Systematics and evolution of the Pan‐Alcidae (Aves,Charadriiformes)

Puffins, auks and their allies in the wing‐propelled diving seabird clade Pan‐Alcidae (Charadriiformes) have been proposed to be key pelagic indicators of faunal shifts in Northern Hemisphere oceans. However, most previous phylogenetic analyses of the clade have focused only on the 23 extant alcid species. Here we undertake a combined phylogenetic analysis of all previously published molecular sequence data (～ 12 kb) and morphological data (n = 353 characters) with dense species level sampling that also includes 28 extinct taxa. We present a new estimate of the patterns of diversification in the clade based on divergence time estimates that include a previously vetted set of twelve fossil calibrations. The resultant time trees are also used in the evaluation of previously hypothesized paleoclimatic drivers of pan‐alcid evolution. Our divergence dating results estimate the split of Alcidae from its sister taxon Stercorariidae during the late Eocene (～ 35 Ma), an evolutionary hypothesis for clade origination that agrees with the fossil record and that does not require the inference of extensive ghost lineages. The extant dovekie Alle alle is identified as the sole extant member of a clade including four extinct Miocene species. Furthermore, whereas an Uria + Alle clade has been previously recovered from molecular analyses, the extinct diversity of closely related Miocepphus species yields morphological support for this clade. Our results suggest that extant alcid diversity is a function of Miocene diversification and differential extinction at the Pliocene–Pleistocene boundary. The relative timing of the Middle Miocene climatic optimum and the Pliocene–Pleistocene climatic transition and major diversification and extinction events in Pan‐Alcidae, respectively, are consistent with a potential link between major paleoclimatic events and pan‐alcid cladogenesis.     

A Plains‐wanderer (Pedionomidae) that did not wander plains: a new species from the Oligocene of South Australia

The remarkable fauna of Australia evolved in isolation from other landmasses for millions of years, yet understanding the evolutionary history of endemic avian lineages on the continent is confounded by the ability of birds to disperse over geographical barriers even after vicariance events. The Plains‐wanderer Pedionomus torquatus (Charadriiformes) is an enigmatic, predominantly sedentary, quail‐like bird that occurs exclusively in sparse native grasslands of southeastern Australia. It is the only known species of its family (Pedionomidae), and its closest relatives are the South American seedsnipes (Thinocoridae). Here we describe a further representative of this lineage, Oligonomus milleri gen. et sp. nov., from the Late Oligocene of South Australia (26–24 Ma), which pre‐dates the earliest record of P. torquatus by c. 22 Ma and attests to the presence of this lineage during Australia's period of isolation (50–15 Ma). Based on the morphology of the coracoid and the palynological record, we propose that O. milleri and P. torquatus were ecologically disparate taxa and that, similar to coeval marsupials, O. milleri inhabited well‐wooded habitats, suggesting that the preference for grassland in the extant P. torquatus and thinocorids is likely to be convergent and not ancestral. The speciation event leading to the evolution of the extant Plains‐wanderer was probably triggered by the spread of grasslands across Australia in the Late Miocene–Pliocene, which this record pre‐dates. The presence of a pedionomid in the Late Oligocene of Australia strengthens the hypothesis of a Gondwanan divergence of the lineages giving rise to Thinocoridae and Pedionomidae.     

A temporally dynamic approach for cladistic biogeography and the processes underlying the biogeographic patterns of North American deserts

We implemented a temporally dynamic approach to the cladistic biogeographic analysis of 13 areas of North American deserts and several plant and animal taxa. We undertook a parsimony analysis of paralogy‐free subtrees based on 43 phylogenetic hypotheses of arthropod, vertebrate and plant taxa, assigning their nodes to three different time slices based on their estimated minimum ages: Early‐Mid‐Miocene (23?7 Ma), Late Miocene/Pliocene (6.9?2.5 Ma) and Pleistocene (2.4?0.011 Ma). The analyses resulted in three general area cladograms, one for each time slice, showing different area relationships. They allowed us to detect influences of different geological and palaeoclimatological events of the Early‐Mid‐Miocene, Late Miocene/Pliocene and Pleistocene that might have affected the diversification of the desert biota. Several diversification events in the deserts of North America might have been driven by Neogene uplift, marine incursion and the opening of the California Gulf during the Miocene–Pliocene, whereas climatic fluctuations had the highest impact during the Pleistocene.     

