Three new species of Callulina (Amphibia: Anura: Brevicipitidae) highlight local endemism and conservation plight of Africa's Eastern Arc forests

Material ascribed to the genus Callulina from north‐east Tanzania and south‐east Kenya is assessed. Three new species of Callulina are described from the North ( Callulina laphami sp. nov. ) and South ( Callulina shengena sp. nov. and Callulina stanleyi sp. nov. ) Pare Mountains in Tanzania. The species are diagnosed based on morphological, acoustic, and molecular data. A new key to the species of Callulina is provided. Based on an interpretation of the International Union for Conservation of Nature (IUCN) red list, we suggest that the three species will qualify as critically endangered, because of their small distributions and the ongoing threat to their habitat. We reveal the high local endemism of Callulina in the northern part of the Eastern Arc Mountains, with each species restricted to no more than one mountain (fragment) block. This high local endemism in Callulina is probably widespread across the Eastern Arc, raising further conservation concern for this group of amphibians. Based on new molecular phylogenetic data for Callulina, we discuss biogeographical relationships among north‐east Tanzanian mountains, and evolutionary patterns in Eastern Arc breviciptids. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 496–514.     

Analysis of the mitochondrial genome of Schistocerca gregaria gregaria (Orthoptera: Acrididae)

In the present study, we present the full sequence of the mitochondrial genome of the African desert locust Schistocerca gregaria gregaria. The size of 15625 bp reported matches very well with mitochondrial genomes of other Orthopteriodea. The mitochondrial genome comprises 13 protein‐coding genes, two ribosomal RNAs and 22 t‐RNAs with two t‐RNA (trnD and trnK) rearrangements that are typical for the taxon Caelifera. We compared the sequence with 12 mitochondrial genes of Schistocerca gregaria flaviventris and Schistocerca americana and used some of these data to construct phylogenetic trees, which confirm the close relationship between the two subspecies S. g. flaviventris and S. g. gregaria. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 296–305.     

Phylogenetic relationships of the pygmy rice rats of the genus Oligoryzomys Bangs, 1900 (Rodentia: Sigmodontinae)

Sequences from two mitochondrial genes (cytochrome b and NADH1) were used to produce a molecular phylogeny for 12 named and two undescribed species of the genus Oligoryzomys. All analyses placed Oligoryzomys microtis as the most basal taxon, a finding consistent with previous studies that suggested the west‐central Amazon as a centre of origin for the tribe Oryzomyini to which Oligoryzomys belongs. Biogeographically, this suggests that Oligoryzomys had a South American origin, and later advanced northwards, entering Central America and Mexico more recently. Different analyses have provided consistent support for several additional clades that did not necessarily agree with the species groups hypothesized by previous studies. A molecular clock derived for these data suggests an origin for the genus of 6.67 Mya, with most speciation within the genus occurring between 3.7 and 1.5 Mya. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 551–566.     

Androdioecy in Kryptolebias killifish and the evolution of self‐fertilizing hermaphroditism

Kryptolebias marmoratus is an important experimental fish, and is considered to represent the only vertebrate species comprising self‐fertilizing hermaphroditic individuals. Subsequent to the discovery of this unusual mode of reproduction, approximately 50 years ago, K. marmoratus has been the focus of a series of studies. However, little is known about the evolution of this rare reproduction mode, and data on the biology of closely related species are still unavailable. The present study aimed to histologically analyse the gonads of three K. marmoratus congeners (i.e. Kryptolebias ocellatus, Kryptolebias caudomarginatus, and Kryptolebias brasiliensis) to check the distribution of features related to hermaphroditism that are useful for forming hypotheses about the origin and evolution of the self‐fertilization mode of reproduction through the available phylogenies. The data obtained demonstrate that populations of K. caudomarginatus consist of males and hermaphrodites, which supports the hypothesis that hermaphroditism arose at the base of the clade containing K. caudomarginatus, K. marmoratus, and K. ocellatus as a first step towards a more advanced condition, uniquely shared by both K. marmoratus and K. ocellatus, in which males are rare or absent in natural populations, with the subsequent occurrence of self‐fertilization. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 344–349.     

