Scale variation in a laboratory colony of amelanistic diamondback rattlesnakes (Crotalus atrox)

Inbreeding for 6 generations has produced a strain of amelanistic western diamondback rattlesnakes (Crotalus atrox) with extremely variable scalation. Forty-four siblings varying from virtually no body scalation to normal scalation have been produced in the latest generation. Two of the 3 most extreme “scaleless” snakes were stillborn; the third was maintained for > 1 year. Two of 7 snakes with greatly reduced head and body scales have died; however, the remaining 5 are being maintained and appear to be growing and healthy at 5 or 6 years of age. All snakes with reduced body scales have abnormal ventral scutes. Fifteen snakes with predominantly normal body scales have anomalous head scales, head scale patterns, and ventral scutes. One snake has predominantly normal scalation except for aberrant ventrals. A total of 18 Generation VI snakes have been classified as “normal,” although all snakes were not closely examined before disposition. Nine of the more normally scaled generation VI snakes are also being maintained in the laboratory. It is difficult to separate the genetic and environmental components of these phenotypes with existing information; however, it seems apparent that more than a single locus is involved. © 1994 Wiley-Liss, Inc.     

Adaptive significance of food income in European snakes: body size is related to prey energetics

Feeding strategies and diet patterns have been extensively investigated in vertebrates and, more specifically, in snakes. Although it has been hypothesized that prey species may differ in terms of energy content, almost no theoretical or practical study has been carried out to determine actual nutritional values of the common prey types of wild snakes. Our model taxa were a selection of widely distributed and well known European snake species, which have all been studied in depth: approximately 76% of their diet is composed of mammals, reptiles, and insects. We therefore selected a single model species for each of these categories and proceeded with the analyses. Nutritional values were determined using a standard procedure: lizards and mice were richer in proteins than insects (crickets); insects and mice were richer in lipids than lizards, and mice and crickets have a higher energy content than lizards; lizards were rich in ashes. We then applied our experimental results to a selected sample of European terrestrial snakes (11 populations, ten species, seven genera, two families) characterized by different body size (50–160 cm total length) and reproductive strategies (oviparous versus viviparous), aiming to correlate these parameters with patterns of energy income. A direct relationship was found between body mass/body length ratio (BCI, body condition index) and meal energetics: the higher the BCI, the higher was the metabolic requirement, whereas BCI was independent of species or of reproductive system effect. Large‐sized snakes thus need a highly diversified and more energy‐rich diet than smaller snakes, supporting previous hypotheses. The simple applicability of this method could be of valuable support in further comparative research work, reducing experimental costs and stimulating further ecological, behavioural, and, possibly, phylogenetic comparisons. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 307–317.     

Phylogeny,evolutionary morphology,and hemipenis descriptions of the Middle American jumping pitvipers (Serpentes: Crotalinae: Atropoides)

The jumping pitvipers, genus Atropoides, occur at low to middle elevations throughout Middle America. Recent molecular phylogenetic analyses have included all six species of Atropoides, but only two studies have found Atropoides to be monophyletic and questions persist about relationships within the A. nummifer complex. In this study, our phylogenetic analyses of morphological data provide strong support for the monophyly of Atropoides and recover relationships within the genus that are mostly congruent with those of recent molecular studies, further supporting the evolutionary and biogeographic hypotheses proposed in those studies. Our analyses find support for a sister relationship between A. picadoi and the other Atropoides species and an A. occiduus–A. indomitus clade sister to an A. nummifer–A. mexicanus–A. olmec clade. Within the A. nummifer complex, we find A. mexicanus and A. olmec to be sister species to the exclusion of A. nummifer. We include morphological synapomorphies to support each clade within Atropoides and describe and illustrate the hemipenes of each species. In addition, we discuss the importance of morphological phylogenetics and the functionality and limitations of hemipenial data in systematics.     

