Lower Detection Probability of Avian Plasmodium in Blood Compared to Other Tissues.

We tested whether the probability of detecting avian haemosporidia (Plasmodium and Haemoproteus) using molecular techniques differs among blood, liver, heart, and pectoral muscle tissues. We used a paired design, sampling the 4 tissue types in 55 individuals of a wild South American suboscine antbird, the white-shouldered fire-eye (Pyriglena leucoptera). We also identified parasites to cytochrome b lineage. Detection probability was significantly lower in blood compared to the other 3 tissue types combined. Eight of 22 infections were not detected in blood samples; 4-7 infections were not detected in the other individual tissues. The same parasite lineage was recovered from different tissues.

Increased lipid peroxidation occurs during development in Japanese quail (Coturnix japonica) embryos.

The growth of the avian embryo is paralleled by an exponential increase in the rate of whole-embryonic oxygen consumption, which potentially increases oxidative damage. Age-related patterns of tissue lipid peroxidation were characterised in brain, liver and heart tissue of developing Japanese quail (Coturnix japonica) embryos between 9 and 15 d of age, over which embryo mass increased by a factor of 6. Lipid peroxidation was quantified in each tissue by spectrophotometric measurement of malondialdehyde using the thiobarbituric acid reactive substances assay. In all tissues, lipid peroxidation increased greatly as development proceeded. Concentrations of malondialdehyde increased in parallel with the cumulative amount of oxygen consumed by the developing embryo consistent with the hypothesis that oxidative stress results from the production of free radicals due to oxidative metabolism. This study describes in vivo oxidative stress in developing avian embryos and suggests that rates of embryonic growth, oxidative metabolism and oxidative damage likely vary in parallel.

Specialized avian Haemosporida trade reduced host breadth for increased prevalence.

Parasite specialization on one or a few host species leads to a reduction in the total number of available host individuals, which may decrease transmission. However, specialists are thought to be able to compensate by increased prevalence in the host population and increased success in each individual host. Here, we use variation in host breadth among a community of avian Haemosporida to investigate consequences of generalist and specialist strategies on prevalence across hosts. We show that specialist parasites are more prevalent than generalist parasites in host populations that are shared between them. Moreover, the total number of infections of generalist and specialist parasites within the study area did not vary significantly with host breadth. This suggests that specialists can infect a similar number of host individuals as generalists, thus compensating for a reduction in host availability by achieving higher prevalence in a single host species. Specialist parasites also tended to infect older hosts, whereas infections by generalists were biased towards younger hosts. We suggest that this reflects different abilities of generalists and specialists to persist in hosts following infection. Higher abundance and increased persistence in hosts suggest that specialists are more effective parasites than generalists, supporting the existence of a trade-off between host breadth and average host use among these parasites.

Niche conservatism constrains Australian honeyeater assemblages in stressful environments.

The hypothesis of phylogenetic niche conservatism proposes that most extant members of a clade remain in ancestral environments because expansion into new ecological space imposes a selectional load on a population. A prediction that follows is that local assemblages contain increasingly phylogenetically clustered subsets of species with increasing difference from the ancestral environment of a clade. We test this in Australian Meliphagidae, a continental radiation of birds that originated in wet, subtropical environments, but subsequently spread to drier environments as Australia became more arid during the late Cenozoic. We find local assemblages are increasingly phylogenetically clustered along a gradient of decreasing precipitation. The pattern is less clear along a temperature gradient. We develop a novel phyloclimatespace to visualise the expansion of some lineages into drier habitats. Although few species extend into arid regions, those that do occupy larger ranges and thus local species richness does not decline predictably with precipitation.

Rivers, refuges and population divergence of fire-eye antbirds (Pyriglena) in the Amazon Basin.

The identification of ecological and evolutionary mechanisms that might account for the elevated biotic diversity in tropical forests is a central theme in evolutionary biology. This issue is especially relevant in the Neotropical region, where biological diversity is the highest in the world, but where few studies have been conducted to test factors causing population differentiation and speciation. We used mtDNA sequence data to examine the genetic structure within white-backed fire-eye (Pyriglena leuconota) populations along the Tocantins River valley in the south-eastern Amazon Basin, and we confront the predictions of the river and the Pleistocene refuge hypotheses with patterns of genetic variation observed in these populations. We also investigated whether these patterns reflect the recently detected shift in the course of the Tocantins River. We sampled a total of 32 individuals east of, and 52 individuals west of, the Tocantins River. Coalescent simulations and phylogeographical and population genetics analytical approaches revealed that mtDNA variation observed for fire-eye populations provides little support for the hypothesis that populations were isolated in glacial forest refuges. Instead, our data strongly support a key prediction of the river hypothesis. Our study shows that the Tocantins River has probably been the historical barrier promoting population divergence in fire-eye antbirds. Our results have important implications for a better understanding of the importance of large Amazonian rivers in vertebrate diversification in the Neotropics.

