. In contrast, C.calculus occupies a broader niche and exhibits isolation-by-distance without a habitat effect. We suggest that edaphic specialisation, coupled with highly restricted seed and pollen dispersal in heterogeneous landscapes, has played a major role in driving rapid diversification at small spatial scales in this system. However, the contrasting patterns in our study species show that these factors do not influence all organisms uniformly, being strongly modulated by lineage-specific traits that influence both the spatial scale of gene flow and habitat specificity.

Genome-wide RAD sequencing resolves the evolutionary history of serrate leaf Juniperus and reveals discordance with chloroplast phylogeny

Juniper (Juniperus) is an ecologically important conifer genus of the Northern Hemisphere, the members of which are often foundational tree species of arid regions. The serrate leaf margin clade is native to topologically variable regions in North America, where hybridization has likely played a prominent role in their diversification. Here we use a reduced-representation sequencing approach (ddRADseq) to generate a phylogenomic data set for 68 accessions representing all 22 species in the serrate leaf margin clade, as well as a number of close and distant relatives, to improve understanding of diversification in this group. Phylogenetic analyses using three methods (SVDquartets, maximum likelihood, and Bayesian) yielded highly congruent and well-resolved topologies. These phylogenies provided improved resolution relative to past analyses based on Sanger sequencing of nuclear and chloroplast DNA, and were largely consistent with taxonomic expectations based on geography and morphology. Calibration of a Bayesian phylogeny with fossil evidence produced divergence time estimates for the clade consistent with a late Oligocene origin in North America, followed by a period of elevated diversification between 12 and 5 Mya. Comparison of the ddRADseq phylogenies with a phylogeny based on Sanger-sequenced chloroplast DNA revealed five instances of pronounced discordance, illustrating the potential for chloroplast introgression, chloroplast transfer, or incomplete lineage sorting to influence organellar phylogeny. Our results improve understanding of the pattern and tempo of diversification in Juniperus, and highlight the utility of reduced-representation sequencing for resolving phylogenetic relationships in non-model organisms with reticulation and recent divergence.

The impact of rainforest area reduction in the Guineo‐Congolian region on the tempo of diversification and habitat shifts in the Berlinia clade (Leguminosae)

AbstractAimThe Guineo‐Congolian region in Africa constitutes the second largest area of tropical rainforest (TRF) in the world. It covered an estimated 15–22 million km2 during the late Miocene (55–11 Ma) and it has experienced since a declining trend, currently reaching 3.4 million km2, associated with increasing aridification and the replacement of TRF by savanna habitats. Here, we examine whether rainforest area contraction led to a decrease in net diversification rates linked to increasing extinction, or if it is associated with increasing opportunities for allopatric or ecological speciation during periods of forest fragmentation.LocationTropical Africa, Guineo‐Congolian region.TaxonAnthonotha, Englerodendron, Berlinia clade (Leguminosae).MethodsWe used a target enrichment approach combined with a complete data set representing all genera within the Berlinia clade. We combined phylogenomic, dating estimates, habitat reconstruction and diversification rate analyses to infer the effect of change in rainforest area coverage at two taxonomic levels: among genera, and within Anthonotha and Englerodendron.ResultsWe recovered fully resolved and well‐supported relationships among all genera and among species within the two genera. Most genera (87.5%) diverged before the Pleistocene, but Anthonotha and Englerodendron diversified recently, during the most recent cycles of forest contraction and expansion of the Pleistocene.Main conclusionsOur results suggest that the Berlinia clade displays an overall trend of accumulation of species over evolutionary time, suggesting the reduction in TRF area has not decreased net diversification rates. Most habitat shifts to savanna occurred in the Miocene, with no major habitat shifts during the most recent phases of forest expansion–contraction in the Pleistocene. Shifts in habitat from lowland forest to savanna did not trigger diversification rates, but habitat fragmentation might have increased diversification rates through allopatric speciation.

Comparing the rates of speciation and extinction between phylogenetic trees.

