First checklist of the ants (Hymenoptera: Formicidae) of French Guiana.

We provide here a checklist of the ants of French Guiana, an overseas department of France situated in northern South America, with a very low human population density and predominantly covered by old-growth tropical rainforests. Based on 165 scientific papers, specimens deposited in collections, and unpublished surveys, a total of 659 valid species and subspecies from 84 genera and 12 subfamilies is presented. Although far from complete, these numbers represent approximately 10% of the ant diversity known to occur in the Neotropical realm. Additionally, three ant genera and 119 species are reported for the first time for French Guiana. Finally, five species are recognized as erroneous records for the the department in the literature. This checklist significantly expands the basic knowledge of the ants in the Guiana Shield, one of the world's most important biodiversity hotspots.

The Guild Concept: From Feudalism to Community Ecology / El concepto de gremio: del feudalismo a la ecología de comunidades

RESUMEN Se analiza el origen y evolución del término gremio, así como su aplicación en ecología, considerando las múltiples connotaciones que se le ha dado y la confusión que se ha generado por utilizarlo de forma indebida. De igual forma, se discute la importancia de homogenizar los términos y definir de manera clara a los gremios, a fin de tener un leguaje que permita entender los alcances del término sin ambigüedades. El uso del término, así como su persistencia en estudios ecológicos, sugiere que el mismo tiene relevancia considerable dependiendo de la forma y el modo en que es empleado. El uso inadecuado o derivado de este término es arriesgado y peligroso, dado que tiende a reducir el término a una palabra vacía con múltiples significados. Más que nada, esta trivialización constituye una amenaza al uso y significado adecuado del concepto de gremio en ecología.

ABSTRACT The origin and evolution of the term guild are analyzed, as well as its application in ecology, considering the multiple connotations that have been given and the confusion that has been generated by using it improperly. Likewise, the importance of homogenizing the terms and clearly defining the guilds is discussed, to have a language that allows understanding the scope of the term without ambiguities. The use of the term, as well as its persistence in ecological studies, suggests that it has considerable relevance depending on the form and the way it is used. The inappropriate or derivative use of this term is risky and dangerous since it tends to reduce the term to an empty word with multiple meanings. Also, this trivialization constitutes a threat to the proper use of the guild concept in ecology and its meaning as a whole.

Genetic Characterization of Some Neoponera (Hymenoptera: Formicidae) Populations Within the foetida Species Complex.

The foetida species complex comprises 13 Neotropical species in the ant genus Neoponera. Neoponera villosa Fabricius (1804) , Neoponera inversa Smith (1858), Neoponera bactronica Fernandes, Oliveira & Delabie (2013), and Neoponera curvinodis (Forel, 1899) have had an ambiguous taxonomic status for more than two decades. In southern Bahia, Brazil, these four species are frequently found in sympatry. Here we used Bayesian Inference and maximum likelihood analyses of COI and 16S mtDNA sequence data and conventional cytogenetic data together with observations on morphology to characterize sympatric populations of N. villosa, N. inversa, N. bactronica, and N. curvinodis. Our results showed marked differences in the karyotype of these ants. Both N. curvinodis and N. inversa have chromosome number of 2n = 30. Their chromosome composition, however, is distinct, which indicates that N. curvinodis is more closely related to N. bactronica. These four species clustered into three distinct groups. The close relationship between N. bactronica and N. curvinodis deserves further investigation since it has not been fully resolved here. Our results confirm that N. inversa, N. villosa, N. bactronica + N. curvinodis indeed represent four distinct taxa within the foetida species complex.

Highly modular pattern in ant-plant interactions involving specialized and non-specialized myrmecophytes.

