Phylogeny and systematics of Melanesia’s most diverse gecko lineage (Cyrtodactylus,Gekkonidae, Squamata)

Oliver, P.M., Richards, S.J. & Sistrom, M. (2012). Phylogeny and systematics of Melanesia’s most diverse gecko lineage (Cyrtodactylus, Gekkonidae, Squamata). —Zoologica Scripta, 41, 437–454. The systematics and biogeographical history of the diverse fauna of New Guinea and surrounding islands (Melanesia) remain poorly known. We present a phylogeny for 16 of the 21 recognised Melanesian bent‐toed geckos in the genus Cyrtodactylus based on mitochondrial sequence data. These analyses reveal two divergent lineages of Cyrtodactylus within Melanesia. One includes a single recognised species with clear affinities to sampled taxa from Asia. The other comprises a relatively diverse radiation (likely 30+ species), not closely related to sampled extralimital taxa and centred on the Melanesian region (including Australia). Many taxa within this second lineage are endemic to islands surrounding New Guinea, and dispersal and speciation on peripheral islands appears to have played an important role in the accumulation of species diversity within this clade. In contrast, little diversity is centred upon montane areas, although we do identify at least one lineage closely associated with hill and lower montane forest that probably dates to at least the late Miocene. Our phylogenetic analyses also reveal numerous divergent lineages that require taxonomic attention, including at least two widespread taxa that are likely to be composite, additional specimens of Cyrtodactylus capreoloides (until recently known only from the holotype) and several divergent and completely novel lineages, two of which we introduce herein: Cyrtodactylus arcanus sp. n. and Cyrtodactylus medioclivus sp. n.     

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.     

Systematics of the Saharo‐Arabian clade of the Palearctic naked‐toed geckos with the description of a new species of Tropiocolotes endemic to Oman

Geckos are one of the most species‐rich, abundant, and widely distributed of all Squamata lineages and present several characteristics that have made them favorite model organisms for biogeographical, ecological, physiological, and evolutionary studies. One of the key aspects of any comparative study is to have a robust, comprehensive phylogeny, and an updated taxonomy. Recently, the Infraorder Gekkota has been the subject of several phylogenetic analyses and taxonomic revisions at different levels. Despite all these phylogenetic and taxonomic advances, there are still some groups whose systematics and taxonomy remain highly problematic. Maybe one of the most poorly resolved groups in spite of decades of intensive research by many herpetologists are the so‐called Palearctic naked‐toed geckos of the family Gekkonidae. This group of nocturnal geckos distributed from Mauritania across North Africa, Arabia, southwestern and central Asia to northern India, western China and southern Mongolia is characterized by the synapomorphy of lack of adhesive subdigital pads. Within the Palearctic naked‐toed geckos, the Saharo‐Arabian clade comprised by the genera Pseudoceramodactylus, Stenodactylus, and Tropiocolotes is the clade with the largest distribution range. At the same time, it is one of the problematic groups, presenting poorly supported phylogenetic relationships, with the genus Tropiocolotes being recovered non‐monophyletic in all analyses despite its morphological uniformity. To reassess the phylogeny of the Palearctic naked‐toed geckos with a special interest in the systematics of Tropiocolotes, we assembled a dataset comprising 298 gecko specimens from 283 different species (including all Tropiocolotes species but one) belonging to 122 of a total of 124 described gecko genera. This dataset included the nuclear c‐mos, ACM4, RAG1, RAG2, and PDC and the mitochondrial ND2 gene. To further investigate the relationships within Tropiocolotes and to revise the systematics of the south Arabian endemic species Tropiocolotes scorteccii, we used an integrative approach including information from the nuclear MC1R and c‐mos, the mitochondrial 12S, 16S, cytb genes, and morphological data from nine of the 10 described Tropiocolotes species. The phylogenetic analyses of the Gekkota dataset recovered a similar topology for the Palearctic naked‐toed geckos to previous studies, but in this case, Tropiocolotes was recovered monophyletic in all analyses, with high support in two of them. The results of the analyses of three datasets specifically assembled to test the effect of both gene sampling and taxon sampling in the monophyly of Tropiocolotes, and the internal relationships of the Palearctic naked‐toed geckos clearly showed that both the number and kind of characters (nuclear or mitochondrial data) and the number of taxa played a fundamental role in recovering the correct phylogenetic relationships. The phylogenetic analyses within Tropiocolotes suggested the existence of high levels of undescribed diversity in the south Arabian T. scorteccii, including a new genetically and morphologically distinct species endemic to Oman (Tropiocolotes confusus sp. nov. ). Our study using a large dataset, including several loci and a dense taxon sampling within Gekkota and especially within Tropiocolotes, has proved a valuable strategy to address the monophyly of Tropiocolotes and the relationships within the Saharo‐Arabian Palearctic naked‐toed geckos. The integrative systematic approach including several samples of south Arabian T. scorteccii based on many years of fieldwork has, once more, uncovered a new species endemic to this region. This highlights the importance of this area of Arabia as a reservoir of reptile endemicity and biodiversity, which is likely linked to the high degree of habitat heterogeneity and the effect of the monsoons. Obviously, based on this and previously published evidence, south Arabia represents an area with still high levels of undiscovered diversity.     