Establishing species' environmental requirements to understand how the southernmost species of South American pitvipers (Bothrops,Viperidae) are distributed: A niche‐based modelling approach

Three Bothrops species are known to be present along an extensive and generally xeric band of the south‐eastern portion of South America. Yet, the environmental factors responsible for the maintenance of the structure of this community have remained undetermined. To have a better understanding of snakes' geographic range limits in Argentina, we modelled the ecological niche of Bothrops alternatus, B. ammodytoides and B. diporus to identify areas of their occurrence and sympatry. We used snakes presence records and pseudo‐absence data with topographic (altitude and slope), vegetation and 19 bioclimatic variables as predictors. These variables were evaluated for each species, using Multiple Logistic Regression Analysis. The distribution of B. alternatus was correlated with temperature, precipitation and vegetation; that of B. ammodytoides with altitude, slope, precipitation, temperature and vegetation; and that of B. diporus related only to precipitation and temperature variables. Five variables appeared in all three models (i.e. annual precipitation, precipitation seasonality, precipitation of the driest month, mean diurnal temperature range and temperature range), although with different effects in different species. Probable sympatry areas were identified, for B. alternatus – B. ammodytoides in a narrow area, for B. diporus – B. alternatus in a wide area and B. ammodytoides – B. diporus in a small area. While some factors constrain the distribution of the southernmost pitvipers community in South America to spatially non‐overlapping ranges, others facilitate sympatry by allowing spatial coexistence. We found high sympatry between phylogenetically distantly related pitvipers. Niche‐based Geographic Information Systems modelling allowed us to identify the environmental factors correlated with each pitviper range in Argentina and in its contact zones, and to understand species distributional limits.     

On the congruence of morphology,trophic ecology,and phylogeny in Neotropical treeboas (Squamata: Boidae: Corallus)

Nine members of the Neotropical treeboa genus Corallus occur from Guatemala to south‐eastern Brazil and recent studies have provided an inconclusive picture about the relationship between morphology and trophic ecology in these snakes. To construct a more complete picture, we conducted the first study of morphology and diet to consider all nine species. Using adult specimens from museum collections, we examined several morphometric and meristic variables and their possible relationship to Corallus diets. Broadly, we found three basic morphologies within the genus: a short, narrow head and a slender body (C. cookii, C. grenadensis, C. hortulanus, and C. ruschenbergerii), useful for exploiting a wide variety of prey; a relatively stout body with a long, wide head (C. batesii, C. caninus, and C. cropanii) associated with feeding on larger mammalian prey; and an intermediate morphology, found in C. annulatus and C. blombergii, which may be indicative of endotherm generalists. These morphological and dietary patterns exhibit a strong degree of congruence with a recent molecular phylogeny of Corallus and highlight a heretofore unexamined ecological diversification within Corallus. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109, 466–475.     

Plastid genome evolution of a monophyletic group in the subtribe Lauriineae (Laureae,Lauraceae)

Litsea, a non-monophyletic group of the tribe Laureae (Lauraceae), plays important roles in the tropical and subtropical forests of Asia, Australia, Central and North America, and the islands of the Pacific. However, intergeneric relationships between Litsea and Laurus, Lindera, Parasassafras and Sinosassafras of the tribe Laureae remain unresolved. In this study, we present phylogenetic analyses of seven newly sequenced Litsea plastomes, together with 47 Laureae plastomes obtained from public databases, representing six genera of the Laureae. Our results highlight two highly supported monophyletic groups of Litsea taxa. One is composed of 16 Litsea taxa and two Lindera taxa. The 18 plastomes of these taxa were further compared for their gene structure, codon usage, contraction and expansion of inverted repeats, sequence repeats, divergence hotspots, and gene evolution. The complete plastome size of newly sequenced taxa varied between 152,377 bp (Litsea auriculata) and 154,117 bp (Litsea pierrei). Seven of the 16 Litsea plastomes have a pair of insertions in the IRa (trnL-trnH) and IRb (ycf2) regions. The 18 plastomes of Litsea and Lindera taxa exhibit similar gene features, codon usage, oligonucleotide repeats, and inverted repeat dynamics. The codons with the highest frequency among these taxa favored A/T endings and each of these plastomes had nine divergence hotspots, which are located in the same regions. We also identified six protein coding genes (accD, ndhJ, rbcL, rpoC2, ycf1 and ycf2) under positive selection in Litsea; these genes may play important roles in adaptation of Litsea species to various environments.     

