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Showing papers in "Biological Journal of The Linnean Society in 2010"


Journal ArticleDOI
TL;DR: A study on the most exhaustive taxonomic sample of amniotes of any quantitative work on this topic demonstrates a strong relationship between lifestyle (aquatic, amphibious or terrestrial) and humeral microanatomy, and proposes an alternative method of palaeobiological inference for hypothetical ancestors.
Abstract: A study on the most exhaustive taxonomic sample of amniotes (75 extant and nine extinct taxa) of any quantitative work on this topic published so far demonstrates a strong relationship between lifestyle (aquatic, amphibious or terrestrial) and humeral microanatomy. We suggest that corrections for multiple testing be used to check for statistical artefacts in the context of a phylogenetic independent contrast analysis, and we use the false discovery rate procedure for this. Linear discriminant models segregate the various lifestyles with excellent success rate of up to 98.5%. Lifestyle was thus inferred for six extinct taxa of uncertain habitat. The results obtained suggest that Captorhinus, Claudiosaurus, and Placodus were amphibious, whereas Neusticosaurus and Mesosaurus were aquatic. Lystrosaurus may have been more aquatic than previously suggested, although the results of our inference models have to be integrated with other sources of data, which suggest that it may have been amphibious, rather than aquatic (as a literal interpretation of the models would suggest). Finally, we propose an alternative method of palaeobiological inference for hypothetical ancestors. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 384–406.

140 citations


Journal ArticleDOI
TL;DR: In this paper, the authors show widespread discordance between phylogenies based on mtDNA (two genes) and nuclear DNA (nucDNA; six loci) in a phylogenetic analysis of the turtle family Emydidae and find an unusual type of discordance involving the unexpected homogeneity of mtDNA sequences across species within genera.
Abstract: Do phylogenies and branch lengths based on mitochondrial DNA (mtDNA) provide a reasonable approximation to those based on multiple nuclear loci? In the present study, we show widespread discordance between phylogenies based on mtDNA (two genes) and nuclear DNA (nucDNA; six loci) in a phylogenetic analysis of the turtle family Emydidae. We also find an unusual type of discordance involving the unexpected homogeneity of mtDNA sequences across species within genera. Of the 36 clades in the combined nucDNA phylogeny, 24 are contradicted by the mtDNA phylogeny, and six are strongly contested by each data set. Two genera (Graptemys, Pseudemys) show remarkably low mtDNA divergence among species, whereas the combined nuclear data show deep divergences and (for Pseudemys) strongly supported clades. These latter results suggest that the mitochondrial data alone are highly misleading about the rate of speciation in these genera and also about the species status of endangered Graptemys and Pseudemys species. In addition, despite a strongly supported phylogeny from the combined nuclear genes, we find extensive discordance between this tree and individual nuclear gene trees. Overall, the results obtained illustrate the potential dangers of making inferences about phylogeny, speciation, divergence times, and conservation from mtDNA data alone (or even from single nuclear genes), and suggest the benefits of using large numbers of unlinked nuclear loci. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 445‐461.

139 citations


Journal ArticleDOI
TL;DR: A comprehensive molecular phylogenetic analysis based on ∼4.0 kb of DNA from four loci across the entire tribe Euglossini indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera, and divergence time estimates indicate that extant orchid bees lineages shared a most recent common ancestor at 27–42 Mya.
Abstract: The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on ∼4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1-α (EF1-α), arginine kinase (ArgK) and RNA polymerase II (Pol-II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil-calibrated molecular clocks and the evolution of morphological traits using disparity-through-time plots. In addition, we inferred past biogeographical events by implementing model-based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27–42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 552–572.

130 citations


Journal ArticleDOI
TL;DR: It is shown that large relative brain size in birds is associated with pair-bond strength, itself a likely consequence of cooperation and negotiation between partners under high levels of parental investment, and that the evolution of both traits is contingent on biparental care.
Abstract: In birds, large brains are associated with a series of population-level phenomena, including invasion success, species richness, and resilience to population decline. Thus, they appear to open up adaptive opportunities through flexibility in foraging and anti-predator behaviour. The evolutionary pathway leading to large brain size has received less attention than behavioural and ecological correlates. Using a comparative approach, we show that, independent of previously recognized associations with developmental constraints, relative brain size in birds is strongly related to biparental care, pair-bonding, and stable social relationships. We also demonstrate correlated evolution between large relative brain size and altricial development, and that the evolution of both traits is contingent on biparental care. Thus, biparental care facilitates altricial development, which permits the evolution of large relative brain size. Finally, we show that large relative brain size is associated with pair-bond strength, itself a likely consequence of cooperation and negotiation between partners under high levels of parental investment. These analyses provide an evolutionary model for the evolution of and prevalence of biparental care, altricial development, and pair-bonding in birds. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 111‐123.