Molecular evidence for Gondwanan origins of multiple lineages within a diverse Australasian gecko radiation

Aim Gondwanan lineages are a prominent component of the Australian terrestrial biota. However, most squamate (lizard and snake) lineages in Australia appear to be derived from relatively recent dispersal from Asia (< 30 Ma) and in situ diversification, subsequent to the isolation of Australia from other Gondwanan landmasses. We test the hypothesis that the Australian radiation of diplodactyloid geckos (families Carphodactylidae, Diplodactylidae and Pygopodidae), in contrast to other endemic squamate groups, has a Gondwanan origin and comprises multiple lineages that originated before the separation of Australia from Antarctica. Location Australasia. Methods Bayesian (beast ) and penalized likelihood rate smoothing (PLRS) (r 8s ) molecular dating methods and two long nuclear DNA sequences (RAG‐1 and c‐mos) were used to estimate a timeframe for divergence events among 18 genera and 30 species of Australian diplodactyloids. Results At least five lineages of Australian diplodactyloid geckos are estimated to have originated > 34 Ma (pre‐Oligocene) and basal splits among the Australian diplodactyloids occurred c. 70 Ma. However, most extant generic and intergeneric diversity within diplodactyloid lineages appears to post‐date the late Oligocene (< 30 Ma). Main conclusions Basal divergences within the diplodactyloids significantly pre‐date the final break‐up of East Gondwana, indicating that the group is one of the most ancient extant endemic vertebrate radiations east of Wallace’s Line. At least five Australian lineages of diplodactyloid gecko are each as old or older than other well‐dated Australian squamate radiations (e.g. elapid snakes and agamids). The limbless Pygopodidae (morphologically the most aberrant living geckos) appears to have radiated before Australia was occupied by potential ecological analogues. However, in spite of the great age of the diplodactyloid radiation, most extant diversity appears to be of relatively recent origin, a pattern that is shared with other Australian squamate lineages.     

The development of cephalic armor in the tokay gecko (Squamata: Gekkonidae: Gekko gecko)

Armored skin resulting from the presence of bony dermal structures, osteoderms, is an exceptional phenotype in gekkotans (geckos and flap-footed lizards) only known to occur in three genera: Geckolepis, Gekko, and Tarentola. The Tokay gecko (Gekko gecko LINNAEUS 1758) is among the best-studied geckos due to its large size and wide range of occurrence, and although cranial dermal bone development has previously been investigated, details of osteoderm development along a size gradient remain less well-known. Likewise, a comparative survey of additional species within the broader Gekko clade to determine the uniqueness of this trait has not yet been completed. Here, we studied a large sample of gekkotans (38 spp.), including 18 specimens of G. gecko, using X-rays and high-resolution computed tomography for visualizing and quantifying the dermal armor in situ. Results from this survey confirm the presence of osteoderms in a second species within this genus, Gekko reevesii GRAY 1831, which exhibits discordance in timing and pattern of osteoderm development when compared with its sister taxon, G. gecko. We discuss the developmental sequence of osteoderms in these two species and explore in detail the formation and functionality of these enigmatic dermal ossifications. Finally, we conducted a comparative analysis of endolymphatic sacs in a wide array of gekkotans to explore previous ideas regarding the role of osteoderms as calcium reservoirs. We found that G. gecko and other gecko species with osteoderms have highly enlarged endolymphatic sacs relative to their body size, when compared to species without osteoderms, which implies that these membranous structures might fulfill a major role of calcium storage even in species with osteoderms.     

