Habitat complexity facilitates coexistence in a tropical ant community

The role of habitat complexity in the coexistence of ant species is poorly understood. Here, we examine the influence of habitat complexity on coexistence patterns in ant communities of the remote Pacific atoll of Tokelau. The invasive yellow crazy ant, Anoplolepis gracilipes (Smith), exists in high densities on Tokelau, but still coexists with up to seven other epigeic ant species. The size-grain hypothesis (SGH) proposes that as the size of terrestrial walking organisms decreases, the perceived complexity of the environment increases and predicts that: (1) leg length increases allometrically with body size in ants, and (2) coexistence between ant species is facilitated by differential habitat use according to body size. Analysis of morphological variables revealed variation inconsistent with the morphological prediction of the SGH, as leg length increased allometrically with head length only. We also experimentally tested the ability of epigeic ants in the field to discover and dominate food resources in treatments of differing rugosity. A. gracilipes was consistently the first to discover food baits in low rugosity treatments, while smaller ant species were consistently the first to discover food baits in high rugosity treatments. In addition, A. gracilipes dominated food baits in planar treatments, while smaller ant species dominated baits in rugose treatments. We found that the normally predictable outcomes of exploitative competition between A. gracilipes and other ant species were reversed in the high rugosity treatments. Our results support the hypothesis that differential habitat use according to body size provides a mechanism for coexistence with the yellow crazy ant in Tokelau. The SGH may provide a mechanism for coexistence in other ant communities but also in communities of other terrestrial, walking insects that inhabit a complex landscape.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

The effects of food presentation and microhabitat upon resource monopoly in a ground-foraging ant (Hymenoptera: Formicidae) community

In Neotropical wet forests several species of omnivorous, resource-defending ants, live and forage in close proximity to one another. Although the forest floor is heterogeneous in microhabitat and food quantity, little is known about the impact of microhabitat and food variation upon resource monopoly among ants. We investigated how food type and microhabitat influence food monopoly in resource-defending ants in old-growth tropical wet forest in the Caribbean lowlands of Costa Rica. We measured several microhabitat characteristics at 66 points in a 0.5 hectare plot, and baited each point with two categories of tuna bait. These baits were presented in "split" and "clumped" arrangements. We measured the frequency of bait monopoly by a single species, as well as the number of recruited ant foragers at a bait. Out of five common species, two (Wasmannia auropunctata and Pheidole simonsi) more frequently monopolized one bait type over the other, and one (P. simonsi) recruited more ants to the split baits. We then considered the recruitment response by all ant species in the community. We found that the frequency of monopoly, sharing, and the absence of ants at a given point in the rainforest differed with bait type. The frequency of monopoly was associated with microhabitat type in two out of eight microhabitat variables (leaf litter depth and palms); variation in two other types (canopy tree distance and leafcutter ant trails) was associated with changes in forager number. In at least two ant species, food presentation affected monopoly at baits; among all resource-defending ants, the microhabitats where ants foraged for food and the type of food located determined in part the frequency of monopoly and the number of foragers at the food item. These results suggest that the location and presentation of food items determines in part which ant species will utilize the resource.     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

Competition for food between the exotic wasp <Emphasis Type="Italic">Vespula germanica</Emphasis> and the native ant assemblage of NW Patagonia: evidence of biotic resistance?

The success of a biological invasion may depend on the interactions between the invader and the native biota. However, little experimental evidence demonstrates whether local species can successfully compete with exotics. We experimentally determined the existence of competition for food between the exotic wasp Vespula germanica, one of the most recent Patagonian invaders, and the native ant assemblage. Both wasps and ants are generalist predators and scavengers, sharing habitat and food resources. We selected 30 sites within scrubland habitats where both ants and wasps were present. At each site, we placed containers with protein baits under three treatments: wasp exclusion, ant exclusion, and control (i.e., free access for wasps and ants). Ant exclusion increased the number of wasps (with regard to a control), but wasp exclusion did not affect ant abundance. This result suggests that native ants affect the foraging activity of exotic wasps but not vice versa. Aggressive behaviors and worker aggregation may explain the competitive advantage of ants. Ants bite wasp legs and massively aggregate on food sources, physically limiting the landing of wasps on baits. If the outcome of interactions at baits reported here influence wasp population-level parameters, this competitive interaction could be one of the factors explaining the low abundance of this exotic wasp in NW Patagonia in comparison with other invaded regions.     