129 citations


Journal ArticleDOI
TL;DR: This work was partially funded by the ‘EU Marie Curie Mobility and Training Program (FP7)’ and project CGL2008-04164 of the Spanish Ministry of Science and Innovation.
Abstract: This work was partially funded by the ‘EU Marie Curie Mobility and Training Program (FP7)’ and project CGL2008-04164 of the Spanish Ministry of Science and Innovation (I. De la Riva, principal investigator).

122 citations


Journal ArticleDOI
TL;DR: In this paper, the authors investigated the functional morphology of shark teeth and found that the majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types.
Abstract: The performance of an organism’s feeding apparatus has obvious implications for its fitness and survival. However, the majority of studies that focus on chondrichthyan feeding have largely ignored the role of teeth. Studying the functional morphology of shark teeth not only elucidates the biological role that teeth play in feeding, but also provides insight specifically into the evolution of shark feeding because teeth are often the only structures available in the fossil record. In the present study, we investigate the puncture and draw performance of three general categories of extant teeth, tearing-type, cutting-type, and cutting–clutching type, as well as three fossil morphologies, utilizing a universal testing system. Differences in puncturing performance occurred among different prey items, indicating that not all ‘soft’ prey items are alike. The majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types; however, few patterns emerged. In some cases, broader triangular teeth were less effective at puncturing than narrow-cusped teeth. There were no differences between the maximum draw forces and maximum puncture forces. Many of the shark teeth in the present study were not only able to perform draw and puncture equally well, but also many tooth morphologies were functionally equivalent to each other. The findings obtained in the present study lend little support to the belief that shark tooth morphology is a good predictor of biological role. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 271–286.

100 citations


Journal ArticleDOI
TL;DR: These results provide independent evidence that climate cooling following the Middle Miocene Climatic Optimum led to important vegetational shifts in eastern Brazil, causing an increase in the dominance of open versus closed habitats.
Abstract: The Brazilian Cerrado is the most species-rich tropical savanna in the world. Within this biome, the Campos Rupestres (‘rocky savannas’) constitute a poorly studied and highly threatened ecosystem. To better understand how plants characteristic of this vegetation have evolved and come to occupy the now widely-separated patches of rocky formations in eastern Brazil, we reconstruct the biogeographical history of the rare orchid genus Hoffmannseggella. We apply parsimony and Bayesian methods to infer the phylogenetic relationships among 40 out of the 41 described species. Absolute divergence times are calculated under penalized likelihood and compared with estimates from a Bayesian relaxed clock. Ancestral ranges are inferred for all nodes of the phylogeny using Fitch optimization and statistical dispersal vicariance analysis. In all analyses, phylogenetic uncertainty is taken into account by the independent analysis of a large tree sample. The results obtained indicate that Hoffmannseggella underwent rapid radiation around the Middle/Late Miocene (approximately 11–14 Mya). The region corresponding today to southern Minas Gerais acted as a main source area for several independent range expansions north- and eastwards via episodic corridors. These results provide independent evidence that climate cooling following the Middle Miocene Climatic Optimum (approximately 15 Mya) led to important vegetational shifts in eastern Brazil, causing an increase in the dominance of open versus closed habitats. Polyploidy following secondary contact of previously isolated populations may have been responsible for the formation of many species, as demonstrated by the high ploidy levels reported in the genus. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 597–607.

99 citations


Journal ArticleDOI
TL;DR: This type of classification function provides a means of integrating the complex morphological differences between morphotypes into a single score that reflects the position of a population along the benthic-limnetic axis and can be used to relate that position to other aspects of stickleback biology.
Abstract: Many species of fish display morphological divergence between individuals feeding on macroinvertebrates associated with littoral habitats (benthic morphotypes) and individuals feeding on zooplankton in the limnetic zone (limnetic morphotypes). Threespine stickleback (Gasterosteus aculeatus L.) have diverged along the benthic-limnetic axis into allopatric morphotypes in thousands of populations and into sympatric species pairs in several lakes. However, only a few well known populations have been studied because identifying additional populations as either benthic or limnetic requires detailed dietary or observational studies. Here we develop a Fisher's linear discriminant function based on the skull morphology of known benthic and limnetic stickleback populations from the Cook Inlet Basin of Alaska and test the feasibility of using this function to identify other morphologically divergent populations. Benthic and limnetic morphotypes were separable using this technique and of 45 populations classified, three were identified as morphologically extreme (two benthic and one limnetic), nine as moderately divergent (three benthic and six limnetic) and the remaining 33 populations as morphologically intermediate. Classification scores were found to correlate with eye size, the depth profile of lakes, and the presence of invasive northern pike (Esox lucius). This type of classification function provides a means of integrating the complex morphological differences between morphotypes into a single score that reflects the position of a population along the benthic-limnetic axis and can be used to relate that position to other aspects of stickleback biology.

