The adaptive significance of sexually dimorphic scale rugosity in sea snakes

In terrestrial snakes, rugose scales are uncommon and (if they occur) generally are found on both sexes. In contrast, rugose scales are seen in most sea snakes, especially in males. Why has marine life favored this sex-specific elaboration of scale rugosity? We pose and test alternative hypotheses about the function of rugose scales in males of the turtle-headed sea snake (Emydocephalus annulatus) and conclude that multiple selective forces have been involved. First, rugosities may aid male positioning during courtship, because histology shows that tubercles are more highly innervated than adjacent flat areas of each scale and hence are presumably more sensitive to tactile cues, and because biomechanical tests show that rugosities enhance friction between the bodies of males and females. Second, the occurrence of rugosities over the entire body of males and (albeit less well developed) in females as well suggests that rugosities also play a hydrodynamic role by modifying water flow across the snake's surface. Flow tank tests show that rugosities reduce the thickness of the boundary layer by almost 50% and create turbulent flow that should massively enhance rates of cutaneous oxygen uptake and hence prolong maximal courtship duration by males.     

The origin of snakes (Serpentes) as seen through eye anatomy

Snakes evolved from lizards but have dramatically different eyes. These differences are cited widely as compelling evidence that snakes had fossorial and nocturnal ancestors. Their eyes, however, also exhibit similarities to those of aquatic vertebrates. We used a comparative analysis of ophthalmic data among vertebrate taxa to evaluate alternative hypotheses concerning the ecological origin of the distinctive features of the eyes of snakes. In parsimony and phenetic analyses, eye and orbital characters retrieved groupings more consistent with ecological adaptation rather than accepted phylogenetic relationships. Fossorial lizards and mammals cluster together, whereas snakes are widely separated from these taxa and instead cluster with primitively aquatic vertebrates. This indicates that the eyes of snakes most closely resemble those of aquatic vertebrates, and suggests that the early evolution of snakes occurred in aquatic environments.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 469–482.     

Length–mass allometry in snakes

Body size and body shape are tightly related to an animal's physiology, ecology and life history, and, as such, play a major role in understanding ecological and evolutionary phenomena. Because organisms have different shapes, only a uniform proxy of size, such as mass, may be suitable for comparisons between taxa. Unfortunately, snake masses are rarely reported in the literature. On the basis of 423 species of snakes in 10 families, we developed clade‐specific equations for the estimation of snake masses from snout–vent lengths and total lengths. We found that snout–vent lengths predict masses better than total lengths. By examining the effects of phylogeny, as well as ecological and life history traits on the relationship between mass and length, we found that viviparous species are heavier than oviparous species, and diurnal species are heavier than nocturnal species. Furthermore, microhabitat preferences profoundly influence body shape: arboreal snakes are lighter than terrestrial snakes, whereas aquatic snakes are heavier than terrestrial snakes of a similar length. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Effects of oceanic salinity on body condition in sea snakes

Since the transition from terrestrial to marine environments poses strong osmoregulatory and energetic challenges, temporal and spatial fluctuations in oceanic salinity might influence salt and water balance (and hence, body condition) in marine tetrapods. We assessed the effects of salinity on three species of sea snakes studied by mark-recapture in coral-reef habitats in the Neo-Caledonian Lagoon. These three species include one fully aquatic hydrophiine (Emydocephalus annulatus), one primarily aquatic laticaudine (Laticauda laticaudata), and one frequently terrestrial laticaudine (Laticauda saintgironsi). We explored how oceanic salinity affected the snakes' body condition across various temporal and spatial scales relevant to each species' ecology, using linear mixed models and multimodel inference. Mean annual salinity exerted a consistent and negative effect on the body condition of all three snake species. The most terrestrial taxon (L. saintgironsi) was sensitive to salinity over a short temporal scale, corresponding to the duration of a typical marine foraging trip for this species. In contrast, links between oceanic salinity and body condition in the fully aquatic E. annulatus and the highly aquatic L. laticaudata were strongest at a long-term (annual) scale. The sophisticated salt-excreting systems of sea snakes allow them to exploit marine environments, but do not completely overcome the osmoregulatory challenges posed by oceanic conditions. Future studies could usefully explore such effects in other secondarily marine taxa such as seabirds, turtles, and marine mammals.     

A synopsis of Goniothalamus species (Annonaceae) in Peninsular Malaysia, with a description of a new species

The species of Goniothalamus (Annonaceae) occurring in Peninsular Malaysia and Singapore are revised, updating the previous taxonomic treatment by James Sinclair, published in 1955. A total of 18 species is recognized, including a new species, G. tomentosus . Collections referrable to G. tomentosus were previously determined as ' G. marcanii '; examination of the types of G. marcanii reveals that the two taxa are not conspecific, however, and a new name is accordingly validated here. Goniothalamus tomentosus has closest affinities with the Javan/Sumatran species G. costulatus . Other important nomenclatural changes include the reduction of G. umbrosus to synonymy with G. tapis .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 142 , 321−339.     