Over the past decade or so it has become increasingly popular to use reconstructed evolutionary trees to investigate questions about the rates of speciation and extinction. Although the methodology of this field has grown substantially in its sophistication in recent years, here I will take a step back to present a very simple model that is designed to investigate the relatively straightforward question of whether the tempo of diversification (speciation and extinction) differs between two or more phylogenetic trees, without attempting to attribute a causal basis to this difference. It is a likelihood method, and I demonstrate that it generally shows type I error that is close to the nominal level. I also demonstrate that parameter estimates obtained with this approach are largely unbiased. As this method can be used to compare trees of unknown relationship, it will be particularly well‐suited to problems in which a difference in diversification rate between clades is suspected, but in which these clades are not particularly closely related. As diversification methods can easily take into account an incomplete sampling fraction, but missing lineages are assumed to be missing at random, this method is also appropriate for cases in which we have hypothesized a difference in the process of diversification between two or more focal clades, but in which many unsampled groups separate the few of interest. The method of this study is by no means an attempt to replace more sophisticated models in which, for instance, diversification depends on the state of an observed or unobserved discrete or continuous trait. Rather, my intention is to provide a complementary approach for circumstances in which a simpler hypothesis is warranted and of biological interest.

Macroevolutionary analysis of the tempo of diversification in snappers and fusiliers (Percomorpha: Lutjanidae)

The percomorph fish family Lutjanidae (snappers and fusiliers) includes about 135 reef-dwelling species, mainly confined to tropical and subtropical marine waters. The great majority of snappers are active predators feeding on fishes or crustaceans, even though some species, including the fusiliers (Caesioninae), have evolved zooplanktivory. Lutjanids show a great diversity of habitat preferences, based on depth segregation and distribution across reef and associated habitats (e.g., mangroves, seagrass beds, estuaries). In spite of their great ecological and economic importance little is known about the tempo of evolution in this group. The present study provides the most comprehensive molecular phylogeny to date for lutjanids, including 70% of extant species and 19 of the 21 currently described genera. We time-calibrated our molecular tree using the oldest described lutjanid fossils, and show how this group most likely originated during the Late Cretaceous or Early Paleocene. Lutjanids experienced a significant radiation during the Late Eocene and Early Oligocene, in contrast to a pattern of Late Oligocene/Miocene radiation observed in many other reef-associated groups. The time-tree allows us to investigate the tempo of diversification, and our results suggest a variation in the rate of speciation during the evolution of the major clade formed by “lutjanins and caesionins”. Variation in diet and life history strategies could explain this clade-specific dynamic, although future phylogenetic comparative studies combining additional ecological and morphological data are needed to test this hypothesis.

Unraveling the biogeographical history of Chrysobalanaceae from plastid genomes.

The complex geological and climatic history of the Neotropics has had major implications on the diversification of plant lineages. Chrysobalanaceae is a pantropical family of trees and shrubs with 75% of its 531 species found in the Neotropics, and a time-calibrated phylogeny of this family should shed light on the tempo of diversification in the Neotropical flora. Previously published phylogenetic hypotheses of this family were poorly supported, and its biogeography remains unclear. We assembled the complete plastid genome of 51 Chrysobalanaceae species, and increased taxon sampling by Sanger-sequencing of five plastid regions for an additional 88 species. We generated a time-calibrated tree including all 139 Chrsyobalanaceae species and 23 outgroups. We then conducted an ancestral area reconstruction analysis and estimated diversification rates in the family. The tree generated with the plastid genome alignment was almost fully resolved. It supports the polyphyly of Licania and Hirtella. The family has diversified starting around the Eocene-Oligocene transition. An ancestral area reconstruction confirms a Paleotropical origin for Chrysobalanaceae with several transoceanic dispersal events. The main Neotropical clade likely resulted from a single migration event from Africa around 28 mya ago, which subsequently underwent rapid diversification. Given the diverse ecologies exhibited by extant species, we hypothesize that the rapid diversification of Chrysobalanaceae following the colonization of the Neotropics was triggered by habitat specialization during the complex geological and paleoclimatic history of the Neotropics.

