Experimental evidence that adult antipredator behaviour is heritable and not influenced by behavioural copying in a wild bird.

Knowledge of the relative importance of genetics and behavioural copying is crucial to appraise the evolvability of behavioural consistencies. Yet, genetic and non-genetic factors are often deeply intertwined, and experiments are required to address this issue. We investigated the sources of variation of adult antipredator behaviour in the Alpine swift (Apus melba) by making use of long-term behavioural observations on parents and cross-fostered offspring. By applying an 'animal model' approach to observational data, we show that antipredator behaviour of adult Alpine swifts was significantly repeatable over lifetime (r = 0.273) and heritable (h(2) = 0.146). Regression models also show that antipredator behaviours differed between colonies and sexes (females were more tame), and varied with the hour and year of capture. By applying a parent-offspring regression approach to 59 offspring that were exchanged as eggs or hatchlings between pairs of nests, we demonstrate that offspring behaved like their biological parents rather than like their foster parents when they were adults themselves. Those findings provide strong evidence that antipredator behaviour of adult Alpine swifts is shaped by genetics and/or pre-hatching maternal effects taking place at conception but not by behavioural copying.

Philippe Garrel: Gramàtica de la llum i de l’ombra

En el següent estudi s’analitza en profunditat el cinema de Philippe Garrel a través dels paràmetres fotogràfics de la seva obra. Centrat fonamentalment en les primeres pel∙lícules de la seva trajectòria underground –de Le Révélateur (1968) a L’Enfant secret (1979)‐ aquesta recerca s’articula al voltant de la repercussió simbòlica, plàstica i narrativa dels usos lumínics, definits com la gramàtica de la llum i de l’ombra. La memòria presenta, en primer lloc, la llum com a essència reveladora, abordant el paradigma solar, les fonts de llum artificials –els canelobres i les làmpades‐ i la plasticitat blanca derivada d’una sobreexposició lluminosa. En segon lloc, s’introdueix el rol de la ombra com a contrapunt fotogràfic de natura englotidora per, finalment, exposar la dualitat visual interactiva del blanc i negre així com la dimensió física de la matèria pel∙licular.

Evaluation of Muscardinus avellanarius population density by nest box and by trap checking

Population densities of marked common dormice Muscardinus avellanarius are generally based on nest box checks. As dormice also use natural cavities and leaf nests, we tried to answer the question "what proportion of the population cannot be monitored by nest boy checks", using parallel trapping sessions. We selected a forest of 1.7ha where a 5-year nest box survey revealed an annual mean of 3.4 ± 1.4 dormice per check. The trap design (permanent grid of 77 hanging platforms) was developed in June. During July and August the traps were set every second week (4 sessions of two nights = 8 nights) resulting in a total of 75 captures with mean of 9.4 dormice per night and the presence of 16 different individuals. The grid of 60 nest boxes was checked weekly (8 times) which allowed the recapture of 19 dormice with a mean of 2.4 dormice, per control day and the presence of 6 different individuals. Population density estimated by calendar of capture and the minimal number of dormice alive methods gave for nest-box checks a value of 2.4 animals/ha and the trap checks 6.6 animals/ha with the conclusion that 63% of the population were not being monitored by nest box checks.

Born to be bee, fed to be worker? The caste system of a primitively eusocial insect.