92 citations


Journal ArticleDOI
TL;DR: The observed relationship between male coloration and water transparency is not mediated by environmental variation alone, and female mating preferences are indicated to have changed in response to this variation, constituting the first evidence for intraspecific preference-trait co-evolution in cichlid fish.
Abstract: Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexual selection: female mate choice exerts sexual selection on male nuptial coloration within species and maintains reproductive isolation between species. However, declining water transparency coincides with increasingly dull coloration and increasing hybridization. In the present study, we investigated the mechanism underlying this pattern in Pundamilia nyererei, a species that interbreeds with a sister species in turbid but not in clear water. We compared measures of intraspecific sexual selection between two populations from locations that differ in water transparency. First, in laboratory mate-choice experiments, conducted in clear water and under broad-spectrum illumination, we found that females originating from turbid water have significantly weaker preferences for male coloration than females originating from clear water. Second, both the hue and body coverage of male coloration differ between populations, which is consistent with adaptation to different photic habitats. These findings suggest that the observed relationship between male coloration and water transparency is not mediated by environmental variation alone. Rather, female mating preferences are indicated to have changed in response to this variation, constituting the first evidence for intraspecific preference-trait co-evolution in cichlid fish. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 398-406.

89 citations


Journal ArticleDOI
TL;DR: Evidence from taxa as divergent as sparrowhawks and great tits indicates that protoporphyrin pigments responsible for maculation have a structural function in compensating for eggshell thinning, as caused by calcium stress, and, more recently, dichlorodiphenyltrichloroethane.
Abstract: Recent work suggests that the evolution of egg coloration may have been constrained in three important ways that have not yet been critically synthesized in any review First, on account of birds being able to see in the ultraviolet spectrum, the interaction between the properties of avian vision and the light environment of nests imply different perceptions of egg coloration from those experienced by humans Second, a new hypothesis to explain blue–green egg coloration interprets it as a sexually selected signal to males of the laying female's genetic quality Third, evidence from taxa as divergent as sparrowhawks and great tits indicates that protoporphyrin pigments responsible for maculation (spotting patterns) have a structural function in compensating for eggshell thinning, as caused by calcium stress, and, more recently, dichlorodiphenyltrichloroethane We consider this to be the most convincing explanation for the primary function of spotting, although an important secondary function might arise through the fact that individual patterns of maculation may allow birds to identify their own eggs, effectively serving as signatures in the face of inter- or intra-specific brood parasitism These constraints or hypotheses are not mutually exclusive, and should not be taken to imply that one, but not other, agents of selection might apply to any one species However, the sexually-selected eggshell coloration hypothesis is least plausible for hole-nesting birds because of the poor light quality available, although such species have been the focus of research in this area, and only a single experimental study has shown a link between egg coloration and male provisioning Furthermore, the observed relationships between female phenotypic quality and egg traits do not necessarily imply that they have signalling functions © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 753–762

88 citations


Journal ArticleDOI
TL;DR: The data obtained in the present study suggest a mechanism by which environmental change as a result of human activities could influence the health and fitness of individuals in natural populations.
Abstract: Breeding success is often dictated by the degree to which parents can synchronize the maximum food requirements of offspring to the peak in abundance of invertebrate prey. Less studied is how the nutritional quality of individual diet items impacts on breeding. In the present study, we assessed the abundance and antioxidant concentrations of arboreal arthropods from oak woodland and provisioning behaviour of the blue tit Cyanistes caeruleus. Dietary antioxidants are important during development because they defend against oxidative stress. Operophtera caterpillars, Erannis caterpillars, and spiders contained significantly different levels of individual carotenoids and a-tocopherol. Concentrations of lutein and b-carotene in Operophtera caterpillars did not vary seasonally, although concentrations of zeaxanthin declined and a-tocopherol increased with date. Blue tit broods hatched later in the season received significantly fewer caterpillars and more spiders per chick compared to earlier broods. Reflecting changes in prey composition, blue tit nestling plasma showed decreases in zeaxanthin and increases in a-tocopherol with date. Thus, processes that shift the timing of breeding in birds and/or prey composition are likely to alter antioxidant intake and thus potentially influence the oxidative stress status of animals. The data obtained in the present study suggest a mechanism by which environmental change as a result of human activities could influence the health and fitness of individuals in natural populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 708–717. ADDITIONAL KEYWORDS: a-tocopherol – carotenoids – caterpillars – neonatal nutrition – oxidative stress – spiders.

