Skull size and shape variation in Psammomys spp. (Rodentia,Gerbillinae) from Tunisia,with emphasis on the impact of allometric variation on species recognition

In Tunisia both Psammomys obesus and P. vexillaris are found. These taxa have been the subject of taxonomic controversy for some time, due to variability in the classical morphological characters used for taxonomic recognition. In this study we investigated skull size and shape variation in the genus Psammomys by using geometric morphometrics to evaluate the extent of intra- and interspecific shape variation and explicitly tested for the impact of allometric shape variation on species discrimination. Eleven populations of the two species from 10 localities in Tunisia were studied. Statistical analyses of size and shape showed large size variation within P. obesus, but no shape differences were revealed among populations of this species. Interspecific analysis revealed that P. vexillaris had the smallest skull. Principal component analysis and Procrustes distances showed good discrimination between the two species after removal of the allometric component of shape variation. The results obtained show that allometric-related shape variation could mask discrimination between the two Psammomys species. This finding might explain the uncertainty in classification of these species in the past. The interspecific allometric-free phenotypic differences observed may be associated with adaptive processes linked to the different environmental and trophic preferences of the two species.     

Comparative multivariate analysis of biometric traits of West African Dwarf and Red Sokoto goats

The population structure of 302 randomly selected West African Dwarf (WAD) and Red Sokoto (RS) goats was examined using multivariate morphometric analyses. This was to make the case for conservation, rational management and genetic improvement of these two most important Nigerian goat breeds. Fifteen morphometric measurements were made on each individual animal. RS goats were superior (P < 0.05) to the WAD for the body size and skeletal proportions investigated. The phenotypic variability between the two breeds was revealed by their mutual responses in the principal components. While four principal components were extracted for WAD goats, three components were obtained for their RS counterparts with variation in the loading traits of each component for each breed. The Mahalanobis distance of 72.28 indicated a high degree of spatial racial separation in morphology between the genotypes. The Ward’s option of the cluster analysis consolidated the morphometric distinctness of the two breeds. Application of selective breeding to genetic improvement would benefit from the detected phenotypic differentiation. Other implications for management and conservation of the goats are highlighted.     

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.     

Rapid divergent coevolution of Sinopotamon freshwater crab genitalia facilitates a burst of species diversification

One of the most striking radiations in brachyuran evolution is the considerable morphological diversification of the external reproductive structures of primary freshwater crabs: the male first gonopod (G1) and the female vulva (FV). However, the lack of quantitative studies, especially the lack of data on female genitalia, has seriously limited our understanding of genital evolution in these lineages. Here we examined 69 species of the large Chinese potamid freshwater crab genus Sinopotamon Bott, 1967 (more than 80% of the described species). We used a landmark‐based geometric morphometric approach to analyze variation in the shape of the G1 and FV, and to compare the relative degree of variability of the genitalia with non‐reproductive structures (the third maxillipeds). We found rapid divergent evolution of the genitalia among species of Sinopotamon when compared to non‐reproductive traits. In addition, the reconstruction of ancestral groundplans, together with plotting analyses, indicated that the FV show the most rapid divergence, and that changes in FV traits correlate with changes in G1 traits. Here we provide new evidence for coevolution between the male and female external genitalia of Sinopotamon that has likely contributed to rapid divergent evolution and an associated burst of speciation in this lineage.     

Foetal Growth Curves and Seasonal Breeding in the Natal Clinging Bat Miniopterus Schreibersi Natalensis

Growth of the foetus of Miniopterus schreibersi natalensis is described. The gestation period in this subspecies is 240 days of which 120 days is the period of delayed implantation. Birth mass is taken as 2,7 g from measurements of near-term foetuses. Foetal growth follows a typical J-shaped curve during the last half of the gestation period. Young of M. s. natalensis are proportionately lighter at birth than many other mammals.     

No evidence for avoidance of black rat scent by the presumably less competitive Natal multimammate mouse in a choice experiment

In Africa, indigenous multimammate mice (Mastomys natalensis) only appear to live commensally in houses when invasive black rats (Rattus rattus) are absent, yet little is known about the underlying mechanism. Avoidance through smell may cause the absence of M. natalensis from areas occupied by R. rattus, but this hypothesis has not yet been tested. We conducted a Y-maze choice experiment where 15 M. natalensis were offered a choice between corridors containing conspecific scent, R. rattus scent and a control scent. Residence time in the R. rattus corridor was greater than that in the control corridor but equal to that in the M. natalensis corridor, suggesting that multimammate mice do not actively avoid the scent of their invasive competitor.     

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.     

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.     