ABSTRACT: INTRODUCTION: Primitively eusocial halictid bees are excellent systems to study the origin of eusociality, because all individuals have retained the ancestral ability to breed independently. In the sweat bee Halictus scabiosae, foundresses overwinter, establish nests and rear a first brood by mass-provisioning each offspring with pollen and nectar. The mothers may thus manipulate the phenotype of their offspring by restricting their food provisions. The first brood females generally help their mother to rear a second brood of males and gynes that become foundresses. However, the first brood females may also reproduce in their maternal or in other nests, or possibly enter early diapause. Here, we examined if the behavioural specialization of the first and second brood females was associated with between-brood differences in body size, energetic reserves and pollen provisions. RESULTS: The patterns of variation in adult body size, weight, fat content and food provisioned to the first and second brood indicate that H. scabiosae has dimorphic females. The first-brood females were significantly smaller, lighter and had lower fat reserves than the second-brood females and foundresses. The first-brood females were also less variable in size and fat content, and developed on homogeneously smaller pollen provisions. Foundresses were larger than gynes of the previous year, suggesting that small females were less likely to survive the winter. CONCLUSIONS: The marked size dimorphism between females produced in the first and second brood and the consistently smaller pollen provisions provided to the first brood suggest that the first brood females are channelled into a helper role during their pre-imaginal development. As a large body size is needed for successful hibernation, the mother may promote helping in her first brood offspring by restricting their food provisions. This pattern supports the hypothesis that parental manipulation may contribute to promote worker behaviour in primitively eusocial halictids.

Diversity, prevalence and virulence of fungal entomopathogens in colonies of the ant Formica selysi

The richness of the parasitic community associated with social insect colonies has rarely been investigated. Moreover, understanding how hosts and pathogens interact in nature is important to interpret results from laboratory experiments. Here, we assessed the diversity, prevalence and virulence of fungal entomopathogens present around and within colonies of the ant Formica selysi. We detected eight fungal species known to be entomopathogenic in soil sampled from the habitat of ants. Six of these entomopathogens were found in active nests, abandoned nests, and corpses from dump piles or live ants. A systematic search for the presence of three generalist fungal entomopathogens in ant colonies revealed a large variation in their prevalence. The most common of the three pathogens, Paecilomyces lilacinus, was detected in 44% of the colonies. Beauveria bassiana occurred in 17% of the colonies, often in association with P. lilacinus, whereas we did not detect Metarhizium brunneum (formerly M. anisopliae) in active colonies. The three fungal species caused significant mortality to experimentally challenged ants, but varied in their degree of virulence. There was a high level of genetic diversity within B. bassiana isolates, which delineated three genetic strains that also differed significantly in their virulence. Overall, our study indicates that the ants encounter a diversity of fungal entomopathogens in their natural habitat. Moreover, some generalist pathogens vary greatly in their virulence and prevalence in ant colonies, which calls for further studies on the specificity of the interactions between the ant hosts and their fungal pathogens.

Altruism, Equity, and Reciprocity in a Gift-Exchange Experiment: An Encompassing Approach

Considerable experimental evidence suggests that non-pecuniary motives must be addressed when modeling behavior in economic contexts. Recent models of non-pecuniary motives can be classified as either altruism- based, equity-based, or reciprocity-based. We estimate and compare leading approaches in these categories, using experimental data. We then offer a flexible approach that nests the above three approaches, thereby allowing for nested hypothesis testing and for determining the relative strength of each of the competing theories. In addition, the encompassing approach provides a functional form for utility in different settings without the restrictive nature of the approaches nested within it. Using this flexible form for nested tests, we find that intentional reciprocity, distributive concerns, and altruistic considerations all play a significant role in players' decisions.

The effects of tourism, beachfront development and increased light pollution on nesting Loggerhead turtles Caretta caretta (Linnaeus, 1758) on Sal, Cape Verde Islands

Loggerhead Caretta caretta is now the only species of marine turtle nesting on the island of Sal, Cape Verde Islands. Since 2008, ADTMA - SOS Tartarugas has patrolled all the southern beaches of the island in order to protect nesting females and to collect nesting data. Although hunting is still a major issue, with 90 turtles killed in 2009, habitat loss and light pollution are becoming an ever more serious threat. Construction sites, hotels, apartment buildings and restaurants close to beaches, bright lights and illegal removal of sand are contributing to a marked decrease in the total number of nesting turtles on some beaches. In 2009, beaches on Sal experienced an average increase in nests of 200%, while the beach most affected by construction (Tortuga Beach) saw a decrease of nests of 7.3% (from 19.1% of total number of nests in 2008 to 11.8% in 2010). This beach also recorded a much lower nest to emergence ratio than normal (17.6% of emergences resulting in nests compared to 29.9% in other areas), indicating reluctance to nest due to light pollution and other disturbances.

