Genus Pogonomyrmex Research Articles

Peptide toxins that target vertebrate voltage-gated sodium channels underly the painful stings of harvester ants

Harvester ants (genus Pogonomyrmex) are renowned for their stings which cause intense, long-lasting pain, and other neurotoxic symptoms in vertebrates. Here, we show that harvester ant venoms are relatively simple and composed largely of peptide toxins. One class of peptides is primarily responsible for the long-lasting local pain of envenomation via activation of peripheral sensory neurons. These hydrophobic, cysteine-free peptides potently modulate mammalian voltage-gated sodium (NaV) channels, reducing the voltage threshold for activation and inhibiting channel inactivation. These toxins appear to have evolved specifically to deter vertebrates.

Population genomics supports multiple hybrid zone origins of socially hybridogenetic lineages of Pogonomyrmex harvester ants.

Reproductive division of labor in the social insects is typically determined by environmental cues; however, genetic effects on caste have been discovered in a growing set of ant taxa. An extreme form of genetic caste determination is "social hybridogenesis," in which co-occurring genetic lineages obligately interbreed to produce workers, whereas daughter queens are of pure-lineage ancestry. In this study, we tested the hypothesis that social hybridogenesis in the genus Pogonomyrmex resulted from one or more interspecific hybridization events, and if so, whether individual lineages were of hybrid ancestry. We reconstructed evolutionary relationships of four lineage pairs to populations of two closely related non-hybridogenetic species, Pogonomyrmex barbatus and Pogonomyrmex rugosus, using nuclear SNP loci and mitochondrial sequencing. The nuclear phylogeny supported a hybridization hypothesis, with one member of each pair nested within P. rugosus, whereas the other was nested within P. barbatus. The source populations corresponded with two distinct geographic areas at the eastern and western edges of a zone of contact. Relatively little gene flow was detected between interbreeding lineages, either historically or currently. This suggests that shifts in reproductive caste determination may reinforce reproductive incompatibility, in a manner similar to the evolution of hybridogenesis in nonsocial systems.

A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part II

This paper provides a taxonomic revision and reviews natural history for 35 South American species of the seed-harvesting ant genus Pogonomyrmex. Species treated herein mostly comprise the P. rastratus-group; four species are revived from synonomy, three taxa are elevated from subspecies to species, five taxa are synonymized, and 20 new species are described. The following taxa are revived from synonomy: P. intermedia Menozzi, P. semistriata Emery, P. spinolae Emery, and P. weiseri Santschi. The following taxa are raised from subspecies to species: P. leonis Kusnezov, P. pulchellus Santschi, and P. sanmartini Kusnezov. The following new synonymies are proposed, with the senior synonym listed first, and the junior synonym(s) in parentheses: P. carbonarius Mayr (= P. kusnezovi Cuezzo Claver, = P. weiseri var. neuquensis Santschi, = P. variabilis Santschi); P. vermiculatus Emery (= P. vermiculatus var. chubutensis Forel, = P. vermiculatus var. jorgenseni Forel). The following new species are described: P. apterogenos, P. araucania, P. atacama, P. bolivianus, P. colca, P. cusquena, P. excelsior, P. forelii, P. granulatus, P. lagunabravensis, P. loaensis, P. mapuche, P. maulensis, P. pichachen, P. propinqua, P. santschii, P. strioligaster, P. tafi, P. varicolor, and P. wilsoni. One species treated herein has brachypterous queens (P. atacama), one species has dimorphic queens (winged and brachypterous in P. longibarbis), and two species have ergatoid (permanently wingless) queens and ergatoid males (P. apterogenos, P. laguanbravensis); the latter two are the only known ant species in which both sexual castes are only ergatoid. I also provide keys for workers and queens (in English and Spanish), diagnoses for males, photographs of known castes, distribution maps, and a summary of known biology.

Genome assembly and annotation of the California harvester ant Pogonomyrmex californicus.

