Vertical organization of the division of labor within nests of the Florida harvester ant, Pogonomyrmex badius.

In the Florida harvester ant, Pogonomyrmex badius, foragers occur only in the top 15 cm of the nest, whereas brood and brood-care workers reside mostly in the deepest regions, yet the food and seeds foragers collect must be transported downward 30 to 80 cm to seed chambers and up to 2 m to brood chambers. Using mark-recapture techniques with fluorescent printer's ink, we identified a class of workers that ranges widely within the vertical structure of the nest, rapidly moving materials dropped by foragers in the upper regions downward, and excavated soil from deeper upward. Within the nest, only 5% of foragers were recovered below 20 cm depth, but about 30% of transfer workers and 82% of unmarked workers were found there. Below 70 cm depth, 90% of workers were unmarked, and were probably involved mostly in brood care. During the summer, the transfer workers comprise about a quarter of the nest population, while foragers make up about 40%. Workers marked as transfer workers later appear as foragers, while those marked as foragers die and disappear from the foraging population, suggesting that transfer workers are younger, and age into foraging. The importance of these findings for laboratory studies of division of labor are discussed. The efficient allocation of labor is a key component of superorganismal fitness.

An illustrated guide to seeds found in nests of the Florida harvester ant, Pogonomyrmex badius.

The Florida harvester ant, Pogonomyrmex badius collects the seeds of many plant species and stores them in underground nest chambers for later consumption. Seeds taken from multiple nests in 1989, 2014 and 2015 were separated by size and species and identified through published keys, comparison with herbarium specimens and with identified seed collections. Harvester ants stored at least 58 species of seeds from 20 plant families in their chambers. This paper presents images of each seed species in several aspects, their relative abundance in P. badius nests, their size relative to the smallest, and links to online data and images of the parent plant species, as well as to herbarium specimens. A number of seeds and plant families present at the site were not found in ant nests. These data and images will be valuable for future studies and experiments to untangle the choices the ants make in relation to what the plants and the seasons offer them.

The Florida Harvester Ant, Pogonomyrmex badius, Relies on Germination to Consume Large Seeds.

The Florida harvester ant, Pogonomyrmex badius, is one of many ant species and genera that stores large numbers of seeds in damp, underground chambers for later consumption. A comparison of the sizes of seeds recovered from storage chambers with those of seed husks discarded following consumption revealed that the used seeds are far smaller than stored seeds. This difference in use-rate was confirmed in field and laboratory colonies by offering marked seeds of various sizes and monitoring the appearance of size-specific chaff. Because foragers collect a range of seed sizes but only open small seeds, large seeds accumulate, forming 70% or more of the weight of seed stores. Major workers increase the rates at which small and medium seeds are opened, but do not increase the size range of opened seeds. Experiments limiting ant access to portions of natural seed chambers showed that seeds germinate during storage, but that the ants rapidly remove them. When offered alongside non germinating seeds, germinating seeds were preferentially fed to larvae. The rate of germination during the annual cycle was determined by both burial in artificial chambers at various depths and under four laboratory temperatures. The germination rate depends upon the species of seed, the soil/laboratory temperature and/or the elapsed time. The seasonal soil temperature cycle generated germination patterns that vary with the mix of locally-available seeds. Taken together, exploitation of germination greatly increases the resources available to the ants in space and time. While the largest seeds may have the nutritional value of 15 small seeds, the inability of workers to open large seeds at will precludes them from rapid use during catastrophic events. The harvester ant's approach to seed harvesting is therefore two-pronged, with both immediate and delayed payoffs arising from the tendency to forage for a wide variety of seeds sizes.

Bioturbation by the Fungus-Gardening Ant, Trachymyrmex septentrionalis.

Soil invertebrates such as ants are thought to be important manipulators of soils in temperate and tropical ecosystems. The fungus gardening ant, Trachymyrmex septentrionalis, is an important agent of biomantling, that is, of depositing soil excavated from below onto the surface, and has been suggested as an agent of bioturbation (moving soil below ground) as well. The amount of bioturbation by this ant was quantified by planting queenright colonies in sand columns consisting of 5 layers of different colored sand. The amount of each color of sand deposited on the surface was determined from April to November 2015. In November, colonies were excavated and the color and amount of sand deposited below ground (mostly as backfill in chambers) was determined. Extrapolated to one ha, T. septentrionalis deposited 800 kg of sand per annum on the surface, and an additional 200 kg (17% of the total excavated) below ground. On average, this mixes 1.3% of the sand from other layers within the top meter of soil per millennium, but this mixing is unlikely to be homogeneous, and probably occurs as "hotspots" in both horizontal and vertical space. Such mixing is discussed as a challenge to sediment dating by optically stimulated luminescence (OSL).

