Phylogeographic analysis of the nocturnal velvet ant genus Dilophotopsis (Hymenoptera: Mutillidae) provides insights into diversification in the Nearctic deserts

Understanding the history of diversification in the North American deserts has long been a goal of biogeographers and evolutionary biologists. Although it appears that a consensus is forming regarding the patterns of diversification in the Nearctic deserts in vertebrate taxa, little work has been done exploring the historical biogeography of widespread invertebrate taxa. Before a robust model of geobiotic change in the North American deserts can be proposed, it needs to be determined whether the same historical events affected vertebrate and invertebrate taxa in the same way. We explore the phylogeographic patterns in a widespread nocturnal wasp genus Dilophotopsis using two rDNA loci, the internal transcribed spacer regions 1 and 2 (ITS1 and ITS2). We use Bayesian phylogenetic analysis and haplotype network analysis to determine whether a consistent geographic pattern exists among species and populations within Dilophotopsis. We also used molecular dating techniques to estimate divergence dates of the major phylogenetic clades. Our analyses indicates that the species‐level divergences in Dilophotopsis occurred in the Neogene, and likely were driven by mountain building during the Miocene–Pliocene boundary (approximately 5 Mya) similar to the divergences in many vertebrate taxa. The population‐level divergences within species occurred during the Pleistocene (0.1–1.8 Mya). The present study shows that similar patterns of diversification exist in vertebrate and invertebrate taxa. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 360–375.     

New observations on the Late Miocene–Early Pliocene Lutrinae (Mustelidae: Carnivora,Mammalia) from the Middle Awash,Afar Rift,Ethiopia

New observations on the Late Miocene and Earliest Pliocene mustelids from the Middle Awash of Ethiopia are presented. The Middle Awash study area samples the last six million years of African vertebrate evolutionary history. Its Latest Miocene (Asa Koma Member of the Adu-Asa Formation, 5.54–5.77 Ma) and Earliest Pliocene (Kuseralee and Gawto Members of the Sagantole Formation, 5.2 and 4.85 Ma, respectively) deposits sample a number of large and small carnivore taxa among which mustelids are numerically abundant. Among the known Late Miocene and Early Pliocene mustelid genera, the Middle Awash Late Miocene documents the earliest Mellivora in eastern Africa and its likely first appearance in Africa, a new species of Plesiogulo, and a species of Vishnuonyx. The latter possibly represents the last appearance of this genus in Africa. Torolutra ougandensis is known from both the Late Miocene and Early Pliocene deposits of the Middle Awash. The genus Sivaonyx is represented by at least two species: S. ekecaman and S. aff. S. soriae. Most of the lutrine genera documented in the Middle Awash Late Miocene/Early Pliocene are also documented in contemporaneous sites of eastern Africa. The new observations presented here show that mustelids were more diverse in the Middle Awash Late Miocene and Early Pliocene than previously documented.     

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.     

Out of Africa: biogeographic history of the open‐habitat chats (Aves,Muscicapidae: Saxicolinae) across arid areas of the old world

Arid and semi‐arid areas constitute a prominent feature of the earth today, especially in Asia and Africa. Their formation started in the middle Miocene with increased stepwise aridification since the Pliocene. This aridification had strong ecological and evolutionary consequences and not only led to fragmentation of moist‐adapted biota, but also fostered the evolution of arid‐adapted taxa from mesic ancestors and triggered speciation within arid areas. The open‐habitat chats, a clade within Saxicolinae (Aves, Muscicapidae), constitute one of the most significant arid‐adapted passerine groups of Africa and Eurasia. Here, we present a temporal and spatial framework for the diversification of open‐habitat chats, using probabilistic approaches for the reconstruction of their biogeographic history based on a time‐calibrated multilocus molecular phylogenetic hypothesis. The diversification of open‐habitat chats was initiated in the late Miocene at around 7.4 Ma, most likely in sub‐Saharan Africa. Southern Africa and the Horn of Africa acted as centres of diversification and biogeographic expansion. From the latter area, the Arabo‐Sindic region and subsequently further parts of Eurasia and North Africa were colonized. The colonization history out of sub‐Saharan Africa contrasts with that of several other songbird clades, where a biogeographic expansion from Eurasia or northern Africa to southern Africa was prevalent. Habitat fragmentation through forest expansions during intermittent wetter periods in Africa influenced diversification in several clades. However, phases of increased aridity, with hyperarid regions acting as drivers of vicariance, seem to have also been important in radiations of the Arabo‐Sindic region and the Horn of Africa during the Pleistocene. Different processes such as colonization of new areas followed by vicariance or speciation across ecotones might have played a role throughout the radiation of open‐habitat chats.     