Because Tachia guianensis (Gentianaceae) is a "non-specialized myrmecophyte" associated with 37 ant species, we aimed to determine if its presence alters the ant guild associated with sympatric "specialized myrmecophytes" (i.e., plants sheltering a few ant species in hollow structures). The study was conducted in a hilly zone of a neotropical rainforest where two specialized myrmecophytes grow at the bottom of the slopes, another at mid-slope, and a fourth on the hilltops. Tachia guianensis, which occurred everywhere, had its own guild of associated ant species. A network analysis showed that its connections with the four other myrmecophytes were rare and weak, the whole resulting in a highly modular pattern of interactions with one module (i.e., subnetwork) per myrmecophyte. Three ant species parasitized three out of the four specialized myrmecophytes (low nestedness noted), but were not or barely associated with T. guianensis that therefore did not influence the parasitism of specialized myrmecophytes.

Methuselah's daughters: Paternal age has little effect on offspring number and quality in Cardiocondyla ants.

Male age may directly or indirectly affect the fitness of their female mating partners and their joint progeny. While in some taxa of insects, old males make better mates and fathers, young males excel in others. Males of most social Hymenoptera are relatively short lived and because of testis degeneration have only a limited sperm supply. In contrast, the wingless fighter males of the ant Cardiocondyla obscurior live for several weeks and produce sperm throughout their lives. Wingless males engage in lethal combat with rival males and the winner of such fights can monopolize mating with all female sexuals that emerge in their nests over a prolonged timespan. Here, we investigate if male age has an influence on sperm quality, the queen's lifespan and productivity, and the size and weight of their offspring. Queens mated to one-week or six-week-old males did not differ in life expectancy and offspring production, but the daughters of young males were slightly heavier than those of old males. Our data suggest negligible reproductive senescence of C. obscurior males even at an age, which only few of them reach. This matches the reproductive strategy of Cardiocondyla ants, in which freshly emerging female sexuals rarely have the option to mate with males other than the one present in their natal nest.

Hollow Internodes Permit a Neotropical Understory Plant to Shelter Multiple Mutualistic Ant Species, Obtaining Protection and Nutrient Provisioning (Myrmecotrophy).

The Neotropical understory plant Tachia guianensis (Gentianaceae)-known to shelter the colonies of several ant species in its hollow trunks and branches-does not provide them with food rewards (e.g., extrafloral nectar). We tested whether these ants are opportunistic nesters or whether mutualistic relationships exist as for myrmecophytes or plants sheltering ant colonies in specialized hollow structures in exchange for protection from enemies and/or nutrient provisioning (myrmecotrophy). We noted 37 ant species sheltering inside T. guianensis internodes, three of them accounting for 43.5% of the cases. They protect their host plants from leaf-cutting ant defoliation and termite damage because individuals devoid of associated ants suffered significantly more attacks. Using the stable isotope 15N, we experimentally showed that the tested ant species furnish their host plants with nutrients. Therefore, a mutualism exists. However, because it is associated with numerous ant species, T. guianensis can be considered a nonspecialized myrmecophyte.

Rediscovery and reclassification of the dipteran taxon Nothomicrodon Wheeler, an exclusive endoparasitoid of gyne ant larvae.

The myrmecophile larva of the dipteran taxon Nothomicrodon Wheeler is rediscovered, almost a century after its original description and unique report. The systematic position of this dipteran has remained enigmatic due to the absence of reared imagos to confirm indentity. We also failed to rear imagos, but we scrutinized entire nests of the Brazilian arboreal dolichoderine ant Azteca chartifex which, combined with morphological and molecular studies, enabled us to establish beyond doubt that Nothomicrodon belongs to the Phoridae (Insecta: Diptera), not the Syrphidae where it was first placed, and that the species we studied is an endoparasitoid of the larvae of A. chartifex, exclusively attacking sexual female (gyne) larvae. Northomicrodon parasitism can exert high fitness costs to a host colony. Our discovery adds one more case to the growing number of phorid taxa known to parasitize ant larvae and suggests that many others remain to be discovered. Our findings and literature review confirm that the Phoridae is the only taxon known that parasitizes both adults and the immature stages of different castes of ants, thus threatening ants on all fronts.

Arthropod Distribution in a Tropical Rainforest: Tackling a Four Dimensional Puzzle.