Small snails in a big place: a radiation in the semi‐arid rangelands in northern Australia (Eupulmonata,Camaenidae, Nanotrachia gen. nov.)

Continuing the revision of the Camaenidae in the Australian Monsoon Tropics, we employed comparative analyses of morphological features (with a focus on shell and penial anatomy) and genetic markers (with a focus on mitochondrial COI and 16S sequences) to address the systematic relationships of land snails from the Victoria River District, Northern Territory, and adjacent East Kimberley (Western Australia). These analyses revealed that the species under study represented the previously undescribed genus Nanotrachia. This genus differs from all other camaenid genera known from north‐western Australia most conspicuously by its small, flat, and ribbed shell. Six species are identified as members of the new genus, four of them new species ( Nanotrachia costulata sp. nov. , Nanotrachia carinata sp. nov. , Nanotrachia coronata sp. nov. , Nanotrachia levis sp. nov. ). Two further species have already been described previously but assigned to different genera. These species, Ordtrachia intermedia (as the type species of Nanotrachia) and Mouldingia orientalis, are here transferred to Nanotrachia. Like other camaenids from the Australian Monsoon Tropics, species of Nanotrachia are characterized by essentially allopatric distributions, regional endemism, and a patchy distribution across their range. © 2013 The Linnean Society of London     

Karyotypic evolution in Gehyra (Gekkonidae: Reptilia) IV. Chromosome change and speciation

Karyotypic data are presented for six additional species from the genus Gehyra collected in Australia, New Guinea and Fiji. C and G-banding of three of the very diverse species which all share the ancestral 2n=44 karyotype, further strengthens the phylogenetic model for the evolution of this complex. With 19 Australian species and chromosome races of Gehyra now karyotyped, it has been possible to evaluate the mode of chromosomal evolution and the role that chromosome change has played in speciation in this genus. It is clear that speciation in certain karyomorph groups has occurred allopatrically, without any gross chromosomal changes. However, in the numerous chromosome races and species which have been involved in colonizing radiations, chromosomal rearrangements have been intricately associated with the speciation process.     

Systematics and biogeography of Australian rhopalosomatid wasps (Hymenoptera: Rhopalosomatidae) with a global synopsis of the enigmatic genus Olixon Cameron

Abstract The wasp family Rhopalosomatidae is represented in Australia only by the genus Olixon Cameron. Species of this genus have previously been considered rare based on material in collections and have rarely been observed in the field. All known species of Olixon are brachypterous, solitary ectoparasitoids of crickets (Orthoptera: Gryllidae). Prior to our study the world fauna comprised 11 species of which two were endemic to north‐eastern Australia. Based on specimens collected during recent intensive surveys in Australia we record 17 species for the continent. Fifteen of these species are described as new, which more than doubles the world fauna of Olixon: Olixon abrahami, O. danggari, O. ferrugineum, O. guyim, O. harveyi, O. helgae, O. jandakotae, O. jawoyn, O. jenningsi, O. kakadui, O. pilbara, O. wajuk, O. waldockae, O. wuthathi and O. zonale spp.n . Females of Olixon australiae Perkins and Olixon flavibase Townes are redescribed, and the male of O. flavibase is described for the first time. A key is provided for all 17 Australian species and their distribution is discussed. Results of a cladistic analysis of the world species of Olixon based on 41 morphological characters for 24 ingroup species are presented. The results obtained from equal and implied weighting parsimony analyses indicate that: (i) the Australian species of Olixon are not monophyletic, (ii) about three‐quarters of the Australian species form a monophyletic group, and (iii) a Central/South American Olixon is sister to all other species. The implications of these results for the biogeography of world Olixon species are briefly discussed.     