Journal ArticleDOI
TL;DR: The mitochondrial data support the species status of M. mongolicus as a member of Alexandromys but not of the Microtus arvalis group, thus being concordant with the cytogenetic data, and the maximum likelihood distance showed a tendency for a progressive underestimation of divergence and time for older splits.
Abstract: To examine phylogenetic relationships within the Asian lineage of voles (Microtus) belonging to subgenus Alexandromys, the mitochondrial cytochrome b gene (cytb) was sequenced for its representatives, and the results were compared with the cytogenetic, morphological, and paleontological data. In all the trees inferred from maximum likelihood, parsimony, and Bayesian phylogenetic analyses, the Asian clade is subdivided into highly supported Alexandromys s.s. and moderately supported Pallasiinus lineages. Four subclades are recovered within Alexandromys: (1) Microtus maximowiczii and Microtus sachalinensis; (2) Microtus miiddendorffii s.l., Microtus mongolicus and Microtus gromovi; (3) Microtus fortis; and (4) Microtus limnophilus. Thus, M. limnophilus demonstrates clear affinities to Alexandromys s.s. but not to Microtus oeconomus (subgenus Pallasiinus), which was always regarded as its sibling species. The results obtained indicate M. mongolicus as a member of Alexandromys but not of the Microtus arvalis group, thus being concordant with the cytogenetic data. The mitochondrial data support the species status of M. gromovi; moreover, its placement as a part of a trichotomy with M. miiddendorffii s.l. and M. mongolicus contradicts the traditional affiliation of M. gromovi with M. maximowiczii. The divergence rate of cytb third position transversions in Microtus is estimated at approximately 8% per Myr, which corresponds to approximately 30% per Myr for all substitution types at all codon positions. The maximum likelihood distance based on complete sequence showed a tendency for a progressive underestimation of divergence and time for older splits. According to our molecular clock analysis employing nonlinear estimation methods, the split between Alexandromys and Pallasiinus and basal radiation within Alexandromys date back to approximately 1.2 Mya and 800 Kya, respectively. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 595–613.

Journal ArticleDOI
TL;DR: It is concluded that although Wallacea as a whole is not a natural biogeographical region, neither is it completely artificial as it is formed from a complex of predominantly Australasian exotic fragments linked by geological processes within a complex collision zone.
Abstract: The concepts of biogeographical regions and areas of endemism are briefly reviewed prior to a discussion of what constitutes a natural biogeographical unit. It is concluded that a natural biogeographical unit comprises a group of endemic species that share a geological history. These natural biogeographical units are termed Wallacean biogeographical units in honour of the biogeographer A.R. Wallace. Models of the geological development of Indonesia and the Philippines are outlined. Areas of endemism within Wallacea are identified by distributional data, and their relationship to each other and to the adjacent continental regions are evaluated using molecular phylogenies from the literature. The boundaries of these areas of endemism are in broad agreement with earlier works, but it is argued that the Tanimbar Islands are biologically part of south Maluku, rather than the Lesser Sundas, and that Timor (plus Savu, Roti, Wetar, Damar, and Babar) and the western Lesser Sundas form areas of endemism in their own right. Wallacean biogeographical units within Wallacea are identified by congruence between areas of endemism and geological history. It is concluded that although Wallacea as a whole is not a natural biogeographical region, neither is it completely artificial as it is formed from a complex of predominantly Australasian exotic fragments linked by geological processes within a complex collision zone. The Philippines are argued to be an integral part of Wallacea, as originally intended.

Journal ArticleDOI
TL;DR: It is described the first account of soil- and beetle-associated nematodes on an island setting by investigating the island of Reunion in the Indian Ocean, establishing Reunion as a case study for nematode island biogeography, in which the analysis of nematODE population genetics and population dynamics can provide insight into evolutionary and ecological processes.
Abstract: Pristionchus pacificus has been established as a nematode model system in evolutionary developmental biology and evolutionary ecology. Field studies in North and South America, Asia, Africa and Europe indicated that nematodes of the genus Pristionchus live in association with scarab beetles. Here, we describe the first account of soil- and beetle-associated nematodes on an island setting by investigating the island of Reunion in the Indian Ocean. Reunion has high numbers of endemic insects and is one among several attractive islands for biodiversity studies. Being of volcanic origin, Reunion is 2–3 million years old, making it the youngest of the Mascareigne islands. We show that beetle- and soil-derived nematodes on Reunion are nearly exclusively hermaphroditic, suggesting that selfing is favoured over gonochorism (outcrossing) during island colonization. Among members of four nematode genera observed on Reunion, Pristionchus pacificus was the most prevalent species. A total of 76 isolates, in association with five different scarab beetles, has been obtained for this cosmopolitan nematode. A detailed mitochondrial haplotype analysis indicates that the Reunion isolates of P. pacificus cover all four worldwide clades of the species. This extraordinary haplotype diversity suggests multiple independent invasions, most likely in association with different scarab beetles. Together, we establish Reunion as a case study for nematode island biogeography, in which the analysis of nematode population genetics and population dynamics can provide insight into evolutionary and ecological processes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 170–179.

Journal ArticleDOI
TL;DR: Widespread occurrence of invasive Northern hemisphere blue mussels in the Southern hemisphere is documented for the first time.
Abstract: The taxonomic and evolutionary affinities of Southern hemisphere smooth-shelled blue mussels are unclear, with studies using different marker types having identified different relationships among various geographic regions. Using an existing and a new molecular assay, the present study builds on previous work to test the distribution of blue mussels native to and introduced to the Southern hemisphere. Populations of Mytilus were sampled from New Zealand, Australia, and Chile. The nuclear-DNA marker Me 15/16 was used to identify the taxonomic status of 484 individuals. A new restriction fragment length polymorphism (RFLP) assay was used to identify the hemisphere of origin for a subset of Mytilus galloprovincialis. The Me15/16 marker identified 478 pure M. galloprovincialis from Southern hemisphere sites and six Mytilus edulis/M. galloprovincialis hybrids from the Auckland Islands (New Zealand) and Chile. A cytoplasmic RFLP identified Northern hemisphere M. galloprovincialis in almost every Southern hemisphere region. The presence of native M. galloprovincialis at high latitudes (up to 52°S) has implications for our understanding of environmentally induced selective constraints considered to determine species distributions. Widespread occurrence of invasive Northern hemisphere blue mussels in the Southern hemisphere is documented for the first time. Identification of inter-specific hybrids (M. edulis ¥ M. galloprovincialis) in Chile and in the Auckland Islands (subantarctic New Zealand) illustrates that environments ranging from international ports to remote protected locations are vulnerable to bioinvasion. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 898‐909.

