Morphological variation in the short-tailed fruit bat (Carollia) in Ecuador,with comments on the practical and philosophical aspects of boundaries among species

The mechanisms for the morphological delimitation of species in Carollia remain poorly understood. This is the first study to assess variation in size and shape from strictly geometric terms. Both factors are assessed by statistical perspectives of distribution, overlap and relative distances. Despite its overlap, the size of the skull seems to be the most influential character for the discrimination of species, with shape playing a much smaller role. The smallest species seems to be the most distinct in shape, not only in terms of distance among centroids in morphometric space, but also in the overall trend and direction of variation. Contrary to previous studies, sexual dimorphism is not given by size but by distinct shapes of the skull. Characters such as the shape of the maxilla, previously described qualitatively as discrete with sharp boundaries, appear to be truly continuous with fuzzy borders among species. Because morphometric space is a gamut of continuous variation and overlap, the taxonomic error rate for size characters seems to be substantial for the medium-sized species (Carollia brevicauda Schinz, 1821), with approximately 30–40% of individuals erroneously assigned to a different species after a jackknifed discriminant function. This taxonomic error is higher for shape characters. Morphological, systematic and ecological consequences of the observed patterns of shape and size variation are commented within the context of previously proposed arguments and hypotheses.     

Geographical variation of the skull morphology of the common tree shrew (Tupaia glis).

Geographical variation was examined morphologically in the common tree shrew (Tupaia glis) in some Indochinese and Malayan regions. Osteometrical examination and principal component analysis elucidated the morphological differences among various populations. The populations from southern and western Thailand were distinguished morphologically from the other populations. Variation in males from south Thailand and Kuala Lumpur suggests that the Isthmus of Kra may have an influence on the variation of skull morphology. However, the Isthmus of Kra was not completely considered as a factor of geographical separation in this species, because we could not confirm the separation in skull size and shape between the localities at least in females. While, the Kanchanaburi population in western Thailand was significantly smaller than the other population in skull size, and constituted the morphologically separable group in our study.     

Morphometric variation in the Tunisian green frog,Rana saharica Anura: Ranidae)

Rana saharica is the most widely distributed anuran in Tunisia. We examined morphological variation in 124 specimens as a function of their geographical origin, using univariate and multivariate statistics with traditional morphometrics. Our results supported the existence of three morphotypes of this species, correctly classified 84.7% of specimens, and indicated significant degrees of variation in morphometric characteristics between regions. No significant differences were found between sexes among populations. We observed an increase in body size and a decrease in body mass as a function of latitude. Specimens from the southernmost region had the smallest bodysize and the greatest morphometric divergence. The pattern of morphometric variation probably results from phenotypic plasticity correlated with local environmental factors.     

Size and Shape Variability in the Skull of the Bottlenose Dolphin,Tursiops truncatus (Montagu, 1821)

The bottlenose dolphin (Tursiops truncatus) is a widely spread cetacean species. We present a quantitative analysis of age dimorphism variation in the skull of T. truncatus assessed by geometric morphometrics (GM) methods. Differences in size and shape of skulls were investigated using eight landmarks plotted on 2‐D images of ventral views of 14 museum specimens. The results of GM revealed differences in size, but not in shape. The left side appears more variable, which is probably an effect of telescoping.     

Toxin variation among salamander populations: discussing potential causes and future directions

Amphibians produce defensive chemicals which provide protection against both predators and infections. Within species, populations can differ considerably in the composition and amount of these chemical defenses. Studying intraspecific variation in toxins and linking it to environmental variables may help us to identify the selective drivers of toxin evolution, such as predation pressure and infection risk. Recently, there has been a renewed interest in the unique toxins produced by salamanders from the genus Salamandra: the samandarines. Despite this attention, intraspecific variation has largely been ignored within Salamandra-species. The aim of this study was to investigate whether geographic variation in profiles of samandarines exists, by sampling 4 populations of Salamandra atra over its range in the Dinaric Alps. In addition, we preliminary explored whether potential variation could be explained by predation (counting the number of snake species) and infection risk (cultivation and genomic analyses of collected soil samples). Salamanders from the 4 populations differed in toxin composition and in the size of their poison glands, although not in overall toxin quantity. Nor predation nor infection risk could explain this variation, as populations barely differed in these variables. Sampling over a much broader geographic range, using better estimators for predation and infection risk, will contribute to an improved understanding of how environment may shape variation in chemical defenses. Nevertheless, as the 4 populations of S. atra did differ in their toxin profiles, we propose that this species provides an interesting opportunity for further ecological and evolutionary studies on amphibian toxins.     

