Females better face senescence in the wandering albatross

Sex differences in lifespan and aging are widespread among animals. Since investment in current reproduction can have consequences on other life-history traits, the sex with the highest cost of breeding is expected to suffer from an earlier and/or stronger senescence. This has been demonstrated in polygynous species that are highly dimorphic. However in monogamous species where parental investment is similar between sexes, sex-specific differences in aging patterns of life-history traits are expected to be attenuated. Here, we examined sex and age influences on demographic traits in a very long-lived and sexually dimorphic monogamous species, the wandering albatross (Diomedea exulans). We modelled within the same model framework sex-dependent variations in aging for an array of five life-history traits: adult survival, probability of returning to the breeding colony, probability of breeding and two measures of breeding success (hatching and fledging). We show that life-history traits presented contrasted aging patterns according to sex whereas traits were all similar at young ages. Both sexes exhibited actuarial and reproductive senescence, but, as the decrease in breeding success remained similar for males and females, the survival and breeding probabilities of males were significantly more affected than females. We discuss our results in the light of the costs associated to reproduction, age-related pairing and a biased operational sex-ratio in the population leading to a pool of non-breeders of potentially lower quality and therefore more subject to death or breeding abstention. For a monogamous species with similar parental roles, the patterns observed were surprising and when placed in a gradient of observed age/sex-related variations in life-history traits, wandering albatrosses were intermediate between highly dimorphic polygynous and most monogamous species.     

Breeding abundance of threatened raptors as estimated from occurrence data

A model derived from the negative binomial distribution (NBD) has been proposed to solve the problem of predicting abundance of species from occurrence maps. The viability of NBD was explored for predicting the breeding abundance of five threatened species of raptor: Bonelli's Eagle Hieraaetus fasciatus, Golden Eagle Aquila chrysaetos, Peregrine Falco peregrinus, Lanner Falco biarmicus and Lesser Kestrel Falco naumanni. First, the accuracy of the NBD was tested in a reference area where the species abundance and occurrence were known through intensive field surveys. Next, an estimation of regional abundance derived from NBD was made for each species. These estimates were then compared to the existing regional data for the five raptors. The spatial distributions of the species were strongly aggregated, with F. peregrinus correctly showing the most widespread area of occupancy. The NBD gave a good approximation of the breeding abundance of the raptors, but tended to overestimate real data, particularly the regional data for falcons. Difficulties in species detection, insufficient sampling effort (F. biarmicus) or data collected over long time spans when population size increased (F. naumanni) may have reduced the NBD's resolution power. The ability of the method to predict local abundances over large areas from readily available presence‐absence data, with relatively low fieldwork effort, could have considerable applications in conservation biology.     

Size variation in the postcranium ofAustralopithecus afarensis and extant species of hominoidea

The postcranial sample ofA. afarensis can be divided into two size groups. Among the best preserved elements which are represented by both morphs are the distal femur, proximal ulna, and capitate. The difference between the large and small fossil femora is similar to the difference between average male and femaleG. gorilla andP. pygmaeus. The distal femora ofH. sapiens are less sexually dimorphic while those ofP. paniscus, P. troglodytes, andH. lar are not significantly dimorphic at all. Large and small capitates and proximal ulnae ofA. afarensis differ slightly more than the highly dimorphic species of extant Hominoidea. In my sample of Amerindians, the capitate and proximal ulna are also strongly dimorphic. The two species ofPan have insignificant sexual dimorphism in these traits. There results imply that strong sexual dimorphism in body size is the primitive condition for the large bodied hominoids.     