Quantifying the spatio-temporal distribution of arthropods in tropical rainforests represents a first step towards scrutinizing the global distribution of biodiversity on Earth. To date most studies have focused on narrow taxonomic groups or lack a design that allows partitioning of the components of diversity. Here, we consider an exceptionally large dataset (113,952 individuals representing 5,858 species), obtained from the San Lorenzo forest in Panama, where the phylogenetic breadth of arthropod taxa was surveyed using 14 protocols targeting the soil, litter, understory, lower and upper canopy habitats, replicated across seasons in 2003 and 2004. This dataset is used to explore the relative influence of horizontal, vertical and seasonal drivers of arthropod distribution in this forest. We considered arthropod abundance, observed and estimated species richness, additive decomposition of species richness, multiplicative partitioning of species diversity, variation in species composition, species turnover and guild structure as components of diversity. At the scale of our study (2 km of distance, 40 m in height and 400 days), the effects related to the vertical and seasonal dimensions were most important. Most adult arthropods were collected from the soil/litter or the upper canopy and species richness was highest in the canopy. We compared the distribution of arthropods and trees within our study system. Effects related to the seasonal dimension were stronger for arthropods than for trees. We conclude that: (1) models of beta diversity developed for tropical trees are unlikely to be applicable to tropical arthropods; (2) it is imperative that estimates of global biodiversity derived from mass collecting of arthropods in tropical rainforests embrace the strong vertical and seasonal partitioning observed here; and (3) given the high species turnover observed between seasons, global climate change may have severe consequences for rainforest arthropods.

A new species of Simopelta (Hymenoptera: Formicidae: Ponerinae) from Brazil and Costa Rica.

The genus Simopelta consists of 21 described species restricted to Central America and South America. The present study describes a new cryptobiotic species, Simopelta anomma sp. nov.. The new species is blind, possesses a 3-segmented antennal club, and has the midtibia with several stout setae, a combination of characters unique within the genus. Moreover, some traits of this species require broadening the definition of the genus. The discovery of S. anomma sp. nov. suggests that many undiscovered species, some of which may be important for understanding ant evolution, remain hidden below ground in Neotropical rainforests.

Bat aggregation mediates the functional structure of ant assemblages.

In the Guianese rainforest, we examined the impact of the presence of guano in and around a bat roosting site (a cave). We used ant communities as an indicator to evaluate this impact because they occupy a central place in the functioning of tropical rainforest ecosystems and they play different roles in the food web as they can be herbivores, generalists, scavengers or predators. The ant species richness around the cave did not differ from a control sample situated 500m away. Yet, the comparison of functional groups resulted in significantly greater numbers of detritivorous fungus-growing and predatory ant colonies around the cave compared to the control, the contrary being true for nectar and honeydew feeders. The role of bats, through their guano, was shown using stable isotope analyses as we noted significantly greater δ(15)N values for the ant species captured in and around the cave compared to controls.

Traits allowing some ant species to nest syntopically with the fire ant Solenopsis saevissima in its native range.

Supercolonies of the red fire ant Solenopsis saevissima (Smith) develop in disturbed environments and likely alter the ant community in the native range of the species. For example, in French Guiana only 8 ant species were repeatedly noted as nesting in close vicinity to its mounds. Here, we verified if a shared set of biological, ecological, and behavioral traits might explain how these 8 species are able to nest in the presence of S. saevissima. We did not find this to be the case. We did find, however, that all of them are able to live in disturbed habitats. It is likely that over the course of evolution each of these species acquired the capacity to live syntopically with S. saevissima through its own set of traits, where colony size (4 species develop large colonies), cuticular compounds which do not trigger aggressiveness (6 species) and submissive behaviors (4 species) complement each other.

Arthropod diversity in a tropical forest.