Functional correlation between habitat use and leg morphology in birds (Aves)

Many of the morphological features of animals are considered to be adaptations to the habitat that the animals utilize. The habitats utilized by birds vary, perhaps more than for any other group of vertebrates. Here, we study possible adaptations in the morphology of the skeletal elements of the hind limbs to the habitat of birds. Measurements of the lengths of the femur, tibiotarsus and tarsometatarsus of 323 bird species from 74 families are used together with body mass data, taken from the literature. The species are separated into six habitat groups on the basis of literature data on leg use. A discriminant analysis of the groups based on leg morphology shows that swimming birds, wading birds and ground living species are more easily identified than other birds. Furthermore, functional predictions are made for each group based on ecological and mechanical considerations. The groups were tested for deviation from the norm for all birds for three indices of size- and leg-length-independent measures of the bones and for a size-independent-index of leg length. Several of the groups deviate significantly from the norm for one or more of the indices used, suggesting habitat-related adaptations in the leg morphology of birds. The results indicate that stability is an important factor affecting the leg morphology of primarily long-legged birds. The femur seems to be more important than previously thought because several of the groups have high femur indices, suggesting a positive selection pressure on this bone. On a general basis, the results suggest that the effect of leg length should be taken into consideration when discussing adaptations of mass-independent lengths of the long bones of the legs of birds.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 461–484.     

A complex biogeographic history of diversification in Neotropical lancehead pitvipers (Serpentes,Viperidae)

Based on the literature, we had predicted that the diversification within the Neotropical snake genus Bothrops occurred along a latitudinal gradient from north to south, with diversification into unoccupied niches through ecological opportunity, not correlated with geoclimatic events. Using a dated phylogeny and estimating likelihoods of ancestral states at cladogenesis events, we reconstructed ancestral areas and assessed major events of the diversification of Bothrops clades, and we also discuss systematic implications for this group. Based on the phylogeny we produced, B. lojanus was not considered as part of the genus Bothrops since the results recovered this species nested within the Bothrocophias clade. We infer that the diversification of the Miocene Bothrops pictus and Bothrops alternatus clades may be related to the uplift of the western slopes of the Andes and the Argentinian Patagonian Andes, respectively. The Pliocene Bothrops taeniatus and Bothrops osbornei clades may be related to the uplift of the eastern and northern Andes, respectively. The Plio-Pleistocene Bothrops neuwiedi clade may be related to the habitat transitions from a warmer and forested environment to a cooler and open landscape; the Bothrops jararaca (i.e. island endemic species) and Bothrops lanceolatus clades to over-water dispersal with island speciation; and Bothrops atrox clade to the appearance of the Panamanian land bridge. We found that a multitemporal and multidirectional history of diversification may be correlated with geoclimatic and dispersalist events. We argue that the vacant niche hypothesis by itself does not explain Bothrops diversification.     

Evolution of opercle bone shape along a macrohabitat gradient: species identification using mtDNA and geometric morphometric analyses in neotropical sea catfishes (Ariidae)

Transitions between the marine and freshwater macrohabitat have occurred repeatedly in the evolution of teleost fishes. For example, ariid catfishes have moved from freshwater to marine environments, and vice versa. Opercles, a skeletal feature that has been shown to change during such transitions, were subjected to 2D geometric morphometric analyses in order to investigate evolutionary shape changes during habitat transition in ariid catfishes and to test the influence of habitat on shape changes. A mtDNA marker, which proved useful in previous studies, was used to verify species identities. It greatly improved the assignment of specimens to a species, which are difficult to assign by morphology alone. The application of a mtDNA marker confirmed the occurrence of Notarius biffi in Central America, South of El Salvador. Molecular identification together with principal component analysis (PCA) and further morphological inspection of neurocrania indicated the existence of a cryptic species within Bagre pinnimaculatus. Principal component (PC) scores of individual specimens clustered in morphospace by genus rather than by habitat. Strong phylogenetic structure was detected using a permutation test of PC scores of species means on a phylogenetic tree. Calculation of Pagel's λ suggested that opercle shape evolved according to a Brownian model of evolution. Yet canonical variate analysis (CVA) conducted on the habitat groups showed significant differences in opercle shapes among freshwater and marine species. Overall, opercle shape in tropical American Ariidae appears to be phylogenetically constrained. This verifies the application of opercle shape as a taxonomic tool for species identification in fossil ariid catfishes. At the same time, adaptation to freshwater habitats shows characteristic opercle shape trajectories in ariid catfishes, which might be used to detect habitat preferences in fossils.     

