Geographic range size, life history and rates of diversification in Australian mammals

What causes species richness to vary among different groups of organisms? Two hypotheses are that large geographical ranges and fast life history either reduce extinction rates or raise speciation rates, elevating a clade's rate of diversification. Here we present a comparative analysis of these hypotheses using data on the phylogenetic relationships, geographical ranges and life history of the terrestrial mammal fauna of Australia. By comparing species richness patterns to null models, we show that species are distributed nonrandomly among genera. Using sister-clade comparisons to control for clade age, we then find that faster diversification is significantly associated with larger geographical ranges and larger litters, but there is no evidence for an effect of body size or age at first breeding on diversification rates. We believe the most likely explanation for these patterns is that larger litters and geographical ranges increase diversification rates because they buffer species from extinction. We also discuss the possibility that positive effects of litter size and range size on diversification rates result from elevated speciation rates.

The three phases of the ensemble forecasting of niche models: geographic range and shifts in climatically suitable areas of Utetheisa ornatrix (Lepidoptera, Arctiidae)

Species' geographic ranges are usually considered as basic units in macroecology and biogeography, yet it is still difficult to measure them accurately for many reasons. About 20 years ago, researchers started using local data on species' occurrences to estimate broad scale ranges, thereby establishing the niche modeling approach. However, there are still many problems in model evaluation and application, and one of the solutions is to find a consensus solution among models derived from different mathematical and statistical models for niche modeling, climatic projections and variable combination, all of which are sources of uncertainty during niche modeling. In this paper, we discuss this approach of ensemble forecasting and propose that it can be divided into three phases with increasing levels of complexity. Phase I is the simple combination of maps to achieve a consensual and hopefully conservative solution. In Phase II, differences among the maps used are described by multivariate analyses, and Phase III consists of the quantitative evaluation of the relative magnitude of uncertainties from different sources and their mapping. To illustrate these developments, we analyzed the occurrence data of the tiger moth, Utetheisa ornatrix (Lepidoptera, Arctiidae), a Neotropical moth species, and modeled its geographic range in current and future climates.

Effects of detectability on estimates of geographic range size in bignonieae

Extinction risk has not been evaluated for 96% of all described plant species. Given that the Global Strategy for Plant Conservation proposes preliminary conservation assessments of all described plant species by 2010, herbarium specimens (i.e., primary occurrence data) are increasingly being used to infer threat components from estimates of geographic range size. Nevertheless, estimates of range size based on herbarium data may be inaccurate due to collection bias associated with interspecific variation in detectability. We used data on 377 species of Bignonieae to test the hypothesis that there is a positive relationship between detectability and estimates of geographic range size derived from herbarium specimens. This relationship is expected if the proportion of the true geographic range size of a species that is documented by herbarium specimens is given by the product of the true geographic range size and the detectability of the species, assuming no relationship between true geographic range size and detectability. We developed 4 measures of detectability that can be estimated from herbarium data and examined the relationship between detectability and 2 types of estimates of geographic range size: area of occupancy and extent of occurrence. Our results from regressing estimates of extent of occurrence and area of occupancy on detectability across genera provided no support for this hypothesis. The same was true for regressions of estimated extent of occurrence on detectability across species within genera. Nevertheless, regressions of estimated area of occupancy on detectability across species within genera provided partial support for our hypothesis. We considered 3 possible explanations for this mixed outcome: violation of the assumption of no relationship between true geographic range size and detectability; the relationships between estimated geographic range size and detectability may be an artifact of a negative relationship between estimated area of occupancy and the sampling variance of detectability; detectability may have had 2 opposite effects on estimated species range sizes: one determines the proportion of the true range of a species documented by herbarium specimens and the other determines the distribution of true range size for the species actually observed with herbarium data. Our findings should help improve understanding of the potential biases incurred with the use of herbarium data.