The Early Pliocene re-colonisation of the deep Mediterranean Sea by benthic foraminifera and their pulsed Late Pliocene–Middle Pleistocene decline

Ninety-five species and 19 genera of cosmopolitan, deep-sea benthic foraminifera belonging to the families Pleurostomellidae, Stilostomellidae and Nodosariidae, became extinct during the Late Pliocene–Middle Pleistocene. Only 50% of these (44 species) were present in the Pliocene or Pleistocene of the deep Mediterranean Sea (ODP Sites 654, 966, 967, 975, 976), being those which had successfully migrated in via the Strait of Gibraltar from the deep Atlantic following the annihilation of the Mediterranean deep-sea fauna during the Late Miocene Messinian Crisis. Most colonisation occurred within the first 0.8 myrs (5.3–4.5 Ma) after re-establishment of the Mediterranean–Atlantic link, with possibly a second lesser period of immigration in the Late Pliocene (3.4–3.0 Ma). We infer that colonisations may have been fortuitous and few in number, as some common members of the group in the Atlantic never succeeded in establishing in the Mediterranean Sea. There is no evidence of any new immigration events during the Pleistocene, implying that the present anti-estuarine circulation may have been in place throughout this period. Our studies suggest that these deep-water, low-oxygen-tolerant foraminifera survived the many periods of deep-water sapropel formation in the Pliocene–Early Pleistocene, possibly in somewhat shallower (~ 500 m) refuges with dysoxic, rather than anoxic conditions.The Pliocene–Pleistocene stratigraphic record of this group of elongate, cylindrical benthic foraminifera with constricted and specialised apertures is similar in the west and east Mediterranean basins. The group declined in abundance (flux) and diversity in two pulses, during the Late Pliocene (3.1–2.7 Ma) and the late Early Pleistocene (1.3–1.0 Ma) in concert with global, southern-sourced, deep-water sites (AABW, CPDW) and earlier than the single decline (1.0–0.6 Ma) in global, intermediate water sites (uNADW, AAIW). All species, with one possible exception, disappeared earlier in the Mediterranean than globally. The highest occurrence of any species of this group in Mediterranean sites was 0.8–0.43 Ma, comparable with 0.7–0.2 Ma outside with the youngest survivors being in abyssal, deep-water.Thus, despite the unusual oceanographic conditions and isolation, the deep Mediterranean Sea was in this case neither the centre for the evolution of new species nor a refuge where species survived after they had disappeared elsewhere.     

The Miocene floras of Iceland and their significance for late Cainozoic North Atlantic biogeography

A large number of plant macrofossils from several Middle to Upper Miocene localities from Iceland have been studied. The fossil material includes four ferns and fern allies, seven conifers, and about 40 species of flowering plants. Betula islandica and Salix gruberi are described as new species. Coniferous twigs previously ascribed to the genus Sequoia are shown to belong to Cryptomeria based on macro‐morphological and epidermal features. Fossil plants from Iceland are compared with coeval fossil taxa from Europe and North America and with living plants. The main finding is that the Miocene flora of Iceland belongs to a widespread Neogene northern hemispheric floral type including plants whose representatives are restricted to East Asia, North America and to western Eurasia at the present time. Previously inferred conspicuous similarities to North American modern equivalents appear to be misleading. The type of vegetation in four plant‐bearing sedimentary formations from the late Mid Miocene to Late Miocene, the 12 Ma Brjánslækur‐Seljá Formation, the 10 Ma Tröllatunga‐Gautshamar Formation, the 9–8 Ma Skarðsströnd‐Mókollsdalur Formation, and the 7–6 Ma Hreðavatn‐Stafholt Formation, corresponds to a humid temperate broadleaved (deciduous)–coniferous mixed forest dominated by Betulaceae, Fagaceae and Acer. Changes in species composition in the sedimentary formations reflect a shift from warm temperate (Cfa climate) to cool temperate (Cfb climate) conditions from the late Mid Miocene to the latest Miocene. This shift was connected to repeated phases of extinction and colonization. Specifically, one set of thermophilic taxa including Magnolia, Liriodendron, Sassafras and Comptonia went extinct between 12 and 10 Ma, and appears to have been replaced by another set of thermophilic taxa in the 10 Ma formation (Juglandaceae aff. Pterocarya/Cyclocarya, Rhododendron ponticum type). The 9–8 and 7–6 Ma formations are characterized by taxa that migrated to Iceland from Europe, such as Fagus gussonii, Betula cristata and Pterocarya fraxinifolia type. Although there is convincing evidence that plants colonized Iceland both from North America and Europe until 12 Ma, migration in the younger formations (9–8, 7–6 Ma) is suggested to have occurred mainly from Europe. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149 , 369–417.     

