Gastropod biodiversity at the 'Evolution Canyon' microsite, lower Nahal Oren, Mount Carmel, Israel

Twenty-six species of gastropods (terrestrial, shell-bearing snails and slugs) were recorded at the 'Evolution Canyon' microsite, lower Nahal Oren, Mount Carmel, Israel. Twenty-five species were recorded at the temperate, mesic north-facing slope (NFS) and 20 species at the xeric south-facing slope (SFS). Out of these species, six were NFS specific ( Pilorcula raymondi hebraica , Euchondrus septemdentatus , Monacha crispulata , Pyramidula rupestris hierosolymitana , Truncatellina haasi , and Vitrea contracta ) and one was SFS specific ( Prolimax eustrictus ). The interslope difference was probably partly due to missing forest species at the SFS in comparison with the NFS. Twenty-two species were Levantine endemics (84.6%) and four species were more widely distributed in the Palaearctic region (15.4%). The Levantine species are inhabitants of the mesic and mainly mountainous regions, but four species ( Granopupa granum , Calaxis hierosolymarum , Cecilioides acicula , and Helix engaddensis ) also penetrate the deserts. Seven species ( Buliminus labrosus , H. engaddensis , Levantina spiriplana caesareana , Metafruticicola fourousi , Monacha syriaca , Sphincterochila cariosa , and Xeropicta vestalis joppensis ) were significantly more abundant on the SFS than on the NFS. The local physical microclimatic sharp divergence leads to gastropod adaptive interslope biotic divergence caused by natural selection.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 147–155.     

Microsatellite loci for Australian agamid lizards

We characterize 15 microsatellite loci from two microsatellite libraries developed from the Australian agamid lizards Amphibolurus muricatus and Ctenophorus pictus. All loci were tested for amplification in four other agamids: Ctenophorus fordi, Ctenophorus decresii, Chlamydosaurus kingii, and Physignathus lesueurii. These loci were highly polymorphic within and across species, with nine to 12 loci amplifying for each species tested.     

Complex patterns of morphological and mtDNA variation in Lacerta perspicillata (Reptilia; Lacertidae)

Lacerta perspicillata is a north-west African lacertid lizard that shows considerable intraspecific variation, with three subspecies described on the basis of colour pattern and body size. Recent observations of a population containing two morphological forms and more than one deep genetic lineage, as well as an apparent lack of concordance between forms and genetic lineages, suggest that the complexity is greater than previously thought. To analyse and quantify this variation, we studied the variability within this species at two levels: (1) external morphology (multivariate analysis of scalation, body dimensions, and colour pattern) and (2) mtDNA (sequencing and single-strand confirmation polymorphism analysis). Fifty-two individuals were studied at Taza, northern Morocco. Two morphological groupings (ostensibly representing two previously described subspecies) and two deep mtDNA lineages were detected at this site, with complete correspondence between the two. This, together with an apparent lack of hybrids, would normally support respective full species recognition. However, analysis of 98 individuals from other populations demonstrated that the situation is highly complex with the same genetic lineages having reversed morphotypes in other areas, making such a designation difficult. Across the other studied populations, we found no support for any of the currently recognized subspecies. The lack of congruence between mtDNA lineages and morphometric patterns (in some cases) and the morphological similarity among lizards from different lineages suggest ecophenotypic convergence or multiple introgressive hybridization. The study highlights the tremendous complexity that may exist within a taxon and the inadequacy of older alpha-taxonomy based designations in describing it.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 479–490.     

Convergent body flattening in a clade of tropical rock-using lizards (Scincidae: Lygosominae)

The repeated occurrence of similar morphologies in organisms from similar habitats provides good evidence of convergent selection, and convergent patterns of evolutionary change. In lizards, a flattened morphology has often been noted; however, whether this trait is convergent in specific habitats has never been tested using phylogenetic methods. The present study examined patterns of morphological convergence in 18 species of tropical Lygosomine skinks from three broad habitat categories (generalist, leaf litter-dwelling, and rock-using species). In general, although there where relatively few morphological differences of species from different habitats, phylogenetic analyses revealed that rock-using species have consistently and repeatedly evolved a dorsoventrally flattened head and body. The adaptive basis of this flattened morphology is consistent with both biomechanical predictions of performance (e.g. climbing locomotion) and ecology (e.g. use of rock crevices, camouflage) of species that occupy rocky habitats.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 399–411.     