Ontogenetic Diet Shifts of a Neotropical Catfish, Pimelodus maculatus (Siluriformes, Pimelodidae): An Ecomorphological Approach

This study analyses the ontogenetic diet shifts of a Neotropical catfish, Pimelodus maculatus (Siluriformes, Pimelodidae), from an ecomorphological standpoint. We collected 241 individuals in the Piracicaba River (Brazil) and, in the laboratory, seven morphometric variables were recorded from each specimen: standard length (SL), body depth, head length, snout length, eye diameter, mouth height and mouth width. After standardizing these measurements (as a proportion of SL), linear regressions were run to determine whether their growth was isometric or allometric in relation to SL. The diet analysis shows that the main food item for fish in the smaller size classes is aquatic insects, while the largest individuals feed mainly on other fish. As the great majority of the morphometric variables analyzed showed isometric growth as a function of SL – which means that these fish do not change their shape significantly during their lives – the ontogenetic diet shifts may be seen as a consequence of the absolute size increment of their mouth gape, which allows individuals to maximize their energetic gain by ingesting larger prey with a higher caloric content.     

Early development of the Neotropical fish known as long sardine Triportheus auritus (Valenciennes 1850) (Characiformes,Triportheidae)

Larvae and juveniles of long sardine, Triportheus auritus, from the lower Amazon river was described, evaluating ontogenetic changes in their external mor­phology, pigmentation, fin development, morphometry, and meristics. A total of 93 individuals, 83 larvae and 10 juveniles were analyzed, they were captured monthly between 2014 and 2019 in the Amazon river channel and in macrophytes aquatic stands in the alluvial plains located in the lower Amazon River. From each specimen, morphometric and meristic data were measured and then the growth pattern between morphometric variables was analyzed. The larvae have an elongated body in a fusiform shape, superior mouth, simple nostril, pigmented spherical eyes and long intestine, surpassing the median region of the body. Initial pigmentation is scarce, but intensifies through development forming a pattern composed of three longitudinal bands concentrated in the ventral, cephalo-dorsal and lateral line regions. There are also pigments in the mandible, surrounding the mouth, under the swim bladder, intestine and fins. The sequence of complete fin formation is: caudal, anal, dorsal, pectoral and pelvic. The total number of myomeres ranged from 45 to 48 (25–29 preanal and 17–22 postanal). Morphometric relationships indicated differential growth for measurable morphometric parameters, with abrupt growth of snout length, head length and body height in the transition from flexion to postflexion stages. The pre-dorsal distance showed a decrease in the growth rate at the threshold from the larval to the juvenile period. The pre-pectoral and pre-anal distances showed negative allometric growth. In conclusion, the combination of body shape pigmentation pattern and, the formation sequence of fins allow the identification of the genus and coupled with the number of myomeres, morphometric relationships, and ray numbers of the anal fin ensure the differentiation of T. auritus from the other congeneric species. The metamorphosis occurred mainly at the end of the larval period and it is related to changes in the physiological and ecomorphological characteristics of the species.     

Foraging strategies,craniodental traits,and interaction in the bite force of Neotropical frugivorous bats (Phyllostomidae: Stenodermatinae)

1. Bats in the family Phyllostomidae exhibit great diversity in skull size and morphology that reflects the degree of resource division and ecological overlap in the group. In particular, the subfamily Stenodermatinae has high morphological diversification associated with cranial and mandibular traits that are associated with the ability to consume the full range of available fruits (soft and hard).
2. We analyzed craniodental traits and their relationship to the bite force in 343 specimens distributed in seven species of stenodermatine bats with two foraging strategies: nomadic and sedentary frugivory. We evaluated 19 traits related to feeding and bite force in live animals by correcting bite force with body size.
3. We used a generalized linear model (GLM) and post hoc tests to determine possible relationships and differences between cranial traits, species, and sex. We also used Blomberg''s K to measure the phylogenetic signal and phylogenetic generalized least‐squares (PGLS) to ensure the phylogenetic independence of the traits.
4. We found that smaller nomadic species, A. anderseni and A. phaeotis , have a similar bite force to the large species A. planirostris and A. lituratus; furthermore, P. helleri registered a bite force similar to that of the sedentary bat, S. giannae. Our study determined that all the features of the mandible and most of the traits of the skull have a low phylogenetic signal. Through the PGLS, we found that the diet and several cranial features (mandibular toothrow length, dentary length, braincase breadth, mastoid breadth, greatest length of skull, condylo‐incisive length, and condylo‐canine length) determined bite force performance among Stenodermatiane.
5. Our results reinforce that skull size is a determining factor in the bite force, but also emphasize the importance of its relationships with morphology, ecology, and phylogeny of the species, which gives us a better understanding of the evolutionary adaptions of this highly diverse Neotropical bat group.