Sexual dimorphism in proconsul

One of the more important sources of variability in primate species is sexual dimorphism. Most Primates heavier than five kilos bodyweight are sexually dimorphic, both in body size and in shape of certain hard tissues. Despite these facts, most of the fossil Primates from East African Miocene deposits were originally perceived as being monomorphic, a perception which has propogated through the literature. Re-examination ofProconsul from various sites in Western Kenya results in the view that it was as dimorphic in its splanchonocranium and in bodyweight as chimpanzees and gorillas. The clearest evidence comes from Rusing Island, where adequate samples are known of two morphs, traditionally identified as two species, but more likely to represent two sexes of a single species,P. nyanzae. Co-occurrence of the two morphs is 100% at the various Rusinga sites. Less complete samples have been collected from the Tinderet sites os Koru and Songhor, yet what is available shows that similar patterns of dimorphism characterise the speciesP. africanus andP. major, and that the co-occurrence of the two morphs in each species is 100%. The identification of fossils taking into consideration the role of sexual dimorphism clarifies many of the old debates in which individual specimens frequently shifted between different species, mainly on the basis of metric rather than morphologic evidence. Consequently, the distribution of the species ofProconsul is rather different after accounting for dimorphism, than it was before.     

Differential mortality of wintering shorebirds on the Banc d'Arguin, Mauritania, due to predation by large falcons

Predators may influence many aspects of the daily life and seasonal movements of their prey. Here we quantify direct, and evaluate indirect effects of predation by three falcon species (Lanner Falcon Falco biarmicus , Barbary Falcon Falco pelegrinoides and Peregrine Falcon Falco peregrinus ) on coastal shorebirds wintering on the Banc d'Arguin, Mauritania, an area hosting approximately 30% of the East Atlantic Flyway population of shorebirds. On the basis of 754 h of observation over five winters, 97 witnessed attacks and 585 collected prey remains, we show that shorebirds were safer in larger flocks, which tended to be attacked less often. Furthermore, species that forage relatively close to shore and in small flocks were depredated more often than expected from their relative abundance. In three species, Red Knot Calidris canutus canutus , Bar-tailed Godwit Limosa lapponica taymyrensis and Dunlin Calidris alpina , the juveniles were more vulnerable than adults. We estimated that on average 1% of the juvenile and 0.1% of the adult Red Knots present were killed by large falcons each winter. For Red Knots we simultaneously quantified annual survival on the basis of an individual colour-marking programme: mortality due to predation by falcons accounted for an estimated 6.2% (juveniles) and 0.8% (adults) of annual mortality. We suggest that juvenile Red Knots are 10 times as likely to be killed by falcons because they use riskier habitats, i.e. early and late tide foraging areas closer to shores where surprise attacks are both more common and more successful. These results indicate that the strength of indirect effects of predation operating in a shorebird population largely outweigh the effects of mortality per se .     

Extreme queen-worker dimorphism inRopalidia ignobilis,a small-colony wasp (Hymenoptera: Vespidae)

Summary Ropalidia ignobilis dwells in small colonies of 100 or fewer contemporaneous adults, founded by one or a few females. Variation in size is continuous when a number of colonies are combined, yet each colony produces females in two discrete morphological castes. Nest architecture shows that workers abruptly switch from producing small to large larvae. Large females apparently fulfill the role of the dominant reproductive, but small females are not obligatorily sterile. Castes differ in size, allometric relationships between body parts, number of hamuli per mm. wing length, and coloration in a fashion consistent with what is known for more advanced dimorphic species of social wasps. This dimorphism, rare in Polistinae and previously unknown in small-colony, independent-founding species, challenges our understanding of the evolution of advanced insect sociality.     

Out of Africa? Phylogenetic relationships between Falco biarmicus and the other hierofalcons (Aves: Falconidae)

The phylogeographic history of the lanner falcon ( Falco biarmicus ) and the phylogenetic relationships among hierofalcons ( F. biarmicus , Falco cherrug , Falco jugger and Falco rusticolus ) were investigated using mitochondrial (mt) DNA sequences. Of the two non-coding mt sections tested, the control region (CR) appeared more suitable as phylogenetic marker sequence compared with the pseudo control region (ΨCR). For the comprehensive analysis samples from a broad geographic range representing all four hierofalcon species and their currently recognized subspecies were included. Moreover, samples of Falco mexicanus were analysed to elucidate its phylogenetic relationships to the hierofalcons. The sequence data indicate that this species is more closely related to Falco peregrinus than to the hierofalcons. In the DNA-based trees and in the maximum parsimony network all hierofalcons appear closely related and none of the species represents a monophyletic group. The close relationships among haplotypes suggest that the hierofalcon complex is an assemblage of morphospecies not yet differentiated in the genetic markers used in the present study and that the radiation of the four hierofalcon species took place rather recently. Based on the high intraspecific diversity found within F. biarmicus we assume an African origin of the hierofalcon complex. The observed pattern of haplotype distribution in the extant species may be due to incomplete lineage sorting of ancestral polymorphisms, and interspecific gene flow through hybridization.     