Habitat-specific differences in thermal plasticity in natural populations of a soil arthropod

When populations experience substantial variation in environmental conditions, they may evolve phenotypic plasticity in response to these varying selection pressures. Evolutionary theory predicts differentiation in the level of phenotypic plasticity among different habitats. We evaluated temperature-induced phenotypic responses in juvenile growth rate in natural populations of the springtail Orchesella cincta , inhabiting forest and heathland. These habitats typically co-occur but differ strongly with respect to, for example, thermal regime, relative humidity, and structure. Offspring of females from the two habitats were reared at different temperatures in climate rooms and the temperature response of juvenile growth rate and egg size was measured. We found a habitat-specific difference in plasticity of juvenile growth rate. The reaction norms of the forest populations were steeper than the reaction norms for heath populations at two replicated sampling sites. Egg weight itself was demonstrated to be a plastic trait with a higher egg weight at low temperatures, but the thermal response did not differ between habitats. We conclude that these populations have diverged due to strong local natural selection. Our results support the argument that the level of phenotypic plasticity itself can be under selection and that differentiation in reaction norms can occur even in neighbouring habitats with no barrier to gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 265–271.     

Adaptive spatiotemporal distribution of soil microfungi in 'Evolution Canyon' II, Lower Nahal Keziv, western Upper Galilee, Israel

We describe and interpret spatiotemporal micromycete community structure and adaptive complexes in contrasting xeric and mesic microclimates in the soils of 'Evolution Canyon' II, Lower Nahal Keziv, western Upper Galilee, Israel. A total of 192 species from 60 genera belonging to Zygomycota (nine species), Ascomycota (13 species), and mitosporic fungi (170 species) were isolated. The fungal communities on the south-facing, xeric, 'African' slope (AS) demonstrated significantly greater diversity than on the north-facing, mesic, 'European' slope (ES) and the valley bottom (VB). Seasonally, winter slope communities were less heterogeneous. Forest localities on the ES and the VB in all seasons and the shady localities on the AS in the winter were overwhelmingly dominated by mesophilic Penicillium species. The sunny locality on the AS was characterized by a dominance of melanin-containing micromycetes that was most pronounced in the summer and by high occurrence and abundance of thermotolerant and thermophilic Aspergillus and Fusarium species. Ascomycetes and zygomycetes were the minor components in all local mycobiota studied; sexual ascomycetes, being stress-selected fungi, were more than ten times more abundant in the soil of the AS than in that of the ES, with the peak of abundance in the sunny summer community. The results demonstrated a microscale adaptive spatiotemporal inter- and intraslope divergence in soil mycobiota structure. Microclimatic natural selection appears to be the major factor affecting soil fungus diversity patterns.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78, 527–539.     

Metabolic rate and endurance capacity in Australian varanid lizards (Squamata: Varanidae: Varanus)

In ecomorphological and ecophysiological studies, locomotor performance is often considered to be an intermediate step between the form of an organism and its environment. We examined this premise by measuring morphology, physiology and circular track endurance in the closely related group of Australian varanid lizards. Body size, body mass and relative body proportions were poor indicators of endurance. Body mass was not correlated with endurance and size-free lower forelimb length had only a weak relationship with endurance. Instead, maximal metabolic rate was positively correlated with endurance capacity in varanids. A comparison of varanids with other groups of lizards supported this result as varanids showed both elevated maximal metabolic rate and elevated endurance scores when compared with similar sized non-varanid lizards. There was support for a strong association between endurance with foraging mode and climate. Varanid species with higher endurance tended to be widely foraging and from xeric climates, while sit-and-wait and mesic species showed reduced endurance.   © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 664–676.     