On the phylogenetic relationships of haemosporidian parasites from raptorial birds (Falconiformes and Strigiformes).

Haemosporidian parasite diversity among raptorial birds (hawks and owls), as estimated by DNA sequencing, is proving to be greater than previously anticipated from taxonomic assessments based on parasite morphology. Here, we place raptor parasites in a phylogenetic context, including new parasite cytochrome b (cyt b) sequences from North America and Europe and from a variety of host species not previously sampled. Mitochondrial DNA sequences reveal raptor-specific parasite clades within Parahaemoproteus, but not within Plasmodium. We also recovered a strikingly divergent clade of raptor parasites that aligns with neither genus, but groups with both as a sister clade to Leucocytozoon. Different cyt b primer sets recovered additional sequences from 3 of these samples, which grouped with Parahaemoproteus in 2 cases and with Plasmodium in 1 case. Possible explanations (after excluding contamination) include multiple infections, alternative cyt b copies within the mitochondrial genome, and nuclear copies of mitochondrial genes. We believe the latter 2 explanations are unlikely because these divergent cyt b lineages form a single clade and were also recovered with several additional genomic markers.

The evolution of dioecy, heterodichogamy, and labile sex expression in Acer.

The northern hemisphere tree genus Acer comprises 124 species, most of them monoecious, but 13 dioecious. The monoecious species flower dichogamously, duodichogamously (male, female, male), or in some species heterodichogamously (two morphs that each produce male and female flowers but at reciprocal times). Dioecious species cannot engage in these temporal strategies. Using a phylogeny for 66 species and subspecies obtained from 6600 nucleotides of chloroplast introns, spacers, and a protein-coding gene, we address the hypothesis (Pannell and Verdú, Evolution 60: 660-673. 2006) that dioecy evolved from heterodichogamy. This hypothesis was based on phylogenetic analyses (Gleiser and Verdú, New Phytol. 165: 633-640. 2005) that included 29-39 species of Acer coded for five sexual strategies (duodichogamous monoecy, heterodichogamous androdioecy, heterodichogamous trioecy, dichogamous subdioecy, and dioecy) treated as ordered states or as a single continuous variable. When reviewing the basis for these scorings, we found errors that together with the small taxon sample, cast doubt on the earlier inferences. Based on published studies, we coded 56 species of Acer for four sexual strategies, dioecy, monoecy with dichogamous or duodichogamous flowering, monoecy with heterodichogamous flowering, or labile sex expression, in which individuals reverse their sex allocation depending on environment-phenotype interactions. Using Bayesian character mapping, we infer an average of 15 transformations, a third of them involving changes from monoecy-cum-duodichogamy to dioecy; less frequent were changes from this strategy to heterodichogamy; dioecy rarely reverts to other sexual systems. Contra the earlier inferences, we found no switches between heterodichogamy and dioecy. Unexpectedly, most of the species with labile sex expression are grouped together, suggesting that phenotypic plasticity in Acer may be a heritable sexual strategy. Because of the complex flowering phenologies, however, a concern remains that monoecy in Acer might not always be distinguishable from labile sex expression, which needs to be addressed by long-term monitoring of monoecious trees. The 13 dioecious species occur in phylogenetically disparate clades that date back to the Late Eocene and Oligocene, judging from a fossil-calibrated relaxed molecular clock.

Evolutionary diversification of clades of squamate reptiles.

We analysed the diversification of squamate reptiles (7488 species) based on a new molecular phylogeny, and compared the results to similar estimates for passerine birds (5712 species). The number of species in each of 36 squamate lineages showed no evidence of phylogenetic conservatism. Compared with a random speciation-extinction process with parameters estimated from the size distribution of clades, the alethinophidian snakes (2600 species) were larger than expected and 13 clades, each having fewer than 20 species, were smaller than expected, indicating rate heterogeneity. From a lineage-through-time plot, we estimated that a provisional rate of lineage extinction (0.66 per Myr) was 94% of the rate of lineage splitting (0.70 per Myr). Diversification in squamate lineages was independent of their stem age, but strongly related to the area of the region within which they occur. Tropical vs. temperate latitude exerted a marginally significant influence on species richness. In comparison with passerine birds, squamates share several clade features, including: (1) independence of species richness and age; (2) lack of phylogenetic signal with respect to clade size; (3) general absence of exceptionally large clades; (4) over-representation of small clades; (5) influence of region size on clade size; and (6) similar rates of speciation and extinction. The evidence for both groups suggests that clade size has achieved long-term equilibrium, suggesting negative feedback of species richness on the rate of diversification.