Female-biased mortality in experimentally parasitized Alpine Swift Apus melba nestlings

1. Sex-biased mortality in adult vertebrates is often attributed to lower immunocompetence and higher parasite susceptibility of males. Although sex-specific mortality has also been reported during growth, the importance of sex-specific immunocompetence and parasite susceptibility in explaining male-biased mortality remains ambiguous in growing individuals because of potentially confounding sources of mortality such as sexual dimorphism. 2. Here, we investigated sex-specific susceptibility to the blood-sucking louse fly Crataerina melbae and sex differences in cell-mediated immunity in a bird species that is sexually monomorphic both in size and plumage coloration at the nestling stage, the Alpine Swift, Apus melba. 3. For this purpose, we manipulated ectoparasite loads by adding or removing flies to randomly chosen nests in two years, and injected nestlings with mitogenic phytohaemagglutinin (PHA) in another year. 4. There were no significant differences between male and female offspring in immune response towards PHA, parasite load, and parasite-induced decrease in growth rate. Secondary sex ratios were however biased toward males in parasitized broods, and this was explained by a greater mortality of females in parasitized than deparasitized broods. 5. Our findings are in contrast to the widely accepted hypothesis that males suffer a greater cost of parasitism. We discuss alternative hypotheses accounting for female-specific mortality.

Effect of an ectoparasite on lay date, nest-site choice, desertion, and hatching success in the great tit (Parus major)

Ectoparasites are common in most bird species, but experimental evidence of their effects on life-history traits is scarce. We investigated experimentally the effects of the hematophagous hen flea (Ceratophyllus gallinae) on timing of reproduction, nest-site choice, nest desertion, clutch size, and hatching success in the great tit (Parus major). When great tits were offered a choice on their territory between an infested and a parasite-free nest-box, they chose the one without parasites. When there was no choice, the great tits in a territory containing an infested nest-box delayed laying the clutch by 11 days as compared with the birds that were offered a parasite-free nesting opportunity. The finding that there was no difference in phenotypic traits related to dominance between the birds nesting in infested boxes and birds nesting in parasite-free boxes suggests that the delay is not imposed by social dominance. Nest desertion between laying and shortly after hatching was significandy higher in infested nests. There was no difference between infested and parasite-free nests in clutch size, but hatching success and hence brood size at hatching were significantly smaller in infested nests. Nest-box studies of great tits have been seminal in the development of evolutionary, ecological, and behavioral theory, but recently a polemic has arisen in the literature about the validity of the conclusions drawn from nest-box studies where the naturally occurring, detrimental ectoparasites are eliminated by the routine removal of old nests between breeding seasons. Our study suggests that this criticism is valid and that the evaluation of the effects of ectoparasites may improve our understanding of behavioral traits, life-history traits, or population dynamics

Comunidade de formigas que nidificam em pequenos galhos da serrapilheira em floresta da Amazônia Central, Brasil

Community of ants that nest in dead twigs on the ground of Central Amazonian forest, Brazil. A total area of 2,880 m² in four forest sites, near Manaus, Brazil, was searched for ant colonies nesting in dead twigs on the ground. An amount of 3,706 twigs (0.5-5 cm in diameter) were gathered, of which only 623 (16.8%) had ants, which is equivalent to a density of 0.22 nests per m². Seventy species have been found. The predominant genera were Pheidole (Westwood), Crematogaster (Lund), and Solenopsis (Westwood). For most species, many of the nests found had only workers and brood, suggesting that colonies either use multiple twigs to nest or do not live exclusively in the twigs, using other types of substrate (e.g., leaf-litter, soil, fruit pods) to nest. Most colonized twigs were hollow or partially hollow inside and relatively easy to break apart. There were significant differences among species with respect to the size (diameter) of twig used as nest. No correlation was found between the number of twigs available and the number colonized by ants, suggesting that ant populations were not limited by the amount of nesting sites (twigs). The three most common Pheidole species had small colonies with less than 200 workers. Colony size was not related to twig size (volume), for any of these three species.

