Evolution, maintenance and ecological consequences of genetic caste determination in " Pogonomyrmex " harvester ants

Summary Division of labor between reproducers (queens) and helpers (workers) is the main characteristic of social insect societies and at the root of their ecological success. Kin selection models predict that phenotypic differences between queens and workers should result from environmental rather than from genetic differences. However, genetic effects on queen and worker differentiation were found in two populations-of Pogonomyrmex harvester ants. Each of the two populations is composed of two genetically distinct lineages. Queens (which can be of either lineage) generally mate with males of their own and of the alternate lineage and produce two types of female offspring, those fertilized by males of the queens' lineage which develop into queens and those fertilized by males of the alternate lineage which develop into workers. All four lineages were further suggested to be themselves of hybrid origin between-the species P: barbatus and P. rugosus, in which queens and workers do not differ genetically. In a first set of experiments, we tested if female caste determination (the differentiation into queens and workers) in the lineages was genetically hardwired and if it was associated with costs in terms of the ability to optimally allocate resources to the production of queens and workers. To this end we first mated queens of-two lineages to a single male. Queens mated to a male of the alternate lineage successfully raised worker offspring whereas queens mated to a male of their own lineage almost always failed to produce workers. This reveals that pure-lineage individuals have lost the ability to develop into workers. Second, we analyzed offspring produced by naturally mated queens. During the stage of colony founding when only workers are produced, naturally mated queens laid a high proportion of pure-lineage eggs but the large majority of these eggs failed to develop. As a consequence, the number of offspring produced by incipient colonies decreased linearly with the proportion of pure-lineage eggs laid by queens. Moreover, queens of the lineage most commonly represented in a given population produced more pure-lineage eggs, in line with the view that they mate randomly with the two types of males and indiscriminately use their sperm. Altogether these results predict frequency-dependent founding success for pairs of lineages because queens of the more common lineage will produce more pure-lineage eggs and their colonies be less successful during the stage of colony founding. To describe the distribution of populations characterized with genetic caste determination relative to the populations with environmental caste determination we genotyped queens and workers collected during a large survey of -additional populations. Genetic caste determination associated with pairs of interbreeding lineages was frequent and widespread in the studied range and we identified four additional lineages displaying genetic caste determination. Overall, there were thus eight highly differentiated lineages with genetic caste determination. These lineages always co-occurred in the same complementary lineage pairs. Three of the four lineage pairs appeared to have a common origin, while their relationship with the forth could not be resolved. The genetic survey also revealed that, in addition to being genetically isolated from one another, all eight lineages were genetically distinct from P. rugosus and P. barbatus, even when colonies of interbreeding lineages co-occurred with colonies of either putative parent at the same site. This raised the question of the mechanisms involved in the reproductive isolation between the lineages and the parental species and between the two lineages of a lineage pair. At a site where one lineage pair co-occurred with P. rugosus, we identified two pre-zygotic mechanisms (differences in timing for mating flights between P. rugosus and the lineage pair and assortative mating) and one post-zygotic mechanism (high levels of hybrid unviablility) which in combination may largely account for the reproductive isolation between the lineages and their parental species. The mechanisms accounting for the reproductive isolation between the two lineages of a lineage pair varied across lineage pairs. In one lineage pair, inter-lineage individuals exclusively occurred in the sterile worker caste, raising the possibility that inter-lineage eggs have completely lost the ability to develop into queens in this lineage pair and that there is thus no opportunity for gene flow. In each of the three remaining lineage pairs, inter-lineage queens were produced by a minority of colonies. In these lineage pairs, colonies headed by inter-lineage queens failed to grow sufficiently to produce reproductive individuals which may account for the reproductive isolation between co-occurring lineages in three lineage pairs. In conclusion, the results of this thesis show that genetic caste determination is costly but widespread in Pogonomyrmex harvester ants. Reproductive isolation among the lineages and between the lineages and the parental species as well as frequency-dependent founding success for co-occurring lineages may contribute to the persistence of this extraordinary system. Résumé La division du travail entre individus reproducteurs (les reines) et individus non-reproducteurs (ouvrières) représente la caractéristique principale des sociétés d'insectes et est à la base de leur succès