Evolutionary innovations of squamate reproductive and developmental biology in the family Chamaeleonidae

The availability of molecular phylogenies has greatly accelerated our understanding of evolutionary innovations in the context of their origin and rate of evolution. Here, we assess the evolution of reproductive mode, developmental rate and body size in a group of squamate reptiles: the chameleons. Oviparity is ancestral and viviparity has evolved at least twice: Bradypodion and members of the Trioceros bitaeniatus clade are viviparous. Viviparous species are medium‐sized as a result of convergence from either small‐sized ancestors or large‐sized ancestors, respectively, but do not differ from oviparous species in clutch size, hatchling size or the trade‐off between clutch and hatchling size. Basal chameleons (Brookesia, Rhampholeon and Rieppeleon) are small‐sized and have developmental rates comparable with those of other lizards. Derived chameleons (Calumma, Chamaeleo, Trioceros and Furcifer) are mostly large‐sized and all have relatively slow developmental rates. Several clades of derived chameleons also exhibit developmental arrest (embryonic diapause or embryonic diapause plus cold torpor) and incubation periods extend to 6–10 months or more. Developmental arrest is associated with dry, highly seasonal climates in which the period favourable for oviposition and hatching is short. Long incubation periods thus ensure that hatching occurs during the favourable season following egg laying. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 656–668.     

Morphological and molecular taxonomy of a new Daptonema (Nematoda,Xyalidae) with comments on the systematics of some related taxa

A new species of Daptonema is described based upon morphological characters and 18S rRNA sequence. Daptonema matrona sp. nov. was collected in Pina Basin (north‐eastern Brazil). It differs from all other species of the genus by the presence of reduced cephalic setae and straight spicules. These features require an adaptation of the generic diagnosis. Moreover, the females are characterized by intra‐uterine development of the offspring, considered herein as their major autapomorphic feature. Molecular systematic analyses supported Daptonema matrona sp. nov. as a distinct genetic and evolutionary lineage. The data also indicate hypotheses of taxonomic synonymies amongst some related taxa from Xyalidae as well as the paraphyly of Daptonema. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 1–15.     

Push and bite: trade‐offs between burrowing and biting in a burrowing skink (Acontias percivali)

Trade‐offs are thought to be important in constraining evolutionary divergence, as they may limit phenotypic diversification. Limbless animals that burrow head‐first have been suggested to be evolutionarily constrained in the development of a large head size and sexual head shape dimorphism because of potential trade‐offs associated with burrowing. Here we use an acontiine skink (Acontias percivali) to test for the existence of trade‐offs between traits thought to be important in burrowing (speed and force). As head size dimorphism has been shown to be limited in acontiine lizards, thus suggesting constraints on head size and shape, we additionally explore the potential for trade‐offs between burrowing and biting. Our data show that A. percivali uses a burrowing style different from those previously described for caecilians and amphisbaenians, which relies on the use of extensive lateral and dorsoventral head movements. Our data also show that animals use their entire bodies to generate force, as peak force was determined by total length only. Additionally, both bite force and the time needed to burrow into the substrate were principally determined by relative head width, suggesting a trade‐off between biting and burrow speed. Performance data were indeed suggestive of a correlation between bite force and the time needed to burrow, but additional data are needed to confirm this pattern. In summary, our data suggests that trade‐offs may exist, and may have been of crucial importance in shaping the evolution of head shape in A. percivali, and burrowing lizards more generally. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 91–99.     

Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time

The grasses (Poaceae) are the fifth most diverse family of angiosperms, including 800 genera and more than 10 000 species. Few phylogenetic studies have tried to investigate palaeo‐biogeographical and palaeo‐ecological scenarios that may have led to present‐day distribution and diversity of grasses at the family level. We produced a dated phylogenetic tree based on combined plastid DNA sequences and a comprehensive sample of Poaceae. Furthermore, we produced an additional tree using a supermatrix of morphological and molecular data that included all 800 grass genera so that ancestral biogeography and ecological habitats could be inferred. We used a likelihood‐based method, which allows the estimation of ancestral polymorphism in both biogeographical and ecological analyses for large data sets. The origin of Poaceae was retrieved as African and shade adapted. The crown node of the BEP + PACCMAD clade was dated at 57 Mya, in the early Eocene. Grasses dispersed to all continents by approximately 60 million years after their Gondwanan origin in the late Cretaceous. PACCMAD taxa adapted to open habitats as early as the late Eocene, a date consistent with recent phytolith fossil data for North America. C₄ photosynthesis first originated in Africa, at least for Chloridoideae in the Eocene at c. 30 Mya. The BEP clade members adapted to open habitats later than PACCMAD members; this was inferred to occur in Eurasia in the Oligocene. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 543–557.     