Chromosome counts in the genus Festuca section Eskia (Poaceae) in the Iberian Peninsula

Chromosome numbers of taxa of Festuca L. section Eskia Willk. in the Iberian Peninsula are given. The levels of ploidy for five taxa are confirmed. Idiograms and karyotypic formulae of the five taxa are presented for the first time. Two levels of ploidy occur in this section: diploid and tetraploid. One taxon, Festuca elegans ssp. merinoi is tetraploid and two other taxa have diploid and tetraploid populations. The remaining two taxa are solely diploid.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 331–337.     

Correlated evolution of aquatic prey-capture strategies in European and American natricine snakes

The evolution of aquatic prey-capture strategies in snakes has been suggested as a model system for the study of convergence. However, hypotheses of correlated evolution of prey-capture strategy with different aspects of foraging niche have never been tested quantitatively. Whereas a considerable amount of data is available for North American species, data for European species are scarce. In this study we combine original data on prey-capture strategies and strike velocities for European natricines with data for North American Natricinae obtained from the literature. We did not find any evidence for correlated evolution between prey-capture strategy and strike velocity with diet, but there was a significant correlation with prey density. Thus, our study suggests that prey density, rather than diet, played an important role in the evolution of the different prey-capture strategies and strike velocities of natricine snakes.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 73–83.     

The evolution of venom-delivery systems in snakes

The Colubroidea represents approximately 2300 of the 2700 species of living snakes and includes all venomous taxa. Although many morphological studies of colubroid snakes have been carried over the last hundred years, the phylogenetic relationships within this group are poorly known. In this study, components of the venom-delivery system (VDS) were examined within the context of two conflicting phylogenetic hypotheses proposed in 1988 by Cadle and in 1998 by Kraus & Braun. The results suggest that several major morphological changes occurred early in colubroid evolution: a Duvernoy's gland evolved, the posterior maxillary teeth became specialized relative to the anterior maxillary teeth, and the attachment of the pterygoideus muscle moved forward to a position associated with the posterior maxillary teeth. These innovations may have allowed the great radiation of colubroid snakes that led to the Colubroidea representing such a large percentage of living snakes. More recently, three separate lineages of colubroids have independently evolved highly specialized front-fanged VDSs with large and complex venom glands, venom gland compressor muscles, and tubular fangs.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 137 , 337−354.     

Morphological, vocal and genetic divergence in the Cettia acanthizoides complex (Aves: Cettiidae)

We used morphological, vocal and molecular (one mitochondrial and two nuclear loci) data to re-evaluate the taxonomic status of the taxa acanthizoides , concolor , and brunnescens in the Cettia acanthizoides (J. Verreaux, 1871) complex. We conclude that all three are valid taxa, and that acanthizoides of China and concolor of Taiwan are best treated as conspecific, whereas brunnescens of the Himalayas is better considered as a separate species. The degree of morphological, vocal, and genetic differentiation is variably congruent among all taxa; the recently separated acanthizoides and concolor differ slightly in plumage and structure but are indistinguishable in vocalizations, whereas the earlier diverged brunnescens and acanthizoides/concolor differ only slightly more in morphology but to a much greater degree in vocalizations. We stress the essential nature of taxonomic revisions as a prerequisite for the biodiversity estimates required for conservation planning.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 437–452.     

Relationships between floral characters, pollination mechanisms, life forms, and habitats in Araceae

The floral traits of the inflorescences of angiosperms have coevolved to ensure and maximize pollination success. Other factors believed to influence floral architecture are external (for example, ecological) to the inflorescence. In order to understand the relationships between such factors and floral characters, 12 floral traits were measured in 54 species of Araceae. An analysis was performed to determine how these traits are linked to the following: (1) self-pollination capacity; (2) life form (evergreen versus seasonally dormant); (3) climatic conditions; and (4) type of pollinator (i.e. flies, bees, or beetles). A significant difference was found between the pollen to ovule ratio of the species able to self-pollinate and those unable to self-pollinate. Evergreen and tropical aroids produced a larger number of gametes than did seasonally dormant and temperate taxa. Finally, several floral traits, such as pollen volume and number, number of female flowers, and flower sexual type (unisexual or bisexual), showed clear differences between the three pollinator types. Variations in floral traits between the different life forms and climatic conditions are discussed with respect to pollination efficiency and properties of the growing season. The pollen to ovule ratio cannot be considered as an accurate indicator of breeding systems in aroids because of the particular pollination ecology of the family.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 29–42.     

