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Showing papers in "Biological Journal of The Linnean Society in 2002"


Journal ArticleDOI
TL;DR: A massive collecting effort involving 400 day-persons at 42 discrete stations on a 295-km2 site on the west coast of New Caledonia, south-west Pacific, revealed 2738 species of marine molluscs, several times the number of species recorded from any area of comparable extension anywhere in the world.
Abstract: Earlier studies in the tropical Indo-Pacific have grossly underestimated the richness of macrofauna species at spatial scales relevant to conservation and management as a result of insufficient collecting and sorting effort. A massive collecting effort involving 400 day-persons at 42 discrete stations on a 295-km2 site on the west coast of New Caledonia, south-west Pacific, revealed 2738 species of marine molluscs. This is several times the number of species recorded from any area of comparable extension anywhere in the world. Spatial and habitat heterogeneity is high with 32% of the species collected at a single station. With 20% of the species represented by single specimens (0.4% of all catches), rare species make up a considerable proportion of the fauna. This justifies the parallel drawn between coral reefs and rain forests in terms of species diversity. The real richness of many soft-bodied marine taxa is probably underestimated, as evidenced by the fact that 28.5% of the mollusc species present at the study site are represented in the samples only by empty shells. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 421–436.

395 citations


Journal ArticleDOI
TL;DR: It is argued that so many exceptions to these assumptions have been demonstrated that they cannot be used with any reliability to distinguish different species, and alternative approaches towards solving the species problems normally solved using genetic distance are explored.
Abstract: Degree of genetic divergence is frequently used to infer that two populations belong to separate species, or that several populations belong to a single species. I explore the logical framework of this approach, including the following assumptions: (i) speciation takes place over very long periods of time; (ii) reproductive isolation is based on the slow accumulation many genetic differences throughout the genome; (iii) genetic divergence automatically leads to reproductive isolation between species; and (iv) pre-mating and post-mating reproductive isolation have a similar genetic basis. I argue that so many exceptions to these assumptions have been demonstrated that they cannot be used with any reliability to distinguish different species. In addition, genetic distance as a species criterion is mostly used within the framework of Mayr's Biological Species Concept and is not free of assumptions about the nature of species or of speciation. The use of genetic distance to infer separate species (or the lack of these) is not parsimonious, its theoretical foundations are not well understood, and it cannot be applied over a wide range of plants and animals. I explore alternative approaches towards solving the species problems normally solved using genetic distance. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 509–516.

259 citations


Journal ArticleDOI
TL;DR: Long-term monitoring and further understanding of GF are essential for the conservation of this unique and diverse tropical forest in south-east Asia, especially in a period of global climatic change.
Abstract: The general flowering (GF) events of forests in south-east Asia are perhaps the most spectacular phenomena in tropical biology. GF events occur at multiyear intervals. In GF, most dipterocarp species and many plants of other families come into flower and set fruit massively; these species and plants rarely flower except during GF events. GF is unique, because it can occur over thousands of kilometers and involve hundreds of plant species representing diverse families and lifeforms. It also involves strict mast fruiting. Satiation of generalist seed predators has been considered a primary force for GF. However, recent observations indicate that several selective agents rather than a single major factor may shape GF. In addition to the satiation of generalist predators, promotion of pollination could be one of the selective factors for GF, since synchronized flowering of many species causes an increase in pollinator activity through immigration and population growth. Although environmental prediction for better establishment of seedlings may also be involved in GF, no field data have been reported to support this idea. Long-term monitoring and further understanding of GF are essential for the conservation of this unique and diverse tropical forest in south-east Asia, especially in a period of global climatic change. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 233–247.

218 citations


Journal ArticleDOI
TL;DR: This work investigated the geographical extent of the savannah vegetation and the positions of the refugia, using a combination of palynonlogical, geological, fossil and termite data to show that during the Last Glacial Maximum, most of Thailand, Peninsula Malaysia, western and southern Borneo, eastern and southern Sumatra, and Java were probably covered by savannah.
Abstract: In south-east Asia, during Quaternary glaciations, increased seasonality and sea level decreases of ~120 m caused considerable fragmentation of the rainforest. It is believed that most of the region's core was covered with savannah vegetation, and rainforest was confined to a few refugia. We investigated the geographical extent of the savannah vegetation and the positions of the refugia, using a combination of palynonlogical, geological, fossil and termite data. Our results show that during the Last Glacial Maximum, most of Thailand, Peninsula Malaysia, western and southern Borneo, eastern and southern Sumatra, and Java were probably covered by savannah. Rainforest refugia were probably present in northern and eastern Borneo, northern and western Sumatra and the Mentawai islands. We also discuss a possible complete history of the region. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 453–466.