Most eukaryotic organisms are arthropods. Yet, their diversity in rich terrestrial ecosystems is still unknown. Here we produce tangible estimates of the total species richness of arthropods in a tropical rainforest. Using a comprehensive range of structured protocols, we sampled the phylogenetic breadth of arthropod taxa from the soil to the forest canopy in the San Lorenzo forest, Panama. We collected 6144 arthropod species from 0.48 hectare and extrapolated total species richness to larger areas on the basis of competing models. The whole 6000-hectare forest reserve most likely sustains 25,000 arthropod species. Notably, just 1 hectare of rainforest yields >60% of the arthropod biodiversity held in the wider landscape. Models based on plant diversity fitted the accumulated species richness of both herbivore and nonherbivore taxa exceptionally well. This lends credence to global estimates of arthropod biodiversity developed from plant models.

Cuticular lipid profiles of fertile and non-fertile Cardiocondyla ant queens.

Both mating and reproduction strongly affect the physiology of insect females. In the ant Cardiocondyla obscurior, a comparison among virgin queens, mated queens, and queens mated with sterilized males ("sham-mated") allows to separate the different effects of mating and egg laying. Here, we investigate whether and how different mating status is reflected in the cuticular lipid profiles of queens, i.e., the blend of chemicals that is thought to signal a queen's fertility. Surprisingly, discriminant analyses failed to reliably distinguish among virgin, mated, and sham-mated queens. A generalized linear model on individual substances showed only very subtle differences. While mating appeared to be positively associated with the proportions of 3-MeC(25,) 11-/13-MeC(27), 5-MeC(27), 3-MeC(27), and 12-/14-MeC(28) and negatively with C(27:1), fecundity was negatively associated with C(29:1), C(31:1), and a sterol derivative. We discuss these results in the light of the special life history of C. obscurior, with completely sterile workers and low egg laying rates in queens.

The ecology and feeding habits of the arboreal trap-jawed ant Daceton armigerum.

Here we show that Daceton armigerum, an arboreal myrmicine ant whose workers are equipped with hypertrophied trap-jaw mandibles, is characterized by a set of unexpected biological traits including colony size, aggressiveness, trophobiosis and hunting behavior. The size of one colony has been evaluated at ca. 952,000 individuals. Intra- and interspecific aggressiveness were tested and an equiprobable null model used to show how D. armigerum colonies react vis-à-vis other arboreal ant species with large colonies; it happens that D. armigerum can share trees with certain of these species. As they hunt by sight, workers occupy their hunting areas only during the daytime, but stay on chemical trails between nests at night so that the center of their home range is occupied 24 hours a day. Workers tend different Hemiptera taxa (i.e., Coccidae, Pseudococcidae, Membracidae and Aethalionidae). Through group-hunting, short-range recruitment and spread-eagling prey, workers can capture a wide range of prey (up to 94.12 times the mean weight of foraging workers).

Inherited biotic protection in a neotropical pioneer plant.

Chelonanthus alatus is a bat-pollinated, pioneer Gentianaceae that clusters in patches where still-standing, dried-out stems are interspersed among live individuals. Flowers bear circum-floral nectaries (CFNs) that are attractive to ants, and seed dispersal is both barochorous and anemochorous. Although, in this study, live individuals never sheltered ant colonies, dried-out hollow stems--that can remain standing for 2 years--did. Workers from species nesting in dried-out stems as well as from ground-nesting species exploited the CFNs of live C. alatus individuals in the same patches during the daytime, but were absent at night (when bat pollination occurs) on 60.5% of the plants. By visiting the CFNs, the ants indirectly protect the flowers--but not the plant foliage--from herbivorous insects. We show that this protection is provided mostly by species nesting in dried-out stems, predominantly Pseudomyrmex gracilis. That dried-out stems remain standing for years and are regularly replaced results in an opportunistic, but stable association where colonies are sheltered by one generation of dead C. alatus while the live individuals nearby, belonging to the next generation, provide them with nectar; in turn, the ants protect their flowers from herbivores. We suggest that the investment in wood by C. alatus individuals permitting still-standing, dried-out stems to shelter ant colonies constitutes an extended phenotype because foraging workers protect the flowers of live individuals in the same patch. Also, through this process these dried-out stems indirectly favor the reproduction (and so the fitness) of the next generation including both their own offspring and that of their siblings, all adding up to a potential case of inclusive fitness in plants.

