Intraspecific taxonomy and ecology characterize morphological divergence among cutthroat trout (Oncorhynchus clarkii ssp. Richardson) populations

We compared the proportion of morphological variation accounted for by subspecies categories with the proportion encompassed by ecologically based categories in cutthroat trout ( Oncorhynchus clarkii ssp.), as a means of assessing the relative importance of each approach in identifying intraspecific diversity. We used linear and geometric morphometrics to compare measures of body shape, fin length, and head features between and within subspecies of cutthroat trout. Both categories accounted for a significant proportion of the variation between and within the subspecies; however, the larger proportion was explained by subspecific differences, with the greatest morphological divergence between coastal cutthroat trout ( Oncorhynchus clarkii clarkii ) and interior subspecies. Ecotypic categories within each subspecies also explained significant morphological differences: stream populations had longer fins and deeper, more robust bodies than lake populations. The largest ecotypic differences occurred between stream and lake populations of Yellowstone cutthroat trout ( Oncorhynchus clarkii bouvieri ). Given that many cutthroat trout subspecies are of conservation concern, our study offers a better understanding of intraspecific variation existing within the species, providing precautionary evidence of incipient speciation, and a framework of describing phenotypic diversity that is correlated with ecological conditions.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 266–281.     

Ultraviolet reflectance and cryptic sexual dichromatism in the ocellated lizard, Lacerta (Timon) lepida (Squamata: Lacertidae)

Ultraviolet (UV) colorations have garnered extensive theoretical and empirical treatment in recent years, although the majority of studies have concerned themselves with avian taxa. However, many lizards have acute visual systems with retinal photoreceptors that are sensitive to UV wavelengths, and also display UV-reflecting colour patches. In the present study, we used UV photography and full-spectrum reflectance spectrophotometry to describe intra- and intersexual colour variation in adult ocellated lizards Lacerta ( Timon ) lepida and to obtain evidence of UV-based ornamentation. We also investigated whether any colour traits correlate with morphological traits potentially related to individual quality. The results obtained show that the prominent eyespots and blue outer ventral scales (OVS) that ocellated lizards have on their flanks reflect strongly in the UV range and are best described as UV/blue in coloration. The eyespots of males are larger and cover a larger surface area than those of females. However, these differences can be entirely accounted for by sex differences in body size, with males being generally larger than females. We also found differences in the shape of reflectance curves from males and females, with the eyespots and blue OVS of males being more UV-shifted than those of females. Other body regions have extremely low UV reflectance and are not sexually dichromatic. Eyespot size and the total surface area covered by eyespots increases with body size in males but not in females, suggesting that they may be signalling an intrinsic individual characteristic such as body size or male fighting ability. We also discuss the alternative and non-exclusive hypothesis that eyespots may function in lizards of both sexes as protective markings against predators.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 766–780.     

Discordance in body size, colour pattern, and advertisement call across genetically distinct populations in a Neotropical anuran (Dendropsophus ebraccatus)

Patterns of intraspecific geographic variation in morphology and behaviour, when examined in a phylogenetic context, can provide insight into the microevolutionary processes driving population divergence and ultimately speciation. In the present study, we quantified behavioural and phenotypic variation among populations from genetically divergent regions in the Central American treefrog, Dendropsophus ebraccatus . Our fine-scale population comparisons demonstrated regional divergence in body size, colour pattern frequencies, and male advertisement call. None of the characters covaried with phylogenetic history or geographic proximity among sampled populations, indicating the importance of highly localized selection pressures and genetic drift in shaping character divergence among isolated regions. The study underscores how multiple phenotypic characters can evolve independently across relatively small spatial scales.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 298–313.     

Genome size evolution and polyploidy in the Daphnia pulex complex (Cladocera: Daphniidae)

Genome size was estimated in 49 clones of the Daphnia pulex complex from temperate and subarctic locations using flow cytometry and microsatellite DNA analyses. Significant genome size differences were found in diploid species belonging to the two genetically distinct groups (the pulicaria and the tenebrosa groups), with clones from the tenebrosa group having genome sizes 22% larger than those in the pulicaria group. Combined flow cytometry and microsatellite DNA analyses revealed that nearly all polyploid clones in the D. pulex complex are triploid and not tetraploid, as was previously suggested. Sequencing analyses of the ND5 gene to position clones in their respective clades within the D. pulex complex have uncovered three triploid clones of Daphnia middendorffiana with a D. pulex maternal parent. This result was unexpected because Daphnia pulicaria has always been identified as the maternal parent of these hybrid polyploid clones. Triploid clones likely owe their origins to interactions between sexual and asexual populations. Further interactions in the tenebrosa group have generated tetraploid clones but these events have been rare.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 68–79.     

