Morphological evolution in relationship to sidewinding, arboreality and precipitation in snakes of the family Viperidae

Compared with other squamates, snakes have received relatively little ecomorphological investigation. We examined morphometric and meristic characters of vipers, in which both sidewinding locomotion and arboreality have evolved multiple times. We used phylogenetic comparative methods that account for intraspecific variation (measurement error models) to determine how morphology varied in relationship to body size, sidewinding, arboreality and mean annual precipitation (which we chose over other climate variables through model comparison). Some traits scaled isometrically; however, head dimensions were negatively allometric. Although we expected sidewinding specialists to have different body proportions and more vertebrae than non-sidewinding species, they did not differ significantly for any trait after correction for multiple comparisons. This result suggests that the mechanisms enabling sidewinding involve musculoskeletal morphology and/or motor control, that viper morphology is inherently conducive to sidewinding (‘pre-adapted’) or that behaviour has evolved faster than morphology. With body size as a covariate, arboreal vipers had long tails, narrow bodies and lateral compression, consistent with previous findings for other arboreal snakes, plus reduced posterior body tapering. Species from wetter environments tended to have longer tails, wider heads and reduced anterior tapering. This study adds to the growing evidence that, despite superficial simplicity, snakes have evolved various morphological specializations in relationship to behaviour and ecology.

Taxonomic revision of the Boiga ceylonensis group (Serpentes: Colubridae): re-examination of type specimens, redefinition of nominate taxa and an updated key

We conducted a taxonomic revision of the arboreal snakes of the Boiga ceylonensis-group, all inhabiting the Indian subcontinent. Based on 15 characters, recorded from more than 100 specimens, representing eight recognized taxa, we conducted a multivariate morphometric analysis to redefine the taxonomic boundaries. We provide redescriptions of the nominate taxa in this group based on re-examination of the name-bearing types. The types of B. ceylonensis, B. beddomei, B. barnesii, B. dightoni, B. nuchalis and B. andamanensis are redescribed, where appropriate lectotypes are selected, illustrated and their nomenclature, synonymy and distribution are discussed. Status of the nominate taxon Boiga ceylonensis dakhunensis Deraniyagala, 1955 is fixed as an objective junior synonym of Dipsas nuchalis Günther, 1875. An updated key to the species of this group is presented.

Arboreality constrains morphological evolution but not species diversification in vipers

An increase in ecological opportunities, either through changes in the environment or acquisition of new traits, is frequently associated with an increase in species and morphological diversification. However, it is possible that certain ecological settings might prevent lineages from diversifying. Arboreality evolved multiple times in vipers, making them ideal organisms for exploring how potentially new ecological opportunities affect their morphology and speciation regimes. Arboreal snakes are frequently suggested to have a very specialized morphology, and being too large, too small, too heavy, or having short tails might be challenging for them. Using trait-evolution models, we show that arboreal vipers are evolving towards intermediate body sizes, with longer tails and more slender bodies than terrestrial vipers. Arboreality strongly constrains body size and circumference evolution in vipers, while terrestrial lineages are evolving towards a broader range of morphological variants. Trait-dependent diversification models, however, suggest similar speciation rates between microhabitats. Thus, we show that arboreality might constrain morphological evolution but not necessarily affect the rates at which lineages generate new species.

Population ecology of a cryptic arboreal snake (Hoplocephalus bitorquatus)

Biologists have traditionally been reluctant to study arboreal snakes due to low rates of capture. Overlooking such taxa can mislead interpretations of population trends for data-deficient species. We used regularly spaced transect searches and standard capture–mark–recapture techniques to describe population structure, growth rates, survival and capture probability in a population of the pale-headed snake (Hoplocephalus bitorquatus) in southern Queensland. We obtained data from 194 captures of 113 individual snakes between 2009 and 2015. Using the Cormack–Jolly–Seber method, we estimated apparent annual survival in subadult snakes at 0.23 ± 0.01 (s.e.) and 0.81 ± 0.08 for adults. Capture probability was estimated at 0.16 ± 0.14 per session in subadult snakes and 0.33 ± 0.06 for adults. Within the red gum forests of our study site, we estimate pale-headed snake density at ~13 ± 7 ha–1. Using von Bertalanffy growth modelling, we predict that snakes reach sexual maturity after about four years and may live for up to 20. Our results suggest that the species is a ‘k’ strategist, characterised by slow maturation and low fecundity. These traits suggest that populations will recover slowly following decline, exacerbating the risk of local extinction.

Gripping during climbing of arboreal snakes may be safe but not economical

On the steep surfaces that are common in arboreal environments, many types of animals without claws or adhesive structures must use muscular force to generate sufficient normal force to prevent slipping and climb successfully. Unlike many limbed arboreal animals that have discrete gripping regions on the feet, the elongate bodies of snakes allow for considerable modulation of both the size and orientation of the gripping region. We quantified the gripping forces of snakes climbing a vertical cylinder to determine the extent to which their force production favoured economy or safety. Our sample included four boid species and one colubrid. Nearly all of the gripping forces that we observed for each snake exceeded our estimate of the minimum required, and snakes commonly produced more than three times the normal force required to support their body weight. This suggests that a large safety factor to avoid slipping and falling is more important than locomotor economy.