217 citations


Journal ArticleDOI
TL;DR: Certain leaf structural traits may potentially be used to predict the functional structure of herbivorous insect assemblages, and are significantly negatively correlated with specific leaf weight, lamina and cuticle thickness, vascular tissue depth and stomate length.
Abstract: The functional composition of herbivorous insect assemblages was correlated with aspects of new and mature leaf surface features, anatomy and morphology across 18 co-occurring plant species. Multivariate analyses of insects and leaf traits revealed that the functional composition of the herbivore assemblage was more strongly correlated with leaf structural traits than with leaf constituents. Leaf traits were more strongly correlated with the functional composition of the herbivore assemblage than with its taxonomic composition. Densities of sessile phloem feeders, ­rostrum chewers, and all herbivores were significantly negatively correlated with specific leaf weight, lamina and cuticle thickness, vascular tissue depth and stomate length, and were significantly positively correlated with stomate density. External chewer densities were significantly negatively correlated with percent lignified vein area, and significantly positively correlated with leaf surface area and the distance between lignified tissues. Spine-like leaves were associated with significantly lower densities of sessile phloem feeders, external chewers and all herbivores compared to kite leaves (kite leaves are comprised of unfortified leaf tissue supported by a framework of vascular tissue). The presence of a thickened leaf hypodermis was associated with significantly lower densities of external chewers and rostrum chewers, while midrib protection was associated with significantly lower densities of external chewers. Leaf structural traits may not be the proximal factors influencing herbivorous insects, as leaf structural traits are correlated with many other plant traits such as photosynthetic rate, relative growth rate and leaf life-span. Nonetheless, these results indicate that certain leaf structural traits may potentially be used to predict the functional structure of herbivorous insect assemblages. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 43–65.

169 citations


Journal ArticleDOI
TL;DR: The analyses of morphometric and allozyme data indicate that most previously-recognized Philippine and Bornean subspecies of the Rana signata complex should be recognized as full species in appreciation of their status as independent evolutionary lineages, but none provides support for the monophyly of its Philippine members.
Abstract: We addressed the evolutionary relationships and biogeographical patterns of a model organism of low relative dispersal ability by electrophoretically assaying the products of 42 presumptive gene loci in Philippine and Bornean members of the Rana signata complex of SE Asian stream frogs. Utilizing three distantly related species of ranid frogs to deeply root trees consisting of five more closely-related species and six in-group species of the Rana signata complex, we conducted phylogenetic analyses that produced concordant topologies, regardless of the data coding strategy employed. All analyses support the hypothesis of monophyly for the Rana signata complex on the whole, but none provides support for the monophyly of its Philippine members. Our analyses of morphometric and allozyme data (along with biogeographical information) indicate that (1) most previously-recognized Philippine and Bornean subspecies of the Rana signata complex should be recognized as full species in appreciation of their status as independent evolutionary lineages; (2) Rana picturata Boulenger (until very recently included in the synonymy of Rana signata signata) is deserving of specific rank; (3) the Mindoro Isl. (Philippine) population, previously confused with Rana signata similis of Luzon Isl. is a new species; (4) two major clades (((R. signata (R. grandocula + R. similis)) + (R. picturata (R. mangyanum + R. moellendorffi))) of Bornean + Philippine lineages are recognized, corresponding to two separate faunal exchanges between the Philippines and the edge of the Sunda Shelf; (5) invasions of the oceanic portions of the Philippine islands from the Sunda Shelf have occurred along both the eastern (Sulus–Mindanao–Samar–Leyte–Luzon) arc and the western (Palawan–Busuanga–Mindoro) island arcs; (6) northern reaches of Wallace’s Line (as modified by Huxley) include exceptions to an otherwise discrete faunal separation. These results suggest the need for revision of this biogeographical barrier, increased recognition of temporal patterns of island connectedness and geographical proximity, and/or a greater appreciation of dispersal abilities of ranid frogs. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76, 393–461.

151 citations


Journal ArticleDOI
TL;DR: Hatchlings from the dry and yolk-removed treatments were significantly smaller than those from the wet treatment, however, neither performance nor survival were affected by moisture or yolk removal.
Abstract: To understand the process of natural selection, relationships between phenotype and fitness and sources of phenotypic variation must be known. We examined the importance of incubation moisture conditions, maternal yolk investment, and clutch (genotype) to phenotypic variation in hatchlings of the lizard Sceloporus undulatus. Eggs were distributed among two moisture treatments and a third treatment in which yolk was removed. After hatching, mass, snout‐vent length, tail length, body shape, thermal preference, running speed, desiccation rate, and growth rate were measured for each hatchling in the laboratory. Hatchlings were then released at a field site in order to monitor growth and survival under natural conditions. Hatchlings from the dry and yolk-removed treatments were significantly smaller than those from the wet treatment. However, neither performance nor survival were affected by moisture or yolk removal. All phenotypes were affected by clutch. Clutches that produced relatively large hatchlings had higher survival than clutches that produced relatively small hatchlings. Furthermore, clutches that produced relatively slow growing individuals and fast runners had higher survival rates than clutches that produced relatively rapid growing individuals and slow runners. Our results emphasize the overriding importance of clutch (genotype) to variation in phenotypes and survival in hatchling S. undulatus. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76, 105‐124.