écologique. Des modèles de sélection de parentèle prédisent que les différences phénotypiques entre reines et ouvrières devraient provenir d'effets environnementaux plutôt que de différences génétiques. Malgré ce fait, des effets génétiques sur la différentiation entre reines et ouvrières ont été montrés dans deux populations de fourmis moissonneuses du genre Pogonomyrmex. Chacune des deux populations est composée de deux lignées génétiquement distinctes. Les reines de chaque lignée s'accouplent en général avec des mâles de leur propre lignée ainsi qu'avec des mâles de l'autre lignée et produisent deux types d'oeufs, ceux qui sont fécondés par les mâles de leur propre lignée qui se développent en nouvelles reines et ceux qui sont fécondés par les mâles de l'autre lignée qui se développent en ouvrières. Il a été suggéré que les lignées sont elles-mêmes des hybrides entre les deux espèces P. barbatus et P. rugosus. Dans ces deux espèces, les reines et ouvrières ne sont pas génétiquement distinctes. Dans une première série d'expériences, nous avons testé si la détermination de la caste femelle (le développement en reine ou en ouvrière) est génétiquement rigide et si elle est associée à des coûts en terme de capacité à allouer de façon optimale les ressources pour la production de reines et d'ouvrières. Pour cela nous avons accouplé des reines de deux lignées avec un seul mâle. Les reines accouplées avec un mâle de l'autre lignée ont élevé de nouvelles ouvrières avec succès alors que les reines accouplées avec un mâle de leur propre lignée ont presque toujours échoué à produire des ouvrières. Ceci montre que les individus de lignée pure ont perdu la capacité de se développer en ouvrière. Deuxièmement, nous avons analysé la descendance de reines qui se sont accouplées naturellement. Durant le stade de fondation de la colonie, où seules des ouvrières sont élevées, les reines accouplées naturellement ont pondu une grande proportion d'oeufs de lignée pure mais la majorité de ces derniers ne se sont pas développés. En conséquence, le nombre de descendants produits par des colonies fondatrices diminuait linéairement avec la proportion des oeufs de lignée pure pondus par la reine en accord avec l'hypothèse que les reines s'accouplent au hasard avec les deux types de mâles et utilisent leur sperme aléatoirement. Dans l'ensemble; ces résultats prédisent un succès de fondation fréquence-dépendant pour les deux lignées, car les reines de la lignée la plus fréquente produiront .plus d'oeufs de lignée pure et leurs colonies auront moins de succès lors de la fondation de colonies par rapport aux colonies de la lignée la moins fréquente. Pour décrire la distribution des-populations caractérisées par une détermination génétique des castes par rapport aux populations caractérisées par une détermination environnementale des castes, nous avons génotypé des reines et des ouvrières qui ont été collectées lors d'une analyse de populations supplémentaires. La détermination génétique des castes associée à des croisements entre lignées est fréquente et largement répartie dans l'aire étudiée. Nous avons identifié quatre lignées supplémentaires, ayant une détermination génétique des castes, pour un total de huit lignées. Ces huit lignées forment quatre paires de lignées et on ne trouve jamais deux lignées de paires différentes, dans une population. Trois des quatre paires de lignées s'avèrent avoir une origine commune alors que leur relation avec la quatrième paire de lignées n'a pas pu être résolue. L'analyse génétique de populations supplémentaires a également révélé qu'en plus d'être génétiquement isolées les unes des autres, les huit lignées sont génétiquement distinctes de P. rugosus et P. barbatus même si les colonies d'une paire de lignées se trouvent en sympatrie avec l'une ou l'autre des espèces parentales. Ceci relève la question des mécanismes impliqués dans l'isolation reproductive entre les lignées et les espèces parentales ainsi qu'entre les deux lignées d'une paire. En étudiant un site où une paire de lignées se trouve en sympatrie avec P. rugosus, nous avons identifié deux mécanismes pré-zygotiques (des différences dans le timing du vol nuptial entre P. rugosus et les lignées et des accouplements assortis) ainsi qu'un mécanisme post-zygotique (un niveau élevé de non-viabilité des hybrides). En combinaison, ces mécanismes peuvent largement expliquer l'isolement reproductif entre les lignées et leurs espèces parentales. Les mécanismes contribuant à l'isolement reproductif entre les deux lignées d'une paire variaient entre paires de lignées. Dans une paire, les individus de génotype inter-lignée se trouvent uniquement dans la caste stérile des ouvrières, suggérant qu'il n'y a pas d'opportunité pour avoir du flux de gènes entre les deux lignées ce cette paire. Dans chacune des trois autres paires de lignées des nouvelles reines de génotype inter-lignée sont produites par une minorité de colonies. Par contre, les colonies avec une reine mère de génotype inter-lignée ne se développent pas suffisamment pour produire des individus reproducteurs. Ceci peut donc expliquer pourquoi il n'y a pas de flux de gènes entre les deux lignées de trois paires. En conclusion, les résultats de cette thèse montrent que la détermination génétique de la caste est coûteuse mais très répandue chez les fourmis. moissonneuses du genre Pogonomyrmex. L'isolement reproductif des lignées entre elles et avec les espèces parentales, ainsi qu'un succès de fondation fréquence-dépendant contribuent à la persistance de ce système extraordinaire.