Constant diversification rates of endemic gastropods in ancient Lake Ohrid: ecosystem resilience likely buffers environmental fluctuations

Abstract. Ancient lakes represent key ecosystems for endemic freshwater species. This high endemic biodiversity has been shown to be mainly the result of intra-lacustrine diversification. Whereas the principle role of this mode of diversification is generally acknowledged, actual diversification rates in ancient lakes remain little understood. At least four types are conceivable. Diversification rates may be constant over time, they may fluctuate, rates may be higher in the initial phase of diversification, or there may be a pronounced lag phase between colonization and subsequent diversification. As understanding the tempo of diversification in ancient lake environments may help reveal the underlying processes that drive speciation and extinction, we here use the Balkan Lake Ohrid as a model system and the largest species flock in the lake, the non-pyrgulinid Hydrobiidae, as a model taxon to study changes in diversification rates over time together with the respective drivers. Based on phylogenetic, molecular-clock, lineage-through-time plot, and diversification-rate analyses we found that this potentially monophyletic group is comparatively old and that it most likely evolved with a constant diversification rate. Preliminary data of the SCOPSCO (Scientific Collaboration On Past Speciation Conditions in Lake Ohrid) deep-drilling program do indicate signatures of severe environmental/climatic perturbations in Lake Ohrid. However, so far there is no evidence for the occurrence of catastrophic environmental events. We therefore propose that the constant diversification rate observed in endemic gastropods has been caused by two factors: (i) a potential lack of catastrophic environmental events in Lake Ohrid and/or (ii) a probably high ecosystem resilience, buffering environmental changes. Parameters potentially contributing to the lake's high ecosystem resilience are its distinct bathymetry, ongoing tectonic activities, and karst hydrology. The current study not only contributes to one of the overall goals of the SCOPSCO deep-drilling program – inferring the driving forces for biotic evolution in Lake Ohrid. It might also enhance our understanding of how ecosystem resilience, in general, may promote relatively constant diversification rates in isolated ecosystems. However, we encourage future studies testing hypotheses about the lack of catastrophic events in Lake Ohrid. These studies should be based on high-resolution data for the entire geological history of the lake, and they should potentially involve information from the sediment fossil record, not only for gastropods but also for other groups with a high share of endemic taxa.

Paraphyly changes understanding of timing and tempo of diversification in subtribe Hakeinae (Proteaceae), a giant Australian plant radiation.

Subtribe Hakeinae (526 spp.) represents a large Australian plant radiation central to our understanding of that flora's evolution and ecology. It contains Grevillea-the third largest plant genus in Australia and a group inferred to have among the highest diversification rates in the angiosperms. However, we lack a robust phylogenetic framework for understanding subtribe Hakeinae and recognize that Grevillea lacks an unambiguous synapomorphy supporting its monophyly. We used four plastid and one nuclear DNA region from a taxonomically even sampling of a third of the species to infer a time-calibrated phylogeny of Hakeinae and absolute diversification rates of major clades. We developed the R package addTaxa to add unsampled taxa to the tree for diversification rate inference. Grevillea is paraphyletic with respect to Hakea and Finschia. Under most parameter combinations, Hakea contains the major clade with the highest diversification rate in Hakeinae, rather than Grevillea. The crown age of the Grevillea+Hakea+Finschia crown group is about double that of prior estimates. We demonstrate that the paraphyly of Grevillea considerably enlarges the number of Australian descendants from its most recent common ancestor but has also misled investigators who considered a single operational taxonomic unit as adequate to represent the genus for inferences of diversification rate and timing. Our time-calibrated phylogeny can form the basis of future evolutionary, comparative ecology, and biogeography studies involving this large Australian plant radiation, as well as nomenclatural changes.

Genetic Differentiation in Insular Lowland Rainforests: Insights from Historical Demographic Patterns in Philippine Birds.