Embryology of Harrisonia (Cneoroideae,Rutaceae): a comparison with related genera and families

Harrisonia, a genus of three or more species distributed from tropical Africa and Southeast Asia to northern Australia, has long been considered a member of Simaroubaceae. Recent molecular analyses, however, have shown that the genus is assigned to subfamily Cneoroideae of Rutaceae, in which Cneoroideae (eight genera) are sister to the remainder of the family (‘core Rutaceae’). Here, we report embryological features of Harrisonia based on published and newly obtained data and provide morphological corroboration for the molecular affinities of the genus and Cneoroideae. We compared its embryological features with those of seven other genera of Cneoroideae and with those of the core Rutaceae and related families (Meliaceae, Simaroubaceae and Sapindaceae). Comparisons showed that Harrisonia fits within Cneoroideae through possessing campylotropous seeds with a multi‐cell‐layered endotesta. Embryological evidence, like molecular evidence, further shows that, in Cneoroideae, Harrisonia probably has affinities to a group of Bottegoa, Cedrelopsis, Cneorum and Ptaeroxylon by sharing a micropyle formed by the inner integument alone and solitary oil cells in the testa. Except for Harrisonia, Cneoroideae are still poorly understood embryologically and further investigations are needed.     

Phylogeography of the whelk genus Cominella (Gastropoda: Buccinidae) suggests long‐distance counter‐current dispersal of a direct developer

The gastropod genus Cominella Gray, 1850 consists of approximately 20 species that inhabit a wide range of marine environments in New Zealand and Australia, including its external territory, the geographically isolated Norfolk Island. This distribution is puzzling, however, with apparently closely‐related species occurring either side of the Tasman Sea, even though all species are considered to have limited dispersal abilities. To determine how Cominella attained its current distribution, we derived a dated molecular phylogeny, which revealed a clade comprising all the Australian and Norfolk Island species nested within four clades of solely New Zealand species. This Australian clade diverged well after the vicariant separation of New Zealand from Australia, and implies two long‐distance dispersal events: a counter‐current movement across the Tasman Sea from New Zealand to Australia, occurring at the origination of the clade, followed by the colonization of Norfolk Island. The biology of Cominella suggests that the most likely method of long‐distance dispersal is rafting as egg capsules. Our robust phylogeny also means that the current Cominella classification requires revision. We propose that our clades be recognized as subgenera: Cominella (s.s.), Cominista, Josepha, Cominula, and Eucominia, with each subgenus comprising only of New Zealand or Australian species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 315–332.     

Interactions between native and invasive gecko lizards in tropical Australia

Invasive geckos of the genus Hemidactylus (Gekkonidae) are spreading rapidly through urban environments in many tropical and subtropical parts of the world. The invaders have caused rapid declines in native gecko abundance in some areas, but their interactions with Australian native species remain unknown. In a small rural village near Darwin, we found that Hemidactylus frenatus is abundant around both lighted and unlighted buildings, but rarely found in surrounding bushland. It is sympatric with the larger Gehyra australis (Gekkonidae) in this disturbed site, and often forages on the same buildings, but is active mostly during the dry‐season (vs. wet‐season for G. australis) and is competitively subordinate to the larger native species. In laboratory encounters, H. frenatus fled from G. australis, and modified its refuge‐site use in the presence of the native lizard. In those same trials, the native gecko often attacked and rarely fled; and did not shift its refuge‐site selection. In field surveys, the two taxa frequently co‐occurred. However, substrate use of the invasive H. frenatus was modified by the presence of the native G. australis, consistent with competitive displacement. Our counts of H. frenatus were highest during the dry‐season, when G. australis (like most other small native reptiles) is relatively inactive. The invasive gecko thus appears to be exploiting a ‘vacant niche’ around buildings, rather than displacing the native gecko taxon. This outcome may reflect the size disparity between the native species and the invader; Hemidactylus frenatus may well have significant ecological impacts on smaller native lizards.     

Morphological differentiation correlates with ecological but not with genetic divergence in a Gehyra gecko

Body size affects life history, the ecological niche of an organism and its interactions with other organisms. Resultantly, marked differences in body size between related organisms are often an indication of a species boundary. This is particularly evident in the Gehyra variegata species complex of geckos, which displays differential body sizes between genetically divergent species, but high levels of intraspecific morphological conservatism. We report on a Gehyra population that displays extraordinary body size differentiation in comparison with other G. variegata species. We used morphological and environmental data to show this population is phenotypically and ecologically distinct from its parapatric congener Gehyra lazelli and that morphology and ecology are significantly correlated. Contrastingly, mtDNA analysis indicates paraphyly between the two groups, and allele frequencies at six microsatellite loci show no population structure concordant with morpho‐/ecotype. These results suggest either ecological speciation or environmentally induced phenotypic polymorphism, in an otherwise morphologically conservative group.     