Experimental evidence that dispersal drives ant community assembly in human-altered ecosystems.

A key shortcoming in our understanding of exotic species' success is that it is not known how post-introduction dispersal contributes to the success of exotic species and the reassembly of invaded communities. Exotic and native species face poorly understood competition-colonization trade-offs in heterogeneous landscapes of natural and anthropogenic habitats. We conducted three experiments that tested how ant queen behavior during dispersal affects community composition. Using experimental plots, we tested whether (1) different types of habitat disturbance and (2) different sizes of habitat disturbance affected the abundance of newly mated queens landing in the plots. The three most abundant species captured were the exotic fire ant Solenopsis invicta, and the native species Brachymyrmex depilis, and S. pergandei, respectively. When queens were considered collectively, more queens landed in plowed, sand-added, and roadside plots than in control or mow plots, in other words, in the more heavily disturbed plots. We also tested (3) the effect of habitat manipulations on the survival of newly mated fire ant queens (Solenopsis invicta). Soil disturbance (tilling), lack of shade, and removal (poisoning) of the ant community resulted in the greatest fire ant colony survivorship. Collectively, experiments revealed that both exotic and native newly mated ant queens select open, human-altered ecosystems for founding new colonies. The selection of such habitats by fire ant queens leads to their successful colony founding and ultimately to their dominance in those habitats. Selection of disturbed habitats is therefore advantageous for exotic species but is an ecological trap for native species because they do not often succeed in founding colonies in these habitats.

Experiments Testing the Causes of Namibian Fairy Circles.

The grasslands on the sandy soils of the eastern edge of the Namib Desert of Namibia are strikingly punctuated by millions of mostly regularly-spaced circular bare spots 2 to 10 m or more in diameter, generally with a margin of taller grasses. The causes of these so called fairy circles are unknown, but several hypotheses have been advanced. In October 2009, we set up experiments that specifically tested four hypothesized causes, and monitored these 5 times between 2009 and 2015. Grass exclusion in circles due to seepage of subterranean vapors or gases was tested by burying an impermeable barrier beneath fairy circles, but seedling density and growth did not differ from barrier-less controls. Plant germination and growth inhibition by allelochemicals or nutrient deficiencies in fairy circle soils were tested by transferring fairy circle soil to artificially cleared circles in the grassy matrix, and matrix soil to fairy circles (along with circle to circle and matrix to matrix controls). None of the transfers changed the seedling density and growth from the control reference conditions. Limitation of plant growth due to micronutrient depletion within fairy circles was tested by supplementing circles with a micronutrient mixture, but did not result in differences in plant seedling density and growth. Short-range vegetation competitive feedbacks were tested by creating artificially-cleared circles of 2 or 4 m diameter located 2 or 6 m from a natural fairy circle. The natural circles remained bare and the artificial circles revegetated. These four experiments provided evidence that fairy circles were not caused by subterranean vapors, that fairy circle soil per se did not inhibit plant growth, and that the circles were not caused by micronutrient deficiency. There was also no evidence that vegetative feedbacks affected fairy circles on a 2 to 10 m scale. Landscape-scale vegetative self-organization is discussed as a more likely cause of fairy circles.

Sequential Subterranean Transport of Excavated Sand and Foraged Seeds in Nests of the Harvester Ant, Pogonomyrmex badius.