Avaliação de duas técnicas de translocação de serrapilheirasobre as assembleias de aranhas (Arachnida: Araneae) e formigas (Hymenoptera: Formicidae)

Com o ritmo atual de destruição dos ecossistemas surge a necessidade de se investir em técnicas que visem à conservação dos remanescentes florestais e das suas espécies.Nosso objetivo foi comparar duas técnicas para translocação da serrapilheira (com e sem a retirada da camada de serrapilheira preexistente), observando como varia a riqueza e composição de aranhas e formigas no local da translocação, a fim de definir a melhor forma de realizá-la. O estudo foi realizado em dois remanescentes de Mata Atlântica (fragmento doador e receptor) localizados em Salvador (BA). Antes da translocação foi realizada uma comparação de ambas as faunas, entre os fragmentos, e foi encontradadiferença significativa tanto na riqueza quanto na composição. Assim pôde-se realizar o experimento. Para tanto, foi realizada a caracterização ambiental a fim de translocar a serrapilheira para as unidades mais similares na tentativa de minimizar o estresse dos organismos. A riqueza em espécies tanto de aranhas quanto de formigas aumentou nasunidades que receberam a serrapilheira, sendo para aranhas mais elevada nas unidades onde não foi retirada a serrapilheira preexistente. Já para as formigas as duas formas parecem eficientes. Contudo, levando em consideração os dois grupos, propõe-se que atranslocação deve ser feita mantendo a serrapilheira preexistente.

Due to the current rhythm of ecosystem destruction, the development of new techniques aiming at the conservation of forest remnants and their biota is urgent. Our objective was to compare two techniques of leaf-litter translocation (with and without withdrawing the pre-extent litter layer), through observations on spider and ant richness and assemblage composition, in order to define the best way to perform translocation. The study was carried out in two Brazilian Atlantic rain forest remnants (called releaser and receptor fragments) in Salvador, State of Bahia, Brazil. A comparison between the faunas of both fragments was performed before translocation and a significant difference was found both in their richness and composition, which is an essential condition to carry out the experiment. A small scale environmental characterization] was made in order to choose similar units capable of minimizing the stress on leaf-litter organisms living there. Both for spidersand ants, the species richness increased in units that received leaf-litter and it was higher for spiders in units where the pre-extent litter had not been removed. For the ants, the two forms of translocation seem to be efficient. Nevertheless, taking into account the two groups of organisms, it is suggested that the translocation must be done maintaining the pre-extent leaf-litter.

[Pathogenic bacteria dissemination by ants (Hymenoptera: Formicidae) in two hospitals in northeast Brazil]. / Disseminação de bactérias patogênicas por formigas (Hymenoptera: Formicidae) em dois hospitais do nordeste do Brasil.

Nosocomial infections bring a high risk to the health of hospital patients and employees. Ants are common organisms in Brazilian hospitals, where they can act as dispersers of opportunistic microorganisms in places they forage. The occurrence of multi-resistant bacteria carried by ants was analyzed in two public hospitals (HA and HB) in southeastern Bahia, Brazil. In these two hospitals 132 workers belonging to three ant species were collected. The bacteria associated to these ants were identified and their susceptibility to antibiotics was evaluated. More than half (57.3%) of ants collected in HA were associated with some kind of bacteria, with 26.7% of them being opportunist bacteria, while 84,2% of the ants from HB presented associated bacteria growth, with 61.4% of them being opportunist bacteria. Twenty four species of bacteria were isolated. The Gram-positive bacilli of the genus Bacillus were the most frequent, followed by the Gram-positive cocci, Gram-negative bacilli (family Enterobacteriaceae) and Gram-negative non-fermenters bacilli. The profile of sensitivity of the bacterial isolates to drugs pointed out the existence of multi-resistant isolates carried by ants. For the first time, are reported cases of the same bacterial resistant isolates taken form homospecific ant workers that point out the importance of ants to bacteria dissemination and proliferation in a hospital. Our results suggest that the risk of contamination presented by these ants is similar to the one of any other mechanical vector of bacterial dissemination.

