Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution

Testing models of macroevolution, and especially the sufficiency of microevolutionary processes, requires good collaboration between molecular biologists and paleontologists. We report such a test for events around the Late Cretaceous by describing the earliest penguin fossils, analyzing complete mitochondrial genomes from an albatross, a petrel, and a loon, and describe the gradual decline of pterosaurs at the same time modern birds radiate. The penguin fossils comprise four naturally associated skeletons from the New Zealand Waipara Greensand, a Paleocene (early Tertiary) formation just above a well-known Cretaceous/Tertiary boundary site. The fossils, in a new genus (Waimanu), provide a lower estimate of 61-62 Ma for the divergence between penguins and other birds and thus establish a reliable calibration point for avian evolution. Combining fossil calibration points, DNA sequences, maximum likelihood, and Bayesian analysis, the penguin calibrations imply a radiation of modern (crown group) birds in the Late Cretaceous. This includes a conservative estimate that modern sea and shorebird lineages diverged at least by the Late Cretaceous about 74 +/- 3 Ma (Campanian). It is clear that modern birds from at least the latest Cretaceous lived at the same time as archaic birds including Hesperornis, Ichthyornis, and the diverse Enantiornithiformes. Pterosaurs, which also coexisted with early crown birds, show notable changes through the Late Cretaceous. There was a decrease in taxonomic diversity, and small- to medium-sized species disappeared well before the end of the Cretaceous. A simple reading of the fossil record might suggest competitive interactions with birds, but much more needs to be understood about pterosaur life histories. Additional fossils and molecular data are still required to help understand the role of biotic interactions in the evolution of Late Cretaceous birds and thus to test that the mechanisms of microevolution are sufficient to explain macroevolution.     

Two waves of colonization straddling the K–Pg boundary formed the modern reef fish fauna

Living reef fishes are one of the most diverse vertebrate assemblages on Earth. Despite its prominence and ecological importance, the origins and assembly of the reef fish fauna is poorly described. A patchy fossil record suggests that the major colonization of reef habitats must have occurred in the Late Cretaceous and early Palaeogene, with the earliest known modern fossil coral reef fish assemblage dated to 50 Ma. Using a phylogenetic approach, we analysed the early evolutionary dynamics of modern reef fishes. We find that reef lineages successively colonized reef habitats throughout the Late Cretaceous and early Palaeogene. Two waves of invasion were accompanied by increasing morphological convergence: one in the Late Cretaceous from 90 to 72 Ma and the other immediately following the end-Cretaceous mass extinction. The surge in reef invasions after the Cretaceous–Palaeogene boundary continued for 10 Myr, after which the pace of transitions to reef habitats slowed. Combined, these patterns match a classic niche-filling scenario: early transitions to reefs were made rapidly by morphologically distinct lineages and were followed by a decrease in the rate of invasions and eventual saturation of morphospace. Major alterations in reef composition, distribution and abundance, along with shifts in climate and oceanic currents, occurred during the Late Cretaceous and early Palaeogene interval. A causal mechanism between these changes and concurrent episodes of reef invasion remains obscure, but what is clear is that the broad framework of the modern reef fish fauna was in place within 10 Myr of the end-Cretaceous extinction.     

Heinrichsia cheilanthoides gen. et sp. nov., a fossil fern in the family Pteridaceae (Polypodiales) from the Cretaceous amber forests of Myanmar

Divergence time estimates suggest that most clades constituting the fern family Pteridaceae (Polypodiales) were in existence by the Early Cretaceous. However, fossil evidence to corroborate this remains exceedingly rare. Burmese amber is an important source of new information on the radiation of derived fern lineages during the Cretaceous Terrestrial Revolution. This study describes Heinrichsia cheilanthoides gen. et sp. nov., a fern with suggested affinities to Pteridaceae, based on fertile foliage portions preserved in Early Cretaceous (~100 Ma) amber from Myanmar. Heinrichsia cheilanthoides is characterized by a pinnate‐pinnatifid frond that bears apical, marginal sori protected by a pseudoindusium. Sporangia are of the polypod type and contain tetrahedral‐globose, trilete spores with a striate perine. This discovery provides a new calibration point to test and refine molecular clock‐based concepts of the evolutionary history of the Pteridaceae. Heinrichsia cheilanthoides further substantiates the suggestion that the Cretaceous forests of Myanmar were home to a rich fern flora.     