Phylogenetics of the allodapine bee genus Braunsapis: historical biogeography and long-range dispersal over water

Aim A previous study of the allodapine bee genus Braunsapis suggested an African origin, with dispersal events into Madagascar and Asia, and from Asia into Australia. We re‐examine the phylogeny of this genus, using an expanded set of taxa from Madagascar and Malawi and additional sequence data, in order to determine the number of dispersals and the timeframe over which they occurred. Location Africa, Madagascar, Malawi, Asia and Australia. Methods One nuclear (EF‐1α F2) and two mitochondrial (CO1 and Cyt b) gene regions were sequenced for 36 allodapine bee species (including members of the genera Braunsapis, Nasutapis, Allodape, Allodapula, and Macrogalea) and one ceratinine species (Ceratina japonica). We used Bayesian analyses to examine phylogenetic structure and a penalized likelihood approach to estimate approximate ages for key divergences in our phylogeny. Results Our analyses indicate a tropical African origin for Braunsapis in the early Miocene followed by very early dispersal into Asia and then a subsequent dispersal, following Asian diversification, into Australia during the late Miocene. There have also been two dispersals of Braunsapis from Africa to Madagascar and this result, when combined with phylogenetic and biogeographical data for other allodapines, suggests that these bees have the ability to cross moderately large ocean expanses. These dispersals may have been aided by the West Wind Drift, but rafting across the Mozambique Channel is also possible, and could be aided by the existence of developmental stages that require minimal or no feeding and by tolerance to sea water and spume. Accumulating evidence suggests that many biogeographical patterns in the southern hemisphere may be better explained by dispersal than by Gondwanan vicariance hypotheses. Our results add to this growing body of data and raise the possibility that some puzzling trans‐Indian Ocean distributions may also be explained by historical dispersal events across oceanic barriers that now seem insuperable.     

Phylogeography of the Indo‐West Pacific maskrays (Dasyatidae,Neotrygon): a complex example of chondrichthyan radiation in the Cenozoic

Maskrays of the genus Neotrygon (Dasyatidae) have dispersed widely in the Indo‐West Pacific being represented largely by an assemblage of narrow‐ranging coastal endemics. Phylogenetic reconstruction methods reproduced nearly identical and statistically robust topologies supporting the monophyly of the genus Neotrygon within the family Dasyatidae, the genus Taeniura being consistently basal to Neotrygon, and Dasyatis being polyphyletic to the genera Taeniurops and Pteroplatytrygon. The Neotrygon kuhlii complex, once considered to be an assemblage of color variants of the same biological species, is the most derived and widely dispersed subgroup of the genus. Mitochondrial (COI, 16S) and nuclear (RAG1) phylogenies used in synergy with molecular dating identified paleoclimatic fluctuations responsible for periods of vicariance and dispersal promoting population fragmentation and speciation in Neotrygon. Signatures of population differentiation exist in N. ningalooensis and N. annotata, yet a large‐scale geological event, such as the collision between the Australian and Eurasian Plates, coupled with subsequent sea‐level falls, appears to have separated a once homogeneous population of the ancestral form of N. kuhlii into southern Indian Ocean and northern Pacific taxa some 4–16 million years ago. Repeated climatic oscillations, and the subsequent establishment of land and shallow sea connections within and between Australia and parts of the Indo‐Malay Archipelago, have both promoted speciation and established zones of secondary contact within the Indian and Pacific Ocean basins.     

The Hawaiian Archipelago is a stepping stone for dispersal in the Pacific: an example from the plant genus Melicope (Rutaceae)