Climatic fluctuations shape the phylogeography of a mesic direct‐developing frog from the south‐western Australian biodiversity hotspot

Aim To assess phylogeographic pattern throughout the range of Metacrinia nichollsi in order to develop specific biogeographical hypotheses for the wet forests of south‐western Australia. This was carried out by contrasting a direct‐developing frog species, M. nichollsi, that breeds independently of free surface water with conventional, aquatic breeders and highly specialized direct developers. Location Wet forests of the south‐western Australian biodiversity hotspot – an area of high species richness and endemism for myobatrachid frogs and many other faunal groups. Methods We compiled an extensive phylogeographic data set from field‐collected samples based on mitochondrial ND2 sequences. Phylogenetic analyses combined with estimates of divergence times were used to build a model of major biogeographical events affecting the species. Phylogeographic analyses were used to provide insights into smaller‐scale processes acting within each major lineage. Results Phylogenetic analysis recovered three major lineages, with divergence dates coincident with late Miocene–early Pliocene arid cycles. One lineage was confined to geographically isolated populations in the Stirling Ranges (Stirling Ranges Lineage, SRL). The continuous range of M. nichollsi was split into two: the Main Range Lineage (MRL) and the Southern Coastal Lineage (SCL). The SCL displays a strong drainage‐based population structure, whereas the MRL displays a strong signature of recent expansion, suggesting that these two lineages have had very different biogeographical histories. Main conclusions Late Miocene–Pliocene aridity appears to have isolated populations in the Stirling Ranges and resulted in the formation of two additional lineages on a north–south gradient that are independent of southward‐flowing drainage systems. Our results demonstrate that climatic fluctuations are likely to have generated fine‐scale phylogeographic structure within M. nichollsi and that catchment regions are important refugia during arid cycles.     

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.     

A protocol for temporal calibration of general area cladograms

Aim To describe a protocol for incorporating a temporal dimension into historical biogeographical analysis, while maintaining the essential independence of all datasets, involving the generation of general area cladograms. Location Global. Methods General area cladograms (GACs) are a reconstruction of the evolutionary history of a set of areas and unrelated clades within those areas. Nodes on a GAC correspond to speciation events in a group of taxa; general nodes are those at which multiple unrelated clades speciate. We undertake temporal calibration of GACs using molecular clock estimates of splitting events between extant taxa as well as first appearance data from the fossil record. We present two examples based on re‐analysis of previously published data: first, a temporally calibrated GAC generated from secondary Brooks parsimony analysis (BPA) of six extant bird clades from the south‐west of North America using molecular clock estimates of divergence times; and second, an analysis of African Neogene mammals based on a phylogenetic analysis for comparing trees (PACT) analysis. Results A hypothetical example demonstrates how temporal calibration reveals potentially critical information about the timing of both unique and general events, while also illustrating instances of incongruence between dates generated from molecular clock estimates and fossils. For the African Neogene mammal dataset, our analysis reveals that most mammal clades underwent geodispersal associated with the Neogene climatic optimum (c. 16 Ma) and vicariant speciation in central Africa correlated with increased aridity and cooler temperatures around 2.5 Ma. Main conclusions Temporally calibrated GACs are valuable tools for assessing whether coordinated patterns of speciation are associated with large‐scale climatic or tectonic phenomena.     

The importance of Late Miocene faunal exchanges between Eastern Mediterranean areas and Central Europe

Recent studies of mammal faunas from the Vienna and Pannonian Basins—in particular the assemblage from Kohfidisch in Burgenland (Austria)—provide new data on the faunal turnover at the Vallesian—Turolian transition. They demonstrated a considerable influence of the faunal exchanges between Greco-Iranian, Eastern European and Central European faunal provinces on renewal of mammal communities in Central Europe, particularly at MN10/MN11 boundary around 8.7 Ma. Five new comers from the Balkano-Iranian region (Gazella aff. pigrimi, ?Nisidorcas, Tragoportax gaudryi, Protoryx and Palaeoryx) coexisted in the Early Turolian of Central Europe with the Middle Miocene autochthonous (Orygotherium, Dorcatherium naui, Micromeryx, Euprox, Amphiprox anocerus and Miotragocerus pannoniae) and Late Miocene invaders from Eastern Europe (Procapreolus and Cervavitus). Dispersal events were close related to palaeoenvironmental and climatic changes.     