Shape differences of the Carina sterni in birds of various locomotion types

The anatomy of the sternum in birds varies according to their habitats and type of locomotion. In particular, the carina sterni manifest different shape variations. In this study, the shape of the carina sterni was investigated by means of geometric morphometrics. Birds of different types of locomotion were used in the study: flying, swimming, and terrestrial. Ducks and chickens show a wider variety of shapes. Pigeons are the species with the least differences. The margo cranialis carinae in a turkey is the flattest compared to other species. In chickens, the apex carinae is more caudally than the base of the carina sterni. The margo cranialis of the carina sterni in ducks is concave. The differences in centroid size and shape differences between species collectively are statistically significant (p < 0.0001). The most distinct shape contrast is between the duck and turkey (p: 0.0003). Form differences between the ducks and geese as well as between the chicken and turkey are statistically insignificant. There is less variation in the shape of the carina sterni among avian species representing the same type of locomotion. Although there are many comparative morphological and morphometric studies of birds, shape analysis studies revealing the interspecific differences and similarities of the sternum are very limited. Morphology of the carina sterni can be useful in taxonomic investigations.     

Patterns of morphometric variation in the alpine newt (Mesotriton alpestris) at the southern limit of its distribution: environmental correlates

We applied multivariate analyses to an array of body measures of alpine newt specimens derived from 11 local populations in Greece to describe, analyse and detect patterns and putative causes of within‐population and among‐population morphometric variation. The observed morphometric variation was partitioned into several independently varying aspects of the external phenotype, frequently following variation patterns in different environmental factors. The size and features of the aquatic habitat were found to affect body size, while altitude was found to affect head‐shape variation in both sexes. At the intra‐population level, variation in generalized body size and shape was found to be significantly lower when competitive newt species were present in the habitat, indicating stabilizing selection towards a decrease in inter‐specific competition. No clear discrimination on body size and shape proportions was detected between the two genetic lineages examined, implying ecogenetic or environmentally induced variation rather than phylogeny.     

Unraveling the morphological patterns of a subantarctic eelpout: a geometric morphometric approach

Phenotypic variation in organisms depends on the genotype and the environmental constraints of the habitat that they exploit. Therefore, for marine species inhabiting contrasting aquatic conditions, it is expected to find covariation between the shape and its spatial distribution. We studied the morphology of the head and cephalic sensory canals of the eelpout Austrolycus depressiceps (4.5–22.5 cm TL) across its latitudinal distribution in South Pacific (45°S–55°S). Geometric morphometric analyses show that the shape varied from individuals with larger snout and an extended suborbital canal to individuals with shorter snouts and frontally compressed suborbital canal. There was size variation across the sampled populations, but that size does not have a clear latitudinal gradient. Only 8% of the shape variation relates to this size variation (allometry), represented by a decrease in the relative size of the eye, and a depression of the posterior margin of the head. There were spatial differences in the shape of the head, but these differences were probably caused by allometric effects. Additionally, 2 of the canals of the cephalic lateralis pores and the head shape showed modularity in its development. This study shows that the morphology of marine fish with a shallow distribution varies across distances of hundreds of kilometers (i.e., phenotypic modulation).     

Genetic diversity within two Tunisian wild jirds: Meriones shawi and Meriones libycus (Rodentia,Gerbillinae)