145 citations


Journal ArticleDOI
TL;DR: Eublepharids provide new evidence that male-biased dimorphic heads are weapons used in aggressive encounters and that the ventral glands probably function in territory marking rather than in intersexual communication.
Abstract: Lizards of the family Eublepharidae exhibit interspecific diversity in body size, sexual size dimorphism (SSD), head size dimorphism (HSD), occurrence of male combat, and presence of male precloacal pores. Hence, they offer an opportunity for testing hypotheses for the evolution and maintenance of sexual dimorphism. Historical analysis of male agonistic behaviour indicates that territoriality is ancestral in eublepharid geckos. Within Eublepharidae, male combat disappeared twice. In keeping with predictions from sexual selection theory, both events were associated with parallel loss of male-biased HSD and ventral scent glands. Eublepharids therefore provide new evidence that male-biased dimorphic heads are weapons used in aggressive encounters and that the ventral glands probably function in territory marking rather than in intersexual communication. Male-biased SSD is a plesiomorphic characteristic and was affected by at least three inversions. Shifts in SSD and male combat were not historically correlated. Therefore, other factors than male rivalry appear responsible for SSD inversions. Eublepharids demonstrate the full scope of Rensch's rule (small species tend to be female-larger, larger species male-larger). Most plausibly, SSD pattern hence seems to reflect body size variation.

144 citations


Journal ArticleDOI
TL;DR: A new perspective of macaque dispersal through the Indonesian archipelago is proposed, which takes into account sea-level changes, as well as latitudinal and altitudinal rainforest shifts following climatic cycles during the Quaternary.
Abstract: The evolution of primates is usually approached from the standpoint of adaptation and interspecific competition. However, climatic and eustatic changes associated with periodic glaciations have had a profound influence on their geographical distribution, favouring the intervention of contingency in evolution. This paper deals with the role of chance and competition in the dispersal and stocking of macaques in the islands of south-east Asia. The genus Macaca is unique among non-human primates for the range of habitats colonized, from continents to islands. We first review current knowledge about the zoogeography of macaques in Sundaland. We point out the inconsisten- cies present in the hypotheses thus far proposed to account for the colonization of shallow- and deep-water islands. We then propose a new perspective of macaque dispersal through the Indonesian archipelago, which takes into account sea-level changes, as well as latitudinal and altitudinal rainforest shifts following climatic cycles during the Quaternary. We envision three steps: (i) dispersal and partial disappearance of the first radiation of macaques; (ii) primary mainland recolonization by pigtailed and longtailed macaques; and (iii) secondary sea-rafting colo- nization by longtailed macaques. This model implies that liontailed, Sulawesi and Mentawai macaques stemmed from pre-glacial remnant populations, whereas pigtailed macaques originated in post-glacial populations that diverged later on. The model accounts for the distribution of longtailed macaques throughout the Indonesian archi- pelago. The riverine habits of this species would have favoured its dispersal by sea rafting, which was otherwise extremely rare for other primate species. Stocking would have been successful for longtailed macaques only in islands where no other macaque competitors were already present. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 555-576. ADDITIONAL KEYWORDS: biogeography - colonization - competition - contingency - island - Macaca - Quaternary - Sunda shelf.

139 citations


Journal ArticleDOI
TL;DR: It is suggested that both genetic and maternal effects are important determinants of geographical variation in amphibian life-histories, and that much of the differentiation resulting from maternal effects is mediated through variation in egg size.
Abstract: The relative importance of genetic, environmental, and maternal effects as determinants of geographical variation in vertebrate life-histories has not often been explored. We examined the role of genetic and maternal effects as determinants of population divergence in survival and three important larval life-history traits (growth rate, age, and size at metamorphosis) using reciprocal crosses between two latitudinally separated populations of the common frog (Rana temporaria Linnaeus). Genetic effects were important in all three traits as indicated by the significant effect of male origin, but there was also evidence for nonadditive genetic contributions on metamorphic size and growth rate. Likewise, maternal effect contributions to population divergence were large, partially environment dependent, and apparently acting primarily through egg size in two of three traits. These results suggest that both genetic and maternal effects are important determinants of geographical variation in amphibian life-histories, and that much of the differentiation resulting from maternal effects is mediated through variation in egg size.

132 citations


Journal ArticleDOI
TL;DR: In a fragment of a Mexican rainforest, a cicada community at the end of the dry season consisting of nine species, seven of the nine species formed dense choruses at dawn and at dusk and each species showed preferences in the height of calling sites.
Abstract: Multispecies cicada communities in neotropical rainforests produce a complex and intense acoustic environment. In a fragment of a Mexican rainforest (Veracruz, Mexico), a cicada community at the end of the dry season consisted of nine species (Daza montezuma; Pacarina schumanni; Miranha imbellis; Dorisiana sutori; Fidicinoides picea; Fidicinoides pronoe; Quesada gigas; one species of the genus Neocicada and one uncaught canopy species). Seven of the nine species formed dense choruses at dawn and at dusk. Each species showed preferences in the height of calling sites. Males of the species were solitary or gregarious, and followed a ‘call-fly’ or a ‘call-stay’ calling strategy. Acoustic signals of each species had particular time and frequency patterns. All these specific features appear to separate the nine species acoustically and lead to a partitioning of the acoustic environment. The acoustic partitioning might decrease the risk of heterospecific courting and mating.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 379–394.