Phylogeographic studies of Philippine birds support that deep genetic structure occurs across continuous lowland forests within islands, despite the lack of obvious contemporary isolation mechanisms. To examine the pattern and tempo of diversification within Philippine island forests, and test if common mechanisms are responsible for observed differentiation, we focused on three co-distributed lowland bird taxa endemic to Greater Luzon and Greater Negros-Panay: Blue-headed Fantail (Rhipidura cyaniceps), White-browed Shama (Copsychus luzoniensis), and Lemon-throated Leaf-Warbler (Phylloscopus cebuensis). Each species has two described subspecies within Greater Luzon, and a single described subspecies on Greater Negros/Panay. Each of the three focal species showed a common geographic pattern of two monophyletic groups in Greater Luzon sister to a third monophyletic group found in Greater Negros-Panay, suggesting that common or similar biogeographic processes may have produced similar distributions. However, studied species displayed variable levels of mitochondrial DNA differentiation between clades, and genetic differentiation within Luzon was not necessarily concordant with described subspecies boundaries. Population genetic parameters for the three species suggested both rapid population growth from small numbers and geographic expansion across Luzon Island. Estimates of the timing of population expansion further supported that these events occurred asynchronously throughout the Pleistocene in the focal species, demanding particular explanations for differentiation, and support that co-distribution may be secondarily congruent.

Miocene extensional tectonics explain ancient patterns of diversification among turret‐building tarantulas (Aphonopelma mojave group) in the Mojave and Sonoran deserts

AbstractAimPhylogeographical studies in the Mojave and Sonoran deserts often find genetic discontinuities that pre‐date the Pleistocene. A recent synthesis of phylogeographical data, called the Mojave Assembly Model, provides a hypothesis for the historical assembly of these desert biotas but does not adequately capture the complexity of pre‐Pleistocene vicariance events. We tested this model and assessed pre‐Pleistocene divergences by exploring the phylogeography of the Aphonopelma mojave group, which is composed of turret‐building tarantula species from the Mojave and Sonoran deserts.LocationMojave and Sonoran deserts, south‐western USA.MethodsWe augmented the sampling from a previous study by sequencing mitochondrial DNA (COI) from new material of the A. mojave group. We used phylogenetic and network analyses to identify clades and a molecular clock and lineages‐through‐time plots (LTT plots) to explore the timing and tempo of diversification. We tested for demographic expansion using neutrality tests and mismatch distributions. Species distribution models (SDMs) were constructed to compare current suitable habitat to that at the Last Glacial Maximum (LGM).ResultsPhylogenetic, network and molecular‐clock analyses identified six major clades that probably diverged during the late Miocene. The rate of diversification appears to have slowed during the Pliocene. Most clades exhibit signals of recent demographic expansion. SDMs predicted that suitable habitat shifted south and to lower elevations during the LGM.Main conclusionsPhylogeographical analyses suggest that the A. mojave group experienced a burst of diversification in the late Miocene, followed by population expansions during the Pleistocene. Six major clades with origins in the late Miocene cannot be adequately explained by the Mojave Assembly Model. We propose the novel hypothesis that Miocene extensional tectonics caused populations to diverge in allopatry by producing low‐elevation habitat barriers. Geological models, such as kinematic reconstructions, provide an ideal but underutilized framework for testing biogeographical hypotheses in these deserts and the wider Basin and Range Province.

Speciation in the dark: diversification and biogeography of the deep-sea gastropod genus Scaphander in the Atlantic Ocean.