The harvester ant genus Pogonomyrmex is endemic to arid and semiarid habitats and deserts of North and South America. The California harvester ant Pogonomyrmex californicus is the most widely distributed Pogonomyrmex species in North America. Pogonomyrmex californicus colonies are usually monogynous, i.e. a colony has one queen. However, in a few populations in California, primary polygyny evolved, i.e. several queens cooperate in colony founding after their mating flights and continue to coexist in mature colonies. Here, we present a genome assembly and annotation of P. californicus. The size of the assembly is 241 Mb, which is in agreement with the previously estimated genome size. We were able to annotate 17,889 genes in total, including 15,688 protein-coding ones with BUSCO (Benchmarking Universal Single-Copy Orthologs) completeness at a 95% level. The presented P. californicus genome assembly will pave the way for investigations of the genomic underpinnings of social polymorphism in the number of queens, regulation of aggression, and the evolution of adaptations to dry habitats.

Complex foraging ecology of the red harvester ant and its effect on the soil seed bank

Granivory is an important interaction in the arid and semi-arid zones of the world, since seeds form an abundant and nutritious resource in these areas. While species of the genus Pogonomyrmex have been studied in detail as seed predators, their impact on seed abundance in the soil has not yet been explored in sufficient depth. We studied the impact of the harvesting activities of the ant Pogonomyrmex barbatus on seed abundance in the soil of the Zapotitlán valley, Mexico. We found that P. barbatus activity significantly impacts the abundance of seeds in the soil, which is lower in the sites where P. barbatus forages than it is in sites with no recorded foraging. We also found that P. barbatus distributes intact seeds of three tree species, two of which are nurse plants, and could consequently be promoting the establishment of these species. Using tools derived from graph theory, we observed that the ant-seed interactions exhibit a nested pattern; where more depredated seed species seem to be the more spatially abundant in the environment. This study illustrates the complex foraging ecology of the harvester ant P. barbatus and elucidates its effect on the soil seed bank in a semi-arid environment.

Colony size and foraging strategies in desert seed harvester ants

We summarized the literature on foraging strategies, colony sizes, and body sizes of seed harvesting ants to test hypotheses about the relationship between colony size and foraging strategy, and body size and foraging strategy. We used a Wilcoxon-Mann-Whitney test to test hypotheses about body size, colony size, and foraging strategy with the following results: (1) The maximum length of a worker ant in trunk trail foraging ant species or group foraging ant species is significantly smaller than in individual foraging ants except in the North American genus Pogonomyrmex; and (2) the maximum number of worker ants per nest in trunk trail foraging ant species or group foraging ant species is significantly larger than in individual foraging ants. While most seed harvesting ants can forage as individuals, only seed harvesters with large colonies develop group or trunk-trail foraging strategies. Flexibility in foraging strategy is important in desert seed harvesters because of the unpredictability of food resources among years.

A new ant genus from southern Argentina and southern Chile, Patagonomyrmex (Hymenoptera: Formicidae)

The ant genus Pogonomyrmex (Hymenoptera: Formicidae: Myrmicinae) comprises 71 described species that occur in North America, South America, and Hispaniola, and it is the nominal genus in the recently established tribe Pogonomyrmecini. A molecular phylogeny using 3,647 base pairs from fragments of one mitochondrial gene (cytochrome oxidase I) and five nuclear genes (long-wavelength rhodopsin, elongation factor 1α F1, elongation factor 1α F2, wingless, rudimentary) inferred that Pogonomymrex was not monophyletic. The vast majority of species belonged to a monophyletic clade (Pogonomyrmex sensu stricto), whereas species in the Po. angustus-group formed a second lineage outside of Pogonomyrmex and Hylomyrma, the latter being the only other genus in the tribe. To maintain monophyly of Pogonomyrmex, we create the genus Patagonomyrmex gen. n., which consists of the three angustus-group species (Patagonomyrmex angustus comb. n., Patagonomyrmex laevigatus comb. n., and Patagonomyrmex odoratus comb. n.) that are sister to all other pogonomyrmecines. The genus is restricted to southern Argentina and southern and southcentral Chile. Workers are characterized by: (1) poorly-developed psammophore that consists of short to medium-length hairs scattered over the ventral surface of the head, (2) anterior clypeal margin strongly convex in full-face view, (3) clypeus strongly convex in profile, (4) superior and inferior propodeal spines long, (5) dorsal surface of promesonotum smooth and shiny, lacking rugae <span style="text-decoration: underline;">or if rugae present, then interrugae strongly granulate-punctate, (6) anteroventral margin of peduncle of the petiole with a small, acuminate spine, and (7) palp formula 5,4. We also provide information on biology, distribution maps, and a key to workers, queens, and males in English and Spanish.