Journal ArticleDOI
TL;DR: It is suggested that it is important to consider plumage age when exploring variation in structural plumage coloration, and that it can largely explain the difference between the British and Swedish studies.
Abstract: Recent studies of blue tits, Parus caeruleus, have found sexual selection and a viability-indicating function of the structural ultraviolet and blue crown plumage, but the reasons for this signal variation are not understood. Furthermore, studies in England and Sweden have yielded somewhat different results (particularly with regard to the spectral position of the reflectance peak). Here we investigate whether the blue tit UV/blue ornament varies with time of year since such variation might be relevant to the signalling function as well as the apparent difference between populations. From 400 blue tits captured at two different localities in Sweden, we found that objective measures of ‘hue’ (spectral location), ‘chroma’ (spectral purity) and ‘brightness’ (spectral intensity), varied substantially with season. Just after moult (October), crown ‘hue’ is maximally UV-shifted (359 nm for males and 373 nm for females). Thereafter the peak drifts upwards and by the time of nestling feeding (June) male reflectance peaks at 404 nm and female at 413 nm. This change is probably due to feather wear as well as fat and dirt accumulation, which might constitute an additional male quality cue. Our results suggest that it is important to consider plumage age when exploring variation in structural plumage coloration, and that it can largely explain the difference between the British and Swedish studies.

Journal ArticleDOI
TL;DR: The complex patterns of morphological, allozymic and mtDNA variation associated with the constantly changing geological landscape give insight into the nature of processes responsible for species formation in Batrachoseps.
Abstract: The plethodontid genus Batrachoseps, the slender salamanders, is the most diverse clade of salamanders in western North America, but it has posed taxonomic difficulties because it contains many morphologically cryptic species. A segment of the mitochondrial DNA gene cytochrome b was studied for 278 individuals densely sampled from throughout the range of all 18 described species and several undescribed species. Phylogenetic analyses of the mtDNA data identify six major clades, one corresponding to the subgenus Plethopsis and five within a monophyletic subgenus Batrachoseps. All major clades and most species within these clades display strong phylogeographic structuring. Comparisons of mtDNA and allozyme data show that several allozymically cohesive groups are not monophyletic with respect to mtDNA. We suggest that this phenomenon results from fragmentation of populations, divergence in allopatry, and then recontact and gradual merging of units caused predominantly by male-mediated gene flow. The mtDNA offers evidence that populations were once more isolated than they are now, while the patterns of allozyme variation reflect recent and current interactions among populations. The complex patterns of morphological, allozymic and mtDNA variation associated with the constantly changing geological landscape give insight into the nature of processes responsible for species formation in Batrachoseps. © 2002 The Linnean Society of London. Biological Journal of the Linnean Society, 2002, 76, 361–391.

Journal ArticleDOI
TL;DR: There is some phylogenetic clustering of claw shape and size, but there does not appear to be a significant level of phylogenetic dependence because no conclusions are changed when independent contrasts are used.
Abstract: For decades, the extreme asymmetric chelipeds of male fiddler crabs (genus Uca) have been used as a model system for the description and discussion of allometry. Almost all previous studies, however, have concentrated on intraspecific variation and have not examined claw variation among species. In this study, modern methods of describing shape and size, geometric morphometrics, are used to study claw variation across the genus. These analyses are also performed in a phylogenetic context using independent contrasts. Within and among species, major claws show allometric trends in both shape and size. Minor claw growth is isometric within species; across species, these claws are isometric with respect to size, but allometric with respect to shape. Although the variation is much greater in major claws, the allometric pattern of shape change for both majors and minors can be characterized by a general increase in the length of the pollex relative to the manus and the size of the propodus relative to the carpus. There is some phylogenetic clustering of claw shape and size, but there does not appear to be a significant level of phylogenetic dependence because no conclusions are changed when independent contrasts are used. Regenerated major claws have stronger allometric patterns than unregenerated claws, causing them to have shapes associated with relatively larger claws. Minor claw shape shows a strong correlation with habitat type. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 147‐162. ADDITIONAL KEYWORDS: allometry ‐ regeneration.

Journal ArticleDOI
TL;DR: It is suggested that cryptic speciation is apt to evolve in species, such as myrmecophiles, endoparasites and koinobiont parasitoids, whose life-styles result in strong selection on their physiological or behavioural characters.
Abstract: The myrmecophilous hoverfly, Microdon mutabilis, is listed as a ‘Rare’ or ‘Nationally Notable Species’ in UK Red Data Books. As an obligate social parasite, feeding only from ant colonies, its life-style satisfies theoretical conditions under which cryptic speciation is predicted to evolve; namely, strong selection for nonmorphological adaptations that enhance its exploitation of a local subspecies or populations of its host. Samples of larvae and pupae in Ireland, Scotland and England showed that M. mutabilis exploits a single and different host ant species on different sites across its range. In nine southern English colonies, 95.6% of infested nests were of Myrmica scabrinodis whereas in six Irish and two Scottish colonies 100% and 94.2%, respectively, of the infested nests were of Formica lemani, despite M. scabrinodis being common at all sites. Although the adults from ‘scabrinodis’ (and lemani) populations are cryptic, morphometric measurements of pupae showed consistent diagnostic characters that were sufficiently distinct for these ecotypes to be classed as separate species. We conclude that M. mutabilis is the ‘lemani-type’ and designate the ‘scabrinodis-type’ as a new species, Microdon myrmicae spec. nov. Thus, one of the listed threatened species of the British Isles becomes two species, each possessing about half the number of populations and occupying half the range of the original ‘species’. Each also inhabits a different serai stage within grassland or heathland, and will require a different management regime if its declining populations are to be conserved. ‘M. mutabilis’ is reported with other host ant species on the European continent. In the light of our results, these may prove to be additional cryptic species. We suggest that cryptic speciation is apt to evolve in species, such as myrmecophiles, endoparasites and koinobiont parasitoids, whose life-styles result in strong selection on their physiological or behavioural characters. The implications for Red Data Book classifications and for practical conservation are discussed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 291–300.