Geometric morphometric study of the skull shape diversification in Sciuridae (Mammalia,Rodentia)

It is generally accepted that the high phenotypic diversity of mammals is a combined result of developmental constraint and ecological adaptation, although the influence of these endogenous and exogenous factors varies in different mammal groups. The rodent family Sciuridae represents an ideal candidate for examining phenotypic diversity in relation to phylogeny and ecological adaptations. In the present study, we investigate the effects of phylogeny and lifestyle on the skull shape in different species of Sciuridae by applying geometric morphometric methods. In addition, we investigate the importance of allometry on sciurid skull shape, because results from geometric morphometrics sometimes dispute those of traditional morphometry. Here, we identify significant associations between patristic distances obtained from molecular phylogeny and shape distances in all 3 views of the cranium and the lateral view of the mandible. Multivariate regression demonstrates that shape differences among lifestyle categories are substantial, especially in the dorsal and ventral structures after the influence of phylogeny is taken into account. Allometry plays an important role in the shape variation, although its importance on different skull structures varies. Our results indicate that complex structures of this highly diverse mammal group, which occupies different niches, are affected by ecological factors and developmental constraint.     

Combined approaches to identify genomic regions involved in phenotypic differentiation between low divergent breeds: Application in Sardinian sheep populations

Selective breeding has led to modifications in the genome of many livestock breeds. In this study, we identified the genomic regions that may explain some of the phenotypic differences between two closely related breeds from Sardinia. A total of 44 animals, 20 Sardinian Ancestral Black (SAB) and 24 Sardinian White (SW), were genotyped using the Illumina Ovine 50K array. A total of 68, 38 and 15 significant markers were identified using the case–control genome‐wide association study (GWAS), the Bayesian population differentiation analysis (F_ST) and the Rsb metric, respectively. Comparisons among the approaches revealed a total of 22 overlapping markers between GWAS and F_ST and one marker between GWAS and Rsb. Three markers detected by Rsb were also located near (<2 Mb) to highly significant regions identified by GWAS and F_ST analyses. Moreover, one candidate marker identified by GWAS and F_ST approaches was located in a run of homozygosity island that was shared by both breeds. We identified several genes involved in many phenotypic differences (such as stature and growth, reproduction, ear size, coat colour, behaviour) between the two analysed breeds. This study shows that combining several genome‐wide approaches could improve discovery of regions involved in the variability of breeding traits and responsible for the phenotypic diversity even between closely related breeds. Overall, the combination of such genome‐wide methods can be extended to other livestock breeds that share between them a similar genetic background, to understand the process that shapes the patterns of genetic variability between closely related populations.     

Milk protein polymorphisms in cattle (Bos indicus), mithun (Bos frontalis) and yak (Bos grunniens) breeds and their hybrids indigenous to Bhutan

In the current study, milk protein variation was examined in cattle (Bos indicus), mithun (Bos frontalis), yak (Bos grunniens) and their hybrid populations in Bhutan to estimate genetic variability, conduct genetic characterization and assess the possibility of gene flow between mithun and cattle. Isoelectric focusing of 372 milk samples from 11 populations detected four molecular types of β‐lactoglobulin (A, B, E and M), five molecular types of α_S1‐casein (A, B, C, E and X) and three molecular types of k‐casein (A, B and X). Mithun and yak shared alleles but were found to exhibit different allele frequencies for the proteins studied. The degree of genetic variability within populations was measured by average heterozygosity and ranged from 24–40% in cattle, 26% for yak and 33% for mithun. We also resolved the traditional mithun and cattle hybridization system via principal component analysis. Our results suggested secondary introgression of mithun genes to the village Thrabum population, and a close genetic relationship between Bhutanese indigenous cattle and Indian cattle.     

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.     

Preventing species invasion: A role for integrative taxonomy?

Integrative taxonomy, a multi‐disciplinary approach adding modern techniques to traditional morphology‐based methods (e.g. molecular and morphological criteria), can play an important role in bioinvasion research to identify introduced taxa, discover pathways of introduction and inform authorities to control and prevent future introductions. The present study is the first on introduced populations of Callosciurus, Asiatic tree squirrels, known as potentially invasive species in Europe (Italy, Belgium and France). We combined molecular (mitochondrial DNA markers: CoxI, D‐loop) and morphometric analysis on skulls, comparing them to the widest morphological and molecular datasets ever assembled for Callosciurus. Squirrels collected in Italy and Belgium share the same haplotypes and skull characteristics, but are conspicuously different from the French population in Antibes. Genetic data revealed close similarity between French squirrels and Pallas's squirrels, Callosciurus erythraeus, from Taiwan, China. Italian and Belgian squirrels formed an independent taxonomic lineage in genetic analyses, whose taxonomic rank needs further investigation. The morphological and morphometric characteristics of these 2 populations are, however, similar to known specimens assigned to Callosciurus erythraeus. These results may indicate a common origin for the populations found in Belgium and Italy. In contrast, French specimens suggest an independent introduction event of squirrels originating from Asia.     

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).     