New representatives of the genus Ancorabolina George, 2006 (Copepoda,Harpacticoida, Ancorabolidae) including remarks on ancorabolid phylogeny

Four new species of Ancorabolina George, 2006 (Copepoda: Harpacticoida: Ancorabolidae) are described: Ancorabolina belgicae sp. nov. (from the Porcupine Seabight, north‐east Atlantic Ocean), Ancorabolina anaximenesi sp. nov. and Ancorabolina galeata sp. nov. (both from Anaximenes Mountain, eastern Mediterranean Sea), and Ancorabolina divasecunda sp. nov. (from Cape and Guinea Basins, south‐east Atlantic Ocean). Furthermore, the generic diagnosis for Ancorabolina is provided and the position of the taxon within Ancorabolidae is discussed. The characters originally employed to allocate Ancorabolina to the subfamily Ancorabolinae Sars, 1909, are critically evaluated and two true autapomorphies for a monophyletic Ancorabolinae (including Ancorabolina) are recognized. Two autapomorphies supporting Ancorabolina are proposed: (1) cephalothorax posteriorly with pair of ventrolateral cuticular processes turned backwards and (2) loss of the third setal element subapically on the antennary endopod. The urgent need for a phylogenetic re‐evaluation of the paraphyletic subfamily Laophontodinae Lang, 1944 is stressed and evidenced by three characters widespread in this subfamily but also present in Ancorabolina. This may be an indication of a closer relationship of Ancorabolina with certain members of Laophontodinae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 16–55.     

Morphology,morphogenesis, and molecular phylogeny of a new marine urostylid ciliate (Ciliophora,Stichotrichia) from the South China Sea,and a brief overview of the convergent evolution of the midventral pattern within the Spirotrichea

The morphology, infraciliature, morphogenetic events, and small subunit ribosomal RNA (SSU rRNA) gene‐based phylogeny of a stichotrich ciliate, Bergeriella ovata gen. et sp. nov. , found in coastal waters off Daya Bay, Guangdong, South China, were investigated. The new taxon has an atypical midventral pattern, and is characterized by several rather unusual morphological features: frontal cirri forming an irregular tricorona; one nonmigratory row located right of the midventral rows, which is morphogenetically derived from the posteriormost fronto‐ventral‐transverse (FVT) streak; conspicuously enlarged postoral ventral cirri, whereas the left ventral cirri are delicate and arranged in oblique rows. Based on both the morphological and morphogenetical data, a new family, Bergeriellidae fam. nov. , is proposed. In the SSU rRNA phylogenetic trees, B. ovata gen. et sp. nov. groups with the classic urostylids (urostylids s.s.) with high nodal support. Our phylogenetic analyses have revealed at least seven separate clades for midventral pattern‐bearing taxa (e.g. Pattersoniella, Neokeronopsis, Rigidothrix, urostylids s.s., Uroleptidae, Holostichidae + Psammomitridae, and Pseudoamphisiellidae), suggesting that the order Urostylida Jankowski, 1979 sensu Lynn, 2008 is a polyphyletic assemblage, and that the convergent evolution of the urostylid midventral pattern might have occurred more times among the spirotrichs than has previously been recognized. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 697–710.     