BOOM AND BUST: ANCIENT AND RECENT DIVERSIFICATION IN BICHIRS (POLYPTERIDAE: ACTINOPTERYGII), A RELICTUAL LINEAGE OF RAY‐FINNED FISHES

Understanding the history that underlies patterns of species richness across the Tree of Life requires an investigation of the mechanisms that not only generate young species‐rich clades, but also those that maintain species‐poor lineages over long stretches of evolutionary time. However, diversification dynamics that underlie ancient species‐poor lineages are often hidden due to a lack of fossil evidence. Using information from the fossil record and time calibrated molecular phylogenies, we investigate the history of lineage diversification in Polypteridae, which is the sister lineage of all other ray‐finned fishes (Actinopterygii). Despite originating at least 390 million years (Myr) ago, molecular timetrees support a Neogene origin for the living polypterid species. Our analyses demonstrate polypterids are exceptionally species depauperate with a stem lineage duration that exceeds 380 million years (Ma) and is significantly longer than the stem lineage durations observed in other ray‐finned fish lineages. Analyses of the fossil record show an early Late Cretaceous (100.5–83.6 Ma) peak in polypterid genus richness, followed by 60 Ma of low richness. The Neogene species radiation and evidence for high‐diversity intervals in the geological past suggest a “boom and bust” pattern of diversification that contrasts with common perceptions of relative evolutionary stasis in so‐called “living fossils.”     

Time scale for cyclostome evolution inferred with a phylogenetic diagnosis of hagfish and lamprey cDNA sequences

The Cyclostomata consists of the two orders Myxiniformes (hagfishes) and Petromyzoniformes (lampreys), and its monophyly has been unequivocally supported by recent molecular phylogenetic studies. Under this updated vertebrate phylogeny, we performed in silico evolutionary analyses using currently available cDNA sequences of cyclostomes. We first calculated the GC-content at four-fold degenerate sites (GC(4)), which revealed that an extremely high GC-content is shared by all the lamprey species we surveyed, whereas no striking pattern in GC-content was observed in any of the hagfish species surveyed. We then estimated the timing of diversification in cyclostome evolution using nucleotide and amino acid sequences. We obtained divergence times of 470-390 million years ago (Mya) in the Ordovician-Silurian-Devonian Periods for the interordinal split between Myxiniformes and Petromyzoniformes; 90-60 Mya in the Cretaceous-Tertiary Periods for the split between the two hagfish subfamilies, Myxininae and Eptatretinae; 280-220 Mya in the Permian-Triassic Periods for the split between the two lamprey subfamilies, Geotriinae and Petromyzoninae; and 30-10 Mya in the Tertiary Period for the split between the two lamprey genera, Petromyzon and Lethenteron. This evolutionary configuration indicates that Myxiniformes and Petromyzoniformes diverged shortly after the common ancestor of cyclostomes split from the future gnathostome lineage. Our results also suggest that intra-subfamilial diversification in hagfish and lamprey lineages (especially those distributed in the northern hemisphere) occurred in the Cretaceous or Tertiary Periods.     