Reconstructing biogeographic temporal events in the evolution of the livebearer fish genus Jenynsia based on total evidence analysis (Cyprinodontiformes: Anablepidae)

The current distribution of the Neotropical ichthyofauna has been widely affected by the main geological events that occurred in South and Central America. However, robust biogeographic information is still scarce or absent for most fish families. The biogeographic relationships of the most diverse anablepid genus, Jenynsia, are herein analysed, using temporal and spatial approaches based on a total evidence dataset, including 167 morphological characters and seven nuclear genes, totalling 6075?bp. A time calibrated analysis recovered the origin of Jenynsia at the Miocene and the diversification of its two subgenera between the Late Miocene and Early Pliocene. This result combined with the analysis of reconstruction of ancestral states indicates that the ancestor of Jenynsia colonized the Paranean Sea, and the current distribution of the species of the genus is probably a result of geological events, including: (1) ancient connections between Iguaçu and upper Uruguay River basins; (2) sea level variation along the Pliocene in the South American Atlantic coast; and (3) the decrease of the Paranean Sea along the Pliocene and Pleistocene.     

临夏盆地的新生代地层及其哺乳动物化石证据

临夏盆地的新生代地层相当发育 ,保存了从渐新世至全新世的连续沉积序列。更为重要的是 ,这些沉积物中含有丰富的哺乳动物化石 ,为划分和对比临夏盆地的新生代地层提供了可靠的证据。然而 ,此前关于这个盆地地层层序和时代的认识有许多矛盾之处 ,地层命名繁复 ,化石证据混乱。近年来我们对临夏盆地的野外考察已理清了沉积序列 ,并在充分的哺乳动物化石证据的基础上重新厘定了各个岩石地层单位所对应的地质时代。临夏盆地的新生代哺乳动物化石以晚渐新世的巨犀动物群、中中新世的铲齿象动物群、晚中新世的三趾马动物群和早更新世的真马动物群最为丰富。     

Phylogeographic diversification of antelope squirrels (Ammospermophilus) across North American deserts

We investigated the biogeographic history of antelope squirrels, genus Ammospermophilus, which are widely distributed across the deserts and other arid lands of western North America. We combined range‐wide sampling of all currently recognized species of Ammospermophilus with a multilocus data set to infer phylogenetic relationships. We then estimated divergence times within identified clades of Ammospermophilus using fossil‐calibrated and rate‐calibrated molecular clocks. Lastly, we explored generalized distributional changes of Ammospermophilus since the last glacial maximum using species distribution models, and assessed responses to Quaternary climate change by generating demographic parameter estimates for the three wide‐ranging clades of A. leucurus. From our phylogenetic estimates we inferred strong phylogeographic structure within Ammospermophilus and the presence of three well‐supported major clades. Initial patterns of historical divergence were coincident with dynamic alterations in the landscape of western North America, and the formation of regional deserts during the Late Miocene and Pliocene. Species distribution models and demographic parameter estimates revealed patterns of recent population expansion in response to glacial retreat. When combined with evidence from co‐distributed taxa, the historical biogeography of Ammospermophilus provides additional insight into the mechanisms that impacted diversification of arid‐adapted taxa across the arid lands of western North America. We propose species recognition of populations of the southern Baja California peninsula to best represent our current understanding of evolutionary relationships among genetic units of Ammospermophilus. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 949–967.     