Genome‐wide association studies for two exterior traits in Chinese Dongxiang spotted pigs

Pigs show extensive variation in exterior appearance. This variation has been explored as one of the selection targets to form breeding features in the pig industry. In this study, a customized Affymetrix Axiom Pig1.4M array plate was used to conduct genome‐wide association studies (GWAS) for two exterior traits—spotted coat color (SC) and facial type (FT)—in Chinese Dongxiang spotted pigs. Two and 12 single nucleotide polymorphisms (SNPs) were identified to be associated with SC and FT at the 5% genome‐wide significant level, respectively. Of these SNPs, two associated with SC were located around the DACH1 gene on Sus Scrofa chromosome (SSC)11. Eleven SNPs associated with FT were located within a 7.68 Mb region (29.03–36.71 Mb) on SSC7. Based on the GWAS results and the biological functions of genes, we highlight EDNRB as a candidate gene for SC and HMGA1 and RPS10 as the potential genes affecting facial variation. The findings contribute to the final characterization of causative genes and mutations underlying the effects of the SSC7 and SSC11 loci and improve our understanding of the genetic basis of phenotypic variation in Chinese indigenous pigs.     

Differences in diet between two rodent species,Mastomys natalensis and Gerbilliscus vicinus,in fallow land habitats in central Tanzania

Differences in the ecological niche requirements among rodent species competing in the same habitat may result from differences in the use of one to three resources: space, time and food or some combination of these. Alternatively, differences in resource use utilization among animal species may simply reflect availability of food, and when food is limited, different animal species compete. In this study, the diet of two rodent pest species, Mastomys natalensis and Gerbilliscus vicinus, coexisting in fallow land in central Tanzania were studied to assess the degree of diet differentiation among them. Dietary niche breadth of G. vicinus was greater than that of M. natalensis in all stages of the maize cropping seasons. The rodent species studied overlapped considerably in the food items consumed ranging from niche overlap (O_jk) of 0.77–0.89. Grains/seeds featured high in the diet of M. natalensis while plant material occurrence was high in G. vicinus. These two food categories may have contributed to differences in diet partitioning, which may, in turn, facilitate their coexistence in fallow land.     

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.     

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.     

Signatures of functional constraint at Fgfr1a Genes in schizothoracine fishes (Pisces: Cypriniformes): The dermal skeleton variation adapted to high‐altitude environments

Fishes harbor a huge resource of phenotypic diversity and are useful for understanding the genetic basis of morphological variation. However, it is difficult to transfer classical genetic mapping to most non‐model species. In this study, we performed a comparative sequence analysis of Fgfr1a to first interpret the evolution of this candidate scale‐loss gene in 15 schizothoracine fishes with various scale phenotypes. While considerable amino acid (AA) substitutions were observed, molecular evolution analysis indicates that the overall coding regions were subject to functional constraint. We also identified extra copies of Fgfr1a in 4 scale‐loss fishes and detected accelerated evolution in one AA substitution specific to these duplicates. We speculate that Fgfr1a had accumulated mutations in the ancestral lineage of scale‐loss schizothoracine fishes before experiencing duplication events, which was further followed by the diversification of species. In silico mutation analysis predicted deleterious effects of the mutations while no disruptive molecular mechanism was detected. Collectively, our results highlight the important role of Fgfr1a gene in the adaptive evolution of schizothoracine fishes during their radiation in the Qinghai‐Tibetan Plateau.     

Genetic structure in Atlantic brown trout (Salmo trutta L.) populations in the Iberian peninsula: evidence from mitochondrial and nuclear DNA analysis

Brown trout (Salmo trutta L.) was sampled in rivers belonging to three different Spanish basins in order to analyse the distribution of genetic variability. The genetic analysis was performed by using two systems and techniques: nuclear DNA was screened through random amplified polymorphic DNAs (screening 2 × 10⁵ bp of the whole genome), and mitochondrial DNA (mtDNA) through sequencing of the hypervariable control region. Genetic distances between the populations were similar using either analysis although some differences arise. For example, some populations of the Tajo basin were very close through nuclear analysis but more distant using mtDNA. Differences between the two DNA sources could be the result of a different evolutionary rate, and the fact that mtDNA is maternally transmitted and differences in sex migration rates will influence the patterns of genetic variation between the transmitted DNAs. Total variation was partitioned using amova showing a clear subdivision among basins although intrapopulation variation remained as high as 62%. A correspondence analysis defined the differences in a three‐dimensional way, clustering the populations according to their common basin. When mtDNA was sequenced, higher variability was noted in the segment between 400 and 600bp of the whole D‐loop sequence, suggesting that these 200bp improved the analysis of the variability more than sequencing the t‐RNA ends of the control region. A comparison was made between the t‐RNA^Pro ends of the 10 populations screened here and the rest of the published sequences found in the literature, leading to a concentration of these populations in group IV which includes all trouts which originate in the Atlantic. The analyses performed suggest that a high genetic variability is present in all populations and that although there has been a probable interference from stocked strains introduced to increase population density, this was only detectable through the variance between rivers which reflect different policies according to the region where the basin is located. However, the genetic analysis using the two approaches allows the control of the natural populations avoiding a loss of their genetic potential.