AimThe aim of this work was to improve understanding about the mode, geography and tempo of diversification in deep‐sea organisms, using a time‐calibrated molecular phylogeny of the heterobranch gastropod genus Scaphander.LocationAtlantic and Indo‐West Pacific (IWP) oceans.MethodsTwo mitochondrial gene markers (COI and 16S) and one nuclear ribosomal gene (28S) from six Atlantic species of Scaphander, and four IWP species were used to generate a multilocus phylogenetic hypothesis using uncorrelated relaxed‐clock Bayesian methods implemented in beast and calibrated with the first occurrence of Scaphander in the fossil record (58.7–55.8 Ma).ResultsTwo main clades were supported: clade A, with sister relationships between species and subclades from the Atlantic and IWP; and clade B, with two western Atlantic sister species. Our estimates indicate that the two earliest divergences in clade A occurred between the middle Eocene and late Miocene and the most recent speciation occurred within the middle Miocene to Pleistocene. The divergence between the two western Atlantic species in clade B was estimated at late Oligocene–Pliocene.Main conclusionsThe prevailing mode of speciation in Scaphander was allopatric, but one possible case of sympatric speciation was detected between two western Atlantic species. Sister relationships between IWP and Atlantic lineages suggest the occurrence both of vicariance events caused by the closure of the Tethyan Seaway and of dispersal between the two ocean basins, probably around South Africa during episodic disruptions of the deep‐sea regional current system caused by glacial–interglacial cycles. Cladogenetic estimates do not support comparatively older diversification of deep‐sea faunas, but corroborate the hypothesis of a pulse of diversification centred in the Oligocene and Miocene epochs. Amphi‐Atlantic species were found to occur at deeper depths (bathyal–abyssal) and we hypothesize that trans‐Atlantic connectivity is maintained by dispersal between neighbouring reproductive populations inhabiting the abyssal sea floor and by dispersal across the shelf and slope of Arctic and sub‐Arctic regions.

Dispersal and speciation in purple swamphens (Rallidae:Porphyrio)

Dispersal, when accompanied by reduced gene flow and natural selection, influences speciation rates among groups of organisms. We used molecular phylogenetics, divergence time estimates, and population genetics to reconstruct the mode, pattern, and tempo of diversification within the wide-ranging purple swamphens (genus Porphyrio), with emphasis on the “supertramp” P. porphyrio. Our results suggest that the Porphyrio clade arose during the Middle Miocene in Africa, with a single colonization in the Americas and several other colonizations in Southeast Asia and the Indo-Pacific around 10 mya. We found that the widespread P. porphyrio is not monophyletic. Indeed, several subspecies and subspecies groups may represent species-level lineages. The P. p. melanotus lineage probably reached Australasia during the Pleistocene (600 kya), although some islands were colonized only in the past few hundred years. New Zealand, and some other islands, had previously been colonized (∼2.5 mya) by flying Porphyrio that evolved into flightless endemic species. Early and recent lineages are now sympatric. Widespread occupation of oceanic islands implies high dispersal and colonization rates, but gene flow probably occurs episodically and follows varying routes at different times. This pattern of colonization enables populations to differentiate and, ultimately, speciate.

Are characiform fishes Gondwanan in origin? Insights from a time-scaled molecular phylogeny of the Citharinoidei (Ostariophysi: Characiformes).

Fishes of the order Characiformes are a diverse and economically important teleost clade whose extant members are found exclusively in African and Neotropical freshwaters. Although their transatlantic distribution has been primarily attributed to the Early Cretaceous fragmentation of western Gondwana, vicariance has not been tested with temporal information beyond that contained in their fragmentary fossil record and a recent time-scaled phylogeny focused on the African family Alestidae. Because members of the suborder Citharinoidei constitute the sister lineage to the entire remaining Afro-Neotropical characiform radiation, we inferred a time-calibrated molecular phylogeny of citharinoids using a popular Bayesian approach to molecular dating in order to assess the adequacy of current vicariance hypotheses and shed light on the early biogeographic history of characiform fishes. Given that the only comprehensive phylogenetic treatment of the Citharinoidei has been a morphology-based analysis published over three decades ago, the present study also provided an opportunity to further investigate citharinoid relationships and update the evolutionary framework that has laid the foundations for the current classification of the group. The inferred chronogram is robust to changes in calibration priors and suggests that the origins of citharinoids date back to the Turonian (ca 90 Ma) of the Late Cretaceous. Most modern citharinoid genera, however, appear to have originated and diversified much more recently, mainly during the Miocene. By reconciling molecular-clock- with fossil-based estimates for the origins of the Characiformes, our results provide further support for the hypothesis that attributes the disjunct distribution of the order to the opening of the South Atlantic Ocean. The striking overlap in tempo of diversification and biogeographic patterns between citharinoids and the African-endemic family Alestidae suggests that their evolutionary histories could have been strongly and similarly influenced by Miocene geotectonic events that modified the landscape and produced the drainage pattern of Central Africa seen today.