Host specificity of Lepidoptera in tropical and temperate forests.

For numerous taxa, species richness is much higher in tropical than in temperate zone habitats. A major challenge in community ecology and evolutionary biogeography is to reveal the mechanisms underlying these differences. For herbivorous insects, one such mechanism leading to an increased number of species in a given locale could be increased ecological specialization, resulting in a greater proportion of insect species occupying narrow niches within a community. We tested this hypothesis by comparing host specialization in larval Lepidoptera (moths and butterflies) at eight different New World forest sites ranging in latitude from 15 degrees S to 55 degrees N. Here we show that larval diets of tropical Lepidoptera are more specialized than those of their temperate forest counterparts: tropical species on average feed on fewer plant species, genera and families than do temperate caterpillars. This result holds true whether calculated per lepidopteran family or for a caterpillar assemblage as a whole. As a result, there is greater turnover in caterpillar species composition (greater beta diversity) between tree species in tropical faunas than in temperate faunas. We suggest that greater specialization in tropical faunas is the result of differences in trophic interactions; for example, there are more distinct plant secondary chemical profiles from one tree species to the next in tropical forests than in temperate forests as well as more diverse and chronic pressures from natural enemy communities.

Climatic unpredictability and parasitism of caterpillars: implications of global warming.

Insect outbreaks are expected to increase in frequency and intensity with projected changes in global climate through direct effects of climate change on insect populations and through disruption of community interactions. Although there is much concern about mean changes in global climate, the impact of climatic variability itself on species interactions has been little explored. Here, we compare caterpillar-parasitoid interactions across a broad gradient of climatic variability and find that the combined data in 15 geographically dispersed databases show a decrease in levels of parasitism as climatic variability increases. The dominant contribution to this pattern by relatively specialized parasitoid wasps suggests that climatic variability impairs the ability of parasitoids to track host populations. Given the important role of parasitoids in regulating insect herbivore populations in natural and managed systems, we predict an increase in the frequency and intensity of herbivore outbreaks through a disruption of enemy-herbivore dynamics as climates become more variable.

Temporal stability of insular avian malarial parasite communities.

Avian malaria is caused by a diverse community of genetically differentiated parasites of the genera Plasmodium and Haemoproteus. Rapid seasonal and annual antigenic allele turnover resulting from selection by host immune systems, as observed in some parasite populations infecting humans, may extend analogously to dynamic species compositions within communities of avian malarial parasites. To address this issue, we examined the stability of avian malarial parasite lineages across multiple time-scales within two insular host communities. Parasite communities in Puerto Rico and St Lucia included 20 and 14 genetically distinct parasite lineages, respectively. Lineage composition of the parasite community in Puerto Rico did not vary seasonally or over a 1 year interval. However, over intervals approaching a decade, the avian communities of both islands experienced an apparent loss or gain of one malarial parasite lineage, indicating the potential for relatively frequent lineage turnover. Patterns of temporal variation of parasite lineages in this study suggest periodic colonization and extinction events driven by a combination of host-specific immune responses, competition between lineages and drift. However, the occasional and ecologically dynamic lineage turnover exhibited by insular avian parasite communities is not as rapid as antigenic allele turnover within populations of human malaria.

Detecting avian malaria: an improved polymerase chain reaction diagnostic.

We describe a polymerase chain reaction (PCR) assay that detects avian malarial infection across divergent host species and parasite lineages representing both Plasmodium spp. and Haemoproteus spp. The assay is based on nucleotide primers designed to amplify a 286-bp fragment of ribosomal RNA (rRNA) coding sequence within the 6-kb mitochondrial DNA malaria genome. The rRNA malarial assay outperformed other published PCR diagnostic methods for detecting avian infections. Our data demonstrate that the assay is sensitive to as few as 10(-5) infected erythrocytes in peripheral blood. Results of avian population surveys conducted with the rRNA assay suggest that prevalences of malarial infection are higher than previously documented, and that studies based on microscopic examination of blood smears may substantially underestimate the extent of parasitism by these apicomplexans. Nonetheless, because these and other published primers miss small numbers of infections detected by other methods, including inspection of smears, no assay now available for avian malaria is universally reliable.