A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships

A nearly complete skull and associated osteoderms from the Middle/Upper Triassic Madygen Formation of Kyrgyzstan are referred to a new chroniosuchid genus and species. The new taxon is characterized by a parabolic skull outline, pustular ornamentation, tabular‐squamosal contact, marked postparietal embayments, and the lack of an antorbital fontanelle. The palate is only preserved in part, showing broad palatines and ectopterygoids. Presence of a preorbital fenestra and characteristic osteoderm morphology are synapomorphies shared with all other chroniosuchids. According to the phylogenetic analysis performed, the new chroniosuchid nests with Chroniosaurus, with which it shares the wide, transversely extended osteoderms and pustular ornamentation. The chroniosuchians are robustly supported as a natural group, but their position within the reptiliomorph (stem‐amniote) clade is not adequately understood. Whereas the parasphenoid is similar to that of anthracosaurs, most other characters support a higher nesting of chroniosuchians within the stem‐amniotes. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 515–530.     

Molecular phylogenetics of Uvaria (Annonaceae): relationships with Balonga,Dasoclema and Australian species of Melodorum

An extended molecular phylogenetic analysis of Uvaria (Annonaceae) is presented, using maximum parsimony, maximum likelihood and Bayesian methods, based on sequences of four plastid DNA regions (matK, psbA‐trnH spacer, rbcL and trnL‐F). The additional taxa include the monotypic West African genus Balonga, the monotypic South‐East Asian genus Dasoclema and seven Australian representatives of the genus Melodorum. The results indicate that all of these taxa are nested within a well‐supported clade otherwise consisting of Uvaria species, indicating that their taxonomic treatment needs to be reassessed. The distinguishing morphological characteristics of the taxa are re‐evaluated and interpreted as specialized adaptations of the basic Uvaria structure. The genus Uvaria is accordingly extended following the transfer of these species, necessitating six new nomenclatural combinations and two replacement names. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 33–43.     

Biogeographic anomaly or human introduction: a cryptogenic population of tree skink (Reptilia: Squamata) from the Cook Islands,Oceania

Archaeological and molecular data have revealed that the present day faunas of many island groups in Melanesia, Polynesia, and Micronesia are not representative of the biodiversity generated within this region on an evolutionary timescale. Erroneous inferences regarding the mechanisms of speciation and the significance of long distance dispersal in shaping the present diversity of these island systems have resulted from this incomplete diversity and distributional data. The lizard fauna east of Samoa has been suggested to derive entirely from human‐mediated introductions, a distribution congruent with biogeographic patterns for other Pacific species. Distinguishing between introduced populations and those that result from natural colonization events is difficult, although molecular data provide a useful means for elucidating population history and identifying the likely sources of introductions. We use molecular data (1726 bp of mitochondrial DNA and 286 bp of nuclear DNA) to evaluate a population of arboreal lizards from the Cook Islands and to determine whether this arboreal skink population is the sole endemic component of the lizard fauna east of Samoa or the result of human‐mediated introduction. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 318–328.     

Evaluating hypotheses on the origin and diversification of the ringneck snake Diadophis punctatus (Colubridae: Dipsadinae)

Close affinities recognized between taxa in Mexico and the contiguous USA have led to a variety of biogeographical scenarios. One such hypothesis suggests that species that occur in both countries have an origin in central Mexico followed by dispersal into the USA. This study expands upon previous phylogeographical work of the ringneck snake Diadophis punctatus by incorporating new data from previously unsampled areas appropriate to critically assess hypotheses regarding a Mexican origin for this species. Maximum likelihood and maximum parsimony analyses inferred a derived position for the lineage from southern Mexico with constraint tests for alternate evolutionary hypotheses resulting in significantly worse likelihood values. Ancestral area reconstructions inferred an origin for D. punctatus in the south‐eastern USA followed by a south‐east to north‐east then westward directionality of historical migration. The position within the phylogeny and date estimate for the south‐western + Mexico clade suggests a recent invasion into central Mexico with expansion into the Nearctic/Neotropic transition zone. The extensive lineage diversity inferred from the mtDNA suggests that the genus is a complex of cryptic species whose conservational status should be re‐evaluated on both the national and regional levels. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 629–640.     

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.     

