Morphological patterns and community organization in leaf-litter ant assemblages

We present a new approach to determine the number and composition of guilds, using the hyperdiverse leaf-litter ant fauna as a model, based on appropriate morphological variables and species co-occurrence null models to describe the complex assemblages of interacting Species Community structure at the 1-m(2) scale. We obtained 18 linear morphometric measures from 949 workers of 171 leaf-litter ant species (18762 measurements) surveyed in four Atlantic Forest localities to test whether the assemblages are morphologically structured; the morphological characters were selected to indicate diet and foraging habits. Principal components analysis was used to characterize the morphospace and to describe the guild structure (number of species and composition). The guild proportionality assembly rule (significant tendency toward constant proportion of species in guilds) was assessed at the 1-m(2) scale. Our analysis indicates that the division of leaf-litter ants into guilds is based mainly on microhabitat distribution in the leaf-litter, body size and shape, eye size, and phylogeny. The same guild scheme applied to four more sites shows that different Atlantic Forest areas have the same leaf-fitter ant guilds. The guild proportionality assembly rule was confirmed for most guilds, Suggesting that there are guild-specific limitations on species coexistence within assemblages; on the other hand, in a few cases the variance in guild proportion was greater than expected under the null assumptions. Other studies on ant functional group classification are partially supported by our quantitative morphological analysis. Our results, however, imply that there are more compartments than indicated in previous models, particularly among cryptic species (confined to soil and litter) and tropical climate specialists. We argue that a general null model for the analysis of species association based oil morphology can reveal objectively defined groups and may thus contribute to a robust theory to explain community structure in general and have important consequences on studies of litter ant community ecology in particular.

Revision of the fungus-growing ant genera Mycetophylax Emery and Paramycetophylax Kusnezov rev. stat., and description of Kalathomyrmex n. gen. (Formicidae: Myrmicinae: Attini)

Based on the morphology of workers, gynes and males, we revise the taxonomy of nominal taxa traditionally included by authors in the fungus-growing ant genus Mycetophylax. Our results indicate that Mycetophylax Emery (Myrmicocrypta brittoni Wheeler, 1907, type species, by designation of Emery, 1913; junior synonym of Cyphomyrmex conformis Mayr, 1884 by Kempf, 1962) includes M. conformis, M. simplex (Emery, 1888), and M. morschi (Emery, 1888) new combination (formerly in Cyphomyrmex), with several synonymies. Mycetophylax bruchi (Santschi, 1916) does not belong to the same genus and is diagnosed, in addition to other characters, by a psammophore arising at the anterior margin of the clypeus. For this species we are resurrecting from synonymy Paramycetophylax Kusnezov, 1956 (Mycetophylax bruchi as type species, by original designation, with M. cristulatus as its new synonym). Myrmicocrypta emeryi Forel, 1907 is the only attine in which females lack the median clypeal seta and have the antennal insertion areas very much enlarged and anteriorly produced, with the psammophore setae arising from the middle of the clypeus and not at its anterior margin as in Paramycetophylax. Notwithstanding its inclusion in Mycetophylax by recent authors, it is here recognized as belonging to a hitherto undescribed, thus far monotypic genus, Kalathomyrmex new genus (Myrmicocrypta emeryi as its type species, here designated). We redescribe workers, gynes and males of all species in the three genera and describe for the first time gynes of Mycetophylax conformis and M. simplex, males of M. simplex and M. morschi, and gynes of P. bruchi. Furthermore we present a key to the workers of the taxa treated here (most formerly included under the name Mycetophylax), a key to workers of the Mycetophylax in the revised sense, SEM pictures and high resolution AutoMontage(C) photographs of the species, along with maps of collection records and a summary of biological observations.

Tropidomyrmex elianae, a new myrmicine ant genus and species from Brazil, tentatively assigned to Solenopsidini (Hymenoptera, Formicidae)

A new myrmicine ant, Tropidomyrmex elianae gen. n. & sp. n., is described from southeastern and central Brazil, based on workers, ergatoid gynes, males and larvae. Tropidomyrmex workers are relatively small, monomorphic, characterized mainly by the feebly pigmented and extremely thin integument; subfalcate mandibles bearing a single apical tooth; palpal formula 1,2; clypeus relatively broad and convex; reduced compound eyes; propodeum unarmed and with a strongly medially depressed declivous face; double and bilobed well developed subpostpetiolar processes; and peculiarities in the sting apparatus. A colony fragment of T. elianae containing workers, ergatoid gynes, males, and brood was found inside a ground termite nest (Anoplotermes pacificus Apicotermitinae) in a montane rocky scrubland in the state of Minas Gerais, southeastern Brazil. Tropidomyrmex elianae is known also from two workers collected in leaf litter samples processed with a Winkler extractor, from the state of Tocantins, central-north Brazil. Despite the differences from the accepted solenopsidine genera, Tropidomyrmex is tentatively assigned to this tribe. Within the solenopsidine ants, the genus is apparently related to Tranopelta. Tropidomyrmex is marked by extreme reductions, perhaps reflecting adaptations to particular habits and habitats.