Descriptions and phylogenetic relationships of a new genus and two new species of Oligo‐Miocene cormorants (Aves: Phalacrocoracidae) from Australia

Tertiary cormorant fossils (Aves: Phalacrocoracidae) from Late Oligocene deposits in Australia are described. They derive from the Late Oligocene – Early Miocene (26–24 Mya) Etadunna and Namba Formations in the Lake Eyre and Lake Frome Basins, South Australia, respectively. A new genus, Nambashag gen. nov. , with two new species ( Nambashag billerooensis sp. nov. , 30 specimens; Nambashag microglaucus sp. nov. , 14 specimens), has been established. Phylogenetic analyses based on 113 morphological and two integumentary characters indicated that Nambashag is the sister taxon to the Early Miocene Nectornis miocaenus of Europe and all extant phalacrocoracids. As Nambashag, Nectornis, and extant phalacrocoracids constitute a strongly supported clade sister to Anhinga species, the fossil taxa have been referred to Phalacrocoracidae. Sulids and Fregata were successive sister taxa to the Phalacrocoracoidea, i.e. phalacrocoracids + Anhinga. As phalacrocoracids lived in both Europe and Australia during the Late Oligocene and no older phalacrocoracid taxa are known, the biogeographical origin of cormorants remains unanswered. The phylogenetic relationships of extant taxa were not wholly resolved, but contrary to previous morphological analyses, considerable concordance was found with relationships recovered by recent molecular analyses. Microcarbo is sister to all other extant phalacrocoracids, and all Leucocarbo species form a well‐supported clade. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 277–314.     

The age of immigration of the vertebrate faunas found at Gargano (Apulia,Italy) and Scontrone (l’Aquila,Italy)

Recently a discussion is taking place about the Scontrone (l’Aquila) and Gargano (Apulia, Italy) mammal faunas and the age of their immigration. Mazza and Rustioni (2008) dated the Scontrone mammal fossils as Tortonian on the basis of their position in the Lithothamnium Limestone and came to the conclusion that some elements of the Scontrone and Gargano faunas must have colonised the area in Oligocene or Early Miocene times. Van den Hoek Ostende et al. (2009) disagreed with this interpretation and suggested a Late Miocene (10 Ma) age for the time of immigration. We think the arguments to place Scontrone in the Tortonian are not convincing. An analysis of the potential ancestors of each of the Gargano faunal components shows that a Messinian age for the immigration is fully compatible with the distribution of these ancestors in the European Miocene.     

Environmental control on the biogeographical distribution of Desmanella (Soricomorpha,Mammalia) in the Miocene of the Iberian Peninsula

Abstract: This paper reports the first record of Desmanella (Mammalia, Soricomorpha) from the Granada Basin in southern Iberian Peninsula, which represents its south‐westernmost occurrence in all Eurasia. It is a controversial taxon whose systematic assignment has been discussed for a long time. This genus belongs to the family Talpidae, a group of insectivores that include extant moles, shrew moles and desmans. Desmanella was very abundant in the late Miocene and early Pliocene of Eurasia, including several basins of northern Iberian Peninsula, but it has not been found until now in southern Iberia. Previous studies have shown that the faunal record and the climatic conditions during the Neogene in the Granada Basin were different from other areas of the Iberian Peninsula. The particular distribution of Desmanella supports the hypothesis that specific climatic features (wetter than neighbouring areas) prevailed in the Granada Basin during the late Turolian (late Miocene).     

