scispace - formally typeset
Search or ask a question

Showing papers in "Biological Journal of The Linnean Society in 2008"


Journal ArticleDOI
TL;DR: Bite force performance measures can be used as ‘traits’, and thus be used in integrative studies at multiple levels of organismal biology, and will help the understanding of the functions, capacities, and evolution of jaw–cranial musculoskeletal systems.
Abstract: Measurements of whole-organism performance traits have been useful in studies of adaptation and phenotype– environment correlations. Bite force capacities may be tightly linked to both the type and magnitude of the ecological challenges of food acquisition, mate acquisition, and antipredation in vertebrates. In the present study, we present technical details on bite meters and on measuring bite forces. The ability to take reliable measurements depends on specific features of the measuring device and on where in the mouth the bite is applied. Using both previously available and original data, we demonstrate several ecologically and evolutionarily relevant features of bite force measurements. First, maximal bite forces are repeatable among individuals across all vertebrates studied to date. Second, in ectotherms such as lizards, maximal bite forces are affected by body temperature and motivational states. Third, bite forces are strongly correlated with head size and shape. Fourth, bite forces correlate with features of prey of vertebrates. Finally, bite forces are linked to male dominance and correlated with social-display structures. Thus, bite force performance measures can be used as ‘traits’, and thus be used in integrative studies at multiple levels of organismal biology. Accordingly, bite force data will help our understanding of the functions, capacities, and evolution of jaw–cranial musculoskeletal systems. Moreover, a plethora of opportunities exist for the use of bite force measurements, and if methods are carefully applied, several levels of organismal and ecological organization can be integrated to aid our understanding of the ecology and evolution of vertebrate taxa. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 709–720.

191 citations


Journal ArticleDOI
TL;DR: It was found that the chondrocranium was the only FDM in which shape consistently had a strong and significant phylogenetic signal, implying that the utility of specific FDMs for phylogenetic inference could, in many cases, be hampered by the low reproducibility of results.
Abstract: Form and genes often tell different stories about the evolution of animals, with molecular data generally considered to be more objective than morphological data. However, form provides the basis for the description of organisms, and the study of fossils crucially depends on morphology. Complex organisms tend to evolve as ‘mosaics’, in which parts may be modified at varying rates and in response to different selective pressures. Thus, individual anatomical regions may contain different phylogenetic signals. In the present study, we used computerized methods to ‘dissect’ the skulls of a primate clade, the guenons, into functional and developmental modules (FDM). The potential of different modules as proxies for phylogenetic divergence in modern lineages was investigated. We found that the chondrocranium was the only FDM in which shape consistently had a strong and significant phylogenetic signal. This region might be less susceptible to epigenetic factors and thus more informative about phylogeny. The examination of the topology of trees from the chondrocranium suggested that the main differences evolved at the time of the radiation of terrestrial and arboreal guenons. However, phylogenetic reconstructions were found to be strongly affected by sampling error, with more localized anatomical regions (i.e. smaller/less complex FDMs) generally producing less reproducible tree topologies. This finding, if confirmed in other groups, implies that the utility of specific FDMs for phylogenetic inference could, in many cases, be hampered by the low reproducibility of results. The study also suggested that uncertainties due to sampling error may be larger than those from character sampling. This might have implications for phylogenetic analyses, which typically provide estimates of support of tree nodes based on characters but do not generally take into account the effect of sampling error on the tree topology. Nonetheless, studies of the potential of different FDMs as proxies for phylogenetic divergence in modern lineages, such as the present study, provide a framework that may help in modelling the morphological evolution of present and fossil species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 813–834. ADDITIONAL KEYWORDS: Cercopithecini – geometric morphometrics – morphological evolution – reliability and reproducibility – skull modules.

169 citations


Journal ArticleDOI
TL;DR: The pattern of use of Asteroideae pollen by the Colletes bees supports recent findings that this pollen possesses unfavourable or protective properties, which render its digestion difficult, and suggests that bees need physiological adaptations to successfully utilize it.
Abstract: To assess the pollen hosts of 60 western palaearctic bee species of the genus Colletes (Colletidae), we microscopically analysed 1336 pollen loads of collected females. Twenty-six species (43.3%) were found to be specialized at the level of plant family, subfamily or genus. Thirty-four species (56.7%) proved to be pollen generalists to varying degrees, visiting the flowers of up to 15 different plant families. Flowers of the subfamily Asteroideae (Asteraceae) are by far the most important pollen source, contributing 23.6% to the pollen-plant spectrum of the whole bee genus. The high significance of Asteroideae pollen is due to the large number of specialists: 14 Colletes species belonging to four different taxonomic groups harvest pollen exclusively or predominantly on flowers of the Asteroideae. By striking contrast, Asteroideae pollen plays only a marginal role in the diets of the pollen generalists: it was recorded in only 2.7% of the pollen loads and in seven out of the 34 pollen generalists. Among the few generalists exploiting Asteroideae for pollen, three closely related species have ancestors which were possibly specialized on Asteraceae. The pattern of use of Asteroideae pollen by the Colletes bees supports recent findings that this pollen possesses unfavourable or protective properties, which render its digestion difficult, and suggests that bees need physiological adaptations to successfully utilize it. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 719–733.