A taxonomic revision of the Neotropical myrmicine ant genus Lachnomyrmex wheeler (Hymenoptera : Formicidae)

The Neotropical ant genus Lachnomyrmex Wheeler, 1910 is revised for the first time. A revised generic description is provided for workers and gynes; males remain unknown. Morphological patterns combined with geographical data led to the recognition of 16 species, of which 10 are here described as new: L. amazonicus sp. n., L. fernandezi sp. n., L. grandis Fernandez & Baena, L. haskinsi Smith, L. lattkei sp. n., L. longinodus Fernandez & Baena, L. longinoi sp. n., L. mackayi sp. n., L. nordestinus sp. n., L. pilosus Weber, L. laticeps sp. n., L. platynodus sp. n., L. plaumanni Borgmeier, L. regularis sp. n., L. scrobiculatus Wheeler, and L. victori sp. n. The gynes of L. grandis, L. haskinsi, and L. pilosus are described for the first time. Illustrated identification keys for workers, distribution maps, and high resolution illustrations are supplied for all species. Some Lachnomyrmex species are relatively common in the leaf-litter of submontane wet forests in Central and South America; others remain known by very few individuals. The relatively small and apparently monogynic Lachnomyrmex colonies inhabit small nests in the ground, among rotten leaves and inside fallen logs. Workers generally forage alone on the ground or within the leaf litter, but have been recorded also on tree trunks, and apparently do not recruit nestmates.

Revisionary studies on the enigmatic neotropical ant genus Stegomyrmex Emery, 1912 (Hymenoptera : Formicidae : Myrmicinae), with the description of two new species

The recent increase in leaf litter ants sampling effort in Neotropical wet forests has revealed new and interesting records of the highly specialized myrmicine ant genus Stegomyrmex Emery, previously considered as extremely rare. We present a modified diagnosis for the genus and describe Stegomyrmex bensoni n. sp. and S. olindae n. sp., based on, respectively, workers, males, and gyne (central-north Brazil) and on a single worker (northern Brazil). Stegomyrmex vizottoi Diniz (southeastern Brazil) is redescribed and compared with S. olindae n. sp.; these species present significant differences in size, sculpturation, and sting apparatus morphology. The males of S. vizottoi are described for the first time. A key for workers and queens and a distribution map for the five know Stegomyrmex species are provided.

Composition of the Ant Fauna (Hymenoptera: Formicidae) in Public Squares in Southern Brazil

The composition of the ant fauna was examined in public squares of three municipalities that compose the hydrographic basin of the Upper Tiete River: Biritiba Mirim, Salesopolis, and Mogi das Cruzes. Richness, frequency of occurrence, similarity, and influence of seasons on the species composition were examined. The method was standardized as sampling units consisted of a set of three baits arranged in a triangle with vertices two meters apart. Sardines in oil were used as attractant. A total of 86 species was collected. Myrmicinae and Pheidole were the richest subfamily and genus, respectively. Eighty species were collected in Mogi das Cruzes, 49 in Salesopolis, and 45 in Biritiba Mirim, with 34 species common to the three areas. The ordination analysis (NMDS) revealed the presence of two distinct communities: one in Mogi das Cruzes and another in Biritiba Mirim-Salesopolis. These data were supported by the dendogram based on the Bray-Curtis dissimilarity index. This result might be associated with the distinct geographic and demographic characteristics of the areas. Regarding seasonality, the composition of the fauna of Mogi das Cruzes is independent of the season of the year, unlike the observed in Biritiba Mirim and Salesopolis.

Abiotic factors shape temporal variation in the structure of an ant-plant network

Despite recognition of key biotic processes in shaping the structure of biological communities, few empirical studies have explored the influences of abiotic factors on the structural properties of mutualistic networks. We tested whether temperature and precipitation contribute to temporal variation in the nestedness of mutualistic ant-plant networks. While maintaining their nested structure, nestedness increased with mean monthly precipitation and, particularly, with monthly temperature. Moreover, some species changed their role in network structure, shifting from peripheral to core species within the nested network. We could summarize that abiotic factors affect plant species in the vegetation (e.g., phenology), meaning presence/absence of food sources, consequently an increase/decrease of associations with ants, and finally, these variations to fluctuations in nestedness. While biotic factors are certainly important, greater attention needs to be given to abiotic factors as underlying determinants of the structures of ecological networks.