Aim Pacific biogeographical patterns in the widespread plant genus Melicope J.R. Forst. & G. Forst. (Rutaceae) were examined by generating phylogenetic hypotheses based on chloroplast and nuclear ribosomal sequence data. The aims of the study were to identify the number of colonization events of Melicope to the Hawaiian Islands and to reveal the relationship of Hawaiian Melicope to the Hawaiian endemic genus Platydesma H. Mann. The ultimate goal was to determine if the Hawaiian Islands served as a source area for the colonization of Polynesia. Location Nineteen accessions were sampled in this study, namely eight Melicope species from the Hawaiian Islands, four from the Marquesas Islands, one species each from Tahiti, Australia and Lord Howe Island, two Australian outgroups and two species of the Hawaiian endemic genus Platydesma. To place our results in a broader context, 19 sequences obtained from GenBank were included in an additional analysis, including samples from Australia, Papua New Guinea, New Zealand, Southeast Polynesia and Asia. Methods DNA sequences were generated across 19 accessions for one nuclear ribosomal and three chloroplast gene regions. Maximum parsimony analyses were conducted on separate and combined data sets, and a maximum likelihood analysis was conducted on the combined nuclear ribosomal and chloroplast data set. A broader nuclear ribosomal maximum parsimony analysis using sequences obtained from GenBank was also performed. Geographic areas were mapped onto the combined chloroplast and nuclear ribosomal tree, as well as onto the broader tree, using the parsimony criterion to determine the dispersal patterns. Results Phylogenetic analyses revealed that Platydesma is nested within Melicope and is sister to the Hawaiian members of Melicope. The Hawaiian Melicope + Platydesma lineage was a result of a single colonization event, probably from the Austral region. Finally, Marquesan Melicope descended from at least one, and possibly two, colonization events from the Hawaiian Islands. Main conclusions These data demonstrate a shifting paradigm of Pacific oceanic island biogeography, in which the patterns of long‐distance dispersal and colonization in the Pacific are more dynamic than previously thought, and suggest that the Hawaiian Islands may act as a stepping stone for dispersal throughout the Pacific.     

Morphological and molecular evidence for phylogeny and classification of South American pitvipers,genera Bothrops,Bothriopsis, and Bothrocophias (Serpentes: Viperidae)

Species in the genus Bothrops s. l. are extraordinarily variable in ecology and geography, compared with other genera in the subfamily Crotalinae. In contrast to the trend of splitting large and variable groups into smaller, more ecologically and phenotypically cohesive genera, the genus Bothrops has remained speciose. In addition, previous phylogenetic analyses have found Bothrops to be paraphyletic with respect to the genus Bothriopsis. Taxonomic arguments exist for synonymizing Bothriopsis with Bothrops, and for splitting Bothrops into smaller genera, but the greatest hindrance to taxonomic revision has been incomplete phylogenetic information. We present a phylogeny of Bothrops, Bothriopsis, and Bothrocophias based on 85 characters of morphology and 2343 bp of four mitochondrial gene regions, and with significantly greater taxonomic coverage than previous studies. The combined data provide improved support over independent datasets, and support the existence of discrete species groups within Bothrops. The monophyly and distinctness of these groups warrant recognition at the generic level, and we propose a new taxonomic arrangement to reflect these findings. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 617–640.     

Gobekko cretacicus (Reptilia: Squamata) and its bearing on the interpretation of gekkotan affinities

Gobekko cretacicus, a Cretaceous lizard from the Gobi Desert of Mongolia, is a key fossil for understanding gecko phylogeny. We revisit this fossil using high‐resolution X‐ray computed tomography. The application of this imaging method reveals new information about sutures, bone shape, and structural details of the palate and basicranium. These data were used to assess the phylogenetic affinities of Gobekko in the context of an existing squamate data set. The effects of character ordering, search strategy, and the addition of another putative gekkonomorph (Hoburogekko suchanovi) on inferred gekkonomorph relationships were explored. Available specimens of G. cretacicus are skeletally mature but have unfused nasals, frontals, and parietals, and (possibly) a persistent basicranial fenestra. Some putative gekkonomorphs are not consistently supported as closer to crown clade gekkotans than to autarchoglossans. In a strict consensus both Gobekko and Hoburogekko form a polytomy with extant geckos. Some of the adult character states of Gobekko are observable in embryos of extant species. The evolution of tubular frontals and dentaries in gekkotans may be structurally related to the loss of the postorbital and supratemporal bars in this lineage. The complete lack of a parietal foramen, and presumably a light‐sensitive parietal eye, in this clade is of interest and could indicate an early origin of nocturnality in geckos. © 2013 The Linnean Society of London     

The historical biogeography of Apsilochorema (Trichoptera,Hydrobiosidae) revised,following molecular studies