Small mammal (rodents and lagomorphs) European biogeography from the Late Oligocene to the mid Pliocene

Aim To analyse the fossil species assemblages of rodents and lagomorphs from the European Neogene in order to assess what factors control small mammal biogeography at a deep‐time evolutionary time‐scale. Location Western Europe: 626 fossil‐bearing localities located within 31 regions and distributed among 18 successive biochronological units ranging from c. 27 Ma (million years ago; Late Oligocene) to c. 3 Ma (mid Pliocene). Methods Taxonomically homogenized pooled regional assemblages are compared using the Raup and Crick index of faunal similarity; then, the inferred similarity matrices are visualized as neighbour‐joining trees and by projecting the statistically significant interregional similarities and dissimilarities onto palaeogeographical maps. The inferred biogeographical patterns are analysed and discussed in the light of known palaeogeographical and palaeoclimatic events. Results Successive time intervals with distinct biogeographical contexts are identified. Prior to c. 18 Ma (Late Oligocene and Early Miocene), a relative faunal homogeneity (high interregional connectivity) is observed all over Europe, a time when major geographical barriers and a weak climatic gradient are known. Then, from the beginning of the Middle Miocene onwards, the biogeography is marked by a significant decrease in interregional faunal affinities which matches a drastic global climatic degradation and leads, in the Late Miocene (c. 11 Ma), to a marked latitudinal pattern of small mammal distribution. In spite of a short rehomogenization around the Miocene/Pliocene boundary (6–4 Ma), the biogeography of small mammals in the mid Pliocene (c. 3 Ma) finally closely reflects the extant situation. Main conclusions The resulting biogeographical evolutionary scheme indicates that the extant endemic situation has deep historical roots corresponding to global tectonic and climatic events acting as primary drivers of long‐term changes. The correlation of biogeographical events with climatic changes emphasizes the prevalent role of the climate over geography in generating heterogeneous biogeographical patterns at the continental scale.     

Tropical forests and the genus Homo

Tropical forests constitute some of the most diverse and complex terrestrial ecosystems on the planet. From the Miocene onward, they have acted as a backdrop to the ongoing evolution of our closest living relatives, the great apes, and provided the cradle for the emergence of early hominins, who retained arboreal physiological adaptations at least into the Late Pliocene. There also now exists growing evidence, from the Late Pleistocene onward, for tool‐assisted intensification of tropical forest occupation and resource extraction by our own species, Homo sapiens. However, between the Late Pliocene and Late Pleistocene there is an apparent gap in clear and convincing evidence for the use of tropical forests by hominins, including early members of our own genus. In discussions of Late Pliocene and Early Pleistocene hominin evolution, including the emergence and later expansion of Homo species across the globe, tropical forest adaptations tend to be eclipsed by open, savanna environments. Thus far, it is not clear whether this Early‐Middle Pleistocene lacuna in Homo‐rainforest interaction is real and representative of an adaptive shift with the emergence of our species or if it is simply reflective of preservation bias.     

Climate change and evolution of the New World pitviper genus Agkistrodon (Viperidae)

Aim We derived phylogenies, phylogeographies, and population demographies for two North American pitvipers, Agkistrodon contortrix (Linnaeus, 1766) and A. piscivorus (Lacépède, 1789) (Viperidae: Crotalinae), as a mechanism to evaluate the impact of rapid climatic change on these taxa. Location Midwestern and eastern North America. Methods We reconstructed maximum parsimony (MP) and maximum likelihood (ML) relationships based on 846 base pairs of mitochondrial DNA (mtDNA) ATPase 8 and ATPase 6 genes sequenced over 178 individuals. We quantified range expansions, demographic histories, divergence dates and potential size differences among clades since their last period of rapid expansion. We used the Shimodaira–Hasegawa (SH) test to compare our ML tree against three biogeographical hypotheses. Results A significant SH test supported diversification of A. contortrix from northeastern Mexico into midwestern–eastern North America, where its trajectory was sundered by two vicariant events. The first (c. 5.1 Ma) segregated clades at 3.1% sequence divergence (SD) along a continental east–west moisture gradient. The second (c. 1.4 Ma) segregated clades at 2.4% SD along the Mississippi River, coincident with the formation of the modern Ohio River as a major meltwater tributary. A single glacial refugium was detected within the Apalachicola region of southeastern North America. Significant support was also found for a hypothesis of trans‐Gulf rafting by the common ancestor of A. piscivorus from eastern Mexico (possibly the Yucatan Peninsula) to northern Florida. There, a Mid–Late Pliocene marine transgression separated it at 4.8% SD from mainland North America. Significant range expansions followed compressive glacial effects in three (of four) A. contortrix clades and in two (of three) A. piscivorus clades, with the Florida A. piscivorus clade exhibiting significant distributional stasis. Main conclusions Pliocene glaciations, rapidly developing western aridity, and Pleistocene glacial meltwaters seemingly led to the diversification of A. contortrix and A. piscivorus in North America. Both species were pushed southwards by Pleistocene climate change, with subsequent northward expansions uninhibited topographically. The subspecific taxonomy used for A. contortrix and A. piscivorus today, however, appear non‐representative. The monophyletic Florida subspecies of A. piscivorus may be a distinct species (at 4.8% SD), whereas two western subspecies of A. contortrix also appear to constitute a single distinct species, pending additional analyses. We conclude that both species of Agkistrodon can be used as suitable ectothermic models to gauge impacts of future climate change.     