168 citations


Journal ArticleDOI
TL;DR: A simple method for assessing colour in unrestrained, wild subjects using digital photography that can be used under natural light conditions and can be readily applied to subjects in their natural habitat is presented.
Abstract: Adaptive hypotheses about colour variation are widespread in behavioural ecology, and several methods of objective colour assessment have been proposed and validated for use in a wide variety of taxa. However, to date, the most objective and reliable methods of assessing colour are not readily applied to wild animals. In the present study, we present a simple method for assessing colour in unrestrained, wild subjects using digital photography. The method we describe uses a digital camera, a colour standard, and colour analysis software, and can be used to measure any part of the visible colour spectrum. We demonstrate that the method: (1) is accurate and precise across different light conditions; (2) satisfies previous criteria regarding linearity and red, green, and blue equality; and (3) can be independently validated visually. In contrast with previous digital methods, this method can be used under natural light conditions and can be readily applied to subjects in their natural habitat. To illustrate this, we use the method to measure chest colour in wild geladas (Theropithecus gelada). Unique among primates, geladas have a red patch of skin on their chest and neck, which, for males, is thought to be a sexually selected signal. Offering some support to this hypothesis, we found differences in chest ‘redness’ for males across different age groups, with males in their reproductive prime exhibiting the reddest chests. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 231–240.

156 citations


Journal ArticleDOI
TL;DR: Expanding studies of kleptoparasitism to other taxa will differentiate between the selective pressures and evolutionary responses that are universal among kleptic symbioses and those that are restricted to certain groups.
Abstract: Understanding of the conditions encouraging the evolution of kleptoparasitism has been limited by the preponderance of attention focused on a limited number of taxa (predominantly birds). However, most animal taxa contain kleptoparasitic members. In many ways, adult birds are aberrant because most are acrobatic, capable of three-dimensional movement, and steal from hosts that are capable of similar feats. The mobility of kleptoparasites and their hosts falls along a continuum, with the acrobatic birds at one end and sessile hosts and kleptoparasites at the other. Considering sedentary systems requires amendments to the criteria previously outlined as necessary for kleptoparasitism. Kleptoparasitic interactions between agile participants are ephemeral, involve numerous hosts over the lifetime of the kleptoparasite, and usually rely on vision. Large concentrations of hosts and large, high-quality food items are important in mobile systems. By contrast, the long-term, intimate associations of sedentary interactions demand that the host not consume the kleptoparasite's offspring and that the host must either have a longer life cycle than the kleptoparasite or be located in aggregations that have staggered life cycles (so the kleptoparasite can move to a new host when the original host dies). Expanding studies of kleptoparasitism to other taxa will differentiate between the selective pressures and evolutionary responses that are universal among kleptoparasitic symbioses and those that are restricted to certain groups. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 745–762.

143 citations


Journal ArticleDOI
TL;DR: Tree topology, inferred divergence dates, palaeogeographical and palaeoclimatic data are all consistent with Miocene immigration into Australia from the north by mesic forest ecomorphs, followed by initial diversification in mesic habitats before radiation into xeric habitats facilitated by increasing aridity.
Abstract: Recent mtDNA phylogenies of Australasian agamid lizards are highly incongruent with existing morphological views. To resolve this discrepancy we sequenced two nuclear gene regions, c-mos and brain-derived neurotrophic factor (BDNF). These were highly concordant with each other and the mtDNA phylogeny, but not the morphology. A combined molecular analysis reveals substantial hidden support (additional phylogenetic signal that emerges only when the data sets interact in a combined analysis). Bayesian posteriors, and a partitioned bootstrap procedure introduced here, indicate strong support for most nodes. The resultant tree implies extensive morphological homoplasy, with many genera emerging as non-monophyletic (Amphibolurus, Rankinia, Ctenophorus, Physignathus, Diporiphora). The water and forest dragons (Physignathus and Hypsilurus) form a paraphyletic basal assemblage to the more derived Australian forms such as Amphibolurus and Ctenophorus, which include almost all the xeric taxa. However, the thorny devil Moloch horridus is a basal lineage and not closely related to the other arid forms. Tree topology, inferred divergence dates, palaeogeographical and palaeoclimatic data are all consistent with Miocene immigration into Australia from the north by mesic forest ecomorphs, followed by initial diversification in mesic habitats before radiation into xeric habitats facilitated by increasing aridity. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 343–358.

118 citations


Journal ArticleDOI
TL;DR: The study indicated that species-specific hydrocarbon profiles remained remarkably stable between Finland and the British Isles and were not influenced by ecological factors such as soil or vegetation types.
Abstract: A comparison of the cuticular hydrocarbon profiles of thirteen sympatric species of Formica ants from Finland, along with a subset of five species from the British Isles, was conducted. This permitted a comparison of the evolution of these profiles within a single genus and the stability of these profiles over large geographical and evolutionary distances. The study indicated that species-specific hydrocarbon profiles remained remarkably stable between Finland and the British Isles and were not influenced by ecological factors such as soil or vegetation types. This stability allowed candidate cuticular hydrocarbons or groups of hydrocarbons to be identified that may constitute species-specific recognition cues in these ants. In Formica, cuticular hydrocarbon composition has evolved down two distinct paths, either elevated production of Z-9-alkenes or the production of various dimethylalkanes. Evolution of hydrocarbons can be predicted for the other five Formica species for which the cuticular profiles are known. The Z-9-alkenes, dienes, and various distinctive combinations of dimethylalkanes can be used as unique species recognition cues in the Formica ants and this parallels the evolution of species-specific cuticular hydrocarbon composition in the Drosophila flies. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 131–140.