Detarieae sensu lato (Fabaceae) from the Late Oligocene (27.23 Ma) Guang River flora of north‐western Ethiopia

New species of caesalpinioid legumes, Cynometra sensu lato and Afzelia, are described from the Late Oligocene (27.23 Ma) Guang River flora in north‐western Ethiopia. Both taxa show leaf characteristics that are shared with extant species in the Guineo‐Congolian, Sudanian and/or Zambezian regions of Africa today. The presence of these two species in Ethiopia during the Palaeogene provides further evidence of the importance of the legume tribe Detarieae in northern and north‐eastern Africa throughout much of the Cenozoic, even although the clade is poorly represented in these regions today. The fossil record documents a significant palaeogeographical and evolutionary history of Detarieae in Africa, especially compared with that of Europe and Anatolia. Based on this evidence, it is unlikely that significant diversification of extant African Detarieae took place on the Eurasian landmass. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 44–54.     

Out of Africa: Biogeography and diversification of the pantropical pond skater genus Limnogonus Stål, 1868 (Hemiptera: Gerridae)

Gondwanan vicariance, long‐distance dispersal (LDD), and boreotropical migration have been proposed as alternative hypotheses explaining the pantropical distribution pattern of organisms. In this study, the historical biogeography of the pond skater genus Limnogonus was reconstructed to evaluate the impact of biogeographical scenarios in shaping their modern transoceanic disjunction. We sampled almost 65% of recognized Limnogonus species. Four DNA fragments including 69 sequences were used to reconstruct a phylogram. Divergence time was estimated using a Bayesian relaxed clock method and three fossil calibrations. Diversification dynamics and ancestral area reconstruction were investigated by using maximum likelihood and Bayesian approaches. Our results showed the crown group of Limnogonus originated and diversified in Africa in the early Eocene (49 Ma, HPD: 38–60 Ma), subsequently expanding into other regions via dispersal. The colonization of the New World originated from the Oriental Region probably via the Bering Land Bridge in the late Eocene. Two split events between the Old World and New World were identified: one between Neotropics and Oriental region around the middle Oligocene (30 Ma, HPD: 22–38 Ma), and the other between Neotropics and Africa during the middle Miocene (14 Ma, HPD: 8–21 Ma). The evolutionary history of Limnogonus involved two biogeographical processes. Gondwanan vicariance was not supported in our analyses. The diversification of Limnogonus among Africa, Oriental, and Neotropical regions corresponded with the age of land bridge connection and dispersed as a member associated with the broad boreotropical belt before local cooling (34 Ma). The current transoceanic disjunctions in Limnogonus could be better explained by the disruption of “mixed‐mesophytic” forest belt; however, the direct transoceanic LDD between the Neotropics and Africa could not be ruled out. In addition, the “LDD” model coupled with island hopping could be a reasonable explanation for the diversification of the Oriental and Australian regions during the Oligocene.     

Miocene to Pliocene vegetation reconstruction and climate estimates in the Iberian Peninsula from pollen data

Pollen analysis of Miocene and Pliocene sediments from the Iberian Peninsula shows a progressive reduction in plant diversity through time caused by the disappearance of thermophilous and high-water requirement plants. In addition, an increase in warm-temperate (mesothermic), seasonal-adapted “Mediterranean” taxa, high-elevation conifers and herbs (mainly Artemisia) occurred during the Middle and Late Miocene and Pliocene. This has mainly been interpreted as a response of the vegetation to global and regional processes, including climate cooling related to the development of the East Antarctic Ice Sheet and then the onset of the Arctic Ice Sheet, uplift of regional mountains related to the Alpine uplift and the progressive movement of Eurasia towards northern latitudes as a result of the northwards subduction of Africa. The development of steppe-like vegetation in southern Iberia is ancient and probably started during the Oligocene. The onset of a contrasted seasonality in temperature during the Mid-Pliocene superimposed on the pre-existing seasonality in precipitation, the annual length of which increased southward. The Mediterranean climatic rhythm (summer drought) began about 3.4 Ma and caused the individualization of modern Mediterranean ecosystems. Quaternary-type Mediterranean climatic fluctuations started at 2.6 Ma (Gelasian) resulting in repeated steppe vs. forest alternations. A latitudinal climatic gradient between the southern and the northern parts of the Iberian Peninsula existed since the Middle Miocene.     