Geographic range, habitats, and host plants of bromeliad-living jumping spiders (Salticidae)

Although spiders are a very diverse group on vegetation, their associations with plants are poorly known. Some salticid species specifically use Bromeliaceae as host plants in some regions of South America. In this study, I report the geographic range of these spider-bromeliad associations, and whether the spiders inhabit particular bromeliad species and vegetation types, as well as open areas or interior of forests. Nine salticid species were found to be associated with up to 23 bromeliad species in cerrados (savanna-like vegetation), semideciduous and seasonal forests, coastal sand dune vegetation, restingas, inselbergs, highland forests, chacos, and rain forests at 47 localities in Brazil, Paraguay, Bolivia, and Argentina. Some species were typically specialists, inhabiting almost exclusively one bromeliad species over a large geographic range (e.g., Psecas chapoda on Bromelia balansae), whereas others were generalists, occurring on up to 7-8 bromeliad species (e.g., Psecas sp., Eustiromastix nativo, and Coryphasia sp. 1). The regional availability of bromeliad species among habitats may explain this pattern of host plant use. More jumping spiders were found on bromeliads in open areas than on bromeliads in the interior of forests. These results show that several jumping spider species may be strictly associated with the Bromeliaceae in the Neotropics. This is one of the few studies to show host-specific associations for spiders on a particular plant type over a wide geographic range.

Honey Ants (Genus Myrmecocystus) Macroecology: Effect of Spatial Patterns on the Relationship between Worker Body Size and Geographic Range Size

Macroecology evaluates the partitioning of physical space and resources among organisms through correlation among ecological variables, such as geographical range size and shape, body size, and population density, measured at large geographical and taxonomic scales. In this article, we analyzed the spatial patterns in worker body size and geographic range size for the 27 described species of honey ants, genus Myrmecocystus Wesmael, in the United States and Mexico, and especially the relationship between these 2 variables after statistically removing their spatial patterns. The 2 variables are correlated, but also displayed significant spatial patterns, as detected by trend surface and spatial autocorrelation analyses. After removing these spatial effects, worker body size and geographic range size were still positively correlated. The relationship, therefore, is not a consequence of spatial effects and it does follow Brown's model, which predicts that the geographic range size will have a positive slope on body size. In this model, the lower population densities caused by foraging activities and local territorial competition are associated with a large geographic range, avoiding stochastic extinction. Although this constraint in local population density does not necessarily hold for small organisms such as insects that could achieve high densities even in very small areas and patchy habitats, it may hold for social insects, especially ants, because of the local competition among colonies.

Understanding bias in geographic range size estimates

Aim Estimates of geographic range size derived from natural history museum specimens are probably biased for many species. We aim to determine how bias in these estimates relates to range size. Location We conducted computer simulations based on herbarium specimen records from localities ranging from the southern United States to northern Argentina. Methods We used theory on the sampling distribution of the mean and variance to develop working hypotheses about how range size, defined as area of occupancy (AOO), was related to the inter-specific distribution of: (1) mean collection effort per area across the range of a species (MC); (2) variance in collection effort per area across the range of a species (VC); and (3) proportional bias in AOO estimates (PBias: the difference between the expected value of the estimate of AOO and true AOO, divided by true AOO). We tested predictions from these hypotheses using computer simulations based on a dataset of more than 29,000 herbarium specimen records documenting occurrences of 377 plant species in the tribe Bignonieae (Bignoniaceae). Results The working hypotheses predicted that the mean of the inter-specific distribution of MC, VC and PBias were independent of AOO, but that the respective variance and skewness decreased with increasing AOO. Computer simulations supported all but one prediction: the variance of the inter-specific distribution of VC did not decrease with increasing AOO. Main conclusions Our results suggest that, despite an invariant mean, the dispersion and symmetry of the inter-specific distribution of PBias decreases as AOO increases. As AOO increased, range size was less severely underestimated for a large proportion of simulated species. However, as AOO increased, range size estimates having extremely low bias were less common.

Clutch size in the small-sized lizard Eurolophosaurus nanuzae (Tropiduridae): does it vary along the geographic distribution of the species?

We studied life history traits of females of the lizard Eurolophosaurus nanuzae (Rodrigues, 1981), an endemic species of rock outcrop habitats in southeastern Brazil. During October 2002 and 2003 we sampled three populations in sites that encompass the meridional portion of the geographic range of the species. Clutch size varied from one to three eggs, with most females carrying two eggs. Clutch size did not vary among populations, but was correlated to female body size. Only larger females produced clutches of three eggs. Females of the small-sized E. nanuzae produce eggs as large as those of medium-sized tropidurids, thus investing a considerable amount of energy to produce clutches resulting in high values of relative clutch mass.