117 citations


Journal ArticleDOI
TL;DR: That's it, a book to wait for in this month; even you have wanted for long time for releasing this book the creation an appeal to save life on earth; you may not be able to get in some stress but now, the authors are coming to give you excellent solution.
Abstract: That's it, a book to wait for in this month. Even you have wanted for long time for releasing this book the creation an appeal to save life on earth; you may not be able to get in some stress. Should you go around and seek fro the book until you really get it? Are you sure? Are you that free? This condition will force you to always end up to get a book. But now, we are coming to give you excellent solution.

103 citations


Journal ArticleDOI
TL;DR: It is suggested that, during the early evolution of those branches giving rise to fully subterranean ctenomyids and octodontids, a change in behaviour probably preceded the origin of structural adaptations.
Abstract: To examine the evolution of burrowing specializations in the sister families Octodontidae and Ctenomyidae (Rodentia: Caviomorpha), we produced a synthetic phylogeny (supertree), combining both molecular and morphological phylogenies, and including both fossil and extant genera. We mapped morphological specializations of the digging apparatus onto our phylogenetic hypothesis and attempted to match morphological diversity with information on the ecology and behaviour of octodontoid taxa. Burrowing for sheltering and rearing is the rule among octodontids and ctenomyids, and adaptations for digging have been known from the Early Pliocene onward. However, only a few taxa have evolved fully subterranean habits. Scratch-digging is widespread among both semifossorial and fully subterranean lineages, and morphological changes associated with scratch-digging are not restricted to subterranean lineages. By contrast, various adaptations for chisel-tooth digging are restricted to some subterranean lineages and are combined differently in the octodontid Spalacopus, the fossil ctenomyid Eucelophorus, and some living Ctenomys. Some octodontid taxa are able to dig complex burrows in spite of having no substantial changes in musculoskeletal attributes. Hence, we suggest that, during the early evolution of those branches giving rise to fully subterranean ctenomyids and octodontids, a change in behaviour probably preceded the origin of structural adaptations. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 267–283. ADDITIONAL KEYWORDS: behaviour – Ctenomyidae – evolutionary morphology – Octodontidae – subterranean niche.

97 citations


Journal ArticleDOI
TL;DR: The results suggest that more than 100 bat species occur sympatrically at TBS and about 50 bat species coexist at BOM, and assemblage structure of these three phyllostomid assemblages was influenced by differences in floral diversity at the three sites.
Abstract: We compared the assemblages of phyllostomid bats in three Neotropical rainforests with respect to species richness and assemblage structure and suggested a method to validate estimates of species richness for Neotropical bat assemblages based on mist-netting data. The fully inventoried bat assemblage at La Selva Biological Station (LS, 100 m elevation) in Costa Rica was used as a reference site to evaluate seven estimators of species richness. The Jackknife 2 method agreed best with the known bat species richness and thus was used to extrapolate species richness for an Amazonian bat assemblage (Tiputini Biodiversity Station; TBS, 200 m elevation) and an Andean premontane bat assemblage (Podocarpus National Park; BOM, 1000 m elevation) in Ecuador. Our results suggest that more than 100 bat species occur sympatrically at TBS and about 50 bat species coexist at BOM. TBS harbours one of the most species-rich bat assemblages known, including a highly diverse phyllostomid assemblage. Furthermore, we related assemblage structure to large-scale geographical patterns in floral diversity obtained from botanical literature. Assemblage structure of these three phyllostomid assemblages was influenced by differences in floral diversity at the three sites. At the Andean site, where understorey shrubs and epiphytes exhibit the highest diversity, the phyllostomid assemblage is mainly composed of understorey frugivores and nectarivorous species. By contrast, canopy frugivores are most abundant at the Amazonian site, coinciding with the high abundance of canopy fruiting trees. Assemblage patterns of other taxonomic groups also may reflect the geographical distribution patterns of floral elements in the Andean and Amazonian regions. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 617–629.

96 citations


Journal ArticleDOI
TL;DR: Three methods for delineating species boundaries within the cosmopolitan nematode species Rhabditis (Pellioditis) marina and Halomonhystera disjuncta highlight that classical taxonomy grossly underestimates species diversity within the phylum Nematoda.
Abstract: Integrative taxonomy considers species boundaries from multiple, complementary perspectives, with the main objective being to compare the observed data against the predictions of the methodologies used. In the present study we used three methods for delineating species boundaries within the cosmopolitan nematode species Rhabditis (Pellioditis) marina and Halomonhystera disjuncta. First, phylogenetic relationships among molecular sequences from the mitochondrial cytochrome oxidase c subunit 1 gene (COI), and from two nuclear regions, internal transcribed spacer (ITS) and D2D3, were analysed. Subsequently, multivariate morphometric analysis was used to investigate whether concordant molecular lineages were also morphologically distinct. When morphological differences were found, typological taxonomy was performed to identify fixed or non-overlapping characters between lineages. Interbreeding experiments were conducted between the two closest related lineages of R. (P.) marina to investigate potential reproductive isolation. This integrative approach confirmed the presence of several species within each nominal species: molecular lineages were concordant across two independent loci (COI and ITS), and were characterized by significant morphological divergence. Most lineages were also detectable in the D2D3 region, but were less resolved. The two lineages investigated in our study did not produce offspring. Our results highlight that classical taxonomy grossly underestimates species diversity within the phylum Nematoda. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 737–753.