Journal ArticleDOI
TL;DR: Investigation of life-history syndromes of three common blackbird populations differing in their human influence confirmed that nest predation and nestling starvation rates are consistent with urbanization imposing new challenges on birds, and that they adaptively respond to them.
Abstract: Urbanization, one of the most extreme land-use alterations, is currently spreading, and the number of species confronting these changes is increasing. However, contradictory results of previous studies impede a clear interpretation of which selective pressure (nest predation or food limitation) is more important in urban habitats compared with natural situations, and whether birds can confront them by adjusting their life-history strategies. We investigated life-history syndromes of three common blackbird (Turdus merula) populations differing in their human influence (urban, rural, and woodland). We analysed daily nest predation and nestling starvation rates to assess the relative importance of these selection pressures in each habitat. Simultaneously, several life-history traits were investigated to determine if T. merula seem adapted to their main source of selection. Food limitation was more important in the city, whereas nest predation was the most important selective force in the forest. The rural habitat was characterized by an intermediate influence of these two factors. Life-history syndromes, as the covariation of a suite of traits, confirmed these results because T. merula seem well adapted to the main cause of selection in each habitat. Our results are consistent with urbanization imposing new challenges on birds, and that they adaptively respond to them. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 759–766.

Journal ArticleDOI
TL;DR: The results suggest that whereas in males, sexual selection seems to be operating on bite performance, in the case of females, natural selection seem to be the most likely and most important selective pressure driving the variation in head size.
Abstract: 461..475 Although differential selective pressures on males and females of the same species may result in sex-specific evolutionary trajectories, comparative studies of adaptive radiations have largely neglected within-species varia- tion. In this study, we explore the potential effects of natural selection, sexual selection, or a combination of both, on bite performance in males and females of 19 species of Liolaemus lizards. More specifically, we study the evolution of bite performance, and compare evolutionary relationships between the variation in head morphology, bite performance, ecological variation and sexual dimorphism between males and females. Our results suggest that in male Liolaemus, the variation in bite force is at least partly explained by the variation in the degree of sexual dimorphism in head width (i.e. our estimate of the intensity of sexual selection), and neither bite force nor the morphological variables were correlated with diet (i.e. our proxy for natural selection). On the contrary, in females, the variation in bite force and head size can, to a certain extent, be explained by variation in diet. These results suggest that whereas in males, sexual selection seems to be operating on bite performance, in the case of females, natural selection seems to be the most likely and most important selective pressure driving the variation in head size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 461-475. ADDITIONAL KEYWORDS: diet - ecomorphology - interspecific variation - natural selection - sexual differences - sexual selection.

Journal ArticleDOI
TL;DR: These intraspecific comparisons support the hypothesis that viviparity has evolved in cold-climate squamates because of the direct fitness advantages that warm temperatures provide developing offspring.
Abstract: Interspecific comparisons suggest a strong association between cool climates and viviparity in reptiles. However, intraspecific comparisons, which provide an opportunity to identify causal pathways and to distinguish facultative (phenotypically plastic) effects from canalized (genetically fixed) responses, are lacking. We documented the reproductive traits in an alpine oviparous lizard, and manipulated thermal regimes of gravid females and their eggs to identify proximate causes of life-history variation. Embryonic development at oviposition was more advanced in eggs laid by females from high-elevation populations than in eggs produced by females from lower elevations. In the laboratory, experimentally imposed low maternal body temperatures delayed oviposition and resulted in more advanced embryonic development at oviposition. Warm conditions both in utero and in the nest increased hatching success and offspring body size. Our intraspecific comparisons support the hypothesis that viviparity has evolved in cold-climate squamates because of the direct fitness advantages that warm temperatures provide developing offspring.