Trophic versus geographic structure in stable isotope signatures of pelagic seabirds breeding in the northeast Atlantic

Feeding ecology and geographic location are 2 major factors influencing animal stable isotope signatures, but their relative contributions are poorly understood, which limits the usefulness of stable isotope analysis in the study of animal ecology. To improve our knowledge of the main sources of isotopic variability at sea, we determined δ15N and δ13C signatures in the first primary feather of adult birds from 11 Procellariiform species (n = 609) across 16 northeast Atlantic localities, from Cape Verde (20°N) to Iceland (60°N). Post-breeding areas (where the studied feather is thought to be grown) were determined using light-level geolocation for 6 of the 11 species. Isotopic variability was geographically unstructured within the mid-northeast Atlantic (Macaronesian archipelagos), but trophically structured according to species and regardless of the breeding location, presumably as a result of trophic segregation among species. Indeed, the interspecific isotopic overlap resulting from combining δ15N and δ13C signatures of seabirds was low, which suggests that most species exploited exclusive trophic resources consistently across their geographic range. Species breeding in north temperate regions (Iceland, Scotland and Northern Ireland) showed enriched δ15N compared to the same or similar species breeding in tropical and subtropical regions, suggesting some differences in baseline levels between these regions. The present study illustrates a noticeable trophic segregation of northeast Atlantic Procellariiformes. Our results show that the isotopic approach has limited applicability for the study of animal movements in the northeast Atlantic at a regional scale, but is potentially useful for the study of long-distance migrations between large marine systems

Geographic Variation in Sea Otters, Enhydra lutris

Univariate and multivariate analyses of 20 skull characters of 304 adult sea otters from throughout the geographic range strongly suggest that three subspecies should be recognized. The nominate form, Enhydra lutris lutris, occurs from the Kuril Islands north to the Commander Islands in the western Pacific Ocean. Individuals of E. l. lutris are characterized by large size and wide skulls with short nasal bones. E. 1. nereis is found along the California coast and off San Nicolas Island, where the species recently has been reintroduced from coastal California. Specimens of E. 1. nereis have narrow skulls with a long rostrum and small teeth, and usually lack the characteristic notch in the postorbital region found in most specimens of the other two subspecies. A new subspecies described by Don E. Wilson in this report, occurs throughout the Aleutian Islands and southward in the eastern Pacific to Washington. Specimens of the new subspecies are intermediate in size in most, but not all, characters and have longer mandibles than either of the other two subspecies

Molecular phylogeny of the Myzorhynchella Section of Anopheles (Nyssorhynchus) (Diptera: Culicidae): genetic support for recently described and resurrected species

Phylogenetic relationships among species of the Myzorhynchella Section of Anopheles (Nyssorhynchus) were investigated using the nuclear ribosomal DNA second internal transcribed spacer (ITS2), the nuclear whitegene and mitochondrial cytochrome oxidase subunit I (COI) regions. The recently described Anopheles pristinus and resurrected Anopheles guarani were also included in the study. Bayesian phylogenetic analyses found Anopheles parvus to be the most distantly related species within the Section, a finding that is consistent with morphology. An. pristinus and An. guarani were clearly resolved from Anopheles antunesi and Anopheles lutzii, respectively. An. lutzii collected in the same mountain range as the type locality were found within a strongly supported clade, whereas individuals from the southern state of Rio Grande do Sul, tentatively identified as An. lutzii based on adult female external morphology, were distinct from An. lutzii, An. antunesi and from each other, and may therefore represent two new sympatric species. A more detailed examination of An. lutzii sensu latoalong its known geographic range is recommended to resolve these anomalous relationships.

Phylogeography of the common vampire bat (Desmodus rotundus): Marked population structure, Neotropical Pleistocene vicariance and incongruence between nuclear and mtDNA markers