A taxonomic revision of South American species of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae). Part I.

South American species in the seed-harvester ant genus Pogonomyrmex (subfamily Myrmicinae) are interesting biologically because of their numerous queen phenotypes and life histories. This paper provides a taxonomic revision and reviews the natural history for 21 South American species of Pogonomyrmex so that we can better study their rich and interesting ecology, life history, and evolution. Species treated herein comprise all South American species-groups except for the brevibarbis and rastratus-groups. The following taxa are raised to species: pencosensis Forel 1914 and serpens Santschi 1922. The following new synonomies are proposed: bruchi Forel 1913 is synonomized under coarctatus Mayr 1868 and cunicularius carnivora Santschi 1925 under serpens Santschi 1922. The following new species is described: tinogasta. This paper redescribes workers of all species, and I describe queens and diagnose males for the following species: bispinosus (ergatoid queen, male), inermis (queen, male), laticeps (male), lobatus (queen, male), micans (queen), naegelii (ergatoid queen), pencosensis (ergatoid queen), serpens (ergatoid queen), tinogasta (brachypterous queen), and uruguayensis (queen, male). A neotype was designated for the untraceable or possibly lost type of P. bispinosus, and a holotype or lectotype was designated from syntypes for all other previously described taxa in order to provide a single name-bearing specimen and to facilitate future taxonomic studies. Of the 21 species treated herein, five species have ergatoid (wingless) queens (bispinosus, cunicularius, pencosensis, serpens, mayri), two have brachypterous (short-winged) queens (mendozanus, tinogasta), and two have dimorphic queens (winged and ergatoid in naegelii, brachypterous and ergatoid in laticeps). I also provide keys for workers and queens (in English and Spanish), photographs of all castes, distribution maps, and a summary of known biology.

A taxonomic revision of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae) on Hispaniola.

We revise species of seed-harvester ants in the genus Pogonomyrmex (subfamily Myrmicinae) that occur on the Caribbean island of Hispaniola. Three species are recognized: P. aterrimus Wheeler (new status), P. saucius Wheeler and Mann, and P. schmitti Forel. Pogonomyrmex schmitti sublaevigatus Wheeler (= schmitti) and P. schmitti darlingtoni Wheeler (= aterrimus) are synonomized. We also describe the queen of P. aterrimus and P. saucius, and provide information on biology, distribution maps, and a key to workers and queens.

Doves kleptoparasitize ants

We describe seven instances of kleptoparasitism by the Inca dove (Columbina inca) on the red harvester ant (Pogonomyrmex barbatus) in eastern Mexico, the first documented case in the order Columbiformes and the first for ants of the genus Pogonomyrmex. Doves visited the periphery of the ants' nest entrance and fed on waste material expelled by ants from inside the nest. Doves also took seeds of the grasses Dactyloctenium aegyptium, Digitaria ciliaris, and the shrub Sida directly from Pogonomyrmex workers. During a 5-day period, we estimated that nearly 20% of the seeds harvested by ants and carried toward the nest were taken by the doves. Inca doves can be defined as facultative kleptoparasites.