As abelhas eussociais (Hymenoptera, Apidae) visitantes florais em um ecossistema de dunas continentais no médio Rio São Francisco, Bahia, Brasil

Highly eusocial bees (Hymenoptera, Apidae) flower visitors in a continental sand dune ecosystem from the medium São Francisco River, Bahia, Brazil. A community of highly eusocial bees in sand dunes, covered with caatinga vegetation, in the medium São Francisco River, Bahia (10º47' 37"S and 42º49' 25"W) was studied. The local climate is semi arid and hot, with mean temperature of 25.7 ºC and annual precipitation of 653.8 mm. Censuses took place every two months, from February to December of 2000. The bees were sampled on flowers with entomological nets, from 6:00 a.m. to 5:00 p.m. A total of 2,147 individuals of eight species of Apinae were found, of which Apis mellifera Linnaeus (40.2%), Trigona spinipes (Fabricius) (28.7%) and Frieseomelitta silvestri languida Moure (14.7%) were the predominant species. The diversity was H' = 1.53 and the evenness E' = 0.73. The bees were active during the whole year, but there was a significant variation in the monthly abundance of individuals (c2= 799.55; df= 35; p<0.0001). The daily activity was greater between 6:00 a.m. and 8:00 a.m. The low bee diversity observed is a consequence of the low richness of botanical species and of the small amount of sites for the bees' nests. The community of highly eusocial bees from the dunes presents organization patterns similar to those observed in other caatinga areas, albeit with some particularities.

Notas sobre a socialidade e a biologia de nidificação de Trypoxylon (Trypoxylon) asuncicola Strand, 1910 (Hymenoptera, Sphecidae)

Twenty-two nests of Trypoxylon asuncicola were sampled in Viçosa, State of Minas Gerais, Brazil, in January 2000 and the occupants' behavior of three nests was registered in 2h of direct observation. 528 brood cells were excavated (24±13.84 SD cells per nest), 129 were reused cells, some of them for seven times (meconium deposit count). The mean number of total cells, mean number of open and closed cells, parasitism rate and mean number of reused cells per nest were similar between old and new nests. Parasitism rate and cell reuse were associated with the number of building cells in the nest, but nest aggregate in the sampled area may play some role in the parasitism rate. Brachymeria sp. (Chalcididae) was the most important agent of brood mortality (80%). Other parasites were Melittobia sp. (Eulophidae) (17%) and a species of Icheumonidae (3%). The number of closed cells with immature individuals per nest was 4±4.2SD (N=17) and the mean reproductivity per female was 3±2.4SD (N=5). New nests produced more offspring (0 a 35%) than old nests (0 to 11%). Females and males can be found resting in the nest but copula or guarding behavior by the male was not observed. There is some evidence that in the sampled area the switch of nests by females is great and agonistic behavior between a nest owner and a visitor was not evident. Females were larger (3.9±0.4SD mm) than males (3.1±0.3SD mm) (measured as forewing length). The secondary sex ratio was 1.26 (±0.07 SE) in favor of females, which was not different from 1:1 ratio. The majority (97%) of the sampled larvae of T. asuncicola showed diapause. Some (5.1%) 'anomalous cells' were found.

