Community-level interactions and functional ecology of major workers in the hyperdiverse ground-foraging Pheidole (Hymenoptera, Formicidae) of Amazonian Ecuador

Ants of the genus Pheidole are abundant and hyperdiverse, particularly in Neotropical rainforests. Very little is known, however, about the degree of ecological and behavioral differentiation of coexisting species comprising Pheidole communities. Additionally, the ecological role of the major worker subcaste, thought to be significant to the diversification of Pheidole, is poorly understood. We investigated the ecology and behavior of a ground-foraging Pheidole community of at least 56 species in Amazonian Ecuador. Pheidole species differed strongly in tolerance to flooding, nest site usage, foraging range, major worker foraging, and control of baits, but not in daily activity or ability to discover baits. A molecular phylogeny based on mitochondrial DNA was characterized by poorly resolved basal relationships and long terminal branches, suggesting an ancient diversification of many Pheidole lineages. Comparison of well-supported sister species suggests that both phylogenetic history and ecologically induced differentiation contribute to interspecific variation in Amazonian Pheidole. Ground-nesting species had larger major workers than twig-nesting species, whereas dominant species with stronger recruitment had a higher proportional abundance of major workers at baits. Variation among species suggests the presence of behavioral groups within the Amazonian Pheidole community that appear to segregate according to nest site usage and/or tolerance to flooding disturbance. Our results suggest an important role for major worker differentiation in the diversification of Pheidole.     

Plant killing by Neotropical acacia ants: ecology,decision‐making,and head morphology

Mutualistic species often associate with several partners that vary in the benefits provided. In some protective ant–plant mutualisms, ants vary in the extent at which they kill neighboring vegetation. Particularly, in acacia ants (Pseudomyrmex), the area around the host tree that ants keep free from vegetation (“clearings”) vary depending on the species. This study assessed whether interspecific variation in clearing size corresponds to workers biting on plant tissue of different thickness. As expected, workers from species making the largest clearings bit more often on thicker plant tissues than workers from species making smaller clearings. Because head shape affects mandible force, I also assessed whether pruning on thick tissue in mutualistic ant species or being a predator in non‐mutualistic species correlated with broader heads, which yield stronger mandible force. The species with the broader heads were non‐mutualistic predators or mutualistic pruners of thick tissues, which suggest that pruning neighboring vegetation in non‐predatory species demands force even when the ants do not kill prey with their mandibles. The findings reveal that clearing size variation in mutualistic ant partners of plants can also be observed at the level of individual decision‐making processes among workers, and suggest that head morphology could be a trait under selection in protective ant–plant mutualisms. Abstract in Spanish is available with online material.     

Diet of two sympatric Pheidole spp. ants in the central Monte desert: implications for seed–granivore interactions

Ants of the genus Pheidole are important seed consumers in several desert ecosystems. In South American deserts, although several Pheidole spp. have been characterized as seed harvesters, studies on their diet and ecological role are still missing. Pheidole spininodis (Mayr) and Pheidole bergi (Mayr) are capable of removing seeds in the central Monte desert. The aim of this study was to quantify and compare the diet of these species and to interpret the results in the context of seed–granivore interactions. Diet was estimated during mid-summer by collecting items brought back to the nest by foragers in ten colonies per species. While P. spininodis was mainly granivorous, P. bergi was mainly insectivorous. However, they both collected ~40% of other types of items. Among seeds, the diet of P. spininodis included mostly grass seeds, whereas the diet of P. bergi was mainly made up of shrub and tree seeds, usually retrieved cooperatively. This behavior allowed P. bergi to carry larger seeds, resulting in diet partitioning in terms of seed size. However, diet of P. spininodis is very similar to that of three sympatric Pogonomyrmex species. Thus, specialized harvester ants remove large quantities of grass seeds in the central Monte desert during the summer, potentially affecting their abundance in the soil seed bank. P. bergi directs its feeding pressure to shrub and tree seeds, and although seeds constitute ~10% of its diet, its high colony density and high activity levels, added to the lower proportion of large seeds in the soil seed bank, indicate that their importance as seed consumers cannot be ruled out.     