Compared morphology of the immatures of males of two urban ant species of Camponotus

The immatures of males of two species of Camponotus ants (Hymenoptera: Formicidae) are described and compared by light and electron microscopy. The numbers of larval instars were determined: Camponotus rufipes Fabricius (Hymenoptera: Formicidae) have four instars; and Camponotus vittatus Forel have three. Male larvae of the two species are similar to previously described Camponotus larvae, sharing the following traits: basic shape of body and mandible, presence of 'chiloscleres', 'praesaepium' (some specimens), labial pseudopalps, and ten pairs of spiracles. However, larvae of the two species can be separated by bodily dimensions and based on their hair number and types. Worker larvae of C. vittatus previously described are extensively similar to male larvae, with only a few inconspicuous differences that may result from intraspecific variation or sexual differences.

Social facilitation and food partitioning in the queenless ant Dinoponera quadriceps (Hymenoptera: Formicidae)

Social facilitation occurs when an animal is more likely to behave in a certain way in response to other animals engaged in the same behaviour. For example, an individual returning to the nest with food stimulates other ants to leave and to forage. In the present study we demonstrate the existence of new facets in the colony organization of Dinoponera quadriceps: a positive feedback between the incoming food and the activation of new foragers, and the occurrence of incipient task partitioning during the food sharing. Lower-ranked workers located inside the nest process protein resources and higher-ranked workers handle smaller pieces and distribute them to the larvae. In conclusion, D. quadriceps has a decentralized pattern of task allocation with a double regulatory mechanism, which can be considered a sophisticated aspect of division of labour in ponerine ants.

Natural history of an ant-plant-butterfly interaction in a Neotropical savanna

Non-myrmecophilous lepidopteran larvae using plants bearing ant attractants such as extrafloral nectaries are good models for studying morphological and behavioural mechanisms against ant predation. Udranomia spitzi (Hesperiidae) is a butterfly whose larvae feed on leaves of Ouratea spectabilis (Ochnaceae), a plant with extrafloral nectaries. We described the early stages of U. spitzi, and used field observations and experiments to investigate the defensive strategies of caterpillars against predatory ants. Larvae pass through five instars and pupation occurs inside larval leaf shelters. Ant-exclusion experiments revealed that the presence of ants did not affect significantly caterpillar survival. Predation experiments showed that vulnerability to ant predation decreased with increase in larval size. The present study showed that predatory ants are not as relevant as demonstrated for other systems, and also illustrates how observational data and field experiments can contribute to a better understanding of the biology and ecology of a species of interest.

Reproductive biology of the ghost shrimp Lepidophthalmus bocourti (A. Milne-Edwards, 1870) (Decapoda: Axiidea: Callianassidae): A tropical species with a seasonal reproduction

Based on the hypothesis that reproduction is a continuous process in tropical habitats, we analysed reproductive periodicity and egg production in the callianassid ghost shrimp Lepidophthalmus bocourti, one of the most common species in mangrove systems along the Pacific coast of Central America. During one year (May 2008 to April 2009), individuals of L. bocourti (N = 499) were collected nearshore Gulf of Nicoya, Pacific coast of Costa Rica. Observations were made on presence or absence of incubated embryos, and gonad activity of females was analysed as gonadosomatic index (GSI). Our results revealed that L. bocourti has a marked seasonal breeding period, which contradicts previous reports regarding coastal marine decapods from the tropics. Ovigerous females were found only from June to August, while high GSI values were obtained from March to July. The increase of GSI and appearance of ovigerous females were associated with a concomitant decrease of salinity, but not with temperature. We assume that reproduction of L. bocourti is adapted to local changes of environmental conditions, and that a decrease in salinity during rainy season may serve as a triggering factor for ovarian development. Compared to other ghost shrimps, L. bocourti produced on average more (2002 +/- 1365) and smaller (0.87 +/- 0.109 mm) eggs, which seems to suggest that this species does not have an abbreviated larval development as reported for other species of genus. The deviation from the generalization of constant reproduction in the tropics for shallow water marine invertebrates and its probable cause are adequately discussed.

Influence of Cave Size and Presence of Bat Guano on Ant Visitation

This is the first study which evaluated the influence of cave size and presence of bat guano in ant visitation in Brazilian caves. We provide a list of the ants associated with 27 caves in northeastern Brazil, an area situated in the transition between Cerrado (Brazilian savanna) and Amazon Domain. The study was conducted between January and August 2010. We recorded 24 ant species inserted into 12 genera, 10 tribes, and six subfamilies. The size of the cave and the presence of guano did not influence the richness of ants, and most of the caves had single species. Camponotus atriceps was the species with the larger distribution, being collected in five caves. In addition, we discuss geographic distribution of records and possible ecological roles of ants in cave environments.