Journal ArticleDOI
TL;DR: All alpine and arctic-alpine disjunction types of the western Palearctic are listed using butterflies and moths as a model group and hypotheses of the distribution patterns during the last ice age are formulated.
Abstract: The cyclic changes of the Pleistocene between cold and warm periods resulted in antagonistic responses within two different groups of organisms: one expanding during the warm periods and retracting during the cold phases and another with opposed responses. The latter group is composed of so-called arctic and alpine species. These species have recently become the focus of phylogeographical research. However, we still lack a comprehensive characterization of the different types of alpine and arctic-alpine disjunctions in the western Palearctic. Such an overview might facilitate the selection of different model species to test the different patterns of disjunctions for congruences revealing their past distribution. Therefore, we list all alpine and arctic-alpine disjunction types of the western Palearctic using butterflies and moths as a model group. We distinguish between (1) endemics of the Alps (α) with strongly restricted and (β) with broader distributions; (2) alpine disjunct species (α) with perialpine disjunctions and (β) with a wide western Palearctic distribution; (3) oro-Mediterranean species; and (4) arctic-alpine disjunct species. These types of distributions and further subtypes are exemplified using chorological data of butterfly and moth species. In this context, we discuss the intraspecific differentiation and the differentiation among sibling-species within and among disjunct parts of the distribution area. We also formulate hypotheses of the distribution patterns during the last ice age which might best explain the actual patterns. Finally, we suggest some case studies of genetic analyses to test the above mentioned hypotheses. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 415–430.

Journal ArticleDOI
TL;DR: It is demonstrated that the widespread IMJ occurrence among extant biters to result from a complex convergent evolutionary history, indicating that the IMJ is a major functional innovation that enhances biting strategies in several prominent reef fish groups.
Abstract: 3 We investigated the functional morphology and ecology of biting among the squamipinnes, an assemblage of nine successful and distinctive reef fish families. We demonstrate that an intramandibular joint (IMJ) may have evolved at least three and possibly five times in this assemblage and discuss the impact of this recurring innovation in facilitating prey-capture by biting. Using character mapping on a supertree for the squamipinnes, we reveal up to seven gains or losses of intramandibular flexion, all associated with trophic transitions between free-living and attached prey utilization. IMJs are basal in six of the studied families whereas the origin of intramandibular flexion in the Chaetodontidae (butterflyfishes) coincides with a transition from ram-suction feeding to benthic coral feeding, with flexion magnitude reaching its peak (49 ± 2.7°) in the coral scraping subgenus Citharoedus. Although IMJs generally function to augment vertical gape expansion during biting behaviours to remove small inverte- brates, algae or coral from the reef, the functional ecology of IMJs in the Pomacanthidae (angelfishes) stands in contrast. Pomacanthid IMJs exhibit over 35° of flexion, permitting gape closure when the jaws are fully protruded. We demonstrate the widespread IMJ occurrence among extant biters to result from a complex convergent evolutionary history, indicating that the IMJ is a major functional innovation that enhances biting strategies in several prominent reef fish groups. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 545-555.

Journal ArticleDOI
TL;DR: In this paper, two proposed colonization routes to the northern Philippines (from mainland Asia and Palawan) are difficult to differentiate from each other, except on the basis of outgroup, as are the two routes from the south (Sulu archipelago and Sulawesi).
Abstract: An unusual geological setting and a high level of endemism makes the Philippine islands of great interest to biogeography. These islands lie adjacent to the continental (Sunda) shelf, yet the majority of the islands have never been directly attached to the mainland. Existing hypotheses of colonization and diversification processes have been tested across multiple taxa, but only in ahistorical contexts. We present explicit phylogenetic predictions based on these hypotheses, and then test them using new molecular datasets for four Philippine birds. Two proposed colonization routes to the northern Philippines (from mainland Asia and Palawan) are difficult to differentiate from each other, except on the basis of outgroup, as are the two routes from the south (Sulu archipelago and Sulawesi). We find unique colonization and diversification patterns for each taxon. These results contrast with expectations from Pleistocene geography, with Luzon and Mindoro indistinguishable genetically, and Negros and Panay are often nonsister taxa. Combining these data with a literature search for studies addressing these patterns, Philippine organisms shows some evidence for each proposed colonization route but the greatest support is for the two routes from Borneo. Many taxa exhibit multiple colonization events using several of these routes, contrasting with past assumptions of single colonization events. Island-by-island differentiation within the Philippines follows, with patterns reflecting colonization patterns rather than Pleistocene geography, particularly for highland species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 620–639.

Journal ArticleDOI
TL;DR: 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.
Abstract: 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.