Social phenotype extended to communities: Expanded multilevel social selection analysis reveals fitness consequences of interspecific interactions

In social species, fitness consequences are associated with both individual and social phenotypes. Social selection analysis has quantified the contribution of conspecific social traits to individual fitness. There has been no attempt, however, to apply a social selection approach to quantify the fitness implications of heterospecific social phenotypes. Here, we propose a novel social selection based approach integrating the role of all social interactions at the community level. We extended multilevel selection analysis by including a term accounting for the group phenotype of heterospecifics. We analyzed nest activity as a model social trait common to two species, the lesser kestrel (Falco naumanni) and jackdaw (Corvus monedula), nesting in either single‐ or mixed‐species colonies. By recording reproductive outcome as a measure of relative fitness, our results reveal an asymmetric system wherein only jackdaw breeding performance was affected by the activity phenotypes of both conspecific and heterospecific neighbors. Our model incorporating heterospecific social phenotypes is applicable to animal communities where interacting species share a common social trait, thus allowing an assessment of the selection pressure imposed by interspecific interactions in nature. Finally, we discuss the potential role of ecological limitations accounting for random or preferential assortments among interspecific social phenotypes, and the implications of such processes to community evolution.     

WHY SHOULD LEK-BREEDERS BE MONOMORPHIC?

Approximately one-quarter of all lek-breeding bird species are sexually monomorphic. Understanding the significance, if any, of this exception to the usual correlation between sexual selection and dimorphism requires detailed data on the mating systems of both monomorphic and dimorphic species. The capuchinbird (Perissocephalus tricolor) is a sexually monomorphic, lek-breeding member of the cotinga family. I studied the social and sexual behavior of this species, and compared it with the Guianan cock-of-the-rock (Rupicola rupicola), a dimorphic, lekking member of the same family. Male–male competition in capuchinbirds involved direct contests for dominance, rather than territorial displays as in classic lek species. In each year, one dominant individual was able to control the most desired display site on the 8-male lek, and was the only male that copulated. In contrast to dimorphic lek birds, female as well as male capuchinbirds engaged in frequent and intense aggression at the lek, and both males and females engaged in sexual mimicry. I suggest that plumage monomorphism in lek birds has evolved as a result of social competition affecting both sexes. This hypothesis accounts for the exaggerated plumage characters shared by males and females in capuchinbirds and a number of other monomorphic lek birds. The evolution of plumage can best be analyzed as an arms race, in which the balance of selective forces acting on each sex can produce a variety of equilibrium states, ranging from sexual indistinguishability to extreme dimorphism.     

Colony demographics of rare soldier-polymorphic worker caste systems in <Emphasis Type="Italic">Pheidole</Emphasis> ants (Hymenoptera,Formicidae)

Nearly all species in the ant genus Pheidole have dimorphic workers, with distinct small minors and larger soldiers. The size range of both castes is typically narrow. Just seven described species are soldier-polymorphic, with a broad soldier size range. Here, we characterize worker caste allocation and demography in the soldier-polymorphic P. obtusospinosa, P. rhea, and P. tepicana, and the dimorphic P. spadonia for comparison. The head allometry of soldiers in soldier-polymorphic species is strongly positive and that of dimorphic species is negative. Among soldier-polymorphic species, the soldier castes differ from each other in the degree of positive allometry. In addition, they differ in the number of size modes: P. obtusospinosa and P. rhea have two and P. tepicana has one. During colony ontogeny, P. obtusospinosa first has one mode and develops the second mode much later, while P. rhea produces multiple modes throughout. We also characterize worker caste systems based on the biomass allocation. For all three soldier-polymorphic species, the majority of soldiers are small soldiers. Pheidole obtusospinosa and P. rhea allocate roughly equal biomass to the two soldier classes, while P. tepicana allocates little to supersoldiers based on both biomass and caste ratio. These findings illustrate the interplay among caste ratios, biomass allocation, size frequency distributions, and allometry in the evolution of different worker caste systems. We conclude that soldier-polymorphic species may have evolved convergently in response to broad-scale factors, but differences among them suggest selection pressures in small-scale environments have been different.     