Phylogeny and tempo of diversification in the superradiation of spiny-rayed fishes

Spiny-rayed fishes, or acanthomorphs, comprise nearly one-third of all living vertebrates. Despite their dominant role in aquatic ecosystems, the evolutionary history and tempo of acanthomorph diversification is poorly understood. We investigate the pattern of lineage diversification in acanthomorphs by using a well-resolved time-calibrated phylogeny inferred from a nuclear gene supermatrix that includes 520 acanthomorph species and 37 fossil age constraints. This phylogeny provides resolution for what has been classically referred to as the "bush at the top" of the teleost tree, and indicates acanthomorphs originated in the Early Cretaceous. Paleontological evidence suggests acanthomorphs exhibit a pulse of morphological diversification following the end Cretaceous mass extinction; however, the role of this event on the accumulation of living acanthomorph diversity remains unclear. Lineage diversification rates through time exhibit no shifts associated with the end Cretaceous mass extinction, but there is a global decrease in lineage diversification rates 50 Ma that occurs during a period when morphological disparity among fossil acanthomorphs increases sharply. Analysis of clade-specific shifts in diversification rates reveal that the hyperdiversity of living acanthomorphs is highlighted by several rapidly radiating lineages including tunas, gobies, blennies, snailfishes, and Afro-American cichlids. These lineages with high diversification rates are not associated with a single habitat type, such as coral reefs, indicating there is no single explanation for the success of acanthomorphs, as exceptional bouts of diversification have occurred across a wide array of marine and freshwater habitats.

Not drowning, (hand)waving? Molecular phylogenetics, biogeography and evolutionary tempo of the ‘Gondwanan’ midge Stictocladius Edwards (Diptera: Chironomidae)

Many insect clades, especially within the Diptera (true flies), have been considered classically ‘Gondwanan’, with an inference that distributions derive from vicariance of the southern continents. Assessing the role that vicariance has played in the evolution of austral taxa requires testing the location and tempo of diversification and speciation against the well-established predictions of fragmentation of the ancient super-continent. Several early (anecdotal) hypotheses that current austral distributions originate from the breakup of Gondwana derive from studies of taxa within the family Chironomidae (non-biting midges). With the advent of molecular phylogenetics and biogeographic analytical software, these studies have been revisited and expanded to test such conclusions better. Here we studied the midge genus Stictocladius Edwards, from the subfamily Orthocladiinae, which contains austral-distributed clades that match vicariance-based expectations. We resolve several issues of systematic relationships among morphological species and reveal cryptic diversity within many taxa. Time-calibrated phylogenetic relationships among taxa accorded partially with the predicted tempo from geology. For these apparently vagile insects, vicariance-dated patterns persist for South America and Australia. However, as often found, divergence time estimates for New Zealand at c. 50mya post-date separation of Zealandia from Antarctica and the remainder of Gondwana, but predate the proposed Oligocene ‘drowning’ of these islands. We detail other such ‘anomalous’ dates and suggest a single common explanation rather than stochastic processes. This could involve synchronous establishment following recovery from ‘drowning’ and/or deleteriously warming associated with the mid-Eocene climatic optimum (hence ‘waving’, which refers to cycles of drowning events) plus new availability of topography providing of cool running waters, or all these factors in combination. Alternatively a vicariance explanation remains available, given the uncertain duration of connectivity of Zealandia to Australia–Antarctic–South America via the Lord Howe and Norfolk ridges into the Eocene.