Morphological and ecological convergence in two natricine snakes

Similar morphologies between species may be due to shared ancestry or convergent evolution . Understanding instances of morphological and ecological convergence is central to evolutionary ecology because they help us understand the fit between organism and environment. Two species of stream-dwelling natricine snakes, Thamnophis rufipunctatus and Nerodia harteri present a model system for studying ecological and morphological convergence and adaptation. The species are allopatric and both live in shallow riffles in streams and forage visually for fish. We studied morphological similarity, trait evolution and functional significance of ecologically relevant traits in these and related species, and used mitochondrial DNA sequences for the ND4 gene to estimate their phylogenetic relationships. Character mapping of head length and head width supported the hypothesis of independent evolution of head shape in T .  rufipunctatus and N .  harteri . The elongate snout is a derived trait in these two taxa that is associated with reduced hydrodynamic drag on the snakes' heads when in a swift current, compared to other species with the ancestral blunt snout. We hypothesize that lower hydrodynamic drag facilitates prey capture success in these species that are known to forage by holding their position in currents and striking at fish prey. The elongate snout morphology has also resulted in a diminished binocular vision field in these snakes, contrary to the hypothesis that visually orientated snakes should exhibit relatively greater binocular vision. Convergent evolution of the long snout and reduced hydrodynamic drag in T. rufipunctatus and N. harteri are consistent with the hypothesis that the long snout is an adaptation to foraging in a swift current.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 363–371.     

Not so rare snakes: a revision of the Geophis sieboldi group (Colubridae: Dipsadinae) in lower Central America and Colombia

The Geophis sieboldi species group is composed of 16 currently recognized species distributed from Mexico to Colombia. Within this group, snakes of populations referred to the Geophis brachycephalus complex of lower Central America and Colombia display a remarkable polychromatism and the systematic status of these and other populations is problematic. The present study provides an analysis, including multivariate techniques, of variation in scalation, coloration, relative tail length and hemipenes to clarify the specific allocation of the populations belonging to this clade. Our results confirm the validity of three previously described taxa, namely G. brachycephalus , G. nigroalbus and G. talamancae , with reassignments of several populations previously referred to G. brachycephalus . In addition we recognize as a new species a suite of western Panama Geophis previously of uncertain status. We further provide a review of all other members of the G. sieboldi group in lower Central America and Colombia based on material obtained since the last revision of the group. Basic synonymies, diagnostics and known distributions are included for the treated taxa. Dietary guild, possible venomous coral snake mimicry and distributional anomalies for the group are discussed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 561–599.     

Mitochondrial DNA sequences of the Afro-Arabian spiny-tailed lizards (genus Uromastyx; family Agamidae): phylogenetic analyses and evolution of gene arrangements

Approximately 1.7 kbp of mitochondrial DNA were sequenced from 29 individuals assignable to 11 Uromastyx species or subspecies and two other agamids. U. ocellata and U. ornata had an insertion between the glutamine and isoleucine tRNA genes, which could be folded into a stable stem-and-loop structure, and the insertion for U. ornata additionally retained a sequence similar to the glutamine tRNA gene. This corroborates the role of tandem duplication in reshaping mitochondrial gene arrangements, and supports the idea that the origin of light-strand replication could be relocated within mitochondrial genomes. Molecular phylogeny from different tree-building methods consistently placed African and Arabian taxa in mutually monophyletic groups, excluding U. hardwickii inhabiting India and Pakistan. Unlike previous studies based on morphology , U. macfadyeni did not cluster with morphologically similar Arabian taxa, suggesting convergent evolution to be responsible for the morphological similarities. Divergence times estimated among the Uromastyx taxa, together with geological and palaeontological evidence, suggest that the Uromastyx agamids originated from Central Asia during the Eocene and colonized Africa after its connection with Eurasia in the early Miocene. Their radiation may have been facilitated by repeated aridification of North Africa since the middle Miocene, and geological events such as the expansion of the Red Sea and the East African Rift Valley.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 247–260.     

Chromosome numbers in the tribe Anthemideae (Asteraceae) from north-east Anatolia

Twenty-two chromosome counts of 19 taxa in the tribe Anthemideae of the family Asteraceae are reported from north-east Anatolia, Turkey. The taxa belong to the subtribes Achilleinae (four Achillea taxa), Anthemidinae (five Anthemis taxa), Artemisiinae (two Artemisia species), Leucantheminae (one Lecanthemum species), Matricariinae (two Tripleurospermum species) and Tanacetinae (five Tanacetum taxa). Six counts are new reports, seven are not consistent with previous counts, and the remainder are confirmations of very limited previous data. Most of the populations of Anthemideae studied have the basic number x  = 9, with ploidy levels ranging from 2 x to 7 to 8 x , but dysploidy is also present, with one diploid species, Artemisia austriaca , having x  = 8.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 203–211.     