Nest-box design for the study of diurnal raptors and owls is still an overlooked point in ecological, evolutionary and conservation studies: a review

The use of artificial nest-boxes has led to significant progress in bird conservation and in our understanding of the functional and evolutionary ecology of free-ranging birds that exploit cavities for roosting and reproduction. Nest-boxes and their improved accessibility have made it easier to perform comparative and experimental field investigations. However, concerns about the generality and applicability of scientific studies involving birds breeding in nest-boxes have been raised because the occupants of boxes may differ from conspecifics occupying other nest sites. Here we review the existing evidence demonstrating the importance of nest-box design to individual life-history traits in three falcon (Falconiformes) and seven owl (Strigiformes) species, as well as the extent to which publications on these birds describe the characteristics of exploited artificial nest-boxes in their 'methods' sections. More than 60% of recent publications did not provide any details on nest-box design (e.g. size, shape, material), despite several calls >15 years ago to increase the reporting of such information. We exemplify and discuss how variation in nest-box characteristics can affect or confound conclusions from nest-box studies and conclude that it is of overall importance to present details of nest-box characteristics in scientific publications.

Triatominae as a model of morphological plasticity under ecological pressure

The use of biochemical and genetic characters to explore species or population relationships has been applied to taxonomic questions since the 60s. In responding to the central question of the evolutionary history of Triatominae, i.e. their monophyletic or polyphyletic origin, two important questions arise (i) to what extent is the morphologically-based classification valid for assessing phylogenetic relationships? and (ii) what are the main mechanisms underlying speciation in Triatominae? Phenetic and genetic studies so far developed suggest that speciation in Triatominae may be a rapid process mainly driven by ecological factors.

Entomological and ecological aspects of six sylvatic species of triatomines (Hemiptera, Reduviidae) from the collection of the National Biodiversity Institute of Costa Rica, Central America

A total of 797 specimens of wild adult triatomines, belonging to six species from the entomological collections of the Costa Rican National Biodiversity Institute, was studied from the standpoint of their relative abundance, as reflected by light traps, distribution in the country, seasonal variations and climatic and altitudinal preferences. Triatoma dimidiata was the most abundant species (32.9% of the total specimens), with a very extensive distribution in different ecological zones, being more common between 100 to 400 m above sea level mainly at the end of the dry season. T. dispar was the third in frequency (21.5%), with narrower distribution, more abundant between 600 to 800 m and scarce during the dry season. Panstrongylus geniculatus and P. rufotuberculatus, second and fourth in frequency (22.1% and 15.1%, respectively), were widely distributed on both the Pacific and Caribbean basins, the former being more common between 80 to 270 m all year round and the latter below 800 m mainly during the first semester. Eratyrus cuspidatus which represented only 4.9% of the insects, was also present on both basins mainly below 200 m with a tendency to be scarce during certain months of the year, and was found in all types of ecological zones. Finally, Rhodnius pallescens, the least abundant species (3.6%) was restricted to very humid areas below 20 m, on the north side and Caribbean basin. With the exception of R. pallescens, males were more commonly found than females. Some epidemiological implications related to the six species are discussed.

Conserved non-genic sequences - an unexpected feature of mammalian genomes.

Mammalian genomes contain highly conserved sequences that are not functionally transcribed. These sequences are single copy and comprise approximately 1-2% of the human genome. Evolutionary analysis strongly supports their functional conservation, although their potentially diverse, functional attributes remain unknown. It is likely that genomic variation in conserved non-genic sequences is associated with phenotypic variability and human disorders. So how might their function and contribution to human disorders be examined?