Journal ArticleDOI
TL;DR: A combined approach including molecular phylogeny, molecular dating, and character optimization analysis was applied to study Prosopis diversification and evolution, suggesting that prosopis is not a natural group.
Abstract: The genus Prosopis is an important member of arid and semiarid environments around the world. To study Prosopis diversification and evolution, a combined approach including molecular phylogeny, molecular dating, and character optimization analysis was applied. Phylogenetic relationships were inferred from five different molecular markers (matK-trnK, trnL-trnF, trnS-psbC, G3pdh, NIA). Taxon sampling involved a total of 30 Prosopis species that represented all Sections and Series and the complete geographical range of the genus. The results suggest that Prosopis is not a natural group. Molecular dating analysis indicates that the divergence between Section Strombocarpa and Section Algarobia plus Section Monilicarpa occurred in the Oligocene, contrasting with a much recent diversification (Late Miocene) within each of these groups. The diversification of the group formed by species of Series Chilenses, Pallidae, and Ruscifoliae is inferred to have started in the Pliocene, showing a high diversification rate. The moment of diversification within the major lineages of American species of Prosopis is coincident with the spreading of arid areas in the Americas, suggesting a climatic control for diversification of the group. Optimization of habitat parameters suggests an ancient occupation of arid environments by Prosopis species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 621–640.

Journal ArticleDOI
TL;DR: The Pleistocene predator guild appears to have been characterized by relatively high levels of competition that are rarely observed today, although some extant species have fracture frequencies that approach those of extinct species.
Abstract: 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

Journal ArticleDOI
TL;DR: A detailed study of variation in an anadromous stickleback population provides crucial insight into a key component of adaptive radiation, the variation on which directional selection acts at the onset of the radiation.
Abstract: Ancestral properties can influence patterns of evolutionary diversification, but ancestors can rarely be observed directly. We examined variation and sexual dimorphism of morphological traits in an anadromous threespine stickleback population representing the ancestral form for resident postglacial stickleback populations in the area. A combination of traditional and geometric morphometric methods were used to study variation over multiple years in an anadromous population that breeds in Rabbit Slough, Cook Inlet, Alaska. Major armor anomalies were extremely rare but their occurrence at measurable frequencies suggests that significant standing variation for armor phenotypes exists in anadromous populations. Sexual dimorphism was a major source of variation, and most traits differed significantly between sexes, particularly head length, length of the pelvic girdle, and body shape. Consequently, some degree of sexual dimorphism appears to be the ancestral condition for many traits in derived resident freshwater stickleback radiations. Morphological variation among years, especially in body shape, was significant in both sexes, but the magnitude of annual variation was always less than variation due to other factors. Phenotypic means were relatively stable over short time scales. Postglacial stickleback radiations are among the most enlightening cases of adaptive radiation, and our detailed study of variation in an anadromous stickleback population provides crucial insight into a key component of adaptive radiation, the variation on which directional selection acts at the onset of the radiation. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 465–478. ADDITIONAL KEYWORDS: Alaska – ancestral variation – evolutionary diversification – Gasterosteus aculeatus – geometric morphometrics.

Journal ArticleDOI
TL;DR: The data support an origin for pythons outside Australia, followed by a radiation into Australia during the mid-Tertiary, and strong support for elevated speciation rates during the period when Australia collided with the proto-Indonesian archipelago.
Abstract: We used nucleotide sequences from four mitochondrial genes and structural features of the mitochondrial control region, combined with a revised, previously published, morphological data set to infer phylogenetic relationships among the pythons. We aimed to determine which of two competing hypotheses of relationships of the genera Aspidites and Python best explains the evolutionary and bioegeographical history of the family. All analyses of the combined data recover a set of relationships in which (1) the genus Python is paraphyletic with the two east Asian species, P. reticulatus and P. timoriensis, as the sister lineage to the seven Australo-Papuan python genera. We support recognition of a distinct genus for the P. reticulatus + P. timoriensis clade; (2) the remaining species of the genus Python form a clade which is the sister lineage to the remainder of the family; (3) the genus Aspidites is embedded among the Australo-Papuan genera. The seemingly primitive characteristics of Aspidites may be better interpreted as reversals or specializations that have accompanied a switch to burrowing in this genus. Resolution of the relationships among the Australo-Papuan lineages is weak, possibly because of rapid diversification early in the history of the radiation. We assessed the tempo of the Indo-Australian python radiation using a maximum likelihood framework based on the birth‐death process. We find strong support for elevated speciation rates during the period when Australia collided with the proto-Indonesian archipelago. The data support an origin for pythons outside Australia, followed by a radiation into Australia during the mid-Tertiary. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 603‐619.

Journal ArticleDOI
TL;DR: This paper attempts to connect some opposing views about speciation by integrating Mayr's ideas about the roles of ecology and geography in speciation with current data and theory.
Abstract: Mayr's best recognized scientific contributions include the biological species concept and the theory of geographic speciation. In the latter, reproductive isolation evolves as an incidental by-product of genetic divergence between allopatric populations. Mayr noted that divergent natural selection could accelerate speciation, but also argued that gene flow so strongly retards divergence that, even with selection, non-allopatric speciation is unlikely. However, current theory and data demonstrate that substantial divergence, and even speciation, in the face of gene flow is possible. Here, I attempt to connect some opposing views about speciation by integrating Mayr's ideas about the roles of ecology and geography in speciation with current data and theory. My central premise is that the speciation process (i.e. divergence) is often continuous, and that the opposing processes of selection and gene flow interact to determine the degree of divergence (i.e. the degree of progress towards the completion of speciation). I first establish that, in the absence of gene flow, divergent selection often promotes speciation. I then discuss how population differentiation in the face of gene flow is common when divergent selection occurs. However, such population differentiation does not always lead to the evolution of discontinuities, strong reproductive isolation, and thus speciation per se. I therefore explore the genetic and ecological circumstances that facilitate speciation in the face of gene flow. For example, particular genetic architectures or ecological niches may tip the balance between selection and gene flow strongly in favour of selection. The circumstances allowing selection to overcome gene flow to the extent that a discontinuity develops, and how often these circumstances occur, are major remaining questions in speciation research. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 26–46.