Molecular systematics of flyingfishes (Teleostei: Exocoetidae): evolution in the epipelagic zone

The flyingfish family Exocoetidae is a diverse group of marine fishes that are widespread and abundant in tropical and subtropical seas. Flyingfishes are epipelagic specialists that are easily distinguished by their enlarged fins, which are used for gliding leaps over the surface of the water. Although phylogenetic hypotheses have been proposed for flyingfish genera based on morphology, no comprehensive molecular studies have been performed. In the present study, we describe a species‐level molecular phylogeny for the family Exocoetidae, based on data from the mitochondrial cytochrome b gene (1137 bp) and the nuclear RAG2 gene (882 bp). We find strong support for previous morphology‐based phylogenetic hypotheses, as well as the monophyly of most currently accepted flyingfish genera. However, the most diverse genus Cheilopogon is not monophyletic. Using our novel flyingfish topology, we examine previously proposed hypotheses for the origin and evolution of gliding. The results support the progressive transition from two‐wing to four‐wing gliding. We also use phylogenetic approaches to test the macroecological effects of two life history characters (e.g. egg buoyancy and habitat) on species range size in flyingfishes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102, 161–174.     

Biogeographical,ecological and morphological structure in a phylogenetic analysis of Ateleia (Swartzieae,Fabaceae) derived from combined molecular,morphological and chemical data

A phylogenetic analysis of combined morphological, chemical and ITS/5.8S sequence data reveals that species of Ateleia are often more genetically than morphologically divergent, and that species thought to be most closely related morphologically are distant relatives within the genus. Ateleia shows niche conservatism, with most species confined to seasonally dry tropical forest in Central America and the Caribbean, and fewer species in the same biome in South America. Four independent transitions to wet forests may have occurred in the genus. The estimated ages of Ateleia lineages spanning Central and South America are either older or younger than the estimated age of closure of the Isthmus of Panama. The older dates clearly suggest that over‐water dispersal is responsible for the distribution of Ateleia that includes the Caribbean Islands. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 39–53.     

Failure to cospeciate: an unsorted tale of millipedes and mites

Mites form symbiotic relationships with many animal taxa, including fish, amphibians, reptiles, birds, mammals, mollusks and arthropods. They are often found living on millipedes and it has often been speculated that these two groups of arthropods have, in some cases, undergone coevolution. However, this hypothesis has never been formally tested. Millipedes of the family Xystodesmidae Cook 1895 (Diplopoda: Polydesmida) and their symbiotic mites of the genus Stylochyrus Canestrini & Canestrini 1882 were collected in broadleaf forests of the eastern USA. The DNA from two mitochondrial regions (16S/12S and cox1) was sequenced for all collected millipede and mite specimens. Phylogenetic trees were reconstructed for both millipede and mite taxa using Bayesian inference. Pairwise distance data were used in distance‐based coevolutionary analyses and reconstructed phylogenies were used in tree‐based coevolutionary analyses. The phylogenetic analyses indicate Stylochyrus and xystodesmid millipede evolutionary history is incongruent. Moreover, the evolutionary relationships among mite individuals and populations have very low support values and indicate little to no geographic structuring. The coevolutionary analyses likewise detected no pattern of coevolution among these millipede and mite lineages. Unlike many arthropod species, Stylochyrus mites appear to be highly vagile. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 272–287.     

An invasive species imposes selection on life‐history traits of a native frog

As well as their direct ecological impacts on native taxa, invasive species can impose selection on phenotypic attributes (morphology, physiology, behaviour, etc.) of the native fauna. In anurans, body size at metamorphosis is a critical life‐history trait: for most challenges faced by post‐metamorphic anurans, larger size at metamorphosis probably enhances survival. However, our studies on Australian frogs (Limnodynastes convexiusculus) show that this pattern can be reversed by the arrival of an invasive species. When metamorph frogs first encounter invasive cane toads (Bufo marinus), they try to eat the toxic invader and, if they are able to do so, are likely to die from poisoning. Because frogs are gape‐limited predators, small metamorphs cannot ingest a toad and thus survive long enough to disperse away from the natal pond (and thus from potentially deadly toads). These data show that larger size at metamorphosis can reduce rather than increase anuran survival rates, because larger metamorphs are more easily able to ingest (and thus be poisoned by) metamorph cane toads. Our results suggest that patterns of selection on life‐history traits of native taxa (such as size and age at metamorphosis, seasonal timing of breeding and duration of pondside aggregation prior to dispersal) can be modified by the arrival of an invasive species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 329–336.