Journal ArticleDOI
TL;DR: It is hypothesized that the ability to be generalist or specialist in Lamellodiscus is controlled by intrinsic, phylogenetically-related characteristics, and that specialist species tend to use large hosts, which may be more predictable.
Abstract: The evolution and determinants of host specificity in Lamellodiscus species (Monogenea, Diplectanidae) were investigated. The 20 known Mediterranean species were studied, all parasites of fishes from the family Sparidae (Teleostei). An index of specificity, which takes into account the phylogenetic relationships of their fish host species, was defined. The link between specificity and its potential determinants was investigated in a phylogenetic context using the method of independent contrasts. Host specificity in Lamellodiscus species appeared to be highly constrained by phylogeny, but also linked to host size. Mapping specificity onto the parasite phylogenetic tree suggests that specialist species do not represent an evolutionary dead end, and that specialization is not a derived condition. It is hypothesized that the ability to be generalist or specialist in Lamellodiscus is controlled by intrinsic, phylogenetically-related characteristics, and that specialist species tend to use large hosts, which may be more predictable. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 431−443.

Journal ArticleDOI
TL;DR: Body shape was quantified for male and female lizards of 30 species of phrynosomatid lizards, indicating a potentially important contribution of sexual selection to the evolution of differences in limb proportions.
Abstract: With the exception of the well-documented case for anoline lizards, recent studies have found few evolutionary relationships between morphology and habitat use in lizards despite clear-cut biomechanical predictions. One of the factors typically hampering these analyses is the clustering of habitat use within evolutionary lineages. In the present study, body shape was quantified for male and female lizards of 30 species of phrynosomatid lizards. This group was selected as little clustering of ecological variables seemed to be present. The results of traditional analyses indicate that evolutionary correlates of habitat use were prominent in the hindlimbs of both sexes. Species living in open habitats are characterized by longer femurs, and longer hindlimbs relative to the forelimb. Moreover, males from ground-dwelling species utilizing open habitats have longer toes on the hind foot than males from climbing species. Phylogenetic analyses indicated strong evolutionary associations between habitat use and the relative length of front and hindlimbs, with species from open terrestrial habitats having significantly shorter frontlimbs relative to their hindlimb than rock or tree climbing species. Evolutionary associations between morphology and habitat use were generally stronger for male lizards, indicating a potentially important contribution of sexual selection to the evolution of differences in limb proportions. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 149–163.

Journal ArticleDOI
Gary M. Barker1
TL;DR: The utility of PD was demonstrated in four case studies: state of the environment reporting, with changes in avian faunas resulting from extinctions quantified as indicators of the state of biodiversity at Global, New Zealand and Waikato region scales, and changes in available habitat quantifiedAs indicators of pressures on biodiversity in the Waikatos region.
Abstract: The value of biodiversity lies in its option value for the future, the greater the complement of contemporary biodiversity conserved today, the greater the possibilities for future biodiversity because of the diverse genetic resource needed to ensure continued evolution in a changing and uncertain world. From this perspective, biodiversity option value can be equated with richness in the different features expressed by species. An individual species of greater value is one contributing more novel features to a given subset. The feature diversity of species and communities is difficult to estimate directly, but can be predicted by the phylogenetic relationships among the species. The ‘Phylogenetic Diversity’ measure (PD) (Faith, 1992a) estimates the relative feature diversity of any nominated set of species by the sum of the lengths of all those branches spanned by the set. These branch lengths reflect patristic or path-length distances. This study first reviews and expands on some of the properties of PD, and develops simple modifications of the measure (δnPD and enPD) to enable capture of both the phylogenetic relatedness of species and their abundances in each sample. Then the application of PD, δnPD and enPD to a wide range of conservation and resource management issues is demonstrated using avian case studies. Supertree construction procedures (matrix representation using parsimony analysis; average consensus) were used to combine the extensive DNA-DNA hybridization tree of Sibley & Ahlquist (1990) with numerous, recently published phylogenetic reconstructions to derive a phylogenetic tree for the global avian fauna. Using this supertree as a systematic framework, the utility of PD was demonstrated in four case studies: (i) state of the environment reporting, with changes in avian faunas resulting from extinctions quantified as indicators of the state of biodiversity at Global, New Zealand and Waikato region scales, and changes in available habitat quantified as indicators of pressures on biodiversity in the Waikato region; (ii) setting priorities for threatened species management, with PD as a measure of option value integrated with information on survivorship expectations to develop a ranking among threatened New Zealand forest bird species; (iii) monitoring biotic response to management, with data from 5-minute counts used to analyse changes in forest bird communities under three management regimes in New Zealand; and (iv) selection of indicator species, with PD used to objectively identify subsets of species in the Global, New Zealand and Waikato avian faunas that comprise a high proportion of the option value in those faunas.