Body size variation and caste ratios in geographically distinct populations of the invasive big‐headed ant,Pheidole megacephala (Hymenoptera: Formicidae)

Body size is an important life history trait that can evolve rapidly as a result of how species interact with each other and their environment. Invasive species often encounter vastly different ecological conditions throughout their introduced range that can influence relative investment in growth, reproduction and defence among populations. In this study, we quantified variation in worker size, morphology and proportion of majors among five populations of a worldwide invasive species, the big‐headed ant, Pheidole megacephala (Fabricius). The sampled populations differed in ant community composition, allowing us to examine if P. megacephala invests differently in the size and number of majors based on the local ant fauna. We also used genetic data to determine if these populations of P. megacephala represented cryptic species or if morphological differences could be attributed to change following introduction. We found significant variation in worker mass among the populations. Both major and minor workers were largest in Australia, where the ant fauna was most diverse, and minor workers were smallest in Hawaii and Mauritius, where P. megacephala interacted with few to no other ants. We also found differences in major and minor worker morphology among populations. Majors from Mauritius had significantly larger heads (width and length) relative to whole body size than those from Hawaii and Florida. Minors had longer heads and hind tibias in South Africa compared with populations from Australia, Hawaii and Florida. The proportion of majors did not differ among populations, suggesting that these populations may not be subject to trade‐offs in investment in major size versus number. Our molecular data place all samples within the same clade, supporting that these morphologically different populations represent the same species. These results suggest that the variation in shape and morphology of major and minor workers may therefore be the result of rapid adaptation or plastic responses to local conditions. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 423–438.     

Caste specialization in food storage in the dimorphic antColobopsis nipponicus (Wheeler)

Summary Food-storage abilities of both minor and major workers of the dimorphic antColobopsis nipponicus were examined to prove the hypothesis that major workers of this species have a trophic role in addition to a defensive one. Both worker subcastes accumulated water, that was supplied with water-soluble food, as well as fat, that probably originated from food given. However, the major workers accumulated much larger amounts of water and fat than did the minor workers. Difference in water accumulation increased as food supply increased. In spite of their difference in body weight, the residual amount of fat after starvation did not differ between the two subcastes. Although the amount of stored fat increased with colony size in both subcastes, the stored fat per dry weight of the major worker was significantly larger than that of the minor worker regardless of the colony size. When workers of a colony were starved, minor workers with a single major worker survived significantly longer than those without a major worker. In addition, about half the behavioral acts of major workers were regurgitation for minor workers. These results demonstrate that major workers ofC. nipponicus functioned as a trophic caste. On the basis of the results, quantitative relationships of storage abilities between the two worker subcastes are discussed.     

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

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

Improving Planting Stocks for the Brazilian Atlantic Forest Restoration through Community‐Based Seed Harvesting Strategies

High‐diversity reforestation can help jumpstart tropical forest restoration, but obtaining viable seedlings is a major constraint: if nurseries do not offer them, it is hard to plant all the species one would like. From 2007 to 2009, we investigated five different seed acquisition strategies employed by a well‐established tree nursery in southeastern Brazil, namely (1) in‐house seed harvesters; (2) hiring a professional harvester; (3) amateur seed harvesters; or (4) a seed production cooperative, as well as (5) participating in a seed exchange program. In addition, we evaluated two strategies not dependent on seeds: harvesting seedlings from native tree species found regenerating under Eucalyptus plantations, and in a native forest remnant. A total of 344 native tree and shrub species were collected as seeds or seedlings, including 2,465 seed lots. Among these, a subset of 120 species was obtained through seed harvesting in each year. Overall, combining several strategies for obtaining planting stocks was an effective way to increase species richness, representation of some functional groups (dispersal syndromes, planting group, and shade tolerance), and genetic diversity of seedlings produced in forest tree nurseries. Such outcomes are greatly desirable to support high‐diversity reforestation as part of tropical forest restoration. In addition, community‐based seed harvesting strategies fostered greater socioeconomic integration of traditional communities in restoration projects and programs, which is an important bottleneck for the advance of ecological restoration, especially in developing countries. Finally, we discuss some of the limitations of the various strategies for obtaining planting stocks and the way forward for their improvement.     