Journal ArticleDOI
TL;DR: The results obtained show the existence of many different genetic lineages within the studied Apodemus species, suggesting the isolation and differentiation of populations in multiple refuge areas, and higher genetic diversities in some regions such as Yunnan, Sichuan (China), and eastern Russia suggest these areas are potential refuges for these species.
Abstract: The phylogeography of four Apodemus species (Apodemus agrarius, Apodemus peninsulae, Apodemus latronum, and Apodemus draco) was studied in the Far East of Asia, based on sequences of the mitochondrial DNA cytochrome b gene. The results obtained show the existence of many different genetic lineages within the studied Apodemus species, suggesting the isolation and differentiation of populations in multiple refuge areas. Higher genetic diversities in some regions such as Yunnan, Sichuan (China), and eastern Russia suggest these areas are potential refuges for these species. The existence of such complex genetic structures could be linked to the presence of many biogeographic barriers (Himalaya Mountains, Tien-shan Mountains, Altai Mountains, Tibetan Plateau, Gobi desert, Yunnan Guizhou Plateau, Dzungaria basin, and others) in these regions, which were probably reinforced during the Quaternary climate changes. These barriers also played an important role concerning the low dispersal abilities of the two studied Apodemus species adapted to forest habitats (A. latronum and A. draco) with respect to colonizing regions other than China. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 797-821.

Journal ArticleDOI
TL;DR: The results obtained in the present study are consistent with genetic sustainability of current exploitation; local depleted stocks are recurrently replenished by recruits that may have originated from nonharvested areas.
Abstract: Phylogeographical analysis of Paracentrotus lividus was carried out by means of sequencing the mitochondrial cytochrome b gene (1143 bp) of 260 individuals collected at 22 Mediterranean and four Atlantic localities. Against a background of high haplotype diversity and shallow genetic structuring, we observed significant genetic divergence between the Adriatic Sea and the rest of the Mediterranean, as well as between the Mediterranean and the Atlantic sample groups. Furthermore, on the largest spatial scale, isolation by distance was detected. Three main haplogroups were identified by network and Bayesian assignment analyses. The relative proportions of haplogroups were different in the four regions considered, with the exception of Western and Eastern Mediterranean that showed a similar pattern. This result together with the outcome of Snn statistics, analysis of molecular variance and network analyses allowed to identify three weakly differentiated populations corresponding to the Atlantic, Western + Eastern Mediterranean, and Adriatic seas. Analyses of mismatch distribution and neutrality tests were consistent with the presence of genetic structuring and past demographic expansion(s). From a fisheries perspective, the results obtained in the present study are consistent with genetic sustainability of current exploitation; local depleted stocks are recurrently replenished by recruits that may have originated from nonharvested areas. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 910-923.

Journal ArticleDOI
TL;DR: The latitudinal richness gradient is a consequence of speciation and extinction events in unglaciated faunas and an increasing domination of faunaas by generalist, large bodied, large channel, recolonizing species in more northern regions.
Abstract: Published species lists were analysed to determine the contributions of dispersal, habitat preference, river channel size, body size, and glacial history to large-scale patterns in freshwater fish species richness in North America, north of central Mexico. Total species richness declines to the north and west but the pattern for endemics differs from that of widespread species. Mississippi Basin regions are more species rich than more isolated, coastal, regions. Richness declines more rapidly with increasing latitude in riverine specialist than in habitat generalist species. Levels of endemism are greatest in species found in small- to medium-sized river channels. The strong Rapoport effect, more marked in migratory than resident species, is correlated with habitat preference, channel size, and glacial history. Body size increases with latitude, largely as a result of a trend from small resident to large migrant species. In unglaciated regions, ancestral species survived in large habitats because these are longer-lived, more extensive, less isolated and more stable than headwaters, permitting larger populations and lower extinction levels. Reduced levels of gene flow in small, peripheral, channels isolated by larger downstream habitats have resulted in the production of many, small range, small-bodied species. The latitudinal richness gradient is a consequence of speciation and extinction events in unglaciated faunas and an increasing domination of faunas by generalist, large bodied, large channel, recolonizing species in more northern regions. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 46-61.

Journal ArticleDOI
TL;DR: The results suggest that the production of melanin pigments and fitness-related life-history traits are concomitantly regulated in a sex-specific way.
Abstract: Ornament expression fluctuates with age in many organisms. Whether these changes are adaptively plastic is poorly known. In order to understand the ultimate function of melanin-based ornaments, we studied their within-individual fluctuations and their covariation with fitness-related traits. In barn owls (Tyto alba), individuals vary from reddish-brown pheomelanic to white, and from immaculate to marked with black eumelanic spots, with males being less reddish and less spotted than females. During the first molt, both sexes became less pheomelanic, females displayed larger spots and males fewer spots, but the extent of these changes was not associated with reproduction. At subsequent molts, intra-individual changes in melanin-based traits covaried with simultaneous reproduction changes. Adult females bred earlier in the season and laid larger eggs when they became scattered with larger spots, whereas adults of both sexes produced larger broods when they became whiter. These results suggest that the production of melanin pigments and fitness-related life-history traits are concomitantly regulated in a sex-specific way. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 689‐704.