Journal ArticleDOI
TL;DR: Positive contrasts of bandwidth were associated with positive contrasts of habitat, confirming that songs of open-habitat species have a wider bandwidth than those of their more closed habitat relatives.
Abstract: Fil: Bertelli, Sara Beatriz. Universidad Nacional de Tucuman. Facultad de Ciencias Naturales e Instituto Miguel Lillo; Argentina. Consejo Nacional de Investigaciones Cientificas y Tecnicas; Argentina

Journal ArticleDOI
TL;DR: It is argued that the high dispersal capabilities of Brueelia species, e.g. phoresis on hippoboscid flies, are a likely explanation for the incongruence between host and parasite phylogenies in this case.
Abstract: Lice (Insecta: Phthiraptera) are often considered a model group of parasites for studies of cospeciation because of their high host specificity, and louse species in the genus Brueelia are relatively host-specific. To test the extent of cospeciation, we reconstructed a phylogeny for Brueelia from nuclear (EF1α) and mitochondrial (COI) DNA sequences. This phylogeny was generally well resolved and supported. Two major clades within Brueelia (as well as several other lineages) were identified, and these corresponded to major morphological differences in the preantennal region of the head and sclerotization of the abdomen. However, the phylogeny of Brueelia showed little concordance to a published phylogeny of the hosts. In addition, we uncovered four cases (out of 15 species) of one species of Brueelia on two or more bird species. We argue that the high dispersal capabilities of Brueelia species, e.g. phoresis on hippoboscid flies, are a likely explanation for the incongruence between host and parasite phylogenies in this case. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society 2002, 77, 233–247.

Journal ArticleDOI
TL;DR: Historical biogeographical analyses of the genus Xylocopa Latreille are presented, showing that the major splits in the carpenter bee phylogeny occurred well after the final breakup of Gondwanaland, but before important Miocene fusion events.
Abstract: The biogeographical history of major groups of bees with worldwide distributions have often been explained through hypotheses based on Gondwanan vicariance or long distance dispersal events, but until recently these hypotheses have been very difficult, if not impossible, to distinguish. New fossil data, comprehensive information on Mesozoic and Cenozoic coastline positions and the availability of phylogenetically informative DNA markers now makes it feasible to test these hypotheses for some groups of bees. This paper presents historical biogeographical analyses of the genus Xylocopa Latreille, based on phylogenetic analyses of species belonging to 22 subgenera using molecular data from two nuclear genes, elongation factor-1α (EF-1α) and phosphoenolpyruvate carboxykinase (PEPCK), combined with previously published morphological and mitochondrial data sets. Phylogenetic analyses based on parsimony and likelihood approaches resulted in several groups of subgenera supported by high bootstrap values (>85%): an American group with the Oriental/Palaearctic subgenera Nyctomelitta and Proxylocopa as sister taxa; a geographically diverse group (Xylocopa s.l); and a group consisting of African and Oriental subgenera. The relationships among these three clades and the subgenus Perixylocopa remained unresolved. The Oriental subgenus Biluna was found to be the sister group of all other carpenter bee subgenera included in this study. Using a relaxed molecular clock calibrated using fossil carpenter bees, we show that the major splits in the carpenter bee phylogeny occurred well after the final breakup of Gondwanaland (the separation of South America and Africa, 100 Mya), but before important Miocene fusion events. Ancestral area analysis showed that the genus Xylocopa most likely had an Oriental-Palaearctic origin and that the present world distribution of Xylocopa subgenera resulted mainly from independent dispersal events. The influence of Pleistocene glaciations on carpenter bee distributions is also discussed. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 249–266.

Journal ArticleDOI
TL;DR: Differences in morphological divergence are closely related to the differences in the ecological surroundings of each population, which appears to be resource polymorphism, which may lead to population differentiation and speciation.
Abstract: Icelandic threespine sticklebacks show parallel sympatric morphological differences related to different substrate habitats in four Icelandic lakes. The level of morphological diversification varies among the lakes, ranging from a population with a wide morphological distribution to a population with clear resource morphs, where morphological diversification was reflected in diet differences. These differences in morphological divergence are closely related to the differences in the ecological surroundings of each population. This appears to be resource polymorphism, which may lead to population differentiation and speciation. Trophically related sexual dimorphism was also common in these sticklebacks, which is possibly the result of sexual selection or habitat segregation by the sexes.

Journal ArticleDOI
TL;DR: The high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability, associated with differences in prey resources available to the snakes.
Abstract: Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons (Morelia spilota) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 113–125.