Intraspecific vicariant history and the evolution of adaptive morphological diversity in a fish species (Osmerus mordax)

Vicariant geographic isolation and resource partitioning have long been independently identified as processes contributing to the morphological divergence of closely-related species. However, little is known about the extent to which vicariant history influences the adaptive ecological divergence associated with resource partitioning and trophic specialization within species. The present study thus quantified the contribution of vicariant historical genetic divergence to the adaptive contemporary morphological divergence of intraspecific feeding specialists in the Rainbow smelt (Pisces: Osmerus mordax ). This species is characterized by the polyphyletic origin of two lacustrine feeding specialists originating in two intraspecific lineages associated with independent glacial refuges. The historical genetic segregation was initiated approximately 350 000 years ago, whereas the lacustrine trophic segregation arose within the past 10 000 years. Wild caught lacustrine smelt populations were grouped a priori based on known historical genetic identities (Acadian and Atlantic mitochondrial DNA clades) and contemporary feeding specializations (microphageous and macrophageous morphotypes). The present study demonstrated that independent suites of correlated morphological traits are associated with either vicariant history or contemporary feeding specializations. Second, functionally-similar feeding specialists exhibit distinct morphologies resulting largely from vicariant historical processes. Although, the evolutionary processes producing historical phenotypes remains unknown, the results obtained demonstrate how adaptive radiation associated with ecological resource partitioning and feeding specializations can be strongly influenced by intraspecific phenotypic diversification resulting from relatively recent vicariant histories.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 140–151.     

Ontogenetically stable dimorphism in a lacertid lizard (Acanthodactylus boskianus) with tests of methodology and comments on life-history

Recent arguments in the literature prompted us to compare methods for assessing sexual dimorphism in body proportions of lacertid lizards, using Acanthodactylus boskianus . Although expressing body-part measurements as proportional to head length was the most effective method, we recommend using trunk length for the baseline as a general method for lizards. We also argue that, when aiming to assess sexual dimorphism in body proportions of lizards, if the context is ecological, all available adults should be included. However, for morphology and taxonomy, small sub-samples of the largest individuals that maximally express their genetic morphological potential should be used. In A. boskianus , the sexual dimorphism of mensural characters in adults was typical: males were larger, with relatively larger head and appendages. However, the ontogeny of this dimorphism was unusual in that the differences existed already in youth and thereafter persisted isometrically. The sexual dimorphism of meristic characters was male-biased in numbers of femoral pores and of caudal vertebrae, and female-biased in numbers of ventral plates along the trunk and of precaudal vertebrae. Size dimorphism may conceivably play a role in sex recognition because two potential visual cues (i.e. size dimorphism and dichromatism) appear to complement each other.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 275–288.     

Intraspecific variation in wing and pupal melanization in copper butterflies (Lepidoptera: Lycaenidae)

Melanin is a widespread pigment causing variation in skin darkness, with darker phenotypes typically reaching higher equilibrium temperatures than lighter ones. Therefore, selection is expected to favour darker phenotypes in colder environments. In the present study, we show intraspecific variation in pupal (and wing) melanization along an altitudinal gradient in two species of copper butterflies. Both, pupal and wing melanization increased with increasing altitude. Consistent with the thermal melanism hypothesis, darker (high-altitude) pupae reached higher equilibrium temperatures than paler (low-altitude) ones. However, as temperature differences were rather small despite pronounced differences in melanization, we cannot rule out that factors (e.g. ultraviolet protection, disease resistance) other than temperature comprise the principal selective agents. Mechanistically, variation in melanization might be related to variation in hormone titres, as demonstrated by low-altitude pupae showing higher ecdysteroid and juvenile hormone titres compared to high-altitude ones. Furthermore, we report sex differences in wing melanization, with males being darker than females, which is potentially related to a higher flight activity of males.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 301–312.     