Socioecology and the ontogeny of sexual size dimorphism in anthropoid primates

This study examines statistical correlations between socioecological variables (including measures of group composition, intermale competition, and habitat preference) and the ontogeny of body size sexual dimorphism in anthropoid primates. A regression-based multivariate measure of dimorphism in body weight ontogeny is derived from a sample of 37 species. Quantitative estimates of covariation between socioecological variables and this multivariate measure are evaluated. Statistically significant covariation between the ontogeny of dimorphism and socioecological variables, with the possible exception of habitat preference, is observed. Sex differences in ontogeny are lacking in species that exhibit low levels of intermale competition and are classifiable as species with monogamous/polyandrous mating systems. Among dimorphic species, two modes of dimorphic growth are apparent, which seem to be related to different kinds of group compositions. Multimale/multifemale species tend to become dimorphic through bimaturism (sex differences in duration of growth) with minimal sex differences in growth rate. Single-male/multifemale species tend to attain dimorphism through differences in rate of growth, often with limited bimaturism. Measures of intermale competition may also covary with these modes of dimorphic growth, but the relations among these variables are sometimes ambiguous. Correlations between dimorphic growth and behavioral variables may reflect alternative life history strategies in primates. Specifically, the ways in which risks faced by subadult males are distributed and the relations of these risks to growth rates seem to influence the evolution of size ontogenies. The absence of dimorphic ontogeny in some species can be tied to similar distributions of risk in each sex. In taxa that become dimorphic primarily through rate differences in growth, the lifetime distribution of risks for males may change rapidly. In contrast, males may face a pattern of uniformly changing or stable risk in species that become dimorphic through bimaturism. Finally, much variation recorded by this study remains unexplained, providing additional evidence of the need to specially examine female ontogeny before primate body size dimorphism can be satisfactorily explained. © 1995 Wiley-Liss, Inc.     

Temporal distribution of diurnal rapacious birds in "Playón de Mismaloya" Reservation, Jalisco, Mexico

Diurnal raptors were counted monthly in the "Playón de Mismaloya", reserve, Jalisco, Mexico, from November 1997 to October 1998. We identified 11 species; eight of them migrants, one resident and two resident with migrant populations. The highest increase in total number of individuals was from December to March, influenced mainly by the presence of Caracara polancus (Crested Caracara), Falco sparverius (American Kestrel) and Pandion haliaetus (Osprey). The two former species were observed frequently in open areas, where they could detect prey more easily, while ospreys were found only in estuaries, lagoons and beach, where food was available.     

Metric variation and sexual dimorphism in the dentition of Ouranopithecus macedoniensis

The fossil sample attributed to the late Miocene hominoid taxon Ouranopithecus macedoniensis is characterized by a high degree of dental metric variation. As a result, some researchers support a multiple-species taxonomy for this sample. Other researchers do not think that the sample variation is too great to be accommodated within one species. This study examines variation and sexual dimorphism in mandibular canine and postcanine dental metrics of an Ouranopithecus sample. Bootstrapping (resampling with replacement) of extant hominoid dental metric data is performed to test the hypothesis that the coefficients of variation (CV) and the indices of sexual dimorphism (ISD) of the fossil sample are not significantly different from those of modern great apes. Variation and sexual dimorphism in Ouranopithecus M(1) dimensions were statistically different from those of all extant ape samples; however, most of the dental metrics of Ouranopithecus were neither more variable nor more sexually dimorphic than those of Gorilla and Pongo. Similarly high levels of mandibular molar variation are known to characterize other fossil hominoid species. The Ouranopithecus specimens are morphologically homogeneous and it is probable that all but one specimen included in this study are from a single population. It is unlikely that the sample includes specimens of two sympatric large-bodied hominoid species. For these reasons, a single-species hypothesis is not rejected for the Ouranopithecus macedoniensis material. Correlations between mandibular first molar tooth size dimorphism and body size dimorphism indicate that O. macedoniensis and other extinct hominoids were more sexually size dimorphic than any living great apes, which suggests that social behaviors and life history profiles of these species may have been different from those of living species.     