Journal ArticleDOI
TL;DR: Mitochondrial (mt)DNA sequence divergences do not support hypotheses for Pleistocene speciation and instead suggest much more ancient speciation events in the genus Skistodiaptomus, and clear evidence of CO I pseudogenes is found in S. pygmaeus, but their presence did not lead to significant overestimates of sequence divergs for this gene.
Abstract: We used mitochondrial [cytochrome c oxidase subunit I (CO I), cytochrome b, and 16S] and nuclear [internal transcribed spacer (ITS) phylogenies of Skistodiaptomus copepods to test hypotheses of Pleistocene divergence and speciation within the genus. Mitochondrial (mt)DNA sequence divergences do not support hypotheses for Pleistocene speciation and instead suggest much more ancient speciation events in the genus. Skistodiaptomus oregonensis and Skistodiaptomus pygmaeus (i.e. two morphologically similar and parapatric species) exhibited uncorrected mtDNA sequence divergences exceeding 20%. Similarly, we identified three divergent clades of Skistodiaptomus pallidus that exhibited mtDNA sequence divergences exceeding 15%, suggesting that even intraspecific divergence within this morphospecies predates the Pleistocene. We found clear evidence of CO I pseudogenes in S. pygmaeus, but their presence did not lead to significant overestimates of sequence divergences for this gene. Substitution saturation and strong purifying selection have most likely led to underestimates of sequence divergences and divergence times among Skistodiaptomus. The widespread phenomenon of morphological stasis among genetically divergent copepod groups indicates that speciation often occurs with little or no morphological change. Instead, morphological evolution may occur idiosyncratically after speciation and create discordant patterns of morphological similarity, shared ancestry and divergence time. Cryptic species complexes are therefore common in copepods, and morphological species concepts underestimate their true species diversity. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 150‐165. ADDITIONAL KEYWORDS: cryptic species ‐ genitalia ‐ intrinsic incompatibility ‐ molecular clocks ‐ molecular phylogeny ‐ morphological stasis ‐ Pleistoscenario ‐ pseudogenes ‐ reproductive isolation ‐ sexual selection.

Journal ArticleDOI
TL;DR: Remarkably, head length, eye length, and head volume decreased in size, whereas cheek depth increased, and total gill surface in resurgent H. pyrrhocephalus increased by 64%.
Abstract: During the upsurge of the introduced predatory Nile perch in Lake Victoria in the 1980s, the zooplanktivorous Haplochromis (Yssichromis) pyrrhocephalus nearly vanished. The species recovered coincident with the intense fishing of Nile perch in the 1990s, when water clarity and dissolved oxygen levels had decreased dramatically due to increased eutrophication. In response to the hypoxic conditions, total gill surface in resurgent H. pyrrhocephalus increased by 64%. Remarkably, head length, eye length, and head volume decreased in size, whereas cheek depth increased. Reductions in eye size and depth of the rostral part of the musculus sternohyoideus, and reallocation of space between the opercular and suspensorial compartments of the head may have permitted accommodation of larger gills in a smaller head. By contrast, the musculus levator posterior, located dorsal to the gills, increased in depth. This probably reflects an adaptive response to the larger and tougher prey types in the diet of resurgent H. pyrrhocephalus. These striking morphological changes over a time span of only two decades could be the combined result of phenotypic plasticity and genetic change and may have fostered recovery of this species.

Journal ArticleDOI
TL;DR: The results suggest that, although there is large scale genetic differentiation for two species, patterns of population connectivity are remarkably consistent within major regions and do not reflect variation in major oceanographic processes or genetic discontinuity coincident with biogeographic boundaries.
Abstract: Extreme variation in early life-history strategies is considered a moderately good predictor of genetic subdivision and hence dispersal for a range of marine species. In reality, however, a good deal of population differentiation must reflect historical effects, more subtle variation in life histories, and, particularly, the interaction of larvae with oceanographic processes. Using a combination of allozyme and microsatellite markers, we show that the large-scale genetic structure of populations of three species (direct and planktonically developing cushion stars and a planktonic developing sea anemone that is also asexually viviparous) varies consistently, in line with the predicted capacity for dispersal within three geographic regions. We detected high levels of genetic subdivision for the direct developing cushion star (FST = 0.6), low levels for the planktonically developing cushion star (FST = 0.009), and intermediate levels for the sexual/asexual sea anmone (FST = 0.19). These patterns are exhibited despite the highly variable patterns of current movement and the presence of biogeographic barriers. Our results suggest that, although there is large scale genetic differentiation for two species, patterns of population connectivity are remarkably consistent within major regions and do not reflect variation in major oceanographic processes or genetic discontinuity coincident with biogeographic boundaries.