Correlates of ecological dominance within Pheidole ants (Hymenoptera: Formicidae)

1. In any group of organisms, one can almost invariably find some species that are ecologically dominant (i.e. disproportionately more abundant and widespread), whereas others are comparatively less prevalent. Understanding of the causes of variation in ecological dominance has been elusive, particularly given that dominant and subordinate species often lack obvious features that could predict their abundance in nature. 2. In this study, physiological, behavioural, morphological, and phylogenetic information is integrated in an effort to understand the mechanisms underlying ecological dominance in ants using the hyperdiverse ant genus Pheidole (Formicidae: Myrmicinae) as a model system. Field estimates of the relative abundance of 10 Pheidole species were compared with potential correlates, which included behavioural (walking velocity), physiological (tolerance to high and low temperatures and desiccation), and morphological traits (body size and degree of dimorphism in the worker caste). A molecular phylogeny of the tested species was also generated to account for potential confounding effects of phylogenetic non‐independence. 3. Dominant Pheidole species were characterised by higher environmental tolerance with respect to temperature and humidity, as well as faster walking speeds. On the other hand, no morphological correlates of ecological dominance were detected. Interestingly, subordinate species showed no evidence of trade‐off in performance, being both more fragile to environmental challenges and slower in their walking speeds. 4. These results provide important insights into the mechanisms involved in local species coexistence in Pheidole.     

The effect of limited visibility on vigilance behaviour and speed of predator detection: implications for the conservation of granivorous passerines

Ants have been traditionally considered either as predators or dispersers of seeds, but not both. That is, ant dispersal is restricted to myrmecochorous seeds, while almost all seeds removed by seed‐harvesting ants are eaten. However, harvesting ants might be simultaneously antagonistic and mutualistic towards seeds. This study analyzes the predation–dispersal relationship between seed‐harvesting ants and seeds of Lobularia maritima, a non‐myrmechorous perennial herb, in order to disentangle the dual role of ants as dispersers and predators of L. maritima seeds. The results obtained confirm the role of harvesting ants as both predators and dispersers of the non‐myrmechorous seeds of L. maritima. The removal activity of Messor bouvieri on L. maritima seeds is very important, particularly in autumn, which is the flowering and fruiting peak of this plant. It can be estimated that harvesting ants collect more than 85% of seeds, and almost 70% of them are effectively lost to predation. However, these granivorous ants also have drawbacks as seed dispersers. There is a relatively small percent of seeds collected by ants that escape predation, either because they are dropped on the way to the nest (16.4% of seeds harvested), or because they are mistakenly rejected on the refuse pile (0.9%). Abiotic dispersal of L. maritima seeds in the absence of ants occurs over very short distances from the plant stem. As seeds dispersed by ants reach a considerably greater distance than that obtained by gravity, this might represent a real advantage for the species, because it reduces intraspecific adult competition for seedlings, which directly influences seedling survivorship. These results challenge the generalization that seed removal by ants generally leads to successful seed dispersal if done by legitimate seed dispersers, or seed loss if done by seed consumers that eat them, and confirm that harvesting ants might have a dual role as both predators and dispersers of nonmyrmechorous seeds.     

Macroevolutionary integration of phenotypes within and across ant worker castes

Phenotypic traits are often integrated into evolutionary modules: sets of organismal parts that evolve together. In social insect colonies, the concepts of integration and modularity apply to sets of traits both within and among functionally and phenotypically differentiated castes. On macroevolutionary timescales, patterns of integration and modularity within and across castes can be clues to the selective and ecological factors shaping their evolution and diversification. We develop a set of hypotheses describing contrasting patterns of worker integration and apply this framework in a broad (246 species) comparative analysis of major and minor worker evolution in the hyperdiverse ant genus Pheidole. Using geometric morphometrics in a phylogenetic framework, we inferred fast and tightly integrated evolution of mesosoma shape between major and minor workers, but slower and more independent evolution of head shape between the two worker castes. Thus, Pheidole workers are evolving as a mixture of intracaste and intercaste integration and rate heterogeneity. The decoupling of homologous traits across worker castes may represent an important process facilitating the rise of social complexity.     

Head and mandible shapes are highly integrated yet represent two distinct modules within and among worker subcastes of the ant genus Pheidole

Ants use their mandibles for a wide variety of tasks related to substrate manipulation, brood transport, food processing, and colony defense. Due to constraints involved in colony upkeep, ants evolved a remarkable diversity of mandibular forms, often related to specific roles such as specialized hunting and seed milling. Considering these varied functional demands, we focused on understanding how the mandible and head shape vary within and between Pheidole subcastes. Using x‐ray microtomography and 3D geometric morphometrics, we tested whether these structures are integrated and modular, and how ecological predictors influenced these features. Our results showed that mandible and head shape of majors and minor workers tend to vary from robust to slender, with some more complex changes related to the mandibular base. Additionally, we found that head and mandible shapes are characterized by a high degree of integration, but with little correlation with feeding and nesting habits. Our results suggest that a combination of structural (allometric) constraints and the behavioral flexibility conferred by subcaste dimorphism might largely buffer selective pressures that would otherwise lead to a fine‐tuning between ecological conditions and morphological adaptation.     