Hunting flight speeds of five southern African raptors

The flight speeds of hunting falconry birds were determined using global positioning system data loggers. Until now, the hunting flight speed of African raptors has not been directly measured. We predicted that hunting flight speeds would differ between species and that flight dynamics, such as altitude, and bird morphology, particularly wing surface area, would influence maximum and mean flight speeds. This study considered five African raptor species, which included two long-wing species, Lanner Falcon Falco biarmicus and Peregrine Falcon F. peregrinus, one short-wing species, Black Sparrowhawk Accipiter melanoleucus, and two broad-wing species, African Hawk-eagle Aquila spilogaster and Jackal Buzzard Buteo rufofuscus. Maximum and mean hunt speeds differed significantly between the long- and short-wing species. There was no difference in acceleration or deceleration rates between these species, but this could be due to small sample sizes. There was a significant positive correlation between maximum hunt speed and maximum flight height for the long-wing species. Maximum and mean flight speeds were significantly negatively correlated with wing area for all five species in this study. However, following phylogenetic correction, no significant relationship between wing area and maximum hunt speeds was found. This study presents baseline data of hunting speeds in African raptors and further highlights the importance of inter-species variation, which can provide accuracy to flight speed models and the understanding of hunting strategies.     

MATING SYSTEM AND SEXCUL DIMORPHISM IN LARGE TERRESTRIAL, MAMMALIAN HERBIVORES

Sexual dimorphism in mammals is not entirely satisfactorily explained by the models that are advanced to account for it among birds. This may be because species-specific styles of being dimorphic, and of attaining mature dimorphic state, are not clearly recognized. Mature dimorphism is a syndrome involving body size, appearance and weaponry; each facet and the whole syndrome may have functions in both fighting and signalling. The mature dimorphic stage has to be reached by growth and change from juvenile and sub-adult states. The occurrence of the separate facets of the dimorphic syndrome are reviewed in species of Bovidae, Cervidae and Macropodidae, large, diverse families of eutherian (the first two) and metatherian mammals, which have broadly similar ecological adaptations. In each family the smallest species tend to be homomorphic, with small, inconspicuous weapons. Greatest dimorphism in size is found in medium-sized bovids and cervids, and the larger macropodids (in which no species exceeds 100 kg male weight); the range of species showing greatest dimorphism in size also shows the most exaggerated weapons. Mature dimorphism is reached by different patterns of growth, which may be determinate and similar in the sexes (leading to homomorphism), determinate but differing between the sexes, or indeterminate and differing, both of which lead to heteromorphism. The syndromes of dimorphism and patterns of growth are associated and a classification of styles of dimorphism is presented. The adaptiveness of the styles is suggested in terms of what is known of the socio-ecology, in particular the male reproductive strategies, of the species. The various styles of heteromorphy appear to be associated with males' way of achieving polygyny: such as by non-resource-based territoriality, by dominance-determined access to oestrous females, or by wandering and formation of a consortship with pro-oestrous females. The relevance of the species' ecology of use of resources to these styles of dimorphism and mate-acquisition is briefly discussed.     

Spatial size dimorphism in New Zealand's last endemic raptor,the Kārearea Falco novaeseelandiae,coincides with a narrow sea strait

Although New Zealand's avifauna includes many unusual birds, species‐level diversity within lineages is typically low. There are, however, several instances where different allied forms are recognized in each of the two main islands. Among them is the Kārearea Falco novaeseelandiae, which is the only surviving endemic raptor species in New Zealand. Recent analysis confirms it to be a distinct lineage in the global radiation of this genus and most closely related to the Aplomado Falcon Falco femoralis of South America. We examined body size metrics and neutral genetic markers in Kārearea sampled across New Zealand to assess subspecific variation within the species. We found strong evidence using linear modelling and Bayesian clustering for two distinct sizes within Kārearea, in addition to the recognized sexual dimorphism. The boundary between the size clusters coincides closely with the Cook Strait, a narrow seaway between the two largest islands. However, analysis of mitochondrial sequence data and nuclear microsatellites showed no compelling partitioning at neutral loci. These data suggest adaptive change along a stepped environmental cline. Lineage splitting in Kārearea has either yet to become apparent in the distribution of neutral genetic variation and/or regional adaptation is proceeding despite gene flow.     