Phylogeny and adaptive diversity of rodents of the family Ctenomyidae (Caviomorpha): delimiting lineages and genera in the fossil record

Differentiation of genera of the modern (Late Miocene to Recent) South American rodent family Ctenomyidae would have been linked to the acquisition of disparate adaptations to digging and life underground. In accordance with this hypothesis, the delimitation of lineages and genera in the ctenomyid fossil record is evaluated here following an adaptation-rooted criterion that involves both an assessment of the monophyly and of the adaptive profiles of recognized clades. The application of such a criterion, including morphofunctional information, delimited four cohesive lineages among crown ctenomyids (i.e. euhypsodont species of the Late Miocene to Recent): Eucelophorus (Early Pliocene–Middle Pleistocene), Xenodontomys-Actenomys (Late Miocene–Pliocene), Praectenomys (Pliocene) and Ctenomys (including Paractenomys ; Pliocene–Recent); in addition, the results supported the status of Xenodontomys as a paraphyletic ancestor of Actenomys . The cladogenesis that gave rise to the crown group would have occurred immediately after the acquisition of euhypsodonty in a Xenodontomys simpsoni -like ancestor during the Late Miocene. This putative ancestor would have had fossorial habits and moderate digging specializations, an adaptive profile maintained in Xenodontomys-Actenomys . Eucelophorus and Ctenomys would have independently evolved subterranean habits at least since the Pliocene. Although the earliest history of the only living representative, Ctenomys , is known only fragmentarily, remains from Esquina Blanca (Uquía Formation), in north-western Argentina, suggest a minimum age of around 3.5 Ma (Early–Late Pliocene) for the differentiation of the genus. This date agrees with recent molecular estimates.     

中国新生代晚期的剑齿象(剑齿象科,长鼻目)及其扩散事件

对我国新生代晚期的剑齿象(Stegodon)进行再研究,提出了下列几点看法:1)依据Ste-godon的特征,可将已命名的中国剑齿象种分为3类。一类为有效的剑齿象种,包括晚中新世的S.licenti Teilhard & Trasseart,1937;上新世的S.zdanskyi Hopwood,1935和S.officinalis Hopwood,1935;上新世晚期至更新世早期的S.zhaotongensis Chow & Zhai,1962;更新世的S.chiai Chow & Zhai,1962,S.orientalis Owen,1870,S.sinensis Owen,1870和S.huananensis sp.n.等种。另一类是从剑齿象属中分出,归入到其他属或科的种。属于这一类的有可能置入真象类(Elephantoidae)的Stegodon parahypsilophus He,1984,S.guizhouensis Li & Wen,1986,Stegodon cf.S.hypsilophus和S.primitium Liu et al.,1973以及归入Stegolophodon的Ste-godon baoshanensis Yun,1975。第三类为属种分类位置未定的种,以S.wushanensis Huang et al.,1991为代表。2)南亚更新世的S.elephantoides(Clift),1828可能在我国不存在。3)我国的剑齿象化石相当丰富,发现于东经99°以东、北纬38°以南的广大地区(包括山西、陕西和甘肃以及华中、华南、西南和部分华东等地区),生存时代为晚中新世晚期至更新世。4)在新生代晚期我国的剑齿象可能经历了3次大的由北向南的扩散和迁移事件。第一次发生在晚中新世,第二次在晚中新世晚期至早上新世早期,第三次在更新世早期。5)剑齿象可能起源于亚洲。S.licenti是我国出现时代最早的剑齿象,也可能是剑齿象最原始的一种。     

Climate and host‐plant associations shaped the evolution of ceutorhynch weevils throughout the Cenozoic