Age-related patterns of fertility in captive populations of birds and mammals.

We have analyzed birth records from the International Species Inventory System database of zoo animals to describe patterns of fertility as a function of age in birds and mammals. Unlike age at death, fertility in captive populations is partly controlled by animal keepers depending on availability of space and potential mates and on the aims of captive propagation programs. Thus, fertility records must be considered carefully to avoid potential age-specific biases. The zoo data suggest that fertility declines from a peak at young adult ages in most species. The rate of decline in reproductive function is correlated with the rate of somatic aging, indicated by the increase in mortality rate with age, and with the age at peak fertility. Over the sample as a whole, these patterns do not differ systematically between the sexes, nor do they differ between birds and mammals. While it is clear that the demographic consequences of aging of the reproductive system follow a different course than the aging of somatic tissue, the rates of each are correlated either through functional connections or through parallel evolutionary responses to common selective factors.

Host plants influence parasitism of forest caterpillars.

Patterns of association between herbivores and host plants have been thought to reflect the quality of plants as food resources as influenced by plant nutrient composition, defences, and phenology. Host-plant-specific enemies, that is, the third trophic level, might also influence the distribution of herbivores across plant species. However, studies of the evolution of herbivore host range have generally not examined the third trophic level, leaving unclear the importance of this factor in the evolution of plant-insect herbivore interactions. Analysis of parasitoid rearings by the Canadian Forest Insect Survey shows that parasitism of particular Lepidoptera species is strongly host-plant-dependent, that the pattern of host-plant dependence varies among species of caterpillars, and that some parasitoid species are themselves specialized with respect to tree species. Host-plant-dependent parasitism suggests the possibility of top-down influence on host plant use. Differences in parasitism among particular caterpillar-host plant combinations could select for specialization of host plant ranges within caterpillar communities. Such specialization would ultimately promote the species diversification of Lepidoptera in temperate forests with respect to escape from enemies.

Physiological, growth, and immune responses of Japanese quail chicks to the multiple stressors of immunological challenge and lead shot.

Exposure to the combination of a contaminant and an immunological challenge during development may greatly increase the impact of either or both of these stressors on an individual. This study investigated the interacting effects of a nonpathogenic immunological challenge and lead shot exposure early in the development of a precocial species. Seventy-one quail (Coturnix coturnix japonica) chicks orally received either one #9 lead shot (0.05 g), four lead shots (0.2 g), or no lead at the age of 8 days. A third of each of these groups of chicks were intraperitoneally injected with either 0.075 ml of 10% chukar (Alectoris graeca) red blood cells (CRBCs), Newcastle disease virus (NDV), or a placebo oil vaccine at 13 and 35 days of age. There was no difference in the survival between any of the lead or antigen treatment groups. Lead concentrations in blood were greater for the lead-dosed groups on day 49 but not on day 128. Growth curves were approximated by Weibull functions; growth parameters did not differ between lead-treated and antigen-treated birds. CRBC antigen increases fluctuating asymmetry (FA) for primary feather five, while NDV reduced FA. Lead did not affect antibody production or cell-mediated immune response. White blood cell numbers increased 7 days after antigen injection on days 27 and 49 of age. Granulocyte numbers were significantly higher for the lead-treated quail than the control quail, and both antigen-treated groups had lower granulocyte numbers than control quail. The treatment groups of combining NDV and lead shot had an average plasma protein 14% lower than other groups. Lead shot in the highest of these doses lowered asymptotic weights, increased hematocrits, lowered plasma protein, and increased granulocyte numbers of quail.

Nonequilibrium diversity dynamics of the Lesser Antillean avifauna.

MacArthur and Wilson's model of island diversity predicts an increase in the number of species until colonization and extinction are balanced at a long-term steady state. We appraise this model on an evolutionary time scale by molecular phylogenetic analysis of the colonization of the Lesser Antilles by small land birds. The pattern of accumulation of species with time, estimated by genetic divergence between island and source lineages, rejects a homogeneous model of colonization and extinction. Rather, our results suggest an abrupt, roughly 10-fold increase in colonization rate or a 90% mass extinction event 0.55 to 0.75 million years ago.

The molecular basis of an avian plumage polymorphism in the wild: a melanocortin-1-receptor point mutation is perfectly associated with the melanic plumage morph of the bananaquit, Coereba flaveola.