Origin and differentiation of endemism in the flora of China

The present paper analyzed 239 endemic genera in 67 families in the flora of seed plants in China.The results showed that there are five families containing more than ten endemic genera,namely,Gesneriaceae (27),which hereafter refers to the number of endemic genera in China,Composite (20),Labiatae (12),Cruciferae (11),and Umbelliferae (10),15 families with two endemic genera,and another 30 families with only one endemic genus.Four monotypic families (Ginkgoaceae,Davidiaceae,Eucommiaceae and Acanthochlamydaceae)are the most ancient,relict and characteristic in the flora of seed plants in China.Based on integrative data of systematics,fossil history,and morphological and molecular evidence of these genera,their origin,evolution and relationships were discussed.In gymnosperms,all endemic genera are relicts of the Arctic-Tertiary flora,having earlier evolutionary history,and can be traced back to the Cretaceous or to the Jurassic and even earlier.In angiosperms,the endemic genera are mostly relicts,and are represented in all lineages in the"Eight-Class System ofClassification of Angiosperms",and endemism can be found in almost every evolutionary stage of extant angiosperms.The relict genera once occupied huge areas in the northern hemisphere in the Tertiary or the late Cretaceous,while neo-endemism mostly originated in the late Tertiary.They came from Arctic-Tertiary,Paleo-tropical-Tertiary and Tethys-Tertiary florisitic elements,and the blend of the three elements with many genera of autochthonous origin.The endemism was formed when some dispersal routes such as the North Atlantic Land Bridge,and the Bering Bridge became discontinuous during the Tertiary,as well as the climate change and glaciations in the late Tertiary and the Quaternary.Therefore,the late Tertiary is the starting point of extant endemism of the flora in China.     

中国植物区系中的特有性及其起源和分化

对中国植物区系中的239个特有属,分属67个科,进行了分析研究,列出了这些特有属在种子植物各个科的分布,现代地理分布范围。结果表明含特有属在10个以上的有5个科即Gesneriaceae,Compositae,Labiatae,Cruiciferae,Umbelliferae;其中以Gesneriaceae居榜首(27属),Compositae位居第二(20属),Labiatae有12属,居第三。含2属的科有15个,含1属的科有30个;其中Ginkgaceae,Davidiaceae,Eucommiaceae,Acanthochlamydaceae组成了中国植物区系最具古老性、特有性和代表性的4个单型科。在此基础上,从特有属在被子植物八纲系统各个纲的分布特点,以及在各个科组成和系统关系及已有地质、化石历史和系统学,形态,分子证据论述了这些特有属的起源、系统关系及在植物地理上的关系。在裸子植物中,特有属最为丰富,几乎皆是地质历史上北极-第三纪成分的残遗,起源时间较早,可追溯到白垩纪或更早。被子植物中,中国特有属存在于八纲被子植物的所有纲中,几乎在现代被子植物各个演化阶段均有古老残遗的特有类群存在,同时也不乏新特有类群尤其是在演化的高级阶段的类群。从起源上看,被子植物的古特有属主要发生于晚白垩纪和早第三纪,地质历史上大都占有广阔的分布区;新特有属多发生在新第三纪以后。其源头主要是北极第三纪、古热带第三纪(冈瓦纳第三纪)和古地中海第三纪的奇妙结合,不少类群是就地起源的;特有性是在第三纪中晚期以后北半球气候变迁,迁移途径(如北大西洋陆桥和白令陆桥)中断后形成的,这一时期是我国特有属形成发展的起始标志。     

Primate origins: implications of a cretaceous ancestry

It has long been accepted that the adaptive radiation of modern placental mammals, like that of modern birds, did not begin until after the Cretaceous/Tertiary (K/T) boundary 65 million years (Ma) ago, following the extinction of the dinosaurs. The first undoubted fossil relatives of modern primates appear in the record 55 Ma ago. However, in agreement with evidence from molecular phylogenies calibrated with dates from denser parts of the fossil record, a statistical analysis of the primate record allowing for major gaps now indicates a Cretaceous origin of euprimates 80-90 Ma ago. If this interpretation is correct, primates overlapped with dinosaurs by some 20 Ma prior to the K/T boundary, and the initial radiation of primates was probably truncated as part of the major extinction event that occurred at the end of the Cretaceous. Following a review of evidence for an early origin of primates, implications of this are discussed with respect to the likely ancestral condition for primates, including a southern continental area of origin and moderately large body size. The known early Tertiary primates are re-interpreted as northern continental offshoots of a 'second wave' of primate evolution.     