‘Out‐of‐Africa’ dispersal of tropical floras during the Miocene climatic optimum: evidence from Uvaria (Annonaceae)

Aim African–Asian disjunctions are common in palaeotropical taxa, and are typically explained by reference to three competing hypotheses: (1) ‘rafting’ on the Indian tectonic plate, enabling Africa‐to‐Asia dispersal; (2) migration via Eocene boreotropical forests; and (3) transoceanic long‐distance dispersal. These hypotheses are tested using Uvaria (Annonaceae), which is distributed in tropical regions of Africa, Asia and Australasia. Recent phylogenetic reconstructions of the genus show a clear correlation with geographical provenance, indicating a probable origin in Africa and subsequent dispersal to Asia and then Australasia. Ancestral areas and migration routes are inferred and compared with estimates of divergence times in order to distinguish between the prevailing dispersal hypotheses. Location Palaeotropics. Methods Divergence times in Uvaria are estimated by analysing the sequences of four DNA regions (matK, psbA–trnH spacer, rbcL and trnL–F) from 59 Uvaria species and 77 outgroup species, using a Bayesian uncorrelated lognormal (UCLD) relaxed molecular clock. The ancestral area of Uvaria and subsequent dispersal routes are inferred using statistical dispersal–vicariance analysis (s‐diva ). Results Uvaria is estimated to have originated in continental Africa 31.6 Ma [95% highest posterior density (HPD): 38.4–25.1 Ma] between the Middle Eocene and Late Oligocene. Two main migration events during the Miocene are identified: dispersal into Madagascar around 17.0 Ma (95% HPD: 22.3–12.3 Ma); and dispersal into Asia between 21.4 Ma (95% HPD: 26.7–16.7 Ma) and 16.1 Ma (95% HPD: 20.1–12.1 Ma). Main conclusions Uvaria fruits are widely reported to be consumed by primates, and are therefore unlikely candidates for successful long‐distance transoceanic dispersal. The other biogeographical hypotheses, involving rafting on the Indian tectonic plate, and dispersal via the European boreotropical forests associated with the Eocene thermal maximum, can be discounted due to incongruence with the divergence time estimates. An alternative scenario is suggested, involving dispersal across Arabia and central Asia via the tropical forests that developed during the late Middle Miocene thermal maximum (17–15 Ma), associated with the ‘out‐of‐Africa’ dispersal of primates. The probable route and mechanism of overland dispersal between Africa and Asia for tropical plant groups during the Miocene climatic optimum are clarified based on the Uvaria data.     

European Neogene rodent communities: explaining family-level replacements through a spatiotemporal approach

We analyse the changes in rodent regional assemblages based on 657 fossil-bearing western European localities distributed ranging from ca. 27 Ma (Late Oligocene) to ca. 3 Ma (Late Pliocene). We compare temporal and spatial patterns in order to identify the factors that drive the evolutions of communities. Regional assemblages are analysed based on the distribution of species richness among families. First, communities are temporally analysed to identify significant changes in their composition. Second, regional communities are spatially compared to indentify diversity gradients. The temporal analysis reveals that communities' evolution is marked by several breaks in their composition, related to either migration or environmental/climatic events. This evolution can be summarised in terms of shifts in the relative abundances of glirids, cricetids and murids within assemblages. In contrast, spatial analysis shows that only environmental changes induce long-lasting changes in diversity gradients. Some observations made on cricetids and murids extant relatives indicate that they have undergone a large dietary diversification enabled by a specific digestive tract (along with the diversification of other life-history traits), whereas glirids are more specialised. The opposite diversity dynamics of these groups emphasises the importance of family-level adaptive potential in diversity conservation issues when facing environmental changes.     

After the deluge: mitochondrial DNA indicates Miocene radiation and Pliocene adaptation of tree and giant weta (Orthoptera: Anostostomatidae)

Aim New Zealand broke away from the margins of Gondwana c. 75 Ma. Since then, New Zealand taxa derived from the Gondwanan biota are thought to have been exposed first to a subtropical climate on a low lying terrain, then severe land reduction during the Oligocene marine transgression, followed by much cooler climates of the Pliocene and Pleistocene, at which time mountain ranges emerged. The biological consequence of New Zealand's geological and climatic history is not well understood, in particular the extent to which the Oligocene acted as a biological bottleneck remains unresolved. Methods We used mitochondrial cytochrome oxidase I and 12S DNA sequences to examine the extent of diversity and inferred timing of speciation of New Zealand weta (Anostostomatidae), a group of Orthoptera with a Gondwanan distribution generally thought to be ancient inhabitants of New Zealand. Main conclusions We hypothesize that at least three distinct groups of weta survived the Oligocene marine transgression and radiated subsequently. Speciation followed during the Miocene and radiation into new habitats occurred during the Pliocene when mountain building created novel environments. Patterns of genetic diversity within species reflect, in some instances, geographical subdivision in the Pliocene, and in other cases, Pleistocene range changes resulting from climate change.     