Ecological Data of Travassosnema travassosi travassosi (Dracunculoidea: Guyanemidae) from the Humour of the Eyes of Acestrorhynchus lacustris from Tibagi River, Paraná, Brazil

Data obtained between 1990 and 1995 provide, for the first time, ecological information of the parasitic nematode Travassosnema travassosi travassosi Costa, Moreira & Oliveira, 1991 from Acestrorhynchus lacustris (Characiformes: Acestrorhynchidae) collected in the Tibagi River, Sertanópolis, Paraná, Brazil. These nematode occurred with low prevalences (7.7% to 28.6%) and intensities (1 to 3) during almost the whole year. The observation of mature females throughout the year indicate that liberation of larvae can occur during all the year around. This is the first report on the occurrence of T. travassosi travassosi inside the eyes. These nematodes were mostly located in humour of the eyes (87% of cases), being less frequently detected in tissues behind the eyes (13% of cases).

Ecological Aspects of Phebotomine (Diptera, Psychodidae) in an Endemic Area of Tegumentary Leishmaniasis in the Northeastern Argentina, 1993-1998

The first epidemic tegumentary leishmaniasis´ outbreak in the province of Misiones was recorded in 1998, in the locality of Puerto Esperanza. Phlebotominae collected in the region, previously or simultaneously to the outbreak (September 1993-December 1998) showed that the species Lutzomyia intermedia s. l. was prevalent (94%, n 6,150) at all the sites sampled with miniature light trap (10) and Shannon trap (3). L. pessoai, L. whitmani, L. migonei, L. shannoni, L. fischeri, L. misionensis, Brumptomyia avellari and B. guimaraesi were also captured. Sand fly distribution in time and space suggests that in the province of Misiones (1) the species already present before 1990 could give rise to the epidemic by the density/dispersion fluctuation of their local populations; (2) the abundance of L. intermedia s. l. was associated with environments with ecotones of primary-secondary vegetation, close to water bodies and with moderate human disturbance; (3) this species showed, towards the end of 1997, peaks of exceptional abundance, subsequent to rainfall peaks in 1996. This increase in abundance of potential vector sand fly populations close to houses with colonizable surroundings could have generated the 1998 epidemic outbreak.

Parasitic arthropods of some wild rodents from Juréia-Itatins Ecological Station, State of São Paulo, Brazil

A study of the associations between three species of rodents in the Atlantic forest and their parasitic arthropods was undertaken at the Juréia-Itatins Ecological Station, located in the State of São Paulo, Southeastern Brazil, from March 1989 to February 1990. Individuals of three species, Oryzomys russatus, Proechimys iheringi and Nectomys squamipes were captured and examined for ectoparasites. Eleven species of parasitic arthropods were found, including four species of insects and seven of Acari. Parasitism intensity, phenology, and rainfall were positively correlated with the abundance of the ectoparasites and their hosts. The most abundant host was O. russatus (Muridae: Sigmodontinae), and the most common parasite on it was the laelapid mite Gigantolaelaps oudemansi. The cuterebrid Metacuterebra apicalis caused myiasis in O. russatus. A mutualistic association between the staphylinid beetle Amblyopinus sp. and its host P. iheringi (Echimyidae) was observed. The few N. squamipes captured had small numbers of ectoparasites.

Triatoma rubrovaria (Blanchard, 1843) (Hemiptera, Reduviidae, Triatominae) II: trophic resources and ecological observations of five populations collected in the State of Rio Grande do Sul, Brazil

Triatoma rubrovaria has become the most frequently captured triatomine species after the control of T. infestans in the State of Rio Grande do Sul (RS), Brazil. Isoenzymatic and chromatic studies indicate the existence of, at least, two distinct phenotypic patterns of T. rubrovaria in RS. The geographic variation noted through molecular tools may also result in distinct profiles of vectorial potentiality. In order to enhance our understanding of the bionomic knowledge of T. rubrovaria separate batches of the species were collected from different municipalities of RS distant from 72 to 332 km: Santana do Livramento (natural ecotope), Santana do Livramento (artificial ecotope), Santiago (natural ecotope), Canguçu (peridomicile) and Encruzilhada do Sul (natural ecotope). A total of 285 specimens were collected, 85 specimens kept sufficient fecal material in their guts for the precipitin analysis. The results indicated the food eclecticism for this species and the anti-rodent serum showed the highest positivity in most localities. From the total of analyzed samples, only 1.3% of unique positivity for human blood was registered, all of them for Santiago population. This reactivity to human blood may be associated to pastures activities in the field.