Effect of harvester ants of the genus Pogonomyrmex on the soil seed bank around their nests in the central Monte desert, Argentina

1. The abundance and composition of soil seed banks is a key determinant of plant community structure. Harvester ants can remove huge quantities of preferred seeds close to the nest affecting composition and spatial distribution of plants.2. In the central Monte desert (Argentina) ants of the genus Pogonomyrmex have high seed removal rates, especially of the five main grasses. The aim of this study was to establish if their foraging activity affects spatial patterns of the soil seed bank around their nests. Our hypotheses were: (1) removal by ants decreases seed abundance of preferred species in the soil; and (2) the effect varies in time.3. Soil seed abundance was assessed at different distances from Pogonomyrmex nests in the litter and in bare soil at the beginning, the middle, and the end of the season (late spring‐early autumn).4. A lower seed abundance of preferred species was observed close to the nest in the litter at the end of the season. Non‐preferred species showed no distance gradient.5. The lower foraging activity and seed consumption at the beginning of the season could explain the temporal variation of the spatial effect. This was only observed in the litter, probably because of the higher removal frequency in this substrate.6. Colonies of Pogonomyrmex spp. could enhance the heterogeneity of soil seed banks in the central Monte desert from the summer to the beginning of the autumn. Implications for vegetation dynamics depend on the degree to which seed density limits perennial grasses recruitment after ant activity season.

Allergen of the Month—Harvester Ant

The genus Pogonomyrmex of harvester ants contains over seventy species found in arid regions of North and South America. The genus name means “bearded ant” and derives from the Greek “pogon” or beard. Most species have a beard-like structure, the psammophore, which is useful for transporting seeds, sand pebbles, or eggs. The common name derives from the practice of sniping and gathering seeds, which are their primary source of nutrition, although the ants will also scavenge animal protein.1,2 Harvester ants are prevalent in the arid southwest United States, represented by the species P. maricopa, P. rugosus, P. barbatus, and P. californicus among others.

Forager abundance and its relationship with colony activity level in three species of South American Pogonomyrmex harvester ants

The proportion of foragers in ant colonies is a fairly constant species-specific characteristic that could be determined by intrinsic or extrinsic factors. If intrinsic factors are relevant, species with similar life history characteristics (e.g., colony size and foraging strategies) would be expected to have a similar proportion of foragers in their colonies. Within the genus Pogonomyrmex, North American species can vary largely in their colony size, whereas only species with small colonies are known in South America. We studied the characteristics of the foraging subcaste in three sympatric South American species of Pogonomyrmex harvester ants, and compared it with the available information on other species of the same genus. We used two mark-recapture methods and colony excavations to estimate the number and proportion of foragers in the colonies of P. mendozanus, P. inermis, and P. rastratus, and to test the relationship between forager external activity levels and abundance per colony. Forager abundance in the three studied species was lower than in most North American species. The percentage of foragers in their colonies ranged 7–15 %, more similar to North American species with large colonies than to those with small colony size. Foraging activity was positively correlated with forager abundance in all three species, implying that colony allocation to number of foragers allows for higher food acquisition. Further comparative studies involving a wider range of traits in South and North American species would allow to unveil the role of environmental factors in shaping each species’ particular traits.

Quantifying the effect of colony size and food distribution on harvester ant foraging.

Desert seed-harvester ants, genus Pogonomyrmex, are central place foragers that search for resources collectively. We quantify how seed harvesters exploit the spatial distribution of seeds to improve their rate of seed collection. We find that foraging rates are significantly influenced by the clumpiness of experimental seed baits. Colonies collected seeds from larger piles faster than randomly distributed seeds. We developed a method to compare foraging rates on clumped versus random seeds across three Pogonomyrmex species that differ substantially in forager population size. The increase in foraging rate when food was clumped in larger piles was indistinguishable across the three species, suggesting that species with larger colonies are no better than species with smaller colonies at collecting clumped seeds. These findings contradict the theoretical expectation that larger groups are more efficient at exploiting clumped resources, thus contributing to our understanding of the importance of the spatial distribution of food sources and colony size for communication and organization in social insects.