     

Relationship between body size and flying-related structures in Neotropical social wasps (Polistinae,Vespidae, Hymenoptera)

Body size influences wing shape and associated muscles in flying animals which is a conspicuous phenomenon in insects, given their wide range in body size. Despite the significance of this, to date, no detailed study has been conducted across a group of species with similar biology allowing a look at specific relationship between body size and flying structures. Neotropical social vespids are a model group to study this problem as they are strong predators that rely heavily on flight while exhibiting a wide range in body size. In this paper we describe the variation in both wing shape, as wing planform, and mesosoma muscle size along the body size gradient of the Neotropical social wasps and discuss the potential factors affecting these changes. Analyses of 56 species were conducted using geometric morphometrics for the wings and lineal morphometrics for the body; independent contrast method regressions were used to correct for the phylogenetic effect. Smaller vespid species exhibit rounded wings, veins that are more concentrated in the proximal region, larger stigmata and the mesosoma is proportionally larger than in larger species. Meanwhile, larger species have more elongated wings, more distally extended venation, smaller stigmata and a proportionally smaller mesosoma. The differences in wing shape and other traits could be related to differences in flight demands caused by smaller and larger body sizes. Species around the extremes of body size distribution may invest more in flight muscle mass than species of intermediate sizes.     

Microsatellite and major histocompatibility complex variation in an endangered rattlesnake,the Eastern Massasauga (Sistrurus catenatus)

Genetic diversity is fundamental to maintaining the long‐term viability of populations, yet reduced genetic variation is often associated with small, isolated populations. To examine the relationship between demography and genetic variation, variation at hypervariable loci (e.g., microsatellite DNA loci) is often measured. However, these loci are selectively neutral (or near neutral) and may not accurately reflect genomewide variation. Variation at functional trait loci, such as the major histocompatibility complex (MHC), can provide a better assessment of adaptive genetic variation in fragmented populations. We compared patterns of microsatellite and MHC variation across three Eastern Massasauga (Sistrurus catenatus) populations representing a gradient of demographic histories to assess the relative roles of natural selection and genetic drift. Using 454 deep amplicon sequencing, we identified 24 putatively functional MHC IIB exon 2 alleles belonging to a minimum of six loci. Analysis of synonymous and nonsynonymous substitution rates provided evidence of historical positive selection at the nucleotide level, and Tajima's D provided support for balancing selection in each population. As predicted, estimates of microsatellite allelic richness, observed, heterozygosity, and expected heterozygosity varied among populations in a pattern qualitatively consistent with demographic history and abundance. While MHC allelic richness at the population and individual levels revealed similar trends, MHC nucleotide diversity was unexpectedly high in the smallest population. Overall, these results suggest that genetic variation in the Eastern Massasauga populations in Illinois has been shaped by multiple evolutionary mechanisms. Thus, conservation efforts should consider both neutral and functional genetic variation when managing captive and wild Eastern Massasauga populations.     