Journal ArticleDOI
TL;DR: Within-species evidence is found that steeper dominance hierarchies were associated with more grooming being directed up the hierarchy, and that a trade-off occurred between the tendency to groom up the hierarchies and the degree of grooming reciprocation.
Abstract: Primates may trade altruistic behaviours, such as grooming, either for itself or for different rank-related benefits, such as tolerance or agonistic support. Ecological conditions are expected to affect competition and thus the steepness of dominance hierarchies. This, in turn, may influence the value of the different currencies that primates exchange. Thus, it can be hypothesized that, as the dominance hierarchy becomes steeper, more grooming is directed up the hierarchy (in exchange for tolerance or agonistic support) and less grooming is exchanged for other grooming. We assembled a large database of within-group grooming distribution in primates (38 social groups belonging to 16 species and eight genera) and tested these hypotheses both within species (i.e. comparing different groups of the same species) and between species (using comparative methods that control for phylogenetic relatedness). We found within-species evidence that steeper dominance hierarchies were associated with more grooming being directed up the hierarchy, and that a trade-off occurred between the tendency to groom up the hierarchy and the degree of grooming reciprocation (although, in some analyses, only a nonsignificant trend was observed). By contrast, phylogenetically controlled comparisons between species did not reveal evidence of correlated evolution between the steepness of the dominance hierarchy, the tendency to direct grooming up the hierarchy, and the degree of grooming reciprocation.

Journal ArticleDOI
TL;DR: This is the most inclusive phylogenetic analysis of Arvicolinae (Mammalia, Rodentia) to date, combining all published cytochrome b gene sequences of greater than 1097 bp and new sequences from two monotypic genera.
Abstract: Comprehensive taxonomic sampling can vastly improve the accuracy of phylogenetic reconstruction. Here, we present the most inclusive phylogenetic analysis of Arvicolinae (Mammalia, Rodentia) to date, combining all published cytochrome b gene sequences of greater than 1097 bp and new sequences from two monotypic genera. Overall, the phylogenetic relationships between 69 species of voles and lemmings, representing 18 genera and 10 tribes, were studied. By applying powerful modern approaches to phylogenetic reconstruction, such as maximum likelihood and Bayesian analysis, we provide new information on the early pulse of evolution within the Arvicolinae. While the position of two highly divergent lineages, Phenacomys and Ondatra, could not be resolved, the tribe Lemmini, appeared as the most basal group of voles. The collared lemmings (Dicrostonychini) grouped together with all of the remaining tribes. The two previously unstudied monotypic genera Dinaromys and Prometheomys form a moderately well-supported monophyletic clade, possibly a sister group to Ellobius (Ellobiusini). Furthermore, with one exception, all tribes (sensu Musser & Carleton, 2005) proved to be monophyletic and can thus be regarded as meaningful evolutionary entities. Only the tribe Arvicolini emerged as paraphyletic in both analyses because of the unresolved phylogenetic position of Arvicola terrestris. Steppe voles of the genus Lagurus were solidly supported as a sister group to the Microtus and allies clade. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 825–835.

Journal ArticleDOI
TL;DR: Using recent advances in plant and butterfly systematics to trace the evolution of host-plant utilization in the butterfly subfamily Nymphalinae, a clear historical pattern emerges, with an ancestral host- plant theme of 'urticalean rosids' and two major colonizations of novel distantly-related plant clades.
Abstract: How and why the great diversity of phytophagous insects has evolved is not clear but, if the explanation is the diversity of plants as a resource, colonizations of novel plant taxa can be expected to be associated with higher net speciation rates. In the present study, we make use of recent advances in plant and butterfly systematics to trace the evolution of host-plant utilization in the butterfly subfamily Nymphalinae (tribes Nymphalini, Melitaeini, and the probably paraphyletic 'Kallimini'). A clear historical pattern emerges, with an ancestral host-plant theme of 'urticalean rosids' and two major colonizations of novel distantly-related plant clades. The asterid order Lamiales was colonized by an ancestor of 'Kallimini' + Melitaeini and the family Asteraceae in Asterales was later colonized by Melitaeini butterflies. These colonization events appear to have been followed by increases in the rate of net butterfly diversification. Two not mutually exclusive scenarios to explain such patterns have been suggested: (1) adaptive radiation due to release from competition following host-plant shifts or (2) higher rates of net speciation during a relatively long-lasting potentially polyphagous (plastic) state. In support of the 'plasticity scenario', phylogenetic traces of a long-lasting stage with some potential to feed on more than one host-plant clade can still be seen, despite the ancient age of the colonizations. When angiosperm communities changed after the K/T boundary due to extinctions and subsequent diversification, herbivore taxa that could occupy several alternative niches may have had the greatest opportunity to diversify in turn.