Molecular evidence for the age, origin, and evolutionary history of the American desert plant genus Tiquilia (Boraginaceae)

Although the deserts of North America are of very recent origin, their characteristic arid-adapted endemic plant lineages have been suggested to be much older. Earlier researchers have hypothesized that the ancestors of many of these modern desert lineages first adapted to aridity in highly localized arid or semi-arid sites as early as the late Cretaceous or early Tertiary, and that these lineages subsequently spread and diversified as global climate became increasingly arid during the Cenozoic. No study has explicitly examined these hypotheses for any North American arid-adapted plant group. The current paper tests these hypotheses using the genus Tiquilia (Boraginaceae), a diverse North American desert plant group. A strongly supported phylogeny of the genus is estimated using combined sequence data from three chloroplast markers (matK, ndhF, and rps16) and two nuclear markers (ITS and waxy). Ages of divergence events within the genus are estimated using penalized likelihood and a molecular clock approach on the ndhF tree for Tiquilia and representative outgroups, including most of the major lineages of Boraginales. The dating analysis suggests that the stem lineage of Tiquilia split from its nearest extant relative in the Paleocene or Eocene ( approximately 59-48 Ma). This was followed by a relatively long period before the first divergence in the crown group near the Eocene/Oligocene boundary ( approximately 33-29 Ma), shortly after the greatest Cenozoic episode of rapid aridification. Divergence of seven major lineages of Tiquilia is dated to the early-to-mid Miocene ( approximately 23-13 Ma). Several major lineages show a marked increase in diversification concomitant with the onset of more widespread semi-arid and then arid conditions beginning in the late Miocene ( approximately 7 Ma). This sequence of divergence events in Tiquilia agrees well with earlier researchers' ideas concerning North American desert flora assembly.     

The age and diversification of terrestrial New World ecosystems through Cretaceous and Cenozoic time

Eight ecosystems that were present in the Cretaceous about 100 Ma (million years ago) in the New World eventually developed into the 12 recognized for the modern Earth. Among the forcing mechanisms that drove biotic change during this interval was a decline in global temperatures toward the end of the Cretaceous, augmented by the asteroid impact at 65 Ma and drainage of seas from continental margins and interiors; separation of South America from Africa beginning in the south at ca. 120 Ma and progressing northward until completed 90-100 Ma; the possible emission of 1500 gigatons of methane and CO(2) attributed to explosive vents in the Norwegian Sea at ca. 55 Ma, resulting in a temperature rise of 5°-6°C in an already warm world; disruption of the North Atlantic land bridge at ca. 45 Ma at a time when temperatures were falling; rise of the Andes Mountains beginning at ca. 40 Ma; opening of the Drake Passage between South America and Antarctica at ca. 32 Ma with formation of the cold Humboldt at ca. 30 Ma; union of North and South America at ca. 3.5 Ma; and all within the overlay of evolutionary processes. These processes generated a sequence of elements (e.g., species growing in moist habitats within an overall dry environment; gallery forests), early versions (e.g., mangrove communities without Rhizophora until the middle Eocene), and essentially modern versions of present-day New World ecosystems. As a first approximation, the fossil record suggests that early versions of aquatic communities (in the sense of including a prominent angiosperm component) appeared early in the Middle to Late Cretaceous, the lowland neotropical rainforest at 64 Ma (well developed by 58-55 Ma), shrubland/chaparral-woodland-savanna and grasslands around the middle Miocene climatic optimum at ca. 15-13 Ma, deserts in the middle Miocene/early Pliocene at ca. 10 Ma, significant tundra at ca. 7-5 Ma, and alpine tundra (páramo) shortly thereafter when cooling temperatures were augmented by high elevations attained, for example, in the Andes<10 Ma and especially after 7-6 Ma.     