Implications of the mastoid anatomy of larger extant felids for the evolution and predatory behaviour of sabretoothed cats (Mammalia, Carnivora, Felidae)

Muscle attachments in the mastoid region of the skull of extant felids are studied through dissection of two adult tigers Panthera tigris (Linnaeus, 1758) Pocock, 1930, a lion Panthera leo (Linnaeus, 1758) Pocock, 1930 and a puma Puma concolor (Linnaeus, 1771) Jardine, 1834, providing for the first time an adequate reference for the study of the evolution of that region in sabretoothed felids. Our study supports the inference by W. Akersten that the main muscles inserting in the mastoid process in sabretooths were those originating in the atlas, rather than those from the posterior neck, sternum and forelimb. Those inferences were based on the anatomy of the giant panda, Ailuropoda melanoleuca (David, 1869) Milne-Edwards, 1870, raising uncertainties about homology, which were founded, as revealed by our results. The mastoid muscle insertions in extant felids differ in important details from those described for Ailuropoda , but agree with those described for domestic cats, hyenas and dogs. The large, antero-ventrally projected mastoid process of pantherines allows a moderate implication of the m. obliquus capitis anterior in head-flexion. This contradicts the widespread notion that the function of this muscle in carnivores is to extend the atlanto-cranial joint and to flex it laterally, but supports previous inferences about the head-flexing function of atlanto-mastoid muscles in machairodontines. Sabretooth mastoid morphology implies larger and longer-fibred atlanto-mastoid muscles than in pantherines, and that most of their fibres ran inferior to the axis of rotation of the atlanto-occipital joint, emphasizing head-flexing action.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 207–221.     

Modelling differential extinctions to understand big cat distribution on Indonesian islands

Aims To model differential extinction rates for island populations of tigers Panthera tigris and leopards P. pardus. Location Indonesia. Methods We built VORTEX population models of tiger and leopard populations on an island the size of Bali (3632 km²), using data from the literature. Results The tiger populations were less extinction prone than the leopard populations. This was unexpected as tigers had the smaller population sizes and, as such, might be assumed to be more extinction prone. We identified several aspects of tiger breeding biology that explain the result. Main conclusions Sea level reconstructions suggest that both tiger and leopard would have been present in Java, Sumatra and Bali at the end of the last glacial. Our model provides a plausible mechanism based on population ecology to explain why these leopard populations were more extinction prone than the tiger populations. In addition it illustrates the potential utility of population ecology models in understanding historical patterns in biogeography.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Phylogeny, biogeography, and recurrent evolution of divergent bill types in the nectar-stealing flowerpiercers (Thraupini: Diglossa and Diglossopis)

Neotropical tanagers known as flowerpiercers ( Diglossa and Diglossopis ) have a novel feeding adaptation, comprising a downward curved hook on the maxilla that allows these species to obtain floral nectar without pollination. Using mitochondrial DNA sequences, the phylogenetic relationships of all 18 species of flowerpiercers were studied for the first time. Strong support was found for the monophyly of flowerpiercers and for the monophyly of four superspecies within flowerpiercers. However, previously described species-groups, as well as the genus Diglossopis , are not monophyletic. The biogeographic origin of flowerpiercers was identified as Andean, with a single dispersal event from the northern Andes to Central America and a single dispersal event from the northern Andes to the tepuis. The first principal component, representing a contrast between hook size and bill size, was mapped onto the phylogeny to examine the evolution of relative hook size in the group. Across the phylogeny, a relatively large hook and a relatively small hook evolved multiple times in unrelated lineages, indicating lability in bill morphology. Differences in hook size among sympatric species, together with habitat partitioning and behavioural differences, can explain the coexistence of multiple species of flowerpiercers at the same locality.   © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 14–28.     

华南虎、东北虎、孟加拉虎的D-100p和ND5序列及其在系统进化分析中的应用

本文采用PCR和质粒克隆测序方法,获得了华南虎线粒体D-loop区的480bp序列和东北虎、孟加拉虎线粒体D-loop区的503bp序列;同时还获得了这三个虎亚种和金钱豹线粒体ND5基因5’端309bp的部分序列。根据D-loop序列分析,华南虎与孟加拉虎、东北虎的平均距离(p-distance)分别为0．11088和O．11087,而东北虎与孟加拉虎间的平均距离为0．00994;根据ND5序列分析,华南虎与孟加拉虎、东北虎的平均距离分别为0．11434和0．11758,而东北虎与孟加拉虎间的平均距离为0．00324。三个虎亚种的mtDNA D-loop和ND5序列比较表明,华南虎是这三个虎亚种中最为古老的亚种。     