Journal ArticleDOI
TL;DR: It is proposed that P. apollo is best viewed as an atypical glacial invader in southern and western Europe, the isolated, montane populations of which, threatened by climate warming, retain a large fraction of the species evolutionary heritage.
Abstract: Parnassius apollo (Linnaeus, 1758) is probably the most renowned Eurasian montane butterfly. Its specialized ecology makes it very sensitive to habitat and climate changes, so that it is now experiencing range contraction and local extinction across most of its range. We sequenced 869 bp of the mitochondrial DNA (mtDNA) cytochrome oxidase I gene in 78 P. apollo populations (201 individuals) in order to: (1) assess the phylogeographic pattern of the species; (2) shed light on the historical biogeographic processes that shaped the distribution of the species; and (3) identify geographic population units of special value for the conservation of the species' genetic diversity. Our analyses revealed a very strong phylogeographic structure in P. apollo, which displays a number of distinctive mtDNA lineages populating geographically distinct areas. Overall sequence divergence is relatively shallow, and is consistent with a recent (late Pleistocene) colonization of most of the range. We propose that P. apollo is best viewed as an atypical glacial invader in southern and western Europe, the isolated, montane populations of which, threatened by climate warming, retain a large fraction of the species evolutionary heritage. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 169–183.

Journal ArticleDOI
TL;DR: Ecological opportunity, in the form of diverse available resources, along with intraspecific competition, may be a prerequisite for resource polymorphism to evolve, because such polymorphisms require diverse resources onto which each morph can specialize as an adaptive response to minimize competition.
Abstract: Resource polymorphism – the occurrence within a single population of discrete intraspecific morphs showing differential resource use – has long been viewed as an important setting for evolutionary innovation and diversification. Yet, relatively few studies have evaluated the ecological factors that favour resource polymorphism. Here, we combine observations of natural populations with a controlled experiment to assess the role of intraspecific competition (specifically, the density of conspecifics) and ecological opportunity (specifically, the range of resources available) on the expression of resource polymorphism in spadefoot toad tadpoles. We found that greater conspecific densities and a greater range of available resources together promoted the expression of resource polymorphism. We conclude that, ecological opportunity, in the form of diverse available resources, along with intraspecific competition, may be a prerequisite for resource polymorphism to evolve, because such polymorphisms require diverse resources onto which each morph can specialize as an adaptive response to minimize competition. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 73–88.

Journal ArticleDOI
TL;DR: The Caribbean region includes a geologically complex mix of islands, which have served as a backdrop for some significant studies of biogeography, mostly with vertebrates, and here, the tropical/subtropical spider genus Selenops is used to obtain a finer resolution of the role of geology in shaping patterns of species diversity.
Abstract: The Caribbean region includes a geologically complex mix of islands, which have served as a backdrop for some significant studies of biogeography, mostly with vertebrates Here, we use the tropical/subtropical spider genus Selenops (Selenopidae) to obtain a finer resolution of the role of geology in shaping patterns of species diversity We obtained a broad geographic sample from over 200 localities from both the islands and American mainland DNA sequence data were generated for three mitochondrial genes and one nuclear gene for eleven outgroup taxa and nearly 60 selenopid species Phylogenetic analysis of the data revealed several biogeographic patterns common to other lineages that have diversified in the region, the most significant being: (1) a distinct biogeographic break between Northern and Southern Lesser Antilles, although with a slight shift in the location of the disjunction; (2) diversification within the islands of Jamaica and Hispaniola; (3) higher diversity of species in the Greater Antilles relative to the Lesser Antilles However, a strikingly unique pattern in Caribbean Selenops is that Cuban species are not basal in the Caribbean clade Analyses to test competing hypotheses of vicariance and dispersal support colonization through GAARlandia, an Eocene–Oligocene land span extending from South America to the Greater Antilles, rather than over-water dispersal © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 288–322

Journal ArticleDOI
TL;DR: The results show a number of similarities to the evolution of three-spined stickleback morphology, indicating that environmental variables other than those related to gross habitat characteristics might also influence nine-spine sticklebacks in different habitats.
Abstract: Parallel evolution is characterised by repeated, independent occurrences of similar phenotypes in a given habitat type, in different parts of the species distribution area. We studied body shape and body armour divergence between five marine, four lake, and ten pond populations of nine-spined sticklebacks [Pungitius pungitius (Linnaeus, 1758)] in Fennoscandia. We hypothesized that marine and lake populations (large water bodies, diverse fish fauna) would be similar, whereas sticklebacks in isolated ponds (small water bodies, simple fish fauna) would be divergent. We found that pond fish had deeper bodies, shorter caudal peduncles, and less body armour (viz. shorter/absent pelvic spines, reduced/absent pelvic girdle, and reduced number of lateral plates) than marine fish. Lake fish were intermediate, but more similar to marine than to pond fish. Results of our common garden experiment concurred with these patterns, suggesting a genetic basis for the observed divergence. We also found large variation among populations within habitat types, indicating that environmental variables other than those related to gross habitat characteristics might also influence nine-spined stickleback morphology. Apart from suggesting parallel evolution of morphological characteristics of nine-spined sticklebacks in different habitats, the results also show a number of similarities to the evolution of three-spined stickleback (Gasterosteus aculeatus Linnaeus, 1758) morphology. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 403–416.