The impact of predation by birds on bat populations in the British Isles

The role played by predation of birds in the mortality of British bats is assessed. A review of dietary studies and anecdotal accounts revealed eleven species of bird which occasionally feed on bats–Herring Gull Larus argentatus , Black-headed Gull Larus ridibundus , Rook Corvus frugilegus , Carrion Crow Corvus corone , Little Owl Athene noctua , Short-eared Owl Asio flammeus , Kestrel Falco tinnunculus , Hobby Falco subbuteo , Merlin Falco columbarius , Peregrine Falco peregrinus and Sparrowhawk Accipiter nisus. A further three species feed more frequently on bats–Barn Owl Tyto alba , Tawny Owl Strix aluco and Long-eared Owl Asio otus. Rates of predation were very low accounting for only 00034^o_o of items taken by small hawks and falcons ( n items = 29 519) but 0035^o, of prey taken by owls ( n items = 99 479). By multiplying together the average annual prey capture rates of the predators, assessed from their energetic food requirements and direct observations of prey intake rates, the British populations of the predators and the contribution to the diet made by bats, the annual number of bats which die each year as a result of predation was estimated. The total losses of bats to predation might amount to about 201 400 bats/annum. The most significant predators are Tawny Owl ( c. 168 850 bats/annum), Barn Owl (c. 8800 bats/ annum), Long-eared Owl (c. 10 200 bats/annum) and Kestrels (c. 8400 bats/annum). This predation by avian predators would account for about 11 % of the annual mortality of British bats. An assessment of the biases involved in this calculation suggests it is probably a minimum estimate. Despite the apparent low representation of bats in the diets of predatory birds, the effects of this predation on bat behaviour and population dynamics cannot be ignored.     

Evolution of host use in fruit flies of the genus Blepharoneura (Diptera: Tephritidae): cryptic species on sexually dimorphic host plants

Blepharoneura , a neotropical genus of tephritid fruit flies, includes newly discovered morphologically cryptic species that are highly host specific. Several species of fly, each specific to a different host tissue can infest a single species of sexually dimorphic host. Using allozyme electrophoresis, we analysed morphologically similar populations of flies reared from various tissues of five species of host plant representing three subtribes of the Cucurbitaceae. In Venezuela, we found three sympatric species of Blepharoneura feeding on Gurania spinulosa Cogn. (one species on male flowers, another on female flowers, and a third on seeds). In Costa Rica, we found four non-interbreeding populations of Blepharoneura feeding on G. costaricensis Cogn. : two sympatric 'lowland' populations (one feeding on male flowers and another on female flowers) and two sympatric 'highland' populations (also specific to flowers of different sexes). Phylogenetic analyses revealed a species-rich clade of specialists associated with flowers of the Guraniinae, and a species-poor clade of generalists associated with seeds of the Guraniinae. Specialization on male vs. female flowers appears to have occurred repeatedly. Patterns of laryal dispersal may underlie differehces in diversification of seed vs. flower clades: larvae that feed on seeds are dispersed by bats; larvae feeding on flowers are not dispersed.     

Ancestral lineages of arbuscular mycorrhizal fungi (Glomales)

Using new and existing 18S rRNA sequence data, we show that at least five species of glomalean fungi lie outside the previously defined families and diverged very early in the evolution of that group. These five fungi would have been missed by many previous ecological studies because their sequences are not well matched to available taxon-specific primers and they do not stain well with the standard reagents used for morphological analysis. Based upon spore morphology, these species are currently assigned to Glomus and Acaulospora, and two of the species are dimorphic, exhibiting spore stages of both genera. This suggests that dimorphic spores are the ancestral state for the order and that one or the other morphology was lost in various lineages. Our analyses also show that Geosiphon pyriforme, a symbiont with cyanobacteria, is not necessarily a sister group of the Glomales; instead, it may be derived from mycorrhizal ancestors.