It is all in the head: morphological basis for differences in bite force among colour morphs of the Dalmatian wall lizard

Males of the lizard Podarcis melisellensis occur in three distinct colours that differ in bite performance, with orange males biting harder than white or yellow ones. Differences in bite force among colour morphs are best explained by differences in head height, suggesting underlying variation in cranial shape and/or the size of the jaw adductors. To explore this issue further, we examined variation in cranial shape, using geometric morphometric techniques. Additionally, we quantified differences in jaw adductor muscle mass. No significant differences in size corrected head shape were found, although some shape trends could be detected between the colour morphs. Orange males have relatively larger jaw adductors than yellow males. Not only the mass of the external jaw adductors, but also that of the internal jaw adductors was greater for the orange morph. Data for other cranial muscles not related to biting suggest that this is not the consequence of an overall increase in robustness in orange individuals. These results suggest that differences in bite performance among morphs are caused specifically by an increase in the mass of the jaw adductor, which may be induced by differences in circulating hormone levels.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 13–22.     

Differences in environmental predictability underlie divergent competitive abilities in three congeneric hydroids

Species in which individuals experience predictable and uniform environments should be most finely adapted to their environment. Many hydrozoan species in the genus Hydractinia simultaneously occupy similar microhabitats (gastropod shells inhabited by hermit crabs) but experience considerable differences in their immediate environment (size and species of shells and hosts). In the present study, hydroid species experience differences in environmental predictability and traits that mediate competitive ability (growth form and growth rate). The inferred competitive ability was directly proportional to the extent to which the gastropod environment promotes interactions between small, juvenile colonies, which always end in competitive elimination. Extensive intraspecific variation in competitive ability was explained primarily by crab host species or site. Dense host populations impose more severe disturbance regimes that favour competitively inferior, but disturbance-resistant, phenotypes. Interplay between different types of variation (gastropods and hermit crabs) provides a possible mechanism for the maintenance of intraspecific growth form variation.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 322–338.     

西双版纳尚勇自然保护区野生印支虎及其三种主要有蹄类猎物种群现状调查

中国野生印支虎及其猎物种群状况的野外实地研究一直处于空白。本研究使用足迹鉴别法、粪堆计数法,首次对西双版纳尚勇自然保护区野生印支虎种群数量现状及该区域内的虎猎物种群状况进行了调查研究。结果显示:2004 ～ 2009 年间,确认西双版纳保护区存在3 只成年印支虎个体(2 雌1 雄),西双版纳尚勇保护区拥有比较丰富的有蹄类种群,其中虎的主要猎物:水鹿平均密度为7.63 (7.40 ～ 9.23)只/ km² ;赤麂平均密度为17. 39 (11.33 ～24.94)只/ km² ,野猪平均密度为10.26 (7.69 ～ 14.51) 只/ km² ,该区域虎猎物生物量为1 715. 74 kg/ km² 。本研究还探讨了该区域印支虎种群的保护前景以及中国境内开展虎种群调查的适用办法等。     

Spatial and temporal variation in flight morphology in the butterfly Melitaea cinxia (Lepidoptera: Nymphalidae)

Some studies on insects have found a relationship between habitat structure and investment in dispersal-related traits. In this study we compared the morphology of the butterfly Melitaea cinxia from five sites on the large Baltic island Öland that differed markedly in degree of fragmentation and size. Both wild-caught adults and individuals reared in a split-plot design were compared. We found significant site differences in size-adjusted thorax mass and total body mass. Male thorax mass was on average larger among the sites with the highest degree of habitat fragmentation. However, due to significant sex–site interactions, males and females may have adapted differently to the habitat fragmentation. Using museum specimens, we also analysed changes in morphology, finding an increase in size (measured as head and thorax width) over time. Thorax width appears to have increased among females and decreased among males. Possible explanations include increasing fragmentation of the landscape and changes in population density.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 445–453.     