Three Meriones species inhabit Tunisia, namely M. shawi, M. libycus and M. crassus, but little genetic data exist on these gerbils. We collected Meriones from eight localities in Tunisia, and obtained mitochondrial (cytochrome b) and nuclear (IRBP) gene sequence data for 37 and 13 specimens, respectively, belonging to two species: M. shawi and M. libycus. We also optimised three microsatellite markers previously described in M. unguiculatus to obtain a finer analysis of their genetic diversity and geographic structure, given their wide distribution. Phylogenetic inferences of cyt b and IRBP data for these species, in the context of other gerbillin data, corroborate their taxonomic affinities reported by previous studies. High cyt b haplotype diversity was observed in both species (25 haplotypes in 29 and 27 sequences for M. shawi and M. libycus, respectively) with little geographical structure for M. shawi but three divergent groups in M. libycus. The average microsatellite diversity within each population was high (H_O ≥ 0.6, H_E ≥ 0.8) with M. libycus populations attaining the highest values. Population differentiation was moderate for several population pairs (F_st ≥ 0.1), the highest being between M. shawi populations. However, genetic distance among populations was not significantly correlated with geographic distance in either M. shawi or M. libycus. Our results contribute to a better characterisation of Tunisian Meriones species, suggesting high geographic structure in mtDNA of M. libycus populations within North Africa.     

Geographical variation of the skull of the lesser mouse deer

We examined the geographical variation of the skull size and shape of the lesser mouse deer (Tragulus javanicus) from Laos, Thailand, Peninsular Malaysia, Sumatra, Java, Borneo, Langkawi and some Islands of Tenasserim in Myanmar. Although the influence of the climatic condition on skull size was not confirmed in the mainland populations, the skull became rostro-caudally longer in the populations of Tenasserim and Sumatra because of island isolation effect. The skull size was classified into the following three clusters of localities from the matrix of Q-mode correlation coefficients: 1) Langkawi and Tenasserim, 2) Laos and Thailand, 3) Sumatra and Borneo. The skulls in the population of Java belong to the cluster of Langkawi and Tenasserim in male, however were morphologically similar to those in the cluster of Borneo and Sumatra. The canonical discriminant analysis pointed out that the Laos and Tenasserim populations were separated from the other ones and that the populations of Sumatra, Java and Borneo were intermingled each other.     

Morphologic,morphometric and contour shape variations of sagittal otoliths of Lepidorhombus spp. in the Aegean Sea

Otoliths are calcareous anatomical structures in the inner ear of fishes, and they can be used in the discrimination of fish species and stocks due to their species-specific shape. Sagittal otoliths in the Lepidorhombus whiffiagonis and Lepidorhombus boscii, two flatfish species of the same genus distributed in the Aegean Sea, were compared using morphology, morphometry and contour shape analyses. Blind and eyed side otoliths of Lepidorhombus species were evaluated separately due to their morphological features and statistical differences in morphometric measurements (p < 0.05). Four analysis groups were formed: blind side otolith of L. whiffiagonis, eyed side otolith of L. whiffiagonis, blind side otolith of L. boscii and eyed side otolith of L. boscii. Morphometric differences in otolith shape were performed by canonical discriminant analysis, and the first discriminant axis explained 97.4% (Wilks λ = 0.270) of the variance between the groups, and the second axis explained 2.3% (Wilks λ = 0.620). Classification success between Lepidorhombus species based on CDA is 100%, and the overall CDA classification score between groups is 76.5%. The wavelet functions obtained in the contour analysis showed high variability in the anterior, posterior-dorsal, antero-dorsal and ventral regions of the otoliths among these four analysis groups. Consequently, it is thought that this study will contribute to the taxonomic classification of fish with the morphological and morphometric differences in the blind and eyed side otoliths and the analyses made with the high discrimination success obtained in Lepidorhombus species.     

A morphological analysis of the skull size and shape of Kerivoulinae (Chiroptera: Vespertilionidae) from Vietnam

Southeast Asia is a region of high biodiversity, containing species of plants and animals that are yet to be discovered. In this region, bats of the subfamily Kerivoulinae are widespread and diverse with six species recorded in Vietnam. However, the taxonomy of the Kerivoulinae in Asia is complicated. In our study, we used diagnostic characters and multivariate analysis to determine morphological differences between the genera Kerivoula and Phoniscus. We showed that the two genera are distinguishable by the size of second upper incisors, the shape of skull, nasal sinus, canines, second upper and lower premolars. In addition, the two genera can be osteometrically separated by measurements of the braincase height, interorbital width and shape of anterior palatal emargination. Our data clearly revealed the morphological variations in the skull shape of Kerivoula hardwickii in Vietnam. This suggests a possible separation into three morphotypes, representing cryptic species supported by statistical differences with wide variation in skull shape, size and teeth. These results demonstrated Kerivoula hardwickii can be separated three subspecies, and the result will serve as the basis for the future assessment and classification of this group in Southeast Asia.     