Notes on the North American harvesting ants of the genus Pogonomyrmex Mayr.

Notes on the North American harvesting ants of the genus Pogonomyrmex Mayr.

Ant-nest ichnofossils in honeycomb calcretes, Neogene Ogallala Formation, High Plains region of western Kansas, U.S.A.

Two new ant-nest trace fossils are described from calcic sandy paleosols of the Neogene Ogallala Formation in western Kansas. The ichnofossils are preserved within and below calcrete beds weathering in positive relief as carbonate-filled casts or as cavities in negative relief. Daimoniobarax ichnogenus nov. is established for burrow systems composed of vertically tiered, horizontally oriented pancake-shaped chambers connected by predominantly vertical and cylindrical shafts ~ 0.8 cm in diameter. Ichnospecies of Daimoniobarax are differentiated based on differences in the plan view outline of chambers, shaft orientation, and junctions between chambers and shafts. Daimoniobarax nephroides ichnospecies nov. is composed of an ~ 24–76 cm long vertical sequence of distinctly lobed chambers (~ 2–20 cm wide and ~ 1 cm high) arranged along sinuous to helical shafts. Chamber shape in plan view ranges from small teardrops to larger kidney- and U-shaped forms. Shafts intersect at chamber edges such that chambers appear to bud from the central shafts. Daimoniobarax nephroides is most similar to the nests of extant seed-harvester ants of the New World genus Pogonomyrmex. Such ants are specialized granivores and prefer sandy soils in arid to semi-arid grassland and desert regions. Daimoniobarax tschinkeli ichnospecies nov. is ~ 30–80 cm in vertical extent. Chambers (~ 2–30 cm wide and ~ 1 cm high) are circular to elongate or pseudopodial in plan view. Vertical shafts are straight to slightly sinuous and intersect most often toward the center of the chambers. The generalized architecture of D. tschinkeli is similar to that of the nests or nest portions of several extant ant genera, though it does not closely resemble any known modern nest. Ant ichnofossils provide valuable information on hidden biodiversity, paleohydrologic regimes, paleopedogenic processes, and paleoclimate during the time of nest occupation. Depth-related changes in chamber size and vertical spacing may also help interpret paleosurfaces and paleodepths, and serve as geopetal features.

Independent colony founding by ergatoid queens in the ant genus Pogonomyrmex: queen foraging provides an alternative to dependent colony founding

Ant queens exhibit two primary strategies to initiate nests, independent colony founding (ICF) by solitary queens and dependent colony founding (DCF) when the queen starts a nest with a group of workers that disperse on foot from the parent nest. Numerous ant species have wingless (ergatoid) queens, and it is generally assumed that these species exhibit obligate DCF because their lack of wing musculature provides them with few resources to divert towards producing their first brood of workers. Thus, ICF by ergatoid queens is viewed as maladaptive because these queens need to take additional dangerous foraging trips to garner sufficient food to rear their first brood of workers. Contrary to this prediction, I document ICF by ergatoid queens for three species of harvester ants in the genus Pogonomyrmex (subfamily Myrmicinae), P. cunicularius cunicularius, P. cunicularius pencosensis, and P. huachucanus. Queens of P. huachucanus were obligate foragers, i.e., no minim workers could be produced without external food, and one queen of P. cunicularius pencosensis was observed foraging in the field. Abundant and/or predictable food resources likely select for the evolution of semi-claustral nest founding and ICF by these ergatoid queens. Under these conditions, foraging time would be minimized and the number and size of minim workers would be maximized. These benefits should increase founding success, which could compensate for loss of long-range dispersal. Overall, this study demonstrates that care should be taken before concluding that ant colonies employ DCF based solely on queen morphology.