Out of the Neotropics: Late Cretaceous colonization of Australasia by American arthropods

The origins of tropical southwest Pacific diversity are traditionally attributed to southeast Asia or Australia. Oceanic and fragment islands are typically colonized by lineages from adjacent continental margins, resulting in attrition of diversity with distance from the mainland. Here, we show that an exceptional tropical family of harvestmen with a trans-Pacific disjunct distribution has its origin in the Neotropics. We found in a multi-locus phylogenetic analysis that the opilionid family Zalmoxidae, which is distributed in tropical forests on both sides of the Pacific, is a monophyletic entity with basal lineages endemic to Amazonia and Mesoamerica. Indo-Pacific Zalmoxidae constitute a nested clade, indicating a single colonization event. Lineages endemic to putative source regions, including Australia and New Guinea, constitute derived groups. Divergence time estimates and probabilistic ancestral area reconstructions support a Neotropical origin of the group, and a Late Cretaceous (ca 82 Ma) colonization of Australasia out of the Fiji Islands and/or Borneo, which are consistent with a transoceanic dispersal event. Our results suggest that the endemic diversity within traditionally defined zoogeographic boundaries might have more complex evolutionary origins than previously envisioned.     

Paleogene stable isotope events

The oxygen isotope record in Paleogene benthic Foraminifera shows that at the base of the Paleogene the ocean deep waters had a temperature of about 10°C, rising to about 12°C at the base of the Eocene and cooling between 51 Ma and 49 Ma to about 9°C. The most dramatic event occurred just after the Eocene/Oligocene boundary, at about 35.8 Ma, when ocean deep waters cooled by several degrees within 10⁴–10⁵ yr, probably in association with temporary glaciation in the Antarctic region. Another more intense glacial event in Antarctica may have occurred later in the Oligocene, at about 31 Ma and a third near the top of the Oligocene at 24 Ma.In the marine carbon isotope record a very rapid negative excursion occurred precisely at the Cretaceous/Tertiary boundary. A recovery to unusually positive values in the Late Paleocene was followed by a second negative excursion close to the Paleocene/Eocene boundary that was even more extreme in magnitude although it was not as rapid. These major carbon isotope events permit very accurate stratigraphic correlation; there are many other smaller features in the carbon isotope record that will also prove useful for this purpose.     

Origin and differentiation of endemism in the flora of China

The present paper analyzed 239 endemic genera in 67 families in the flora of seed plants in China. The results showed that there are five families containing more than ten endemic genera, namely, Gesneriaceae (27), which hereafter refers to the number of endemic genera in China, Composite (20), Labiatae (12), Cruciferae (11), and Umbelliferae (10), 15 families with two endemic genera, and another 30 families with only one endemic genus. Four monotypic families (Ginkgoaceae, Davidiaceae, Eucommiaceae and Acanthochlamydaceae) are the most ancient, relict and characteristic in the flora of seed plants in China. Based on integrative data of systematics, fossil history, and morphological and molecular evidence of these genera, their origin, evolution and relationships were discussed. In gymnosperms, all endemic genera are relicts of the Arctic-Tertiary flora, having earlier evolutionary history, and can be traced back to the Cretaceous or to the Jurassic and even earlier. In angiosperms, the endemic genera are mostly relicts, and are represented in all lineages in the “Eight-Class System of Classification of Angiosperms”, and endemism can be found in almost every evolutionary stage of extant angiosperms. The relict genera once occupied huge areas in the northern hemisphere in the Tertiary or the late Cretaceous, while neo-endemism mostly originated in the late Tertiary. They came from Arctic-Tertiary, Paleo-tropical-Tertiary and Tethys-Tertiary florisitic elements, and the blend of the three elements with many genera of autochthonous origin. The endemism was formed when some dispersal routes such as the North Atlantic Land Bridge, and the Bering Bridge became discontinuous during the Tertiary, as well as the climate change and glaciations in the late Tertiary and the Quaternary. Therefore, the late Tertiary is the starting point of extant endemism of the flora in China. __________ Translated from Acta Botanica Yunnanica, 2005, 27(6): 577–604 [译自: 云南植物研究]     