Experimental evidence and ecological perspectives for the adaptation of Schistosoma mansoni Sambon, 1907 (Digenea: Schistosomatidae) to a wild host, the water-rat, Nectomys squamipes Brants, 1827 (Rodentia: Sigmodontinae)

Due to the semi aquatic habits and the overlap of the geographical distribution of the water-rat, Nectomys spp., with schistosomiasis endemic areas, these wild rodents are very likely to acquire Schistosoma mansoni infection in their daily activities. The role of the water-rat in the S. mansoni cycle would be substantiated if one could prove that these rodents acquire the parasite during their own activity time, a completely independent time schedule of human activities. To pursue this goal, we performed two field experiments in the municipality of Sumidouro, State of Rio de Janeiro, Brazil, a schistosomiasis endemic area where N. squamipes is found naturally infected. One experiment was devised as a series of observations of activity time of the water-rat. The other experiment was a test of the occurrence of late transmission of S. mansoni to the water-rat. The daily activity pattern showed that the water-rat is active chiefly just after sunset. At both diurnal and late exposition essays the water-rat sentinels got infected by S. mansoni. These findings clarify ecological and behavioral components necessary to the adaptation of S. mansoni to the water-rat as a non human definitive host and the existence of a transmission cycle involving this animals as a reservoir.

Ecological characteristics of Triatoma patagonica at the Southern limit of its distribution (Chubut, Argentina)

This study aimed at identifying the ecological characteristics of Triatoma patagonica Del Ponte 1929 (Hemiptera: Reduviidae) and evaluating the epidemiological importance of this species at its southernmost distribution limit. We carried out two surveys in the Province of Chubut, in summer and in spring, 1998. In each survey, we interviewed local health care agents for triatomine recognition and reports of the bugs, followed by entomological searches in houses and peridomestic and silvatic biotopes. The presence of T. patagonica was confirmed in two of the five localities indicated by interview. In agreement with previous studies, all these positive localities were east of the 11°C isotherm, within the Monte phytogeographic province. Triatomine abundance and infestation levels (in peridomiciles and peridomestic biotopes) were higher in summer than in spring, possibly reflecting adverse environmental conditions such as flood and cold winter weather. In the silvatic environment, we found three adult bugs under rocks in summer. In peridomestic sites bugs were only found associated with chickens, which were also the only blood meal source identified. Infection by Trypanosoma cruzi was not detected. We conclude that T. patagonica at its southernmost distribution limit does not represent a risk to public health due to its low abundance, lack of association with humans, and absence of T. cruzi infection.

Ecological analysis of Acari recovered from coprolites from archaeological site of Northeast Brazil

Coprolite samples of human and animal origin from the excavations performed at the archaeological site of Furna do Estrago, at Brejo da Madre de Deus in the state of Pernambuco, Brazil and sent to the Paleoparasitology Laboratory at Escola Nacional de Saúde Pública-Fiocruz, Rio de Janeiro, were analyzed for mites. After rehydratation and sedimentation of the coprolites, the alimentary contents and the sediments were examined and the mites collected and prepared in definitive whole mounts, using Hoyer's medium. Mites of the following suborders and orders were recovered: suborder Acaridia; order Gamasida; order Ixodida with the familiy Ixodidae (Ixodes sp. and Amblyomma sp. larvae, scutum, idiosoma, gnathosoma); order Oribatida (Aphelacarus sp., Apolohmannia sp., Eophypochthonius sp., Cosmochthonius sp., Pterobates sp., Poronoticae with pteromorphae not auriculate); order Astigmata with the families Atopomelidae (Chirodiscoides caviae), Anoetidae hypopus, Acaridae (Suidasia pontifica), Glycyphagidae (Blomia tropicalis), Pyroglyphidae (Hirstia passericola); order Actinedida with the family Tarsonemidae (Iponemus radiatae). The present work discusses the possibility of the preservation of the mite groups found up to the present day. We also discuss their relationship with the environment and their importance to present populations.