A comprehensive and integrative reconstruction of evolutionary history for Anomura (Crustacea: Decapoda)

BackgroundThe infraorder Anomura has long captivated the attention of evolutionary biologists due to its impressive morphological diversity and ecological adaptations. To date, 2500 extant species have been described but phylogenetic relationships at high taxonomic levels remain unresolved. Here, we reconstruct the evolutionary history—phylogeny, divergence times, character evolution and diversification—of this speciose clade. For this purpose, we sequenced two mitochondrial (16S and 12S) and three nuclear (H3, 18S and 28S) markers for 19 of the 20 extant families, using traditional Sanger and next-generation 454 sequencing methods. Molecular data were combined with 156 morphological characters in order to estimate the largest anomuran phylogeny to date. The anomuran fossil record allowed us to incorporate 31 fossils for divergence time analyses.ResultsOur best phylogenetic hypothesis (morphological + molecular data) supports most anomuran superfamilies and families as monophyletic. However, three families and eleven genera are recovered as para- and polyphyletic. Divergence time analysis dates the origin of Anomura to the Late Permian ~259 (224–296) MYA with many of the present day families radiating during the Jurassic and Early Cretaceous. Ancestral state reconstruction suggests that carcinization occurred independently 3 times within the group. The invasion of freshwater and terrestrial environments both occurred between the Late Cretaceous and Tertiary. Diversification analyses found the speciation rate to be low across Anomura, and we identify 2 major changes in the tempo of diversification; the most significant at the base of a clade that includes the squat-lobster family Chirostylidae.ConclusionsOur findings are compared against current classifications and previous hypotheses of anomuran relationships. Many families and genera appear to be poly- or paraphyletic suggesting a need for further taxonomic revisions at these levels. A divergence time analysis provides key insights into the origins of major lineages and events and the timing of morphological (body form) and ecological (habitat) transitions. Living anomuran biodiversity is the product of 2 major changes in the tempo of diversification; our initial insights suggest that the acquisition of a crab-like form did not act as a key innovation.

Time and tempo of diversification in the flora of New Caledonia

New Caledonia is well known for its rich and unique flora. Many studies have focused on the biogeographical origins of New Caledonian plants but rates of diversification on the island have scarcely been investigated. Here, dated phylogenetic trees from selected published studies were used to evaluate the time and tempo of diversification in New Caledonia. The 12 plant lineages investigated all appear to have colonized the island < 37 Mya, when New Caledonia re-emerged after a period of inundation, and the timing of these arrivals is spread across the second half of the Cenozoic. Diversification rates are not particularly high and are negatively correlated with lineage age. The palms have the fastest diversification rates and also the most recent arrival times. The lineage ages of rainforest plants suggest that this ecosystem has been present for at least 6.9 Myr. The New Caledonian flora is apparently a relatively old community that may have reached a dynamic equilibrium. Colonization by new immigrants has been possible until relatively recently and diversity-dependent processes may still be affecting the diversification rates of the earlier colonizers. Further studies on the diversification of large plant clades with exhaustive sampling should help to clarify this.

EXPLOSIVE RADIATION OF A BACTERIAL SPECIES GROUP

The current diversity of life on earth is the product of macroevolutionary processes that have shaped the dynamics of diversification. Although the tempo of diversification has been studied extensively in macroorganisms, much less is known about the rates of diversification in the exceedingly diverse and species-rich microbiota. Decreases in diversification rates over time, a signature of explosive radiations, are commonly observed in plant and animal lineages. However, the few existing analyses of microbial lineages suggest that the tempo of diversification in prokaryotes may be fundamentally different. Here, we use multilocus and genomic sequence data to test hypotheses about the rate of diversification in a well-studied pathogenic bacterial lineage, Borrelia burgdorferi sensu lato (sl). Our analyses support the hypothesis that an explosive radiation of lineages occurred near the origin of the clade, followed by a sharp decay in diversification rates. These results suggest that explosive radiations may be a general feature of evolutionary history across the tree of life.