The genus Apsilochorema Ulmer, 1907 is unique in the family Hydrobiosidae Ulmer, being widely distributed in the Palaearctic, Oriental and Australian Regions. All other 49 genera in the family, except the New World Atopsyche Banks, 1905, are confined to a single biogeographical Region. This unique distribution has independently stimulated researchers to formulate competing hypotheses about the biogeographical history of the genus. Molecular sequence data from mitochondrial cytochrome oxidase I (COI) and nuclear cadherin (CAD) genes of Apsilochorema species from the Oriental and Australian areas were analysed phylogenetically. The results retain a monophyletic Apsilochorema, which forms the sistergroup to the other genera in the subfamily Apsilochorematinae. The results from the biogeographical analyses dispute the earlier assumptions of an Oriental or northern Gondwana origin for the genus, revealing unambiguously an initial Australian radiation of the ancestral Apsilochorema with a subsequent dispersal into the Oriental Region. All but one of the Apsilochorema species occurring on the Pacific islands had an Oriental ancestor. The exception is the sistergroup to the New Caledonian species, which is found in both Australia and Oriental Regions. The molecular dating analysis, using a relaxed clock model, indicates that the genus Apsilochorema is about 36.4 MY old and that it dispersed from Australia into the Oriental Region about 28.3 Ma. It also gives an estimate of the approximate ages of the dispersals into New Caledonia to about 15.3 Ma; to the Solomon Islands at about 16.2 Ma; to the Fiji Islands at about 16.1 Ma; and to the Vanuatu Islands at about 5.4 Ma.     

TAXONOMY AND MOLECULAR PHYLOGENY OF THE RED ALGAL GENUS LENORMANDIA (RHODOMELACEAE,CERAMIALES)1

The genus Lenormandia Sonder is currently composed of nine species from Australia and New Zealand. Some of these are well known, but others are rare and ill defined. Material of all nine species has been examined and found to fall into three discrete morphological groups forming highly supported clades on analysis of 18S rDNA sequences. The first group contains four Australian‐endemic species and includes the type species L. spectabilis Sonder. Plants have a cleft apex that is not inrolled, a distinctive rhombic surface areolation pattern caused by a one‐ to two‐layered medulla of interlocking cells, lack pseudopericentral cells, and produce their reproductive structures on the blade surfaces. The type species of the genus Lenormandiopsis, L. latifolia (Harvey et Greville) Papenfuss, was found to belong to this group and is thus returned to Lenormandia where it was originally placed. Species falling into the other two groups are removed to new genera that are being described separately. One extremely rare species of Lenormandia from southwestern Australia is transferred to the delesseriacean genus Phitymophora.     

Contrasting fire-related resilience of ecologically dominant ants in tropical savannas of northern Australia

This paper examines the role of fire in mediating the relative abundance of two of the world's major ecologically dominant ant genera, Iridomyrmex and Oecophylla, where they coexist across the tropical savanna landscapes of northern Australia. These taxa have contrasting biogeographical histories, which are predicted to lead to contrasting responses to fire. Iridomyrmex is an autochthonous Australian genus that has radiated primarily in the arid zone; as such, its abundance is predicted to be promoted by frequent fire because this maintains an open habitat. In contrast, Oecophylla is a genus of leaf‐nesting ants occurring in the canopies of Old World tropical rainforest, and is a recent arrival to Australia in geological time; the abundance of these ants is predicted to decline under frequent fire. We test these predictions using results from a landscape‐scale fire experiment, where three experimental fire regimes (including no fire) were applied to replicated subcatchments over a 5‐year period. Using sweep nets, ants were sampled in the grass layer (the habitat layer of greatest overlap between Iridomyrmex and Oecophylla) in eucalypt woodland (canopy cover < 30%) and open eucalypt forest (canopy cover about 50%) habitats. A total of 27 species from 11 genera were collected during the study; eight were common enough for statistical analysis, and the abundances of four of these were significantly affected by fire treatment. As predicted, the abundance of Iridomyrmex was promoted by fire, whereas that of Oecophylla declined. These changes occurred only under late‐season (relatively high intensity) fires, and for Oecophylla occurred only in open forest (not woodland) habitat. This fire‐mediated relationship between Iridomyrmex and Oecophylla mirrors the much broader, ecosystem‐wide dynamic between eucalypt‐dominated savanna and rainforest in tropical Australia, with savannas dominated by fire‐resistant sclerophyll elements of Australian origin, and rainforest dominated by fire‐sensitive mesophyll elements of South‐East Asian origin.