Journal ArticleDOI
TL;DR: It is proposed that the clinal genetic structure in the European eel may be due to (1) isolation by distance, (2) temporal reproductive separation, (3) post-larval selective forces, (4) contact between formerly separated groups or (5) some combination thereof.
Abstract: The genetic variability and structure of the European eel ( Anguilla anguilla L.) in populations throughout Europe was reassessed using 15 allozymic loci, seven of which were polymorphic. Seven sites were sampled on a latitudinal gradient across the natural continental range, extending from southern France to southern Norway. Heterozygosity ( H e = 0.05) and level of polymorphism (P = 0.43) were comparable to other marine fish. Populations were poorly differentiated ( G ST = 0.014, F ST = 0.002), which is not surprising considering the high dispersal capability of the European eel. However, a significant geographical cline was detected at two alleles ( IDH-1 * 100 and GPI-1 * 110 ), and genetic distances ( D CE ) were concordant with geographical coastal distances. Mantel tests, pairwise F ST ’s and multidimensional scaling analyses identify three distinct groups: Northern Europe, Western Europe and the Mediterranean Sea. We propose that the clinal genetic structure in the European eel may be due to (1) isolation by distance (as recently detected with microsatellites), (2) temporal reproductive separation, (3) post-larval selective forces, (4) contact between formerly separated groups or (5) some combination thereof. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 509‐521

Journal ArticleDOI
TL;DR: The feeding ecology of the North American gopher snake, Pituophis catenifer, was studied based on the stomach contents of more than 2600 preserved and free-ranging specimens, and published and unpublished dietary records to assess size-dependent and geographical variation in feeding preferences of gape-limited predators.
Abstract: Studies of food relations are important to our understanding of ecology at the individual, population and community levels. Detailed documentation of the diet of large-bodied, widespread snakes allows us to assess size-dependent and geographical variation in feeding preferences of gape-limited predators. Furthermore, with knowledge of the food habits of sympatric taxa we can explore possible causes of interspecific differences in trophic niches. The feeding ecology of the North American gopher snake, Pituophis catenifer, was studied based on the stomach contents of more than 2600 preserved and free-ranging specimens, and published and unpublished dietary records. Of 1066 items, mammals (797, 74.8%), birds (86, 8.1%), bird eggs (127, 11.9%), and lizards (35, 3.3%) were the most frequently eaten prey. Gopher snakes fed upon subterranean, nocturnal and diurnal prey. The serpents are primarily diurnal, but can also be active at night. Therefore, gopher snakes captured their victims by actively searching underground tunnel systems, retreat places and perching sites during the day, or by pursuing them or seizing them while they rested at night. Gopher snakes of all sizes preyed on mammals, but only individuals larger than 40 and 42 cm in snout–vent length took bird eggs and birds, respectively, possibly due to gape constraints in smaller serpents. Specimens that ate lizards were smaller than those that consumed mammals or birds. Gopher snakes raided nests regularly, as evidenced by the high frequency of nestling mammals and birds and avian eggs eaten. Most (332) P. catenifer contained single prey, but 95 animals contained 2–35 items. Of the 321 items for which direction of ingestion was determined, 284 (88.5%) were swallowed head-first, 35 (10.9%) were ingested tail-first, and two (0.6%) were taken sideways. Heavier gopher snakes took heavier prey, but heavier serpents ingested prey with smaller mass relative to snake mass, evidence that the lower limit of prey mass did not increase with snake mass. Specimens from the California Province and Arid Deserts (i.e. Mojave, Sonoran and Chihuahuan Deserts) took the largest proportion of lizards, whereas individuals from the Great Basin Desert consumed a higher percentage of mammals than serpents from other areas, and P. catenifer from the Great Plains ate a greater proportion of bird eggs. Differences in prey availability among biogeographical regions and unusual circumstances of particular gopher snake populations may account for these patterns. Gopher snakes have proportionally longer heads than broadly sympatric Rhinocheilus lecontei (long-nosed snake), Charina bottae (rubber boa) and Lampropeltis zonata (California mountain kingsnake), which perhaps explains why, contrary to the case in P. catenifer, the smaller size classes of those three species do not eat mammals. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 165–183.

Journal ArticleDOI
TL;DR: This support for locally restricted in situ survival of E. alpinus in the northern Prealps or nearby Jura mountains is considered, as the west-eastern gene-pool probably immigrated postglacially from southern France through the prealpine lowlands, skirting central Switzerland.
Abstract: It has been hypothesized that the subalpine Erinus alpinus survived the Pleistocene glaciation in southern French refugia peripheral to the Alps and postglacially immigrated to its northern Alpine distribution (tabula rasa hypothesis). However, E. alpinus is also known from nunataks in the northern Prealps. PCR–RFLP of the cpDNA revealed no variation, whereas AFLPs identified three phylogeographical groups among the 22 populations: (1) a central Swiss group, (2) a single central Swiss population on Mount Rigi, and (3) all other populations located to the west and east of the central Swiss group (west-eastern populations). The population representing the putative French refugium was not distinct from the west-eastern populations but was different from the populations of central Switzerland. We consider this support for locally restricted in situ survival of E. alpinus in the northern Prealps or nearby Jura mountains (dating back to 60–28 ky BP). However, the west-eastern gene-pool probably immigrated postglacially from southern France through the prealpine lowlands (not before 14.6 kyr BP), skirting central Switzerland. The near-absence of gene flow among the central Swiss and the west-eastern populations may be the reason why these historical genetic patterns are still detectable. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 77, 87–103.