Phylogeny and biogeography Eupholidoptera Mařan (Orthoptera,Tettigoniidae): morphological speciation in correlation with the geographical evolution of the eastern Mediterranean

Recently, the systematics and biogeography of the Mediterranean biota have received much attention. This paper deals with Eupholidoptera Ma?an, a Mediterranean lineage of Tettigoniidae. The genus is restricted to the northern and eastern basin of the Mediterranean, with a significant number of species found on the Aegean islands. To produce a phylogeny and use it to make assumptions about the historical biogeography of Eupholidoptera, material of 46 species from several collections was studied. A phylogenetic analysis based mainly on morphological characters suggested two lineages in the genus: the E. chabrieri and the E. prasina groups. Based on the consistency between historical geographical events and branching events on the phylogenetic tree, Eupholidoptera is assumed to have evolved from an ancestor present in the Aegeid plate in the Mid‐Miocene. The division of the Aegeid plate into Anatolia and Greece in the Tortonian, the reoccurrence of terrestrial corridors between these mainlands in the Messinian, the regression of the Aegean area in the Pliocene and sea level changes in the Pleistocene are assumed to have been the main palaeogeographical events directing speciation in Eupholidoptera. As most of the species are allopatric, vicariance is suggested to be the main pattern. By combining the nature of the characters used in the phylogenetic analysis, the phylogenetic tree produced and the biogeographical assumptions, four tentative conclusions can be made: (i) radiation in the genus is a result of divergence in morphology; (ii) because the main character source is male genitalia, there has possibly been intensive sexual selection, which leads to morphological speciation; (iii) as the difference in temporal parameters of the song is prominent in sympatric/parapatric species pairs only, co‐occurrence is suggested to be the main reason driving divergence in the song; (iv) there seems to be a negative correlation between the size of the distribution range and the evolutionary rate in speciation; this may be the reason why the E. prasina group (restricted to a small part of the range of the genus) is more diverse than the E. chabrieri group, which is distributed over the entire range.     

Out of Australia and back again: the world‐wide historical biogeography of non‐pollinating fig wasps (Hymenoptera: Sycophaginae)

Aim Figs (Ficus, Moraceae) are exploited by rich communities of often host‐specific phytophagous wasps. Among them, gall‐inducing Sycophaginae (Hymenoptera, Chalcidoidea) may share a common history with Ficus and their mutualistic pollinators (Agaonidae). We investigate here, for the first time, the phylogeny and biogeographical history of Sycophaginae and compare the timing of radiation and dispersion of major clades with available data on Ficus and fig pollinators. Reconstructing the history of their host colonization and association over space and time is central to understanding how fig wasp communities were assembled. Location World‐wide. Methods Maximum likelihood and Bayesian analyses were conducted on 4267 bp of mitochondrial and nuclear DNA to produce a phylogeny of all genera of Sycophaginae. Two relaxed clock methods with or without rate autocorrelation were used for date estimation. Analyses of ancestral area were also conducted to investigate the geographical origin of the Sycophaginae. Results The phylogeny is well resolved and supported. Our data suggest a post‐Gondwanan origin for the Sycophaginae (50–40 Ma) and two independent out‐of‐Australia dispersal events to continental Asia. Given palaeoclimatic and palaeogeographic records, the following scenario appears the most likely. The ancestor of Idarnes+Apocryptophagus migrated to Greater India through the Ninetyeast Ridge (40–30 Ma). The ancestor of Anidarnes+Conidarnes dispersed later via Sundaland (25–20 Ma). Idarnes and Anidarnes subsequently reached the New World via the North Atlantic land bridges during the Late Oligocene Warming Event. Apocryptophagus reached Africa c. 20 Ma via the Arabic corridors and returned to Australasia following the expansion of Sundaland tropical forests (20–10 Ma). Main conclusions Sycophaginae probably invaded the fig microcosm in Australia c. 50–40 Ma after the origin of their host plant. Once associated with figs, they dispersed out of Australia and radiated together with their host fig and associated pollinator through the tropics. We recorded a good coincidence of timing between dispersal events of Sycophaginae and continental connections. Furthermore, fruit pigeons that disperse figs probably spread out of Australasia through the Indian Ocean via the Ninetyeast Ridge c. 38 Ma. Therefore, our study highlights the potential for combining molecular phylogenetics with multiple methods of dating of interacting groups to reconstruct the historical biogeography of plant–herbivore associations.     