The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals

Extant terrestrial biodiversity arguably is driven by the evolutionary success of angiosperm plants, but the evolutionary mechanisms and timescales of angiosperm-dependent radiations remain poorly understood. The Scarabaeoidea is a diverse lineage of predominantly plant- and dung-feeding beetles. Here, we present a phylogenetic analysis of Scarabaeoidea based on four DNA markers for a taxonomically comprehensive set of specimens and link it to recently described fossil evidence. The phylogeny strongly supports multiple origins of coprophagy, phytophagy and anthophagy. The ingroup-based fossil calibration of the tree widely confirmed a Jurassic origin of the Scarabaeoidea crown group. The crown groups of phytophagous lineages began to radiate first (Pleurostict scarabs: 108 Ma; Glaphyridae between 101 Ma), followed by the later diversification of coprophagous lineages (crown-group age Scarabaeinae: 76 Ma; Aphodiinae: 50 Ma). Pollen feeding arose even later, at maximally 62 Ma in the oldest anthophagous lineage. The clear time lag between the origins of herbivores and coprophages suggests an evolutionary path driven by the angiosperms that first favoured the herbivore fauna (mammals and insects) followed by the secondary radiation of the dung feeders. This finding makes it less likely that extant dung beetle lineages initially fed on dinosaur excrements, as often hypothesized.     

The complete mitochondrial genome of Alligator mississippiensis and the separation between recent archosauria (birds and crocodiles)

The complete mitochondrial genome of the alligator, Alligator mississippiensis, was sequenced. The size of the molecule is 16,642 nucleotides. Previously reported rearrangements of tRNAs in crocodile mitochondrial genomes were confirmed and, relative to mammals, no other deviations of gene order were observed. The analysis of protein-coding genes of the alligator showed an evolutionary rate that is roughly the same as in mammals. Thus, the evolutionary rate in the alligator is faster than that in birds as well as that in cold-blooded vertebrates. This contradicts hypotheses of constant body temperatures or high metabolic rate being correlated with elevated molecular evolutionary rates. It is commonly acknowledged that birds are the closest living relatives to crocodiles. Birds and crocodiles represent the only archosaurian survivors of the mass extinction at the Cretaceous/Tertiary boundary. On the basis of mitochondrial protein- coding genes, the Haemothermia hypothesis, which defines birds and mammals as sister groups and thus challenges the traditional view, could be rejected. Maximum-likelihood branch length data of amino acid sequences suggest that the divergence between the avian and crocodilian lineages took place at approximately equal to 254 MYA.      

Diversification Times and Biogeographic Patterns in Apiales

This study provides an overview of the historical biogeography of the major clades of Apiales based on extensive taxon sampling from all major lineages of the order, and character sampling of sequence data from the plastid rpl16 intron and trnD-trnY-trnE-trnT intergenic spacers. Divergence times were estimated in BEAST using relaxed molecular clocks and six calibration points from three families. Biogeographic reconstructions were estimated in DIVA and Lagrange using stratified and non-stratified models, addressing alternative scenarios for taxa with conflicting or poorly supported placements. Our analyses in BEAST estimated the origin of Apiales to Australasia in the Early Cretaceous (c.117 Ma). Most major clades also appear to have originated in Australasia, with the youngest family (Apiaceae) originating in the Late Cretaceous, c. 87 Ma. Diversification of the early lineages appears to be influenced by vicariance events related to the break up of Africa and Australasia (Torricelliaceae from Griseliniaceae and Apiineae), Australasia from Zealandia (e.g., Myodocarpaceae and Araliaceae), and Antarctica from South America, Australia, and possibly Africa (main lineages of Apiaceae). Long-distance dispersal appears as the likely explanation for many younger lineages within major clades, including Subantarctic pathways (e.g., Griseliniaceae and Azorelloideae), across the Pacific and Indian Ocean Basins (e.g., Pittosporaceae and Araliaceae), from Asia across Europe into the Americas (Araliaceae).     