During their approximately annual nest relocations, Florida harvester ants (Pogonomyrmex badius) excavate large and architecturally-distinct subterranean nests. Aspects of this process were studied by planting a harvester ant colony in the field in a soil column composed of layers of 12 different colors of sand. Quantifying the colors of excavated sand dumped on the surface by the ants revealed the progress of nest deepening to 2 m and enlargement to 8 L in volume. Most of the excavation was completed within about 2 weeks, but the nest was doubled in volume after a winter lull. After 7 months, we excavated the nest and mapped its structure, revealing colored sand deposited in non-host colored layers, especially in the upper 30 to 40 cm of the nest. In all, about 2.5% of the excavated sediment was deposited below ground, a fact of importance to sediment dating by optically-stimulated luminescence (OSL). Upward transport of excavated sand is carried out in stages, probably by different groups of ants, through deposition, re-transport, incorporation into the nest walls and floors and remobilization from these. This results in considerable mixing of sand from different depths, as indicated in the multiple sand colors even within single sand pellets brought to the surface. Just as sand is transported upward by stages, incoming seeds are transported downward to seed chambers. Foragers collect seeds and deposit them only in the topmost nest chambers from which a separate group of workers rapidly transports them downward in increments detectable as a "wave" of seeds that eventually ends in the seed chambers, 20 to 80 cm below the surface. The upward and downward transport is an example of task-partitioning in a series-parallel organization of work carried out by a highly redundant work force in which each worker usually completes only part of a multi-step process.

Biomantling and bioturbation by colonies of the Florida harvester ant, Pogonomyrmex badius.

In much of the world, soil-nesting ants are among the leading agents of biomantling and bioturbation, depositing excavated soil on the surface or in underground chambers. Colonies of the Florida harvester ant, Pogonomyrmex badius excavate a new nest once a year on average, depositing 0.1 to 12 L (3 L average) of soil on the surface. Repeated surveys of a population of about 400 colonies yielded the frequency of moves (approximately once per year), the distance moved (mean 4 m), and the direction moved (random). The area of the soil disc correlated well with the volume and maximum depth of the nest, as determined by excavation and mapping of chambers. The population-wide frequency distribution of disc areas thus yielded the frequency distribution of nest volumes and maximum depths. For each surveyed colony, the volume of soil excavated from six specified depth ranges and deposited on the surface was estimated. These parameters were used in a simulation to estimate the amount of soil mantled over time by the observed population of P. badius colonies. Spread evenly, P. badius mantling would create a soil layer averaging 0.43 cm thick in a millennium, with 10-15% of the soil deriving from depths greater than 1 m. Biomantling by P. badius is discussed in the context of the ant community of which it is a part, and in relation to literature reports of ant biomantling.

Nest relocation and excavation in the Florida harvester ant, Pogonomyrmex badius.

The Florida harvester ant (Pogonomyrmex badius) excavates deep nests in the sandy soils of the Gulf and Atlantic coastal plains. Nest relocations of over 400 colonies in a north Florida coastal plains pine forest were tracked and mapped from 2010 to 2013. Individual colonies varied from one move in two years to four times a year, averaging about one per year. Almost all moves occurred between May and November peaking in July when more than 1% of the colonies moved per day. Move directions were random, and averaged 4 m, with few moves exceeding 10 m. Distance moved was not related to colony size. Over multiple moves, paths were random walks around the original nest location. Relocation is probably intrinsic to the life history of this species, and the causes of relocation remain obscure--the architecture of old and new nests was very similar, and neither the forest canopy nor the density or size of neighbors was correlated with relocation. Monitoring entire relocations (n = 20) showed that they were usually completed in 4 to 6 days. Moves were diurnal, peaking in the mornings and afternoons dipping during mid-day, and ceasing before sundown. Workers excavated the new nest continuously during the daytime throughout the move and beyond. A minority of workers carried seeds, charcoal and brood, with seeds being by far the most common burden. The proportion of burdened workers increased throughout the move. Measured from year to year, small colonies gained size and large ones lost it. Colonies moving more than once in two years lost more size than those moving less often, suggesting that moving may bear a fitness cost. Colony relocation is a dramatic and consistent feature of the life history of the Florida harvester ant, inviting inquiry into its proximal and ultimate causes.

The morphometry of solenopsis fire ants.