Biogeographical origins and diversification of the exoneurine allodapine bees of Australia (Hymenoptera,Apidae)

Aim Early diversification of allodapine bees occurred in Africa c. 50 Ma. They are most abundant in sub‐Saharan Africa and Australia, and one of the oldest phylogenetic divergences in the tribe involves a split between an African + Malagasy clade and an Australian clade. The historical biogeographical scenario for this has been highly problematic, entailing an Eocene dispersal from Africa to Australia, followed by an unresolved, and apparently rapid, set of bifurcations leading to the Australian ‘exoneurine’ genera. Here we use an expanded taxon set of Australian species to explore the timing and historical biogeography of the exoneurine radiation. Location Australia, Africa, Madagascar. Methods One nuclear gene (F2 copy of elongation factor 1α) and two mitochondrial genes (cytochrome c oxidase subunit I and cytochrome b) were sequenced for 33 Australian exoneurine species from all five genera found on the continent, as well as for an additional 37 species from all non‐parasitic genera in the remainder of the tribe. We used Bayesian inference analyses to study phylogenetic topology and penalized likelihood analyses to infer key dates of divergence within the tribe. We also used lineage‐through‐time (LTT) analyses and Bayesian analyses to explore the tempo of radiations and biogeographical history of the exoneurines. Results Results from the phylogenetic analyses were congruent with previous studies, indicating a single colonization event c. 34 Ma, too late for Gondwanan vicariance models, and too early for a Laurasian dispersal route. In contrast to earlier studies, we show that this colonization event did not result in an ancient rapid radiation. However, LTT patterns indicated a rapid radiation of the temperate‐adapted genera Exoneura and Brevineura, but not of the xeric‐adapted genus Exoneurella, from 10 to 6 Ma. Main conclusions Our results indicate a trans‐oceanic dispersal event from Africa to Australia, most likely via Antarctica, with an accelerated diversification of temperate‐adapted lineages during the major Late Miocene event referred to as the ‘Hill Gap’. This is the first study to link radiations in Australian bee faunal elements to changing climate, and differs from many other plant and insect phylogenetic studies by showing increased radiation of temperate clades, rather than xeric clades, with increasing aridification of Australia.     

Historical biogeography of the hyperdiverse hidden snout weevils (Coleoptera,Curculionidae, Cryptorhynchinae)

The first dated phylogeny of the weevil subfamily Cryptorhynchinae is presented within a framework of Curculionoidea. The inferred pattern and timing of weevil family relationships are generally congruent with previous studies, but our data are the first to suggest a highly supported sister-group relationship between Attelabidae and Belidae. Our biogeographical inferences suggest that Cryptorhynchinae s.s. originated in the Late Cretaceous (c. 86 Ma) in South America. Within the ‘Acalles group’ and the ‘Cryptorhynchus group’, several independent dispersal events to the Western Palaearctic via the Nearctic occurred in the Late Cretaceous and Early Paleogene. A second southern route via Antarctica may have facilitated the colonization of Australia in the Late Cretaceous (c. 82 Ma), where a diverse Indo-Australian clade probably emerged c. 73 Ma. In the Early Eocene (c. 50–55 Ma), several clades independently dispersed from Australia to proto-New Guinea, i.e. the tribe Arachnopodini s.l., the ‘Rhynchodes group’ and the genus Trigonopterus. New Zealand was first colonized in the Late Palaeocene (c. 60 Ma). Divergence time estimations and biogeographical reconstructions indicate that the colonization of New Guinea is older than expected from current geological reconstructions of the region.