Markedly discordant mitochondrial DNA and allozyme phylogenies of tube-nosed fruit bats, Nyctimene, at the Australian–Oriental biogeographical interface

Australo-Papuan tube-nosed fruit bats of the genus Nyctimene reach their most westerly distribution on the island archipelagos of eastern Indonesia. A recent morphological examination indicates three species occur on Moluccan islands. Both allozyme electrophoresis and mitochondrial DNA (mtDNA) sequencing reveal there are only two species present, Nyctimene cephalotes and Nyctimene albiventer , but there is considerable disparity between the results obtained from the two genetic data sets. Allozyme data indicate N. albiventer occurs on Wokam, which sits on the Australian continental shelf and was joined to Australo-Papua during the last glacial maximum, and N. cephalotes on the other Moluccan islands, all of which are off the continental shelf. Divergence of these two species is dated at approximately 1.5 Mya. By contrast, the mtDNA gene tree shows two deep clades, one containing all specimens from Wokam and Yamdena, and the second all the specimens from the other islands. This especially marked incongruence between the two genetic data sets is ascribed to either a single introgression event of N. albiventer mtDNA into N. cephalotes on Yamdena at around 100 000 ya or lineage sorting of very old mtDNA lineages that coalesce a rather long time before the speciation event. These results highlight that caution should be exercised when relying on mtDNA as barcodes in species taxonomy. The continental-associated N. albiventer on Wokam has higher levels of allozyme heterozygosity and mtDNA nucleotide diversity than the N. cephalotes populations occurring on the more remote islands, indicating that this colonizing species has experienced bottlenecks and/or a low effective population size since speciation.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 589–602.     

mtDNA variation and evaluation of phylogenetic relationships among karyotypically polymorphic populations of Microtus (Terricola) thomasi (Arvicolidae, Rodentia) from Greece

The Balkan endemic species Microtus (Terricola) thomasi – Thomas' vole – is widely distributed in the Greek mainland and presents regionally confined karyotypic forms. In order to test the significance of geographical isolation and/or karyological differences in microevolutionary processes at the intraspecific level we used restriction enzymes and partial sequencing to study the polymorphism of the mtDNA d -loop control region in ten Greek populations. An overall high rate of both inter- and intrapopulation polymorphism was observed. The mtDNA diversity was unevenly distributed so that most of the populations studied were very similar and closely related, and only a few of them were clearly differentiated from the others. The mtDNA analysis failed to reveal any clear variation pattern that could be correlated with karyotypic forms or geographical proximity of the populations studied. Instead, the mtDNA phylogeny gave evidence in support of the coexistence of non-polymorphic (2 n  = 44) and polymorphic (2 n  < 44) karyotypic forms within the ancestral Greek M. (T.) thomasi stock. Interestingly, only one population (Agios Stefanos) seemed to significantly diverge from all the others in both methods used, forming a well-supported paraphyletic clade. Allozymic studies have already reported similar results, corroborating the genetic peculiarity of this population.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 55–68.     

Response patterns of net photosynthesis to moisture of mosses in xeric habitats

The response patterns of net photosynthesis to moisture level of mosses in xeric habitats were compared with those in mesic habitats, in order to determine whether the former species are better adapted to the xeric condition with regard to carbon gain. Moss species examined wereRhacomitrium lanuginosum andR. barbuloides in xeric open habitats andDicranum japonicum, Hypnum plicatulum, Ptilium crista-castrensis, Pleurozium schreberi andHylocomium splendens in mesic habitats on the coniferous forest floor in the upper subalpine zone of Mt. Fuji. Three additional xerophytic species collected at other localities,Ptychomitrium polyphylloides, Grimmia pilifera andHedwigia ciliata, were also examined. Five species in the xeric habitats showed an optimum range of moisture level for net photosynthesis, 2 to 3g·g⁻¹. On the other hand, species in the forest showed a wider optimum range, 3 to 8g·g⁻¹. Net photosynthetic rate at the moisture level of 0.5g·g⁻¹ was positive in xerophytic mosses, but negative in most forest mosses. Moisture levels where external capillary water disappeared and drop of water potential began was determined by blotting water-saturated shoots with membrane filters. These moisture levels were low in the xerophytic mosses and high in the forest mosses, although there were some exceptions. It was concluded that mosses in xeric habitats are better adapted for the efficient use of water for photosynthesis than those in mesic habitats.     