BACKGROUND: Evolution depends on natural selection acting on phenotypic variation, but the genes responsible for phenotypic variation in natural populations of vertebrates are rarely known. The molecular genetic basis for plumage color variation has not been described in any wild bird. Bananaquits (Coereba flaveola) are small passerine birds that occur as two main plumage variants, a widespread yellow morph with dark back and yellow breast and a virtually all black melanic morph. A candidate gene for this color difference is the melanocortin-1 receptor (MC1R), a key regulator of melanin synthesis in feather melanocytes. RESULTS: We sequenced the MC1R gene from four Caribbean populations of the bananaquit; two populations of the yellow morph and two populations containing both the yellow morph and the melanic morph. A point mutation resulting in the replacement of glutamate with lysine was present in at least one allele of the MC1R gene in all melanic birds and was absent in all yellow morph birds. This substitution probably causes the color variation, as the same substitution is responsible for melanism in domestic chickens and mice. The evolutionary relationships among the MC1R haplotypes show that the melanic alleles on Grenada and St. Vincent had a single origin. The low prevalence of nonsynonymous substitutions among yellow haplotypes suggests that they have been under stabilizing selection, whereas strong selective constraint on melanic haplotypes is absent. CONCLUSIONS: We conclude that a mutation in the MC1R is responsible for the plumage polymorphism in a wild bird population and that the melanic MC1R alleles in Grenada and St. Vincent bananaquit populations have a single evolutionary origin from a yellow allele.

Comparison of aging-related mortality among birds and mammals.

We use the Weibull model to characterize initial (extrinsic) mortality rates (m(0)) and rate of increase in mortality with age (omega) for natural and captive populations of birds and mammals. Weibull parameters can be estimated for small samples of ages at death by constructing survival curves and fitting the Weibull model by nonlinear least-squares regression. Both m(0) and omega decrease in captivity, on average, and omega bears a strong relationship to m(0), as it does in nature, irrespective of body mass or differences between birds and mammals. Rate of aging is most closely related to brain size in birds and to rate of postnatal growth in mammals. It is not related to duration of embryonic development, body size independently of brain size, or genome size. We suggest that causes of extrinsic mortality in nature may be replaced in captivity by intrinsically controlled causes of mortality related to processes that regulate the rate of aging.

Catabolic capacity of the muscles of shorebird chicks: maturation of function in relation to body size.

Newly hatched precocial chicks of arctic shorebirds are able to walk and regulate their body temperatures to a limited extent. Yet, they must also grow rapidly to achieve independence before the end of the short arctic growing season. A rapid growth rate may conflict with development of mature function, and because of the allometric scaling of thermal relationships, this trade-off might be resolved differently in large and small species. We assessed growth (mass) and functional maturity (catabolic enzyme activity) in leg and pectoral muscles of chicks aged 1-16 d and adults of two scolopacid shorebirds, the smaller dunlin (Calidris alpina: neonate mass 8 g, adult mass 50 g) and larger whimbrel (Numenius phaeopus; neonate mass 34 g, adult mass 380 g). Enzyme activity indicates maximum catabolic capacity, which is one aspect of the development of functional maturity of muscle. The growth rate-maturity hypothesis predicts that the development of catabolic capacity should be delayed in faster-growing muscle masses. Leg muscles of both species were a larger proportion of adult size at hatching and grew faster than pectoral muscles. Pectoral muscles grew more rapidly in the dunlin than in the whimbrel, whereas leg muscles grew more rapidly in the whimbrel. In both species and in both leg and pectoral muscles, enzyme activities generally increased with age, suggesting increasing functional maturity. Levels of citrate synthase activity were similar to those reported for other species, but l-3-hydroxyacyl-CoA-dehydrogenase and pyruvate kinase (PK) activities were comparatively high. Catabolic capacities of leg muscles were initially high compared to those of pectoral muscles, but with the exception of glycolytic (PK) capacities, these subsequently increased only modestly or even decreased as chicks grew. The earlier functional maturity of the more rapidly growing leg muscles, as well as the generally higher functional maturity in muscles of the more rapidly growing dunlin chicks, contradicts the growth rate-maturity function trade-off and suggests that birds have considerable latitude to modify this relationship. Whimbrel chicks, apparently, can rely on allometric scaling of power requirements for locomotion and the thermal inertia of their larger mass to reduce demands on their muscles, whereas dunlin chicks require muscles with higher metabolic capacity from an earlier age. Thus, larger and smaller species may adopt different strategies of growth and tissue maturation.