Journal ArticleDOI
TL;DR: The data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa, and consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus.
Abstract: Genetic variation was studied using protein electrophoresis of 28–38 gene loci in 1347 specimens of Apodemus agrar-ius, A. peninsulae, A. flavicollis, A. sylvaticus, A. alpicola, A. uralensis, A. cf. hyrcanicus, A. hermonensis, A. m. mystacinus and A. m. epimelas, representing 121 populations from Europe, the Middle East, and North Africa. Mean values of heterozygosity per locus for each species ranged from 0.02 to 0.04. Mean values of Nel's genetic distance (D) between the taxa ranged from 0.06 (between A. flavicollis and A. alpicola) to 1.34 (between A. uralensis and A. agrarius). The highest values of D were found between A. agrarius and other Apodemus species (0.62-1.34). These values correspond to those generally observed between genera in small mammals. Our data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa. High genetic distance between A. m. mystacinus and A. m. epimelas leads us to consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus. The data also suggest a recent separation of members of the latter group from a common ancestor, and subsequent rapid radiation, making it difficult to infer phylogenetic relationships. Some taxonomic implications of the results are discussed further.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 395–419.

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TL;DR: Increasing productivity and predictability of environment in respect to rainfall, without concomitant frost stress, was found to select for high reproductive biomass and large seeds, a high fraction of germinating seeds and high vigour of seedlings.
Abstract: Reciprocal introduction of seeds and seedlings of wild barley, Hordeum spontaneum , originating in four different environments of Israel was used to: (1) test for local adaptation, (2) make inferences about environmental effects on life-history and reproductive traits, and (3) identify trait combinations with recognizable ‘strategies’. The four populations examined represented the following environments: (1) desert − low productivity and predictability, drought stress; (2) semi-steppe batha − moderate productivity and predictability; (3) grassland − high productivity and predictability; and (4) mountain − high productivity and predictability but with severe frost stress. Significant genotype-by-environment interactions were observed for yield and reproductive biomass, seedling biomass and percentage germinated and survived seeds, suggesting local ecotype adaptation. Increasing productivity and predictability of environment in respect to rainfall, without concomitant frost stress, was found to select for high reproductive biomass and large seeds, a high fraction of germinating seeds and high vigour of seedlings. The optimal strategy changes with increasing productivity and predictability and involves a trade-off between seed size and number, with reduced yield but increased seed mass, consistent with competition selection (or K-selection sensu MacArthur & Wilson (1967)) type. No specific life-history adaptations to predictable frost stress were detected for the mountain ecotype, but there was higher survival of seedlings in their indigenous (mountain) environment compared with other ecotypes. The latter appears to be a physiological adaptation to frost, which is consistent with selection for stress tolerance (or S-selection sensu Grime (1977)) type. The other stress factor, drought, which is very unpredictable in deserts, was associated with high seed dormancy, small seed size and low vigour of seedlings, but relatively high yield, which is consistent with a stress-escape bet-hedging strategy. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77, 479–490.

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TL;DR: There is strong evidence, through a generalized linear model, that increased habitat generalism, lack of migratory tendency, and sexual monochromatism together explain significant variation in the successful establishment of introduced land bird species.
Abstract: Despite the many studies that have investigated successful establishment of introduced bird species, very little is known about the patterns of success worldwide and the influence of life history and ecological traits. This study describes the analysis of non-native land bird introductions to test existing hypotheses of establishment success using a modern comparative approach to control for phylogenetic relatedness among taxa. I used randomization tests, permutational phylogenetic regressions, and across-taxa and sister-taxa comparisons to examine predicted correlates of introduction success. My analyses confirmed that the variability in establishment success among introduced land bird families is distributed in a manner significantly different from a random process, and that life history and ecological attributes are an important influence of introduction success. I found strong evidence, through a generalized linear model, that increased habitat generalism, lack of migratory tendency, and sexual monochromatism together explain significant variation in the successful establishment of introduced land bird species. This has resulted in a predictive equation for the novel introduction of land bird species.

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TL;DR: This study confirms that considerable intrafamilial variation exists among chameleons and that these traits appear to be evolutionarily quite labile, and investigates whether morphological variation correlates with differences in ecology and whether correlations exist among different aspects of morphology.
Abstract: The lizard family Chameleonidae is one of the most distinctive taxa of all vertebrates. Nonetheless, despite great intrafamilial diversity, little research has been conducted on morphological variation among chameleons. As a first step in this direction, we took morphological measurements on the head, limbs, spines, and tail of 56 species. Our goals were to investigate whether morphological variation correlates with differences in ecology and to examine whether correlations exist among different aspects of morphology. Based on existing information, species were classified either as arboreal or terrestrial, the latter referring to species that are known to use the ground on a regular basis. This study confirms that considerable intrafamilial variation exists among chameleons and that these traits appear to be evolutionarily quite labile. Once the effects of size are removed, functionally related traits tend to covary; however, few correlations are observed between non-functionally related traits. Many differences in the lengths of the limbs and head elements were detected between terrestrial and arboreal species, but the functional and selective significance of these differences is not clear. Further research on chameleon behaviour and ecology is required to understand the factors contributing to chameleon morphological diversity.