Background: The common vampire bat Desmodus rotundus is an excellent model organism for studying ecological vicariance in the Neotropics due to its broad geographic range and its preference for forested areas as roosting sites. With the objective of testing for Pleistocene ecological vicariance, we sequenced a mitocondrial DNA (mtDNA) marker and two nuclear markers (RAG2 and DRB) to try to understand how Pleistocene glaciations affected the distribution of intraspecific lineages in this bat. Results: Five reciprocally monophyletic clades were evident in the mitochondrial gene tree, and in most cases with high bootstrap support: Central America (CA), Amazon and Cerrado (AMC), Pantanal (PAN), Northern Atlantic Forest (NAF) and Southern Atlantic Forest (SAF). The Atlantic forest clades formed a monophyletic clade with high bootstrap support, creating an east/west division for this species in South America. On the one hand, all coalescent and non-coalescent estimates point to a Pleistocene time of divergence between the clades. On the other hand, the nuclear markers showed extensive sharing of haplotypes between distant localities, a result compatible with male-biased gene flow. In order to test if the disparity between the mitochondrial and nuclear markers was due to the difference in mutation rate and effective size, we performed a coalescent simulation to examine the feasibility that, given the time of separation between the observed lineages, even with a gene flow rate close to zero, there would not be reciprocal monophyly for a neutral nuclear marker. We used the observed values of theta and an estimated mutation rate for the nuclear marker gene to perform 1000 iterations of the simulation. The results of this simulation were inconclusive: the number of iterations with and without reciprocal monophyly of one or more clades are similar. Conclusions: We therefore conclude that the pattern exhibited by the common vampire bat, with marked geographical structure for a mitochondrial marker and no phylogeographic structure for nuclear markers is compatible with a historical scenario of complete isolation of refuge-like populations during the Pleistocene. The results on demographic history on this species is compatible with the Carnaval-Moritz model of Pleistocene vicariance, with demographic expansions in the southern Atlantic forest.

CONTRASTING PHYLOGEOGRAPHIC PATTERNS IN MITOCHONDRIAL DNA AND MICROSATELLITES: EVIDENCE OF FEMALE PHILOPATRY AND MALE-BIASED GENE FLOW AMONG REGIONAL POPULATIONS OF THE BLUE-AND-YELLOW MACAW (PSITTACIFORMES: ARA ARARAUNA) IN BRAZIL

Comparing the patterns of population differentiation among genetic markers with different modes of inheritance call provide insights into patterns of sex-biased dispersal and gene flow. The blue-and-yellow Macaw (Ara ararauna) is a Neotropical parrot with a broad geographic distribution ill South America. However, little is known about the natural history and current status Of remaining wild populations, including levels of genetic variability. The progressive decline and possible fragmentation of populations may endanger this species in the near future. We analyzed mitochondrial DNA (mtDNA) control-region sequences and six microsatellite 106 Of Blue-and-yellow Macaws sampled throughout their geographic range ill Brazil to describe population genetic Structure, to make inferences about historical demography and dispersal behavior, and to provide insight for conservation efforts. Analyses of population genetic structure based on mtDNA showed evidence of two major populations ill western and eastern Brazil that share a few low-frequency haplotypes. This phylogeographic pattern seems to have originated by the historical isolation of Blue-and-yellow Macaw populations similar to 374,000 years ago and has been maintained by restricted gene flow and female philopatry. By contrast, variation ill biparentally inherited microsatellites was not structured geographically, Male-biased dispersal and female philopatry best explain the different patterns observed in these two markers. Because females disperse less than males, the two regional populations with well-differentiated mtDNA haplogroups should be considered two different management units for conservation purposes. Received 4 November 2007 accepted 10 December 2008.

Low intraspecific variation in the rRNA internal transcribed spacer 2 (ITS2) of the Australian paralysis tick, Ixodes holocyclus

Ixodes holocyclus has a narrow, discontinuous distribution along the east coast of Australia. We studied ticks from 17 localities throughout the geographic range of this tick. The ITS2 of I. holocyclus is 793 bp long. We found nucleotide variation at eight of the 588 nucleotide positions (1.4%) that were compared for all ticks. There were eight different nucleotide sequences. Most sequences were not restricted to a particular geographic region. However, sequences F, G and H, which had an adenine at position 197, were found only in the far north of Queensland - all other ticks had a guanine at this position. The low level of intraspecific variation in this tick (0.7%) contrasts with the sequence divergence between L holocyclus and its close relative, I. cornuatus (13.1 %). These data indicate that L holocyclus does not contain cryptic species despite possible geographic isolation of some populations. We conclude that variation in the ITS2 is likely to be informative about the phylogeny of the group.

Poracanthodid acanthodian from the Upper Silurian (Pridoli) of Nevada

Dissociated remains of the acanthodian Poracanthodes punctatus are described from Upper Silurian (Pridoli) limestones of the Roberts Mountains Formation at Pete Hanson Creek, Eureka County, Nevada. The vertebrate microremains in sample residues comprise scales, a dentigerous jaw bone fragment, and a fin spine fragment assigned to P. punctatus, plus one possible acanthothoracid placoderm scale. Some macroremains from the same locality are also assigned to P. punctatus. This taxon has been nominated as, a zone fossil for the Silurian vertebrate biozonal scheme, and its presence has been recorded throughout the circum-Arctic region. Identification of the taxon in Nevada extends its known geographic range.