Phylogenetic relationships and biogeographical history of the genus Rhinoclemmys Fitzinger, 1835 and the monophyly of the turtle family Geoemydidae (Testudines: Testudinoidea)

Rhinoclemmys is an interesting genus of turtles biogeographically and ecologically, being the only genus of the family Geoemydidae that occurs in the New World and inhabiting a wide range of habitats from aquatic to highly terrestrial. Here we present a molecular phylogeny of Rhinoclemmys using both mitochondrial and nuclear genes. Our results strongly support the monophyletic and subfamilial status of Rhinoclemmys within the monophyletic family Geoemydidae. Within Rhinoclemmys , two clades are strongly supported, i.e. R. annulata  +  R. pulcherrima and R. areolata  +  R. punctularia  +  R. diademata  +  R. funerea  +  R. melanosterna , but the positions of R. nasuta and R. rubida are still weakly supported. In terms of the biogeographical history, the results of this study, coupled with palaeontological evidence, corroborate the hypothesis that this group migrated from Asia to the Americas across the Bering Strait during the early Eocene. The radiation of Rhinoclemmys in Central and South America corresponds well with vicariance events, including the emergence of the Sierra Madres of Mexico and the Nuclear Highland, and dispersals across the Panama land bridge. Interestingly, our resulting phylogeny suggests this group invaded South America at least four times and that dispersal of R. nasuta to South America probably took place in the early Miocene before the emergence of the Isthmus of Panama. We finally discuss our phylogenetic results with regard to the monophyly of the family Geoemydidae and in the context of previous morphological analyses. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 751–767.     

A new species of Cryptolepis (Apocynaceae: Periplocoideae) from Kenya and Tanzania, Africa

A new species of Cryptolepis is described from Kenya and Tanzania. It is a tropical forest climber and its unique second corona whorl of deeply cleft lobes distinguishes it from all other Cryptolepis species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 343–346.     

Reticulate evolution: ancient introgression of the Adriatic brown trout mtDNA in softmouth trout Salmo obtusirostris (Teleostei: Salmonidae)

Two populations of softmouth trout ( Salmo obtusirostris ) from the rivers Neretva (Bosnia and Herzegovina) and Jadro (Croatia), along with two neighbouring populations of brown trout ( Salmo trutta ) were analysed with a suite of genetic markers (two mtDNA genes, two nuclear genes, and nine microsatellites) as well as morphological characters. The Jadro softmouth trout were fixed for a brown trout mtDNA haplotype of the Adriatic lineage, which is 1.7% divergent from a previously described haplotype characteristic for the Neretva softmouth trout. All other genetic markers, as well as morphological analysis, supported the clear distinction of softmouth trout from the rivers Neretva and Jadro from brown trout in neighbouring populations, and thus a mtDNA capture event is assumed. Population specific microsatellite allele profiles, as well as a high number of private alleles for both populations of softmouth trout, support the hybridization between brown trout and the Jadro softmouth trout most likely being of ancient origin, thus leading to a reticulate evolutionary pattern of mtDNA in this taxon.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 139–152.     

A comprehensive phylogeny of the bumble bees (Bombus)

Bumble bees ( Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear to be similar in morphology throughout their range, suggesting that behavioural adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspecific variation in colour pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographical distribution and the evolution of behavioural and morphological adaptations, including colour pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data, we report the first nearly complete species phylogeny of bumble bees, including most of the 250 known species from the 38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1α, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology-based subgenera. Most subgenera fall into two distinct clades ( short-faced and long-faced ) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one-quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a firm foundation for reclassification and for evaluating character evolution in the bumble bees.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 161–188.     