Investigating South American biogeographic history using patterns of skull shape variation on Cerdocyon thous (Mammalia: Canidae)

Patterns of geographic variation of the canid Cerdocyon thous have historically been obscured by its remarkable intraspecific morphological variability. The observed distribution is highly associated with phytophysiognomy, a feature considered highly dynamic along geological time. In the present study, we tested whether vegetation distribution during the Holocene Glacial Maximum of South America (HGM) explains the patterns of morphological variation within Cerdocyon thous . The species was divided in groups according to paleohabitats that could support their presence during the HGM, and then tested for differences in skull morphometrics. The results obtained demonstrate that the climatic changes during the HGM influenced the population structure of this species, resulting in the establishment of geographical groups with different degrees of morphological cohesion. Higher morphological cohesion found in the Northern group might be explained by the marked discontinuity between its geographical range and the rest of the species'distribution. The Eastern and Southern morphological divergence is less striking and, although this could be related to past vegetation distribution, the disappearance of those barriers leads to a population structure that could be slowly breaking down.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 77–84.     

Heterochronic differences in fin development between latitudinal populations of the medaka Oryzias latipes (Actinopterygii: Adrianichthyidae)

Heterochrony is believed to have played important roles in macroevolutionary morphological changes. However, few studies have focused on intraspecific heterochrony, although interspecific differences ultimately originated from variation within ancestral species. We have demonstrated heterochrony in fin development between two latitudinal populations of the medaka, Oryzias latipes . Comparisons of fin length (anal and dorsal) among wild individuals revealed that fins are shorter with respect to body length in the northern population, indicating that they are 'paedomorphic' compared with the southern population. Observations of fin ray formation and subsequent fin growth in the laboratory revealed that the timing of pterygiophore development occurs later, and that fins start to elongate later with respect to body length in the northern fish, indicating that fin growth is 'post-displaced' compared with the southern population. In addition, the rate of fin growth with respect to body length was lower in the northern males, indicating 'neoteny'. Given that all Oryzias except O. latipes are distributed in the tropics, it is likely that higher-latitude fish have evolved post-displacement and neoteny during northern extension of their geographic range. The delayed development in higher-latitude fish is probably a trade-off for faster body growth, which has evolved as an adaptation to seasonally time-constrained environments.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 571–580.     

Costs of carnivory: tooth fracture in Pleistocene and Recent carnivorans

Large, carnivorous mammals often break their teeth, probably as a result of tooth to bone contact that occurs when carcasses are consumed more fully, a behaviour likely to occur under conditions of food stress. Recent studies of Pleistocene predators revealed high numbers of teeth broken in life, suggesting that carcass utilization and, consequently, food competition was more intense in the past than at present. However, the putative association between diet and tooth fracture frequency was based on a small sample of large, highly carnivorous species. In the present study, a greater diversity of extant carnivorans is sampled, including insectivorous, omnivorous, and carnivorous forms, ranging in size from weasels to tigers. Species that habitually consume hard foods (bones, shells) had the highest fracture frequencies, followed by carnivores, and then insectivorous and/or omnivorous species. Predator and prey sizes were not associated with tooth fracture frequency, but more aggressive species did break their teeth more often. Comparison of the modern sample with five Pleistocene species confirms the previous finding of higher tooth breakage in the past, although some extant species have fracture frequencies that approach those of extinct species. Thus, the Pleistocene predator guild appears to have been characterized by relatively high levels of competition that are rarely observed today.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 68–81.     

Does male secondary sexual trait size reveal fertilization efficiency in Australian Drosophila bipectinata Duda (Diptera: Drosophilidae)?

Males developing relatively large, costly sexually selected traits may be of superior body condition compared to small-ornamented males. Thus, males developing the largest secondary sexual trait in a given environment may also be able to augment their investment into ejaculate quality, and fertilize a larger proportion of a female's eggs. We tested the prediction that the degree of expression of a condition-dependent secondary sexual trait, the male sex comb, in a Cape Tribulation (northeastern Australia) population of Drosophila bipectinata Duda, reveals male ability to fertilize eggs in the absence of sperm competition. This test permitted us also to evaluate whether pre-copulatory sexual selection and fertilization efficiency might act additively to influence male reproductive success because a previous study of the same population demonstrated a positive association between comb size and copulation probability. The results obtained indicate that, although genotypes developing smaller sex combs collectively had a significantly higher rate of insemination failure compared to larger comb genotypes, the hatch rate and the number of eggs laid by females inseminated by the two genotypic categories were not statistically different. The results fail to support the prediction that comb size reveals noncompetitive fertilization efficiency of males in this Australian population.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 406–413.