Cuticular Hydrocarbon Variation of Castes and Sex in the Weaver Ant Camponotus textor (Hymenoptera: Formicidae)

Cuticular hydrocarbons play important roles as chemical signatures of individuals, castes, sex and brood. They also can mediate the regulation of egg laying in ants, by informing directly or indirectly the reproductive status of queens. In this study we asked whether cuticular hydrocarbon profiles are correlated with castes and sex of Camponotus textor. Cuticular hydrocarbons were extracted from part of a mature colony (80 workers, 27 major workers, 27 queens, 27 virgin queens and 27 males). Results showed that cuticular hydrocarbons varied quantitatively and qualitatively among the groups and this variation was sufficiently strong to allow separation of castes and genders. We discuss the specificity of some compounds as possible regulatory compounds of worker tasks and reproduction in C. textor.

No-pole condition in Landau gauge: Properties of the Gribov ghost form factor and a constraint on the 2d gluon propagator

We study general properties of the Landau-gauge Gribov ghost form factor sigma(p(2)) for SU(N-c) Yang-Mills theories in the d-dimensional case. We find a qualitatively different behavior for d = 3, 4 with respect to the d = 2 case. In particular, considering any (sufficiently regular) gluon propagator D(p(2)) and the one-loop-corrected ghost propagator, we prove in the 2d case that the function sigma(p(2)) blows up in the infrared limit p -> 0 as -D(0) ln(p(2)). Thus, for d = 2, the no-pole condition sigma(p(2)) < 1 (for p(2) > 0) can be satisfied only if the gluon propagator vanishes at zero momentum, that is, D(0) = 0. On the contrary, in d = 3 and 4, sigma(p(2)) is finite also if D(0) > 0. The same results are obtained by evaluating the ghost propagator G(p(2)) explicitly at one loop, using fitting forms for D(p(2)) that describe well the numerical data of the gluon propagator in two, three and four space-time dimensions in the SU(2) case. These evaluations also show that, if one considers the coupling constant g(2) as a free parameter, the ghost propagator admits a one-parameter family of behaviors (labeled by g(2)), in agreement with previous works by Boucaud et al. In this case the condition sigma(0) <= 1 implies g(2) <= g(c)(2), where g(c)(2) is a "critical" value. Moreover, a freelike ghost propagator in the infrared limit is obtained for any value of g(2) smaller than g(c)(2), while for g(2) = g(c)(2) one finds an infrared-enhanced ghost propagator. Finally, we analyze the Dyson-Schwinger equation for sigma(p(2)) and show that, for infrared-finite ghost-gluon vertices, one can bound the ghost form factor sigma(p(2)). Using these bounds we find again that only in the d = 2 case does one need to impose D(0) = 0 in order to satisfy the no-pole condition. The d = 2 result is also supported by an analysis of the Dyson-Schwinger equation using a spectral representation for the ghost propagator. Thus, if the no-pole condition is imposed, solving the d = 2 Dyson-Schwinger equations cannot lead to a massive behavior for the gluon propagator. These results apply to any Gribov copy inside the so-called first Gribov horizon; i.e., the 2d result D(0) = 0 is not affected by Gribov noise. These findings are also in agreement with lattice data.

Sodium-specific foraging by leafcutter ant workers (Atta cephalotes, Hymenoptera: Formicidae)

1. Sodium is often a limiting nutrient for terrestrial animals, and may be especially sought by herbivores. Leafcutter ants are dominant herbivores in the Neotropics, and leafcutter foraging may be affected by nutritional demands of the colony and/or the demands of their symbiotic fungal mutualists. We hypothesized that leafcutter colonies are sodium limited, and that leafcutter ants will therefore forage specifically for sodium. 2. Previous studies demonstrated that leafcutter Atta cephalotes Linnaeus workers preferentially cut and remove paper baits treated with NaCl relative to water control baits. Atta cephalotes colonies in this study were presented with baits offering NaCl, Na2SO4, and KCl to test whether leafcutters forage specifically for sodium. Sucrose and water were used as positive and negative controls, respectively. 3. Atta foragers removed significantly more of the baits treated with NaCl and Na2SO4 than the KCl treatment, which did not differ from water. The NaCl and Na2SO4 treatments were collected at similar rates. We conclude A. cephalotes forage specifically for sodium rather than for anions (chloride) or solutes in general. This study supports the hypothesis that leafcutter ants are limited by, and preferentially forage for, sodium.