Ejaculate characteristics over seasons in five species of lancehead pitvipers (Bothrops spp) kept in captivity

Due to their major medical importance in Latin America, lancehead pitvipers are frequently kept and bred in captivity for venom extraction to the production of antivenom serums. Nevertheless, despite the great contribution given to captive breeding, much of the knowledge of Bothrops' reproductive biology derived from sporadic and insufficient data provided by zoological collections. Thus, we aimed to investigate seasonal changes in gonadosomatic index (GSI) and seminal parameters (e.g., volume, concentration, motility, viability, and acrosome integrity) of five species of lancehead pitvipers from different biomes and phylogenetic groups, maintained in the indoors serpentarium at Butantan Institute (Brazil). Patterns of variation in GSI and semen parameters differed from one species to another, suggesting that captive populations should perhaps be managed distinctly to maximize reproductive success. Furthermore, in none of the studied species did changes in GSI occur concomitantly with seminal variations. GSI remained unaltered year-round for Jararaca (Bothrops jararaca) and Brazilian lancehead (Bothrops moojeni), whereas it peaked in the autumn for Common lancehead (Bothrops atrox), Jararacussu (Bothrops jararacussu), and Whitetail lancehead (Bothrops leucurus). But surprisingly, the scenario was inverted when we estimated the total number of motile spermatozoa per season, as Jararaca and Brazilian lancehead displayed seasonal differences and the other species did not vary throughout the year. Potential ecological and evolutionary factors underlying these differences were also discussed in the present article. Together, these findings can help to better define breeding management strategies for each species in captivity, in addition to optimizing the future use of artificial insemination and semen cryopreservation.     

The evolution of locomotor morphology, performance, and anti-predator behaviour among populations of Leiocephalus lizards from the Dominican Republic

Most ecomorphological studies use a comparative approach to examine adaptation by studying variation among species. A question of considerable interest is whether ecomorphological patterns observed among species also exist at the population level. We studied variation in morphology, performance, and behaviour in four populations of Leiocephalus personatus and two populations of Leiocephalus barahonensis in the Dominican Republic. We combined these data with measurements of predation intensity and habitat structure to test for convergence at the population level. We predicted that predation intensity would be higher in open habitats and that lizards in these habitats would have traits conferring higher predator evasion capacity (increased wariness, faster sprint speeds, and longer limbs). Principal components analysis suggests that sites tend to differ with respect to the abundance and spacing of low-lying vegetation (i.e. percentage of shrub cover and distance to nearest vegetation), but we did not detect any striking differences among sites in tail-break frequencies or attacks on clay lizard models. Consistent with predictions we find that in open habitats, lizards tend to have longer limbs, faster sprint speeds (relative to body size), and longer approach distances. These patterns corroborate findings in other ground-dwelling lizard species and indicate that they have evolved at least twice among populations of Leiocephalus lizards. The results of this study also suggest that these traits have evolved rapidly despite recent or ongoing gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 445–456.     

Comparative trophic morphology in eight species of damselfishes (Pomacentridae)

Damselfishes show significant biodiversity in the coral reefs. To better understand such diversity, an ecomorphological approach was investigated in the trophic morphology of eight species of Pomacentridae (Chromis acares, C. margaritifer, Dascyllus aruanus, D. flavicaudus, Pomacentrus pavo, Plectroglyphidodon johnstonianus, Pl. lacrymatus and Stegastes nigricans) belonging to different trophic guilds (zooplankton, algal, coral polyp feeders and omnivores). Geometric morphometrics were used to quantify size and shape variations in four skeletal units: (1) neurocranium, (2) suspensorium and opercle, (3) mandible and (4) premaxilla. This method allowed us to reveal shape and size differences correlated to functional diversity both within and between trophic guilds. Among zooplanktivores, C. margaritifer, D. aruanus and D. flavicaudus have a high and long supraoccipital crest, short mandibles forming a small mouth and high suspensoria and opercles. These three species can be considered to be suction feeders. In the same guild, C. acares shows opposite characteristics (long and thin mandibles, lengthened neurocranium and suspensorium) and can be considered as a ram feeder. Among herbivores and corallivores, the two species of Plectroglyphidodon and S. nigricans can be considered as grazers. Differences in skeletal shape are mainly related to improving the robustness of some skeletal parts (broad hyomandibular, short and high mandibles). The shapes of P. pavo, which feeds largely on algae, strongly differ from that of the other three grazers exhibiting similar morphological characteristics to C. acares (e.g., long and shallow suspensorium, lengthened neurocranium). This highlights likely differences concerning cutting or scraping method. Finally, no strong correlations exist between size and shapes in the eight studied species. Size difference among species having a very similar shape could be viewed as a factor optimizing resource partitioning.