Shift in frequency-dependent selection across the life-cycle in obligately interbreeding harvester ant lineages

The long-term persistence of hybridogenetic systems, in which one genetic lineage is dependent on another as a sperm host, is paradoxical because they require continual coexistence between ecological competitors. Several species of ants display a social form of hybridogenesis, in which two distinct genetic lineages obligately interbreed to produce sterile workers, while intra-lineage progeny become the reproductives. A recent model suggests that persistence of such a system depends on the relative strengths of negative frequency-dependent selection acting during colony growth and positive frequency-dependent selection during reproduction. We used path analysis to investigate the effect of lineage frequency on reproductive output and colony sex ratio over a single reproductive season in a natural population of the H1/H2 lineage pair in the genus Pogonomyrmex. Results suggest that lineage frequency does impact reproduction via two opposing routes: the more common lineage procures more same-lineage mates, resulting in a higher proportion of same-lineage mates increases reproduction through higher queen-egg availability, but the rarer lineage procures more alternate-lineage mates, yielding a more genetically diverse worker caste that increases productivity. As a result, lineage frequency has only a weakly positive overall effect on colony reproduction, and overall frequency-dependence in this system across the entire life cycle is likely to be negative as predicted by the persistence model.

Horned lizards (Phrynosoma) incapacitate dangerous ant prey with mucus

Horned lizards (Iguanidae, Phrynosomatinae, Phrynosoma) are morphologically specialized reptiles characterized by squat, tank-like bodies, short limbs, blunt snouts, spines and cranial horns, among other traits. They are unusual among lizards in the degree to which they specialize on a diet of ants, but exceptional in the number of pugnacious, highly venomous, stinging ants they consume, especially harvester ants (genus Pogonomyrmex). Like other iguanian lizards, they capture insect prey on the tongue, but unlike other lizards, they neither bite nor chew dangerous prey before swallowing. Instead, they employ a unique kinematic pattern in which prey capture, transport and swallowing are combined. Nevertheless, horned lizards consume dozens of harvester ants without harm. We show that their derived feeding kinematics are associated with unique, mucus-secreting pharyngeal papillae that apparently serve to immobilize and incapacitate dangerous ants as they are swallowed by compacting them and binding them in mucus strands. Radially branched esophageal folds provide additional mucus-secreting surfaces the ants pass through as they are swallowed. Ants extracted from fresh-killed horned lizard stomachs are curled ventrally into balls and bound in mucus. We conclude that the pharyngeal papillae, in association with a unique form of hyolingual prey transport and swallowing, are horned lizard adaptations related to a diet of dangerous prey. Harvester ant defensive weapons, along with horned lizard adaptations against such weapons, suggest a long-term, predator-prey, co-evolutionary arms race between Phrynosoma and Pogonomyrmex.

Maternal Effect on Female Caste Determination in a Social Insect

Caste differentiation and division of labor are the hallmarks of social insect colonies [1, 2]. The current dogma for female caste differentiation is that female eggs are totipotent, with morphological and physiological differences between queens and workers stemming from a developmental switch during the larval stage controlled by nutritional and other environmental factors (e.g., [3-8]). In this study, we tested whether maternal effects influence caste differentiation in Pogonomyrmex harvester ants. By conducting crossfostering experiments we identified two key factors in the process of caste determination. New queens were produced only from eggs laid by queens exposed to cold. Moreover, there was a strong age effect, with development into queens occurring only in eggs laid by queens that were at least two years old. Biochemical analyses further revealed that the level of ecdysteroids was significantly lower in eggs developing into queens than workers. By contrast, we found no significant effect of colony size or worker exposure to cold, suggesting that the trigger for caste differentiation may be independent of the quantity and quality of resources provided to larvae. Altogether these data demonstrate that the developmental fate of female brood is strongly influenced by maternal effects in ants of the genus Pogonomyrmex.