Habitat-associated morphological divergence in two Neotropical fish species

We examined intraspecific morphological diversification between river channel and lagoon habitats for two Neotropical fish ( Bryconops caudomaculatus , Characidae; Biotodoma wavrini , Cichlidae). We hypothesized that differences between habitats (e.g. flow regime, foraging opportunities) might create selective pressures resulting in morphological divergence between conspecific populations. We collected fish from four channel-lagoon habitat pairs in the Río Cinaruco, Venezuela, and compared body morphology using geometric morphometrics. There were two aspects of divergence in both species: (1) placement of maximum body depth and (2) orientation of the mouth. For both species, maximum body depth was positioned more anteriorly (i.e. fusiform) in the river channel than in lagoons. Both species exhibited a relatively terminal mouth in lagoons compared to the channel. The mouth of B. caudomaculatus was relatively upturned, whereas the mouth of B. wavrini was relatively subterminal, in channel habitats. Observed morphological patterns are consistent with functional morphological principles suggesting adaptive divergence. We also show that spatial distance between habitats, presumably reflecting rates of population mixing, appears to have constrained diversification. For both species, morphological divergence increased with distance between habitats. Thus morphological divergence between channel and lagoon habitats apparently reflects a balance between diversification driven by natural selection, and homogenization driven by population mixing.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 689–698.     

A new species of Ceratozamia (Zamiaceae) from Oaxaca, Mexico with comments on habitat and relationships

Ceratozamia chimalapensis sp. nov. is described and illustrated. It is related to C. mirandae Vovides, Pérez-Farrera & Iglesias from Chiapas, but differs in trunk and peduncle size as well as in diameter of both megastrobili and microstrobili. Petiole, megasporophyll and indument colour also differ from that of C. mirandae . Ceratozamia chimalapensis forms part of the C. norstogii D.W.Stev. species complex, a group of ceratozamias with narrow leaflets growing in the herbaceous layer of oak forests in southern Mexico. These forests were severely affected by forest fires during 1998 and we recommend an IUCN Red List Category of CR B.1. Speciation in Ceratozamia has been discussed in the light of floristic refugia.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 169–175.     

Sympatric occurrence of diverged mtDNA lineages of Pityogenes chalcographus (Coleoptera, Scolytinae) in Europe

Differentiation among the European populations of the six-toothed spruce bark beetle Pityogenes chalcographus was observed for the first time in the 1970s as mating males from Northern Europe with females from Central Europe led to a significant decline in fecundity. Morphological examination revealed that P. chalcographus can be separated into two European races. Here, we investigated the genetic background of this separation by analysing 39 populations ( n  = 695), sequencing almost the complete cytochrome oxidase I gene (1543 bp) and applying single-strand conformation polymorphism. Phylogenetic analysis of 58 haplotypes yielded three major clades with a maximum sequence divergence of 2.33%, indicating that the demographic events took place in the late Pleistocene. These results support the hypothesis of allopatric divergence of the mtDNA lineages, which postglacially came into sympatric existence in Europe. However, as a result of partial crossing incompatibility the diverged lineages retained their genetic identity during postglacial times.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 331–340.     

Python phylogenetics: inference from morphology and mitochondrial DNA

We used nucleotide sequences from four mitochondrial genes and structural features of the mitochondrial control region, combined with a revised, previously published, morphological data set to infer phylogenetic relationships among the pythons. We aimed to determine which of two competing hypotheses of relationships of the genera Aspidites and Python best explains the evolutionary and bioegeographical history of the family. All analyses of the combined data recover a set of relationships in which (1) the genus Python is paraphyletic with the two east Asian species, P. reticulatus and P. timoriensis , as the sister lineage to the seven Australo-Papuan python genera. We support recognition of a distinct genus for the P. reticulatus  +  P. timoriensis clade; (2) the remaining species of the genus Python form a clade which is the sister lineage to the remainder of the family; (3) the genus Aspidites is embedded among the Australo-Papuan genera. The seemingly primitive characteristics of Aspidites may be better interpreted as reversals or specializations that have accompanied a switch to burrowing in this genus. Resolution of the relationships among the Australo-Papuan lineages is weak, possibly because of rapid diversification early in the history of the radiation. We assessed the tempo of the Indo-Australian python radiation using a maximum likelihood framework based on the birth–death process. We find strong support for elevated speciation rates during the period when Australia collided with the proto-Indonesian archipelago. The data support an origin for pythons outside Australia, followed by a radiation into Australia during the mid-Tertiary.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 603–619.