Secondary seed dispersal by ants in Neotropical cerrado savanna: species‐specific effects on seeds and seedlings of Siparuna guianensis (Siparunaceae)

1. Most woody plant species in tropical habitats are primarily vertebrate‐dispersed, but interactions between ants and fallen seeds and fruits are frequent. This study assesses the species‐specific services provided by ants to fallen arillate seeds of Siparuna guianensis, a primarily bird‐dispersed tree in cerrado savanna. The questions of which species interact with fallen seeds, their relative contribution (versus vertebrates) to seed removal, and the potential effects on seedling establishment are investigated. 2. Seeds are removed in similar quantities in caged and control treatments, suggesting that ants are the main dispersers on the ground. Five ant species attended seeds. Pheidole megacephala (≈0.4 cm) cooperatively transported seeds, whereas the smaller Pheidole sp. removed the seed aril on spot. Large (> 1.0 cm) Odontomachus chelifer, Pachycondyla striata, and Ectatomma edentatum individually carried seeds up to 4 m. Bits of aril are fed to larvae and intact seeds are discarded near the nest entrance. 3. Overall, greater numbers of seedlings were recorded near ant nests than in control plots without nests. This effect, however, was only detected near P. megacephala and P. striata nests, where soil penetrability was greater compared with controls. Soil nutrients did not differ between paired plots. 4. This study confirms the prevalence of ant–seed interactions in cerrado and shows that ant‐derived benefits are species‐specific. Ant services range from seed cleaning on the spot to seed displacement promoting non‐random spatial seedling recruitment. Although seed dispersal distances by ants are likely to be shorter than those by birds, our study of S. guianensis shows that fine‐scale ant‐induced seed movements may ultimately enhance plant regeneration in cerrado.     

Worker size and seed size selection by harvester ants in a neotropical forest

The rules ants follow when selecting seeds are important both to theories of colony organization and to the shaping of their impact as harvesters. Two Costa Rican forests yielded the most diverse harvester ant assemblages yet studied (41 and 44±4 species). I assayed seed size preferences within and between species using milled barley. Seed size selection varied at a number of levels. First, species differed in seed size use: small species mainly carried off small seeds; large species retrieved a broad range of seed sizes. Within species, one-third of those tested yielded inter-colony differences in preferred seed sizes. Finally, workers of polymorphic species tested showed significant worker/seed size correlations. Species, colony, and worker level differences are common among harvester ants. Their significance to community organization and colony ergonomics however is hotly debated and requires a better understanding of the dynamics of food limitation. From the plant's perspective, small seeds will be harvested by a wider variety of ants than large seeds.     

Caste specialization in behavioral defenses against fungus garden parasites in <Emphasis Type="Italic">Acromyrmex octospinosus</Emphasis> leaf-cutting ants

Division of labor and caste specialization plays an important role in many aspects of social insect colony organization, including parasite defense. Within leaf-cutting ant colonies, worker caste specialization permeates colony tasks ranging from foraging, substrate incorporation, brood care, and chemical defenses via glandular secretions and mutualistic bacteria. Leaf-cutting ants rely on protecting a mutualistic fungus they grow for food from microfungi in the genus Escovopsis that parasitizes the ant–fungus relationship. Here, we examine whether Acromyrmex octospinosus leaf-cutter ant castes (minors and majors) display task specialization in two behavioral defenses against Escovopsis: fungus grooming (the removal of Escovopsis spores) and weeding (the removal of large pieces of Escovopsis-infected fungus garden). Using behavioral observations, we show that minors are the primary caste that performs fungus grooming, while weeding is almost exclusively performed by majors. In addition, using a sub-colony infection experimental setup, we show that at the early stages of infection, minors more efficiently remove Escovopsis spores from the fungus garden, thereby restricting Escovopsis spore germination and growth. At later stages of infection, after Escovopsis spore germination, we find that major workers are as efficient as minors in defending the fungus garden, likely due to the increased importance of weeding. Finally, we show, using SEM imaging, that the number of sensory structures is similar between minor and major workers. If these structures are invoked in recognition of the parasites, this finding suggests that both castes are able to sense Escovopsis. Our findings support that leaf-cutter ant behavioral defense tasks against Escovopsis are subject to caste specialization, likely facilitated by worker sizes being optimal for grooming and weeding by minors and majors, respectively, with important consequences for cultivar defense.     