Geographical clines in the size of the herb field mouse (Apodemus uralensis)

Patterns of body size variation along geographical gradients have long been searched for and generalized into eco‐geographical rules. However, no rodent species has yet been analyzed in relation to the 3 dimensions of latitude, longitude and altitude. We analyzed geographical clines and dimorphism of body and skull size in the herb field mouse (Apodemus uralensis) across the species range, based on field data and on data from the literature. Sexual dimorphism in adult A. uralensis was not expressed at a large scale, while local patterns were inconsistent. Age‐dependent size changes were most expressed in adult individuals: most characters of adults exceeded in size those of subadults, while subadult–juvenile size differences were only significant in body weight and length, zygomatic skull width, length of cranial diastema and breadth of braincase. Despite central morphological niches along the clines being separated, A. uralensis populations showed a high degree of size overlap in morphological space. We found the species to be characterized by high size variability, with the largest individuals inhabiting the eastern and southern edges of the distribution range. Tail, hind foot and ear lengths were largest in the southern part of the range, in agreement with Allen's rule. The main measurements that we analyzed, namely body mass, zygomatic skull width and condylobasal skull length, show the presence of 3 clines in the size of adult A. uralensis: (i) a decreasing south–north cline, opposing Bergmann's rule; (ii) an increasing west–east cline, in accordance with Murphy's rule; and (iii) an increasing altitudinal cline.     

Phenotypic plasticity is maintained despite geographical isolation in an African cichlid fish,Pseudocrenilabrus multicolor

Gene flow among populations in different selective environments should favor the evolution of phenotypic plasticity over local adaptation. Plasticity in development is a common response to long‐term hypoxia in some widespread African fishes, including Pseudocrenilabrus multicolor, a cichlid that exploits both normoxic (high oxygen) rivers/lakes and hypoxic (low oxygen) swamps. Previous studies have shown that fish from normoxic and hypoxic sites differ in many traits, including gill size, brain size and body shape, and that much of this variation reflects developmental plasticity. However, these earlier studies focused on areas in Uganda where gene flow between swamp and river or lake populations is high. In this study we tested the hypothesis that P. multicolor from a relatively isolated lake population (Lake Saka, Uganda) exhibit low levels of plasticity in traits related to oxygen uptake. Multiple broods of P. multicolor from Lake Saka were reared under low and high dissolved oxygen, and traits related to gill size, brain mass and body shape were quantified. Surprisingly, both gill size and brain mass showed high levels of developmental plasticity. We suggest that high levels of plasticity, particularly in the gill size of P. multicolor, reflects low costs of maintaining the plastic response, even in relatively isolated populations.     

Allozyme variation amongst populations of the freshwater crab,Potamonautes perlatus (Decapoda: Potamonautidae) in the Berg River system,Western Cape

The Cape river crab, Potamonautes perlatus, is widely distributed in streams and rivers of the Western Cape where it exhibits considerable morphological variation. Recent genetic work on populations in the Olifants River system has demonstrated the existence of a new cryptic species of river crab, while populations of P. perlatus in the nearby Berg River system remained uninvestigated. Six populations of P. perlatus were collected from along the length of the Berg River system (125 km) and the genetic structure was investigated using allozyme electrophoresis. Results from 14 allozyme loci showed that the populations were genetically invariant across the river system. It is suggested that gene flow may be responsible for the poor genetic differentiation amongst populations in the Berg River system. The implication for future management of the system is briefly discussed.     

Geographical variation of skull morphology and its functional significances in the red-cheeked squirrel

Skulls of the red-cheeked squirrel (Dremomys rufigenis) from various geographical locations: Malaysia (peninsular area), Vietnam (south district)-Laos, and Thailand (north district) were osteometrically examined. The skull size of the squirrels in the southern (Malaysia) population was fundamentally larger than that in the northern (Vietnam, Laos and Thailand) populations. The proportion indices indicated that the splanchnocranium was relatively longer in the Malaysia population, and that the interorbital space was narrower in Vietnam-Laos, and Thailand populations. We suggest that the long nose and laterally-oriented orbits in the skull may be better adapted for terrestrial-insectivorous life in the Malaysia population and the binocular sense facilitated by rostrally-oriented eyes contributes to the arboreal-fruit eating behavior in the two northern populations. The Malaysia population was clearly distinguished from the other populations by the principal component analysis. We suggest that the geographical barrier of the Isthmus of Kra influences the morphological variation of the skull among the squirrel populations.     