Meliponini neotropicais: o gênero Partamona Schwarz, 1939 (Hymenoptera, Apidae, Apinae) - bionomia e biogeografia

This work, dedicated to the study of nesting habits of the species of the Neotropical genus Partamona Schwarz, is a sequence to the taxonomic revision recently published elsewhere. A total of 214 nests and nest aggregations of 18 species [Partamona epiphytophila Pedro & Camargo, 2003; P. testacea (Klug, 1807); P. mourei Camargo, 1980; P. vicina Camargo, 1980; P. auripennis Pedro & Camargo, 2003; P. combinata Pedro & Camargo, 2003; P. chapadicola Pedro & Camargo, 2003; P. nhambiquara Pedro & Camargo, 2003; P. ferreirai Pedro & Camargo, 2003; P. pearsoni (Schwarz, 1938); P. gregaria Pedro & Camargo, 2003; P. batesi Pedro & Camargo, 2003; P. ailyae Camargo, 1980; P. cupira (Smith, 1863); P. mulata Moure in Camargo, 1980; P. seridoensis Pedro & Camargo, 2003; P. criptica Pedro & Camargo, 2003; P. helleri (Friese, 1900)] were studied , including data about habitat, substrate, structural characteristics, construction materials and behavior. The descriptions of the nests are illustrated with 48 drawings. Partial data of the nests of P. bilineata (Say, 1837), P. xanthogastra Pedro & Camargo, 1997, P. orizabaensis (Strand, 1919), P. peckolti (Friese, 1901), P. aequatoriana Camargo, 1980, P. musarum (Cockerell, 1917) and P. rustica Pedro & Camargo, 2003 are also presented. Nests of P. grandipennis (Schwarz, 1951), P. yungarum Pedro & Camargo, 2003, P. subtilis Pedro & Camargo, 2003, P. vitae Pedro & Camargo, 2003, P. nigrior (Cockerell, 1925), P. sooretamae Pedro & Camargo, 2003 and P. littoralis Pedro & Camargo, 2003 are unknown. The species of Partamona build notable nest entrance structures, with special surfaces for incoming / exiting bees; some of them are extremely well-elaborated and ornamented, serving as flight orientation targets. All species endemic to western Ecuador to Mexico with known nesting habits (P. orizabaensis, P. peckolti, P. xanthogastra, P. bilineata, P. aequatoriana and P. musarum) build their nests in several substrates, non-associated with termitaria, such as cavities and crevices in walls, among roots of epiphytes and in bases of palm leaves, in abandoned bird nests, under bridges, and in other protected places, except P. peckolti that occasionally occupies termite nests. In South America, on the eastern side of the Andes, only P. epiphytophila and P. helleri nest among roots of epiphytes and other substrates, non-associated with termitaria. All other species studied (P. batesi, P. gregaria, P. pearsoni, P. ferreirai, P. chapadicola, P. nhambiquara, P. vicina, P. mourei, P. auripennis, P. combinata, P. cupira, P. mulata, P. ailyae, P. seridoensis, P. criptica and P. rustica) nest inside active termite nests, whether epigeous or arboreous. The only species that builds obligate subterranean nests, associated or not with termite or ant nests (Atta spp.) is P. testacea. Nests of Partamona have one vestibular chamber (autapomorphic for the genus) closely adjacent to the entrance, filled with a labyrinth of anastomosing pillars and connectives, made of earth and resins. One principal chamber exists for food and brood, but in some species one or more additional chambers are filled with food storage pots. In nests of P. vicina, there is one atrium or "false nest", between the vestibule and the brood chamber, which contains involucral sheaths, cells and empty pots. All structures of the nest are supported by permanent pillars made of earth and resins (another autapomorphy of the genus). The characters concerning nesting habits were coded and combined with morphological and biogeographic data, in order to hypothesize the evolutive scenario of the genus using cladistic methodology. The phylogenetic hypothesis presented is the following: (((((P. bilineata (P. grandipennis, P. xanthogastra)) (P. orizabaensis, P. peckolti)) (P. aequatoriana, P. musarum)) P. epiphytophila, P. yungarum, P. subtilis, P. vitae) (((((P. testacea (P. mourei, P. vicina)) (P. nigrior (P. auripennis, P. combinata))) (P. ferreirai (P. pearsoni (P. gregaria (P. batesi (P. chapadicola, P. nhambiquara)))))) ((((P. ailyae, P. sooretamae) P. cupira, P. mulata) P. seridoensis) P. criptica, P. rustica, P. littoralis)) P. helleri))). One area cladogram is presented. Dates of some vicariance / cladogenesis events are suggested. For bilineata / epiphytophila group, which inhabits the Southwestern Amazonia and the Chocó-Mexican biogeographical components, the origin of ancestral species is attributed to the Middle Miocene, when the transgressions of the Maracaibo and Paranense seas isolated the tropical northwestern South America from the eastern continental land mass. The next cladogenic event in the history of the bilineata / epiphytophila group is attributed to the Plio-Pleistocene, when the Ecuadorian Andes reached more than 3000 m, and the ancestral species was fragmented in two populations, one occupying the western Andes (ancestral species of the bilineata subgroup) and other the southwestern Amazon (ancestral species of the epiphytophila subgroup). Other aspects of the history of Partamona are also discussed.