Size-related changes of body shape were explored in 15 polymorphic species of Solenopsis fire ants by analyzing body weight along with linear measurements of 24 body parts. Log regression slopes were used to detect changes of shape with increasing size. Within species, the largest workers weighed from about 5 to 30-fold as much as the smallest. The range of within-species body lengths varied from 1.6 mm to 4 mm. As worker size increased, the gaster tended to make up a larger proportion of body length, usually at the cost of the petiole, and rarely at the cost of head length or mesosoma length. In most, the relative volume of the gaster increased and that of the head and mesosoma decreased. Most also showed an increasingly "humped" mesosoma. For all species, head shape changed from barrel-shaped to heart-shaped as worker size increased. Antennae became relatively shorter as the relative size of the club decreased. Shape changes of the legs were more variable. S. geminata was exceptional in the extreme nature of its head shape change, and was the only species in which relative head volume increased and gaster volume decreased with increasing body size. With the exception of S. geminata, the allometric rules governing shape are remarkably similar across species, suggesting a genus-level developmental scheme that is not easily modified by evolution. It also suggests that the evolution of shape is highly constrained by these conserved growth rules, and that it acts primarily (perhaps only) through allometric growth. The results are discussed in light of the growth of imaginal discs in a resource-limited body (the pupa). The substantial variation of allometries within species and across localities is also discussed in relation to using allometric patterns to identify species or to construct phylogenies.

The role of habitat in the persistence of fire ant populations.

The association of the exotic fire ant, Solenopsis invicta with man-modified habitats has been amply demonstrated, but the fate of such populations if ecological succession proceeds has rarely been investigated. Resurvey of a fire ant population in a longleaf pine plantation after 25 years showed that the recovery of the site from habitat disturbance was associated with a large fire ant population decline. Most of the persisting colonies were associated with the disturbance caused by vehicle tracks. In a second study, mature monogyne fire ant colonies that had been planted in experimental plots in native groundcover of the north Florida longleaf pine forest had mostly vanished six years later. These observations and experiments show that S. invicta colonies rarely persist in the native habitat of these pine forests, probably because they are not replaced when they die. A single site harbored a modest population of polygyne fire ants whose persistence was probably facilitated by reproduction through colony fission.

Florida harvester ant nest architecture, nest relocation and soil carbon dioxide gradients.

Colonies of the Florida harvester ant, Pogonomyrmex badius, excavate species-typical subterranean nests up the 3 m deep with characteristic vertical distribution of chamber area/shape, spacing between levels and vertical arrangement of the ants by age and brood stage. Colonies excavate and occupy a new nest about once a year, and doing so requires that they have information about the depth below ground. Careful excavation and mapping of vacated and new nests revealed that there was no significant difference between the old and new nests in any measure of nest size, shape or arrangement. Colonies essentially built a replicate of the just-vacated nest (although details differed), and they did so in less than a week. The reason for nest relocation is not apparent. Tschinkel noted that the vertical distribution of chamber area, worker age and brood type was strongly correlated to the soil carbon dioxide gradient, and proposed that this gradient serves as a template for nest excavation and vertical distribution. To test this hypothesis, the carbon dioxide gradient of colonies that were just beginning to excavate a new nest was eliminated by boring 6 vent holes around the forming nest, allowing the soil CO2 to diffuse into the atmosphere and eliminating the gradient. Sadly, neither the nest architecture nor the vertical ant distribution of vented nests differed from either unvented control or from their own vacated nest. In a stronger test, workers excavated a new nest under a reversed carbon dioxide gradient (high concentration near the surface, low below). Even under these conditions, the new and old nests did not differ significantly, showing that the soil carbon dioxide gradient does not serve as a template for nest construction or vertical worker distribution. The possible importance of soil CO2 gradients for soil-dwelling animals is discussed.

The life cycle and life span of Namibian fairy circles.

In Namibia of southwestern Africa, the sparse grasslands that develop on deep sandy soils under rainfall between 50 and 100 mm per annum are punctuated by thousands of quasi-circular bare spots, usually surrounded by a ring of taller grass. The causes of these so-called "fairy circles" are unknown, although a number of hypotheses have been proposed. This paper provides a more complete description of the variation in size, density and attributes of fairy circles in a range of soil types and situations. Circles are not permanent; their vegetative and physical attributes allow them to be arranged into a life history sequence in which circles appear (birth), develop (mature) and become revegetated (die). Occasionally, they also enlarge. The appearance and disappearance of circles was confirmed from satellite images taken 4 years apart (2004, 2008). The frequency of births and deaths as a fraction of the total population of circles allowed the calculation of an approximate turnover rate, and from this, an estimate of circle lifespan. Lifespan appeared to vary with circle size, with small circles averaging about 24 years, and larger ones 43-75 years. Overall lifespan averaged about 41 yr. A second, independent estimate of lifespan was made by revisiting circles 2 to 9 years after their clear status had been confirmed. This resulted in a lifespan estimate of about 60 years. Any causal explanation of fairy circles must include their birth, development and death, their mean lifespan and the variation of their features under different conditions.