Using molecular phylogenetic data and methods we inferred divergence times and diversification patterns for the weevil subfamily Ceutorhynchinae in the context of host‐plant associations and global climate over evolutionary time. We detected four major diversification shifts that correlate with both host shifts and major climate events. Ceutorhynchinae experienced an increase in diversification rate at ～53 Ma, during the Early Eocene Climate Optimum, coincident with a host shift to Lamiaceae. A second major diversification phase occurred at the end of the Eocene (～34 Ma). This contrasts with the overall deterioration in climate equability at the Eocene‐Oligocene boundary, but tracks the diversification of important host plant clades in temperate (higher) latitudes, leading to increased diversification rates in the weevil clades infesting temperate hosts. A third major phase of diversification is correlated with the rising temperatures of the Late Oligocene Warming Event (～26.5 Ma); diversification rates then declined shortly after the Middle Miocene Climate Transition (～14.9 Ma). Our results indicate that biotic and abiotic factors together explain the evolution of Ceutorhynchinae better than each of these drivers viewed in isolation.     

Late neogene biostratigraphy and stable isotope stratigraphy of a drilled core from the Gulf of Mexico

Late Neogene stable isotope stratigraphy and planktonic foraminiferal biostratigraphy have been examined in a high sedimentation rate core (E67-135, Shell Oil Co.) drilled at 725 m water depth in the De Soto Canyon, Gulf of Mexico. The 305 m core contains sections that are Late Miocene, Early Pliocene, Late Pliocene, and Quaternary in age, and is rich in well-preserved assemblages of planktonic foraminifera.A biostratigraphy has been established based on the ranges of 34 selected species of foraminifera. The core 3orrelates with sections from the Gulf of Mexico, the Caribbean Sea, and the subtropical North and South Atlantic Oceans using, as datums, the evolutionary appearances of Globorotalia miocenica Palmer and Globorotalia margaritae evoluta Cita, the extinction of Globorotalia miocenica and the first appearance of Globorotalia truncatulinoides (d'Orbigny).Oxygen and carbon isotope stratigraphy is based on analysis of the benthonic foraminifer, Uvigerina d'Orbigny. Isotopic trends are similar to those observed in the Pacific and Atlantic Oceans. From Early Pliocene to Late Pleistocene time, average δ¹⁸O values increase (2.42‰ to 3.36‰) and exhibit a wider range of values (0.71‰ in Early Pliocene compared to 1.65‰ in Late Pleistocene sediments), probably reflecting Late Neogene climatic deterioration. The ratio ${}^{13}\text{C}{}^{12}\text{C}$ decreases significantly by ?0.21‰ from the Late Miocene to the Early Pliocene. A decrease in δ¹³C is observed in other cores and is probably related to changing oceanic circulation patterns in Late Miocene time.     

Cryptic species and co‐diversification in sand scorpions from the Karakum and Kyzylkum deserts of Central Asia

Little is known about species diversification within the deserts of Central Asia. For example, the degree of lineage divergence and timing of population differentiation, as well as potential biogeographic barriers driving diversification, are nearly unknown. Here, we analysed a multi‐locus data set for a widespread sand scorpion (Mesobuthus gorelovi) to evaluate cryptic species diversity and phylogeographic patterns across the Karakum and Kyzylkum deserts. We also combined these data with previously published sequence data to test for a signal of co‐diversification. A consensus species delimitation approach indicated that the widespread M. gorelovi is likely composed of up to five distinct species that began to diversify at the Miocene–Pliocene boundary. We observed shared patterns of lineage divergence across the Amu Darya River region in three scorpion taxa and found support for a shared history of assemblage diversification across this biogeographic barrier. Thus, major river systems appear to facilitate diversification among desert scorpions.     

Historical biogeography of the genus Rhadinaea (Squamata: Dipsadinae)