Journal ArticleDOI
TL;DR: The present study investigates the post-copulatory interactions between male and female sexual traits of a highly promiscuous simultaneous hermaphrodite, the free-living flatworm Macrostomum lignano, and derive hypotheses about how the different traits may represent evolutionary responses to such sexual conflicts.
Abstract: Different interests between mating partners regarding the fate of their gametes can lead to sexual conflicts in many species. Although these conflicts can sometimes be dealt with pre-copulatorily (e.g. by choosing with which partners to mate), they often extend beyond copulation. Post-copulatory sexual conflicts are expected to be particularly strong in simultaneous hermaphrodites because an individual may have to accept sperm in order to obtain an opportunity to donate sperm, reducing the effectiveness of pre-copulatory conflict resolution. The present study investigates the post-copulatory interactions between male and female sexual traits of a highly promiscuous simultaneous hermaphrodite, the free-living flatworm Macrostomum lignano. Using light and electron microscopy, we show the different levels of complexity of the sperm and the genitalia, and derive hypotheses about how the different traits may represent evolutionary responses to such sexual conflicts. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 370–383.

Journal ArticleDOI
TL;DR: The deep history hypothesis suggests that divergences deep in the evolutionary history of organisms resulted in niche preferences that are maintained, for the most part, in species represented in present-day assemblages.
Abstract: The structure of animal communities has long been of interest to ecologists. Two different hypotheses have been proposed to explain origins of ecological differences among species within present-day communities. The competition–predation hypothesis states that species interactions drive the evolution of divergence in resource use and niche characteristics. This hypothesis predicts that ecological traits of coexisting species are independent of phylogeny and result from relatively recent species interactions. The deep history hypothesis suggests that divergences deep in the evolutionary history of organisms resulted in niche preferences that are maintained, for the most part, in species represented in present-day assemblages. Consequently, ecological traits of coexisting species can be predicted based on phylogeny regardless of the community in which individual species presently reside. In the present study, we test the deep history hypothesis along one niche axis, diet, using snakes as our model clade of organisms. Almost 70% of the variation in snake diets is associated with seven major divergences in snake evolutionary history. We discuss these results in the light of relevant morphological, behavioural, and ecological correlates of dietary shifts in snakes. We also discuss the implications of our results with respect to the deep history hypothesis. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 476–486.

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TL;DR: In this article, the ontogenetic background for paedomorphosis in phocoenids was evaluated and assessed in terms of the development of epiphyseal fusion, cranial suture fusion, and ontogeny of cranial shape.
Abstract: To evaluate and assess the ontogenetic background for paedomorphosis in phocoenids, samples of 144 harbour porpoises, 81 white-beaked dolphins, and 130 Commerson’s dolphins were compared in terms of the development of epiphyseal fusion, cranial suture fusion, and ontogeny of cranial shape. Harbour porpoises and Commerson’s dolphins terminated growth and development of all investigated traits sooner than white-beaked dolphins, leading to lesser degrees of fusion of skeletal elements and less postnatal allometric development. The latter occurred even though shape in the two paedomorphic species developed at twice the rate relative to the size of white-beaked dolphins. These observations imply that progenetic evolution has occurred convergently in phocoenid and Cephalorhynchus ancestors. The truncated ontogenies allow sexual maturity to be attained earlier and provide a greater reproductive potential. Both species inhabit similar temperate productive habitats and, hence, ecological factors are proposed to have supplied the selection pressures leading to progenesis. Constant prey availability must be a prerequisite for the observed phenomena because frequent food-intake is necessitated by the limited capacity for energy storage and high heat-loss entailed by the resulting small body sizes. Progenesis has rarely been proposed in mammal species. This may reflect rarity or that mammalian expressions of progenesis are less obvious. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99, 278–295.

Journal ArticleDOI
TL;DR: The results suggest the existence of glacial refugia for Sus scrofa in the Iberian Peninsula during the Last Glacial Maximum, as well as a significant genetic difference among and within populations.
Abstract: The Eurasian wild boar, Sus scrofa, is an important game species and is widely distributed in Eurasia. The recent demographic expansions allied to man mediated translocations and reintroductions, reshaped the genetic variability of wild boar populations, and gave rise to management problems. Genetic variability and the population structure of the European populations are not well-known, in particular in the Iberian Peninsula. In the present study, we assessed the phylogenetic relationship among 17 Iberian wild boar populations and several Eurasian wild and domestic pigs by sequencing 660 bp of the mitochondrial (mt)DNA control region. This analysis was extended to the two autochthonous Portuguese pig breeds, Bisaro and Alentejano, and their relationship with Eurasian pigs was also established. High levels of gene diversity were found in Iberian wild boar and Portuguese domestic pigs, as well as a significant genetic difference among and within populations. Iberian wild boar and Portuguese pig breeds belong to the main European clade but diverge with animals from Central Europe, showing a high proportion of private (Iberian) haplotypes. These results suggest the existence of glacial refugia for Sus scrofa in the Iberian Peninsula during the Last Glacial Maximum. No signs of Asian mtDNA introgression from Asian populations were found in the present survey. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 797–822.