Removal of seeds from frugivore defecations by ants in a Costa Rican rain forest

At our Costa Rican field site, seeds defecated by frugivorous birds usually do not remain where they have been deposited. Many species of ants are attracted to frugivore defecations and remove seeds and/or pulp. Pheidole species selectively remove seeds, fungus-growing species (tribe Attini) remove both pulp and seeds. Seeds of many Melastomataceae have an appendage, which we hypothesized is an elaiosome. Indeed, preference trials demonstrated that two species of Pheidole selected seeds with the appendage over seeds of the same species in which the appendage had been removed. However, we found that these ants did not take the appendage when it was offered by itself. We conclude that the appendage is not an elaiosome. In further trials, different ant species preferentially selected different seed species. These ants consumed some seeds and deposited others unharmed in refuse piles. We conclude that because the composition of leaf-litter ant communities is highly variable between neighboring square meter plots, and the probability of seed predation depends upon the species of ant, the over-all effect of ants on seed shadows and seed banks is spatially unpredictable. Addendum: The names of the two Pheidole emphasized in this study. P. nebulosa and P. nigricula, are unpublished names from a generic revision being prepared by E. O. Wilson and W. L. Brown. Their use here is not intended to constitute taxonomic publication but is solely for more precise indentification in future ecological research of similar nature     

Relationships between intra‐specific variation in seed size and recruitment in four species in two contrasting habitats

Large seeds contain more stored resources, and seedlings germinating from large seeds generally cope better with environmental stresses such as shading, competition and thick litter layers, than seedlings germinating from small seeds. A pattern with small‐seeded species being associated with open habitats and large‐seeded species being associated with closed (shaded) habitats has been suggested and supported by comparative studies. However, few studies have assessed the intra‐specific relationship between seed size and recruitment, comparing plant communities differing in canopy cover. Here, seeds from four plant species commonly occurring in ecotones between open and closed habitats (Convallaria majalis, Frangula alnus, Prunus padus and Prunus spinosa) were weighed and sown individually (3200 seeds per species) in open and closed‐canopy sites, and seedling emergence and survival recorded over 3 years. Our results show a generally positive, albeit weak, relationship between seed size and recruitment. In only one of the species, C. majalis, was there an association between closed canopy habitat and a positive seed size effect on recruitment. We conclude that there is a weak selection gradient favouring larger seeds, but that this selection gradient is not clearly related to habitat.     

Supplementing Seed Banks to Rehabilitate Disturbed Mojave Desert Shrublands: Where Do All the Seeds Go?

Revegetation of degraded arid lands often involves supplementing impoverished seed banks and improving the seedbed, yet these approaches frequently fail. To understand these failures, we tracked the fates of seeds for six shrub species that were broadcast across two contrasting surface disturbances common to the Mojave Desert—sites compacted by concentrated vehicle use and trenched sites where topsoil and subsurface soils were mixed. We evaluated seedbed treatments that enhance soil‐seed contact (tackifier) and create surface roughness while reducing soil bulk density (harrowing). We also explored whether seed harvesting by granivores and seedling suppression by non‐native annuals influence the success of broadcast seeding in revegetating degraded shrublands. Ten weeks after treatments, seeds readily moved off of experimental plots in untreated compacted sites, but seed movements were reduced 32% by tackifier and 55% through harrowing. Harrowing promoted seedling emergence in compacted sites, particularly for the early‐colonizing species Encelia farinosa, but tackifier was largely ineffective. The inherent surface roughness of trenched sites retained three times the number of seeds than compacted sites, but soil mixing during trench development likely altered the suitability of the seedbed thus resulting in poor seedling emergence. Non‐native annuals had little influence on seed fates during our study. In contrast, the prevalence of harvester ants increased seed removal on compacted sites, whereas rodent activity influenced removal on trenched sites. Future success of broadcast seeding in arid lands depends on evaluating disturbance characteristics prior to seeding and selecting appropriate species and seasons for application.     