Ant distribution in relation to ground water in north Florida pine flatwoods.

Longleaf pine savannas are one of the most threatened ecosystems in the world, yet are understudied. Ants are a functionally important and diverse group of insects in these ecosystems. It is largely unknown how local patterns of species diversity and composition are determined through the interaction of this dominant animal group with abiotic features of longleaf pine ecosystems. Here we describe how an important abiotic variable, depth to water table, relates to ant species distributions at local scales. Pitfall trapping studies across habitat gradients in the Florida coastal plains longleaf pine flatwoods showed that the ant community changed with mild differences in habitat. In this undulating landscape, elevation differences were less than 2 m, and the depth to the water table ranged from < 20 cm to 1.2 m. The plant species composing the ground cover were zoned in response to depth to water, and shading by canopy trees increased over deeper water tables. Of the 27 ant species that were analyzed, depending on the statistical test, seven or eight were significantly more abundant over a deep water table, eight to ten over a shallow one, and nine to eleven were not significantly patterned with respect to depth to water. Ant species preferring sites with shallow groundwater also preferred the shadier parts of the sites, while those preferring sites with deeper groundwater preferred the sunnier parts of the sites. This suggests that one group of species prefers hot-dry conditions, and the other cooler-moist. Factor analysis and abundance-weighted mean site characteristics generally confirmed these results. These results show that ant communities in this region respond to subtle differences in habitat, but whether these differences arise from founding preferences, survival, competition, or some combination of these is not known.

The organization of foraging in the fire ant, Solenopsis invicta.

Although natural selection in ants acts most strongly at the colony, or superorganismal level, foraging patterns have rarely been studied at that level, focusing instead on the behavior of individual foragers or groups of foragers. The experiments and observations in this paper reveal in broad strokes how colonies of the fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), allocate their available labor to foraging, how they disperse that force within their territory, and how this force changes with colony size, season and worker age. Territory area is positively related to colony size and the number of foragers, more so during the spring than fall. Changes of colony size and territory area are driven by seasonal variation of sexual and worker production, which in turn drive seasonal variation of worker age-distribution. During spring sexual production, colonies shrink because worker production falls below replacement. This loss is proportional to colony size, causing forager density in the spring to be negatively related to colony and territory size. In the fall, colonies emphasize worker production, bringing colony size back up. However, because smaller colonies curtailed spring worker production less than larger ones, their fall forager populations are proportionally greater, causing them to gain territory at the expense of large colonies. Much variation of territory area remains unexplained and can probably be attributed to pressure from neighboring colonies. Boundaries between territories are characterized by "no ants' zones" mostly devoid of fire ants. The forager population can be divided into a younger group of recruitable workers that wait for scouts to activate them to help retrieve large food finds. About one-third of the recruits wait near openings in the foraging tunnels that underlie the entire territory, while two-thirds wait in the nest. Recruitment to food is initially very rapid and local from the foraging tunnels, while sustained recruitment gradually involves the recruits waiting in the nest. As recruits age, they become scouts searching for food on the surface, and die about two weeks later. Foraging tunnels decrease in cross-sectional area with distance from the nest, in keeping with the gradual bleeding off of workers to the surface with distance. Foragers lack route-faithfulness, and having been marked and released at one point within the territory, they can be recaptured at any other point a day later. The size of the territory actually occupied may be limited during dry weather, resulting in very large no-ants' zones.

The nest architecture of three species of north Florida Aphaenogaster ants.