Cretaceous origin and repeated tertiary diversification of the redefined butterflies

Although the taxonomy of the ca 18 000 species of butterflies and skippers is well known, the family-level relationships are still debated. Here, we present, to our knowledge, the most comprehensive phylogenetic analysis of the superfamilies Papilionoidea, Hesperioidea and Hedyloidea to date based on morphological and molecular data. We reconstructed their phylogenetic relationships using parsimony and Bayesian approaches. We estimated times and rates of diversification along lineages in order to reconstruct their evolutionary history. Our results suggest that the butterflies, as traditionally understood, are paraphyletic, with Papilionidae being the sister-group to Hesperioidea, Hedyloidea and all other butterflies. Hence, the families in the current three superfamilies should be placed in a single superfamily Papilionoidea. In addition, we find that Hedylidae is sister to Hesperiidae, and this novel relationship is supported by two morphological characters. The families diverged in the Early Cretaceous but diversified after the Cretaceous-Palaeogene event. The diversification of butterflies is characterized by a slow speciation rate in the lineage leading to Baronia brevicornis, a period of stasis by the skippers after divergence and a burst of diversification in the lineages leading to Nymphalidae, Riodinidae and Lycaenidae.     

Early diversification trend and Asian origin for extent bat lineages

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

Dating the evolutionary origins of wrasse lineages (Labridae) and the rise of trophic novelty on coral reefs

We estimated ages of divergence between major labrid tribes and the timing of the evolution of trophic novelty. Sequence data for 101 labrid taxa and 14 outgroups consisting of two mitochondrial gene regions (12s, 16s), and two nuclear protein-coding genes (RAG2, TMO4c4), a combined 2567 bp of sequence, were examined using novel maximum likelihood, maximum parsimony and mixed model Bayesian inference methods. These analyses yielded well supported trees consistent with published phylogenies. Bayesian inference using five fossil calibration points estimated the minimum ages of lineages. With origins in the late Cretaceous to early tertiary, the family diversified quickly with both major lineages (hypsigenyine and julidine) present at approximately 62.7 Ma, shortly after the K/T boundary. All lineages leading to major tribes were in place by the beginning of the Miocene (23 Ma) with most diversification in extant lineages occurring within the Miocene. Optimisation of trophic information onto the chronogram revealed multiple origins of novel feeding modes with two distinct periods of innovation. The Palaeocene/Eocene saw the origins of feeding modes that are well represented in other families: gastropod feeders, piscivores and browsing herbivores. A wave of innovation in the Oligocene/Miocene resulted in specialized feeding modes, rarely seen in other groups: coral feeding, foraminifera feeding and fish cleaning. There is little evidence of a general relationship between trophic specialization and species diversity. The current trophic diversity of the Labridae is a result of the accumulation of feeding modes dating back to the K/T boundary at 65 Ma, with all major feeding modes on present day reefs already in place 7.5 million years ago.     

Bees diversified in the age of eudicots

Reliable estimates on the ages of the major bee clades are needed to further understand the evolutionary history of bees and their close association with flowering plants. Divergence times have been estimated for a few groups of bees, but no study has yet provided estimates for all major bee lineages. To date the origin of bees and their major clades, we first perform a phylogenetic analysis of bees including representatives from every extant family, subfamily and almost all tribes, using sequence data from seven genes. We then use this phylogeny to place 14 time calibration points based on information from the fossil record for an uncorrelated relaxed clock divergence time analysis taking into account uncertainties in phylogenetic relationships and the fossil record. We explore the effect of placing a hard upper age bound near the root of the tree and the effect of different topologies on our divergence time estimates. We estimate that crown bees originated approximately 123 Ma (million years ago) (113–132 Ma), concurrently with the origin or diversification of the eudicots, a group comprising 75 per cent of angiosperm species. All of the major bee clades are estimated to have originated during the Middle to Late Cretaceous, which is when angiosperms became the dominant group of land plants.