Orchid seed removal by ants in Neotropical ant‐gardens

• Most plants that inhabit ant‐gardens (AGs) are cultivated by the ants. Some orchids occur in AGs; however, it is not known whether their seeds are dispersed by AG ants because most orchid seeds are tiny and dispersed by wind.
• We performed in situ seed removal experiments, in which we simultaneously provided Azteca gnava ants with seeds of three AG orchid species and three other AG epiphyte species (Bromeliaceae, Cactaceae and Gesneriaceae), as well as the non‐AG orchid Catasetum integerrimum.
• The seeds most removed were those of the bromeliad Aechmea tillandsioides and the gesneriad Codonanthe uleana, while seeds of AG orchids Coryanthes picturata, Epidendrum flexuosum and Epidendrum pachyrachis were less removed. The non‐AG orchid was not removed. Removal values were positively correlated with the frequency of the AG epiphytes in the AGs, and seeds of AG orchids were larger than those of non‐AG orchids, which should favour myrmecochory.
• Our data show that Azt. gnava ants discriminate and preferentially remove seeds of the AG epiphytes. We report for the first time the removal of AG orchid seeds by AG ants in Neotropical AGs.

     

Cryptically dimorphic caste differences in a Neotropical, swarm-founding paper wasp genus, Parachartergus (Hymenoptera: Vespidae)

Morphological differences in body shape between females of different reproductive conditions (in terms of insemination and ovarian development) were examined in two species of the Neotropical polistine genus Parachartergus: P. smithii and P. fraternus. The present study shows, for the first time, that non‐size‐based morphological divergence between queens and workers occurs in Parachartergus, an epiponine genus once believed to have little or no morphological caste differences. In the P. smithii colony examined, queens were significantly larger than workers in five of the eight body parts measured (head width, eye width, genal width, mesosomal length, wing length, first metasomal tergum width, and width and length of the second tergum), but the mean values of wing length and first and second tergum widths were not significantly different between them. The queen : worker size ratios tended to be greater anteriorly and smaller posteriorly, although the size ratio was greatest in second tergum length. Analysis of covariance (ancova ) with mesosomal length as covariate showed that queens had proportionally wider heads and narrower first terga than did workers. In the P. fraternus colony, size differences between queens and workers were not significant, and there was little or no difference in shape, but queens had significantly proportionally wider first terga than did workers.     

Differences in sNPF Receptor-Expressing Neurons in Brains of Fire Ant (Solenopsis invicta Buren) Worker Subcastes: Indicators for Division of Labor and Nutritional Status?

In the red imported fire ant, Solenopsis invicta Buren, the neuronal and molecular mechanisms related to worker division of labor are poorly understood. Workers from different subcastes (major, medium and minors) perform different tasks, which are loosely associated with their size. We hypothesized that the short neuropeptide F (sNPF) signaling system (NPY-like) could be involved in mechanisms of worker division of labor and sensing or responding to colony nutritional requirements. Thus, we investigated the expression of the short neuropeptide F receptor (sNPFR) in the brain and subesophageal ganglion (SEG) of workers from colonies with and without brood. Across worker subcastes a total of 9 clusters of immunoreactive sNPFR cells were localized in the brain and the subesophageal ganglion (SEG); some of these cells were similar to those observed previously in the queen. Worker brain sNPFR cell clusters were found in the protocerebrum near mushroom bodies, in the central complex and in the lateral horn. Other sNPFR immunoreactive cells were found at the edge of the antennal lobes. Across subcastes, we observed both a constant and a differential pattern of sNPFR clusters, with a higher number of sNPFR cells found in minor than in major workers. Those sNPFR cells detected in all worker subcastes appear to be involved in olfaction or SEG functions. The differential expression of clusters in subcastes suggests that sNPFR signaling is involved in regulating behaviors associated with specific subcastes and thus, division of labor. Some sNPFR cells appear to be involved in nutrient sensing and/or brood care, feeding behavior and locomotion. In colonies without brood, workers showed a lower cluster number, and an overall reduced sNPFR signal. Our results suggest the sNPF signaling system is a candidate for the neurobiological control of worker division of labor and sensing brood presence, perhaps correlating with protein requirements and availability.