Sex allocation and sex-dependent selection for body size in Trypoxylon rogenhoferi Kohl (Hymenoptera, Sphecidae)

Two populations of the wasp Trypoxylon rogenhoferi Kohl, 1884 from São Carlos and Luís Antônio, State of São Paulo, Brazil, were observed and sampled from May 1999 to February 2001 using trap-nests. This mass-provisioning wasp was used to test some aspects of optimal sex allocation theory. Both populations fit all the predictions of the models of Green and Brockmann and Grafen. Maternal provisions determined the size of each offspring, and females allocated well-stocked brood cells to daughters, the sex that benefits most being large. This strategy resulted in a difference in size between the sexes. In São Carlos, female weight at emergence was 1.18 times that of males, in Luís Antônio this value was 1.13. The brood cell volume was correlated with both wing length and weight at emergence in both sexes, and the chance that a given brood cell contained a male offspring decreased with increased brood cell volume. In T. rogenhoferi female body size was related to fitness. Larger females were able to collect more mass of spiders per day, the spiders they captured were heavier, and they provisioned more brood cells per day. They also produced larger daughters. For males, no relationship between body size and fitness was found, but the data were scarce. Since the patterns of provisioning were variable among different females in both study sites, it is possible that the females not follow a unique strategy for sex allocation. The sex ratio and/or investment ratio in the São Carlos population was female-biased and in Luís Antônio, male-biased. In spite of the influence of trap-nests diameters on male production in Luís Antônio, there is some evidence that in São Carlos population the local availability of prey and/or lower rate of parasitism may be major forces in determining the observed sex ratio, but further studies are necessary to verify such hypothesis.

Expression of foraging and Gp-9 are associated with social organization in the fire ant Solenopsis invicta.

The aim of this study was to investigate levels of expression of two major genes, the odorant binding protein Gp-9 (general protein-9) and foraging, that have been shown to be associated with behavioural polymorphisms in ants. We analysed workers and young nonreproductive queens collected from nests of the monogyne (single reproductive queen per nest) and polygyne (multiple reproductive queens) social forms of Solenopsis invicta. In workers but not young queens, the level of foraging expression was significantly associated with social form and the task performed (ie localization in the nest or foraging area). The level of expression of Gp-9 was also associated with social form and worker localization. In addition there was a higher level of expression of the Gp-9(b) allele compared with the Gp-9(B) allele in the heterozygote workers and the young nonreproductive queens. Finally, in the polygyne colonies the level of expression of foraging was not significantly associated with the Gp-9 genotype for either workers or young nonreproductive queens, suggesting that both genes have independent non-epistatic effects on behaviour in S. invicta.