A temporary social parasite of tropical plant-ants improves the fitness of a myrmecophyte.

Myrmecophytes offer plant-ants a nesting place in exchange for protection from their enemies, particularly defoliators. These obligate ant-plant mutualisms are common model systems for studying factors that allow horizontally transmitted mutualisms to persist since parasites of ant-myrmecophyte mutualisms exploit the rewards provided by host plants whilst providing no protection in return. In pioneer formations in French Guiana, Azteca alfari and Azteca ovaticeps are known to be mutualists of myrmecophytic Cecropia (Cecropia ants). Here, we show that Azteca andreae, whose colonies build carton nests on myrmecophytic Cecropia, is not a parasite of Azteca-Cecropia mutualisms nor is it a temporary social parasite of A. alfari; it is, however, a temporary social parasite of A. ovaticeps. Contrarily to the two mutualistic Azteca species that are only occasional predators feeding mostly on hemipteran honeydew and food bodies provided by the host trees, A. andreae workers, which also attend hemipterans, do not exploit the food bodies. Rather, they employ an effective hunting technique where the leaf margins are fringed with ambushing workers, waiting for insects to alight. As a result, the host trees' fitness is not affected as A. andreae colonies protect their foliage better than do mutualistic Azteca species resulting in greater fruit production. Yet, contrarily to mutualistic Azteca, when host tree development does not keep pace with colony growth, A. andreae workers forage on surrounding plants; the colonies can even move to a non-Cecropia tree.

Disseminação de bactérias patogênicas por formigas (Hymenoptera: Formicidae) em dois hospitais do nordeste do Brasil / Pathogenic bacteria dissemination by ants (Hymenoptera: Formicidae) in two hospitals in northeast Brazil

Nosocomial infections bring a high risk to the health of hospital patients and employees. Ants are common organisms in Brazilian hospitals, where they can act as dispersers of opportunistic microorganisms in places they forage. The occurrence of multi-resistant bacteria carried by ants was analyzed in two public hospitals (HA and HB) in southeastern Bahia, Brazil. In these two hospitals 132 workers belonging to three ant species were collected. The bacteria associated to these ants were identified and their susceptibility to antibiotics was evaluated. More than half (57.3 percent) of ants collected in HA were associated with some kind of bacteria, with 26.7 percent of them being opportunist bacteria, while 84,2 percent of the ants from HB presented associated bacteria growth, with 61.4 percent of them being opportunist bacteria. Twenty four species of bacteria were isolated. The Gram-positive bacilli of the genus Bacillus were the most frequent, followed by the Gram-positive cocci, Gram-negative bacilli (family Enterobacteriaceae) and Gram-negative non-fermenters bacilli. The profile of sensitivity of the bacterial isolates to drugs pointed out the existence of multi-resistant isolates carried by ants. For the first time, are reported cases of the same bacterial resistant isolates taken form homospecific ant workers that point out the importance of ants to bacteria dissemination and proliferation in a hospital. Our results suggest that the risk of contamination presented by these ants is similar to the one of any other mechanical vector of bacterial dissemination.

Worldwide invasion by the little fire ant: routes of introduction and eco-evolutionary pathways.

Biological invasions are generally thought to occur after human aided migration to a new range. However, human activities prior to migration may also play a role. We studied here the evolutionary genetics of introduced populations of the invasive ant Wasmannia auropunctata at a worldwide scale. Using microsatellite markers, we reconstructed the main routes of introduction of the species. We found three main routes of introduction, each of them strongly associated to human history and trading routes. We also demonstrate the overwhelming occurrence of male and female clonality in introduced populations of W. auropunctata, and suggest that this particular reproduction system is under selection in human-modified habitats. Together with previous researches focused on native populations, our results suggest that invasive clonal populations may have evolved within human modified habitats in the native range, and spread further from there. The evolutionarily most parsimonious scenario for the emergence of invasive populations of the little fire ant might thus be a two-step process. The W. auropunctata case illustrates the central role of humans in biological change, not only due to changes in migration patterns, but also in selective pressures over species.