Filogenia e evolução de roedores Echimyidae na Mata Atlântica

Os roedores Echimyidae tem distribuição Neotropical e são a família mais diversa de roedores Caviomorpha. Apesar da grande diversidade, pouco se sabe sobre a distribuição geográfica, história natural e evolução de vários grupos de equimídeos. O histórico taxonômico dessa família é confuso, sendo alguns grupos raramente coletados e, consequentemente, inferências sobre aspectos evolutivos e biológicos são pouco conclusivas e limitadas à análise de poucos exemplares. Filogenias moleculares não corroboram a classificação taxonômica para a família baseada em dados morfológicos, evidenciando a complexidade da história evolutiva desse grupo. Na Mata Atlântica são registrados cinco gêneros de Echimyidae: o rato-do-bambu, Kannabateomys; os arborícolas Phyllomys e Callistomys; o terrestre Trinomys, e o semi-fossorial Euryzygomatomys. O presente trabalho se baseou na utilização de sequências de DNA para abordar aspectos da evolução e filogenia de roedores equimídeos da Mata Atlântica em três níveis taxonômicos: família, gênero e espécie. O primeiro capítulo aborda a posição filogenética do gênero Callistomys dentro da família, utilizando sequências de 1 marcador mitocondrial (CitB) e 3 nucleares (GHR, RAG1 e vWF). Os resultados mostram que Callistomys forma um clado com o ratão-do-banhado (Myocastor), roedor semi-aquático das regiões abertas no cone sul da América do Sul e com o rato-de-espinho terrestre Proechimys com ocorrência na Amazônia. Esse clado é irmão de Thrichomys, um equimídeo terrestre que ocupa as áreas secas do centro da América do Sul. O agrupamento encontrado é inesperado, uma vez que seus membros apresentam aspectos morfológico, ecológicos e distribuição geográfica distintos e contrastantes. A filogenia resultante indica que Callistomys não é proximamente relacionado aos outros equimídeos arborícolas e sugere que o hábito arborícolas evoluiu mais de uma vez na família. O segundo capítulo investiga aspectos da filogenia, evolução e limites entre espécies de Phyllomys utilizando dois marcadores mitocondriais (CitB e COI) e três nucleares (GHR, RAG1 e vWF). Foram identificados três grupos principais de espécies: um com distribuição longitudinal pela porção central da Mata Atlântica (P. pattoni (P. mantiqueirensis, Phyllomys sp. 4)); e a partir daí dois outros grupos, um com distribuição na porção norte da Mata Atlântica (Phyllomys sp. 2 (P. blainvilii (P. brasiliensis, P. lamarum))); e outro na porção sul (Phyllomys sp. 3 ((Phyllomys sp. 1, P. lundi), (Phyllomys sp. 5 (P. dasythrix (P. nigrispinus (P. sulinus, Phyllomys sp. 6)))))). Foram identificadas duas linhagens independentes representando possíveis espécies novas, elevando o potencial número de espécies do gênero de 17 para 19. As filogenias associadas aos dados de distribuição geográfica sugerem que a diversificação e distribuição das espécies de Phyllomys foi influenciada pela ação conjunta de vários fatores como atividade neotectônica, gradientes altitudinais e latitudinais e mudanças climáticas que atuaram desde o Mioceno, marcando os primeiros eventos de diversificação do gênero até as especiações mais recentes, no Pleistoceno. O terceiro capítulo avalia a variação genética, distribuição geográfica e status taxonômico da espécie Euryzygomaotmys spinosus utilizando dois marcadores mitocondriais (CitB e D-loop). Os resultados mostraram que E.spinosus apresenta distribuição em áreas de Mata Atlântica e adjacências ao sul do Rio Doce, no Brasil, Paraguai e Argentina, incluindo um registro confirmado no Cerrado. A espécie ocupa habitats muito diversos e pode ser considerada generalista. As populações são geneticamente estruturadas ao longo da sua distribuição e os dados genéticos corroboram a taxonomia atual que considera apenas uma espécie, E. spinosus, para o gênero.