Journal ArticleDOI
TL;DR: Congruence between the acoustic affinities among species and that obtained by means of morphological and genetic data is suggested, and first hypotheses on the evolution of acoustic communication and the associated mechanisms in this fish group are suggested.
Abstract: Sound production of 11 Mediterranean goby species, belonging to five different genera, have been comparatively analysed on the basis of the quantitative properties of the acoustic signal emitted by the male in both the reproductive and aggressive context The results obtained showed that three groups of species can be recognized on the basis of signal similarity: the larger sized species (genus Padogobius and Gobius paganellus) producing tonal sounds, showing high values of pulse rate and low values of duration; the larger-sized species producing grunt sounds (genus Gobius and Zosterisessor) with low pulse rate and low duration; and the small-sized species producing grunt sounds (genus Pomatoschistus and Knipowitschia) with low pulse rate and high duration The comparison between these results and those found in previous studies suggests congruence between the acoustic affinities among species and that obtained by means of morphological and genetic data Furthermore, first hypotheses on the evolution of acoustic communication and the associated mechanisms in this fish group are suggested © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 763–778

Journal ArticleDOI
TL;DR: The results support the argument that the level of phenotypic plasticity itself can be under selection and that differentiation in reaction norms can occur even in neighbouring habitats with no barrier to gene flow.
Abstract: When populations experience substantial variation in environmental conditions, they may evolve phenotypic plasticity in response to these varying selection pressures. Evolutionary theory predicts differentiation in the level of phenotypic plasticity among different habitats. We evaluated temperature-induced phenotypic responses in juvenile growth rate in natural populations of the springtail Orchesella cincta, inhabiting forest and heathland. These habitats typically co-occur but differ strongly with respect to, for example, thermal regime, relative humidity, and structure. Offspring of females from the two habitats were reared at different temperatures in climate rooms and the temperature response of juvenile growth rate and egg size was measured. We found a habitat-specific difference in plasticity of juvenile growth rate. The reaction norms of the forest populations were steeper than the reaction norms for heath populations at two replicated sampling sites. Egg weight itself was demonstrated to be a plastic trait with a higher egg weight at low temperatures, but the thermal response did not differ between habitats. We conclude that these populations have diverged due to strong local natural selection. Our results support the argument that the level of phenotypic plasticity itself can be under selection and that differentiation in reaction norms can occur even in neighbouring habitats with no barrier to gene flow. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 265‐271.

Journal ArticleDOI
TL;DR: The obtained molecular data clearly support the subdivision of S. paniculata into three subspecies: ssp.
Abstract: The molecular biogeography of the disjunctly distributed and morphologically highly variable species Saxifraga paniculata Mill. was analysed using amplified fragment length polymorphism (AFLP) and chloroplast microsatellites. The study comprised 77 samples from mountain regions in Europe and North America throughout the complete range of distribution. AFLP data revealed clear genetic differentiation between samples from the Arctic, the Caucasus, and the eastern European mountains. Samples from the Alps were divided into two groups. One group clustered with the samples from central Europe and the Pyrenees, whereas another group with individuals from southern Norway. AFLP diversity was lowest in the Arctic and highest in the Alps. Chloroplast microsatellite analysis revealed eight haplotypes but no unequivocal phylogeographical pattern. However, haplotype diversity was highest in the Alps and central Europe whereas, in the Arctic, only few widespread haplotypes could be found. The results indicate in situ survival of S. paniculata in the Caucasus, the eastern European mountains, and the Alps. The Arctic has presumably been colonized postglacially from North American refugia south of the ice shield. Southern Norway and the Pyrenees have most likely been colonized from two phylogeographically different groups in the Alps. The origin of the central European samples remains ambiguous. In situ survival seems to be as possible as several postglacial recolonization events from the Alps. The obtained molecular data clearly support the subdivision of S. paniculata into three subspecies: ssp. cartilaginea from the Caucasus, ssp. laestadii from northern Norway, Iceland, and North America, and ssp. paniculata from the other geographical regions. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 93, 385–398.

Journal ArticleDOI
TL;DR: E Ernst Mayr's arguments were more convincing than Huxley's, and this caused a delay in the acceptance of a more balanced view of speciation for many decades, so it is only now, with new molecular evidence, that the authors are beginning to appreciate more fully the expected Darwinian intermediates between coexisting species.
Abstract: We commonly read or hear that Charles Darwin successfully convinced the world about evolution and natural selection, but did not answer the question posed by his most famous book, ‘On the Origin of Species …’. Since the 1940s, Ernst Mayr has been one of the people who argued for this point of view, claiming that Darwin was not able to answer the question of speciation because he failed to define species properly. Mayr undoubtedly had an important and largely positive influence on the study of evolution by stimulating much evolutionary work, and also by promoting a ‘polytypic species concept’ in which multiple, geographically separated forms may be considered as subspecies within a larger species entity. However, Mayr became seduced by the symmetry of a pair of interlocking ideas: (1) that coexistence of divergent populations was not possible without reproductive isolation and (2) reproductive isolation could not evolve in populations that coexist. These beliefs led Mayr in 1942 to reject evidence of the importance of intermediate stages in speciation, particularly introgression between hybridizing species, which demonstrates that complete reproductive isolation is not necessary, and the existence of ecological races, which shows that ecological divergence can be maintained below the level of species, in the face of gene flow. Mayr's train of thought led him to the view that Darwin misunderstood species, and that species were fundamentally different from subspecific varieties in nature. Julian Huxley, reviewing similar data at the same time, came to the opposite conclusion, and argued that these were the intermediate stages of speciation expected under Darwinism. Mayr's arguments were, however, more convincing than Huxley's, and this caused a delay in the acceptance of a more balanced view of speciation for many decades. It is only now, with new molecular evidence, that we are beginning to appreciate more fully the expected Darwinian intermediates between coexisting species. © The Author. Journal compilation © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 3–16.