Multiple geological and climatic events have created geographical or ecological barriers associated with speciation events, playing a role in biological diversification in North and Central America. Here, we evaluate the influence of the Neogene and Quaternary geological events, as well as the climatic changes in the diversification of the colubrid snake genus Rhadinaea using molecular dating and ancestral area reconstruction. A multilocus sequence dataset was generated for 37 individuals of Rhadinaea from most of the biogeographical provinces where the genus is distributed, representing 19 of the 21 currently recognized species, and two undescribed species. Our analyses show that the majority of the Rhadinaea species nest in two main clades, herein identified as “Eastern” and “Southern”. These clades probably diverged from each other in the early Miocene, and their divergence was followed by 11 divergences during the middle to late Miocene, three divergences during the Pliocene, and six divergences in the Pleistocene. The ancestral distribution of Rhadinaea was reconstructed across the Sierra Madre del Sur. Our phylogenetic analyses do not support the monophyly of Rhadinaea. The Miocene and Pliocene geomorphology, perhaps in conjunction with climate change, appears to have triggered the diversification of the genus, while the climatic changes during the Miocene probably induced the diversification of Rhadinaea in the Sierra Madre del Sur. Our analysis suggests that the uplifting of the Trans‐Mexican Volcanic Belt and Chiapan–Guatemalan highlands in this same period resulted in northward and southward colonization events. This was followed by more recent, independent colonization events in the Pliocene and Pleistocene involving the Balsas Basin, Chihuahuan Desert, Pacific Coast, Sierra Madre Occidental, Sierra Madre Oriental, Sierra Madre del Sur, Trans‐Mexican Volcanic Belt, and Veracruz provinces, probably driven by the climatic fluctuations of the time.     

The Taxonomic and Evolutionary History of Fossil and Modern Balaenopteroid Mysticetes

Balaenopteroids (Balaenopteridae + Eschrichtiidae) are a diverse lineage of living mysticetes, with seven to ten species divided between three genera (Megaptera, Balaenoptera and Eschrichtius). Extant members of the Balaenopteridae (Balaenoptera and Megaptera) are characterized by their engulfment feeding behavior, which is associated with a number of unique cranial, mandibular, and soft anatomical characters. The Eschrichtiidae employ suction feeding, which is associated with arched rostra and short, coarse baleen. The recognition of these and other characters in fossil balaenopteroids, when viewed in a phylogenetic framework, provides a means for assessing the evolutionary history of this clade, including its origin and diversification. The earliest fossil balaenopterids include incomplete crania from the early late Miocene (7–10 Ma) of the North Pacific Ocean Basin. Our preliminary phylogenetic results indicate that the basal taxon, “Megaptera” miocaena should be reassigned to a new genus based on its possession of primitive and derived characters. The late late Miocene (5–7 Ma) balaenopterid record, except for Parabalaenoptera baulinensis and Balaenoptera siberi, is largely undescribed and consists of fossil specimens from the North and South Pacific and North Atlantic Ocean basins. The Pliocene record (2–5 Ma) is very diverse and consists of numerous named, but problematic, taxa from Italy and Belgium, as well as unnamed taxa from the North and South Pacific and eastern North Atlantic Ocean basins. For the most part Pliocene balaenopteroids represent extinct species and genera and reveal a greater degree of morphological diversity than at present. The Pleistocene record is very limited and, unfortunately, fails to document the evolutionary details leading to modern balaenopteroid species diversity. It is evident, however, that most extant species evolved during the Pleistocene. Morphological and molecular based phylogenies support two competing hypotheses concerning relationships within the Balaenopteroidea: (1) balaenopterids and eschrichtiids as sister taxa, and (2) eschrichtiids nested within a paraphyletic Balaenopteridae. The addition of fossil taxa (including a new Pliocene species preserving a mosaic of balaenopterid and eschrichtiid characters) in morphological and “total evidence” analyses, offers the potential to resolve the current controversy concerning the possible paraphyly of Balaenopteridae.     

THE EARLY DIVERSIFICATION HISTORY OF DIDELPHID MARSUPIALS: A WINDOW INTO SOUTH AMERICA'S “SPLENDID ISOLATION”

The geological record of South American mammals is spatially biased because productive fossil sites are concentrated at high latitudes. As a result, the history of mammalian diversification in Amazonia and other tropical biomes is largely unknown. Here we report diversification analyses based on a time‐calibrated molecular phylogeny of opossums (Didelphidae), a species‐rich clade of mostly tropical marsupials descended from a Late Oligocene common ancestor. Optimizations of habitat and geography on this phylogeny suggest that (1) basal didelphid lineages inhabited South American moist forests; (2) didelphids did not diversify in dry‐forest habitats until the Late Miocene; and (3) most didelphid lineages did not enter North America until the Pliocene. We also summarize evidence for an Early‐ to Middle‐Miocene mass extinction event, for which alternative causal explanations are discussed. To the best of our knowledge, this study provides the first published molecular‐phylogenetic evidence for mass extinction in any animal clade, and it is the first time that evidence for such an event (in any plant or animal taxon) has been tested for statistical significance. Potentially falsifying observations that could help discriminate between the proposed alternative explanations for didelphid mass extinction may be obtainable from diversification analyses of other sympatric mammalian groups.