Nomenclatural changes in preparation for a World Rubiaceae Checklist

During the preparation of a World Rubiaceae Checklist , numerous unplaced taxa were encountered, including illegitimate and invalid names, and species for which generic placement is uncertain. In this contribution, 35 new combinations and 20 new names are proposed, and the names of three taxa are validated.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 115–124.     

Sexual size dimorphism and sex ratios in dragonflies (Odonata)

Sexual size dimorphism and biased sex ratios are common in animals. Rensch's rule states that sexual size dimorphism (SSD) would increase with body size in taxa where males are larger than females and decrease with body size in taxa where females are larger. We tested this trend in dragonflies (Odonata) by analysing body size of 21 species and found support for Rensch's rule. The increase in SSD with increasing size among species can be explained by sexual selection favouring large males. We also estimated the slope of the relationship between sex ratio and size ratio in populations of the 21 species. A negative slope would suggest that the larger sex suffers from high mortality in the larval stage, consistent with riskier foraging. The slope of this relationship was negative, but after correcting for phylogentic non-independence with independent contrasts the relationship was no longer statistically significant, perhaps because of phylogenic inertia or low sample size.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 507–513.     

Determinants of feeding performance in free-ranging pit-vipers (Viperidae: Ovophis okinavensis): key roles for head size and body temperature

Feeding performance (handling time, capture success) in numerous animal species is well known to be influenced by a variety of ecological, functional, and physiological factors. Nonetheless, few studies have tested which factors are the strongest determinants of animal feeding performance in the wild. Using a field-based experiment, we examined the relationships among a number of functionally important variables and the predatory behaviour of free-ranging pit-vipers ( Ovophis okinavensis ) from Okinawa Island, Japan. Our main findings were: (1) strike latency was negatively related to snake body temperature and, hence, hotter snakes struck at frogs more readily than colder snakes; (2) initial bite position was correlated with ingestion direction (headfirst versus hindfirst) but ingestion direction was not correlated with ingestion duration; and (3) both snake head length and body temperature were negatively related with ingestion duration and, thus, snakes with longer heads and higher body temperatures had shorter ingestion durations. In O. okinavensis , head size and body temperature are therefore likely to have direct ecological consequences in terms of its feeding rate on explosively breeding frogs. More generally, however, this field-based study adds to the growing body of literature demonstrating that temperature has a pervasive influence on the feeding performance of ectotherms in general.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 53–62.     

Polyploidy in Aconitum subgenus Lycoctonum (Ranunculaceae)

Chromosome numbers and morphology in 22 populations belonging to 11 taxa of Aconitum subgenus Lycoctonum (Ranunculaceae) from China were studied. Some taxa were diploid, with 2 n  = 16, but four species and two varieties were found to be tetraploid, with 2 n  = 32. They are concentrated in the Hengduan Mountains region in south-west China, indicating that polyploidy could have played an important role in the speciation of the subgenus in this region, one of the areas of the world with a high concentration of endemic species. The relationships of some of the species are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 343–353.     

Karyotype differentiation between Koelreuteria bipinnata and K. elegans ssp. formosana (Sapindaceae) based on chromosome banding patterns

Koelreuteria Laxm., a genus in the tribe Koelreuterieae (Sapindaceae), includes three tree species that are widely recognized as having horticultural merit. The two more closely related species, K. bipinnata Franch. and K. elegans (Seem.) A. C. Sm., are easily distinguished from K. paniculata Laxm. by their bipinnate leaves. In this study, both species were investigated cytogenetically and their karyotypes and heterochromatic patterns were compared. Koelreuteria bipinnata and K. elegans ssp. formosana (Hayata) F. G. Mey. both have 2 n  = 32 but their karyotypes present slight morphological differences when observed using conventional staining. Chromosome banding patterns are reported for the first time for this genus. Both species exhibit terminal heterochromatic blocks, as revealed by C-Giemsa and C-CMA₃, but the band size varies between the species. Koelreuteria bipinnata has larger heterochromatic blocks and more GC-rich segments, while in K. elegans ssp. formosana these bands are smaller. The relationship between the karyotype features in these closely related species is discussed.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 451–455.     

Leaf teeth in certain Salicaceae and 'Flacourtiaceae'

The leaf teeth glands in four taxa from Salicaceae and six from 'Flacourtiaceae' were examined using both light and scanning electron microscopes. There appears to be a progression from glands of simple structure in the flacourtiaceous taxa and a tendency to a more complicated development in morphology and anatomy of the salicaceous species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 241–256.