The architecture of the subterranean nests of Aphaenogaster floridana Smith (Hymenoptera: Formicidae), A. treatae Forel and A. ashmeadi (Emery), was studied from plaster, wax, or metal casts. After structural features were quantified from digital images, the entombed ants were retrieved from the plaster by dissolution or wax casts by melting and counted. Nests of all three species were rather simple, small and vertical, with horizontal chambers connected by vertical shafts. Shafts descending to lower chambers tended to arise from chamber edges, whereas those connecting to a chamber above tended to arise from chamber centers. A. floridana had the largest nests and colonies, and multiple shafts commonly connected upper chambers, a feature lacking in the other two species. In A. floridana nests a higher proportion of chamber area and greater spacing between chambers occurred in the deeper parts of the nest, regardless of nest size. The other two species showed no vertical differentiation of any size-free measure at any nest size. In all three species, nest size increased more slowly than the worker population, so crowding was greater in large colonies than in small, in contrast to the situation in three other ant species for which data were available. An appendix with stereo images of all casts is provided.

Early climate change consensus at the National Academy: the origins and making of "Changing Climate".

The 1983 National Academy of Sciences report entitled "Changing Climate," authored by a committee of physical and social scientists chaired by William Nierenberg, was an early comprehensive review of the effects of human-caused increases in the levels of atmospheric CO2. Study of the events surrounding the committee's creation, deliberations, and subsequent report demonstrates that the conclusions of the report were the consensus of the entire committee and in line with the scientific consensus of the time. This result contraverts a 2008 paper in which Naomi Oreskes, Erik M. Conway, and Matthew Shindell asserted that the report contradicted a growing consensus about climate change, and that Nierenberg for political reasons deliberately altered the summary and conclusions of the report in a way that played down the concerns of the other physical scientists on the committee. Examining the production of the report and contextualizing it in contemporaneous scientific and political discussion, we instead show how it was a multi-year effort with work divided among the various members of the committee according to their expertise. The synthesis and conclusions were expressly a joint statement of the committee and were consistent with other assessments of that time expressing deep concern over the potential issues while stopping short of recommending major policy changes due to the uncertainties, and to a lack of good alternatives.

The nest architecture of the ant Odontomachus brunneus.

The architecture of the subterranean nests of the ant Odontomachus brunneus (Patton) (Hymenoptera: Formicidae) was studied by means of casts with dental plaster or molten metal. The entombed ants were later recovered by dissolution of plaster casts in hot running water. O. brunneus excavates simple nests, each consisting of a single, vertical shaft connecting more or less horizontal, simple chambers. Nests contained between 11 and 177 workers, from 2 to 17 chambers, and 28 to 340 cm(2) of chamber floor space and reached a maximum depth of 18 to 184 cm. All components of nest size increased simultaneously during nest enlargement, number of chambers, mean chamber size, and nest depth, making the nest shape (proportions) relatively size-independent. Regardless of nest size, all nests had approximately 2 cm(2) of chamber floor space per worker. Chambers were closer together near the top and the bottom of the nest than in the middle, and total chamber area was greater near the bottom. Colonies occasionally incorporated cavities made by other animals into their nests.

The foraging tunnel system of the Namibian desert termite, Baucaliotermes hainesi.

The harvester termite, Baucaliotermes hainesi (Fuller) (Termitidae: Nasutitermitinae), is an endemic in southern Namibia, where it collects and eats dry grass. At the eastern, landward edge of the Namib Desert, the nests of these termites are sometimes visible above ground surface, and extend at least 60 cm below ground. The termites gain access to foraging areas through underground foraging tunnels that emanate from the nest. The looseness of the desert sand, combined with the hardness of the cemented sand tunnels allowed the use of a gasoline-powered blower and soft brushes to expose tunnels lying 5 to 15 cm below the surface. The tunnels form a complex system that radiates at least 10 to 15 m from the nest with cross-connections between major tunnels. At 50 to 75 cm intervals, the tunnels are connected to the surface by vertical risers that can be opened to gain foraging access to the surrounding area. Foraging termites rarely need to travel more than a meter on the ground surface. The tunnels swoop up and down forming high points at riser locations, and they have a complex architecture. In the center runs a smooth, raised walkway along which termites travel, and along the sides lie pockets that act as depots where foragers deposit grass pieces harvested from the surface. Presumably, these pieces are transported to the nest by a second group of termites. There are also several structures that seem to act as vertical highways to greater depths, possibly even to moist soil. A census of a single nest revealed about 45,000 termites, of which 71% were workers, 9% soldiers and 6% neotenic supplementary reproductives. The nest consisted of a hard outer "carapace" of cemented sand, with a central living space of smooth, sweeping arches and surfaces. A second species of termite, Promirotermes sp. nested in the outer carapace.