紫花苜蓿复合体的形态变异式样及分类学研究

紫花苜蓿复合体(Medicago sativa complex)是极为重要的牧草资源,其成员间具有明星的形态渐渡特征.为摸清其种质资源状况并明确其分类学问题,通过查阅馆藏腊叶标本和野外居群采样,观测了39个形态性状,并利用主成分分析和聚类分析的方法进行运算.结果表明,紫花苜蓿复合体的花器官有形成"大花类型"和"小花类型...     

Geographical variation in the skeletal morphology of red jungle fowl

1. The skulls and postcranial skeletons of the red jungle fowl (Gallus gallus) were compared osteometrically between the populations from North and South Vietnam, North and Central Laos and Southeast Bangladesh. The populations include the three subspecies of G. g. spadiceus, G. g. gallus and G. g. murghi and were sampled to reveal the geographical morphological variations among populations in G. gallus.

2. The morphometric characteristics of subspecies murghi could be clearly distinguished from those of the other subspecies using a canonical discriminant analysis. However, the size and shape of the skull of the gallus population from South Vietnam were not statistically different from that of the subspecies spadiceus from North Laos. The canonical discriminant scores also clearly indicated that there were morphological similarities in the skulls of the populations from North Laos and South Vietnam.

3. From the results, therefore, it is concluded that red jungle fowls do not exhibit high levels of osteometric variation between geographical localities at least within the Indochinese Peninsula.

4. This contrasts with previous studies which have described these subspecies as having various external morphological differences and have argued that zoogeographical barriers exist between the north and south areas of the Indochinese Peninsula.     

Geographical variation of skull size and shape in various populations in the black giant squirrel

We osteometrically examined the skulls of the black giant squirrel (Ratufa bicolor) from three mainland populations (M. Malayan Peninsula, V. South Vietnam, and B. Burma, India and North Thailand) and from two island populations (T. Tioman, and S. Sumatra Islands). The skull in the Malayan peninsula population was significantly smaller than that of the two other mainland populations. It is consistent with Bergmann's rule as shown in the gray-bellied squirrel. The two island populations did not show obvious differences in comparison with the Malayan population in many measurements. In the proportion analysis eliminating the size factor, the differences among populations were not easily confirmed and we concluded that the osteological characters peculiar to each population could not be shown in this species. The first and second principal component scores of M, S, and T populations were intermingled, whereas the V and B populations of V and B were not separated in the chart. We pointed out that the morphological differences were demonstrated between northern and southern groups of the Isthmus of Kra in the mainland populations, and that the two island populations did not show the island-isolation effect in comparison with the M population. The adaptational variation related to feeding and locomotion could not be confirmed among populations of the black giant squirrel as shown in the proportion analysis.     

Problems presented by geographical variation in the African vine snakes,genus Thelotornis

Analysis of variation in 771 specimens of the genus Thelotornis A. Smith (Colubridae) confirms the absence of intergradation between the taxa kirtlandii (Hallowell) and oatesii (Gunther) in northern Angola and southern Zaire, so 7. capensis A. Smith is accepted as a full species. The western race 7. capensis oatesii shows very little variation, but the eastern populations of 7. capensis are extremely variable and it is convenient to revive the name mossambicanus (Bocage) for the populations which occur from central Mocambique northwards. T. c. oatesii hybridizes extensively with T. c. capensis and 7. c. mossambicanus in southeastern Rhodesia and adjacent Mocambique. Northern populations of 7. c. mossambicanus show features of 7. kirtlandii, but these are apparently primitive characters retained since the original partition of Thelotornis into western and eastern forms. The highlands of Kenya and northern Tanzania isolate these northern mossambicanus from the nearest kirtlandii populations in Uganda.