Litter Biomass and Nutrient Determinants of Ant Density, Nest Size, and Growth in a Costa Rican Tropical Wet Forest

The nutritional demands of animals vary by taxon. Across landscapes, communities of animals experience variability in the stoichiometry of carbon and nutrients within their resource base. Thus, we expect stoichiometry to contribute to the spatial variance in the demographic parameters of animal communities. Here, we measure how the composition of a litter-nesting tropical rainforest ant community is influenced by spatial variation in environmental stoichiometry relative to litter biomass, a known predictor of ant density. We found the density of ants and their nests were strongly related to litter biomass and carbon: phosphorus stoichiometry. The spatial variation in soil nutrients, which determines leaf litter stoichiometry, was an excellent predictor of nest size in the two most common genera of ants. We found a negative relationship between species' growth rate and local soil stocks of phosphorus. Overall, the density of litter-dwelling ants varied greatly across this tropical forest landscape and environmental stoichiometry can account for limits on ant density independent of the biomass of the leaf litter resource base.     

The size–grain hypothesis and interspecific scaling in ants

1. The size–grain hypothesis maintains that as terrestrial walking organisms decrease in size, their environment becomes less planar and more rugose. The benefits of long legs (efficient, speedy movement over a planar environment) may thus decrease with smaller body size, while the costs (larger cross-sectional area limiting access to the interstitial environment) are enhanced.
2. A prediction from this hypothesis – that leg size should increase proportionately with body mass – is examined. Ants are among the smallest walking animals and extend the size gradient five orders of magnitude beyond the traditional 'mouse to elephant' curve. The mass of 135 species of worker ants spans 3·7 orders of magnitude (0·008–53 mg). Larger ants tended to be slimmer and longer legged. Ant subfamilies varied in their scaling relationships, but four out of five showed a positive allometry for hind leg length ( b > 0·33). Mammals, in contrast, show isometry for leg length over six orders of magnitude.
3. It is suggested that ants make a transition from living in an interstitial environment when small to a planar environment when large, a habit continued by most terrestrial mammals. Head length and pronotum width are robust estimators of mass in ants.     

Soil surface complexity has a larger effect on food exploitation by ants than a change from grassland to shrubland

1. Changes in vegetation community composition, such as a transition from grassland to shrubland (woody encroachment), are associated with reductions in plant cover and increases in bare ground. Encroachment‐driven changes in surface cover at small spatial scales can alter ant community assemblages by changing their foraging behaviour and their ability to locate and monopolise resources. 2. Artificial arenas with three levels of complexity were used to examine changes in ant foraging efficiency, body size and ability to monopolise food. The three levels of complexity included a control (no substrate), low‐complexity treatment (woody debris) and high‐complexity treatment (leaf litter). 3. No difference was found in ant species composition within the complexity arenas between grassland and shrubland, but ant functional groups ‘generalised Myrmicinae’ and ‘subordinate Camponotini’ were more abundant in grassland arenas, whereas ‘opportunists’ were more abundant in shrubland arenas. Ants took twice as long to find baits in high‐complexity treatments, and 1.5 times as long in low‐complexity treatments, than in control treatments, which were bare arenas with no substrate. Ant body size declined with increasing surface complexity, suggesting that larger ants are discouraged from foraging in complex habitats. 4. There was also significantly greater monopolisation of the protein bait (tuna) in low‐ and high‐complexity treatments, but there were no differences between tuna and carbohydrate (honey) in the control treatment. Consistently, no differences were found in ant behaviour between grasslands and shrublands. 5. The present study shows that ants are more responsive to small‐scale alterations in soil surface complexity than to changes in vegetation community composition. Changes in soil surface complexity select for ants based on body size, which in turn influences their foraging success. Changes in vegetation complexity at small spatial scales are therefore likely to influence ant behaviour and abundance of some functional groups, potentially having an effect on the many ecosystem functions carried out by ants.     

The size-grain hypothesis: a phylogenetic and field test

Abstract.  1. The size-grain hypothesis predicts that environmental rugosity results in positive allometric scaling of leg length on body length because of changes in locomotion costs.
2. The scaling of leg length and body length in ants was re-examined using phylogenetic independent contrast methods, and the allometric relationship found by Kaspari and Weiser ( Functional Ecology , 13 , 530–538, 1999) was supported.
3. The size-grain hypothesis was tested further by comparing the body sizes of ants from areas of contrasting habitat complexity in two different savanna habitats. No support for the size-grain hypothesis was found. Small body size classes were no more speciose in the rugose than in the more planar environment, and small ants were more abundant in the planar environment.     

Detrimental effects of highly efficient interference competition: invasive Argentine ants outcompete native ants at toxic baits

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. Previous studies that examined the mechanisms by which Argentine ants attain ecological dominance showed that superior interference and exploitation competition are key to the successful displacement of native ant species. The objective of this research was to test the hypothesis that effective interference competition by Argentine ants may also be detrimental to the survival of Argentine ant colonies where Argentine ants and native ants compete at toxic baits used to slow the spread of Argentine ants. To study this hypothesis, we examined the competitive interactions between Argentine ants and native odorous house ants, Tapinoma sessile, in the presence and absence of toxic baits. Results showed that Argentine ants aggressively outcompete T. sessile from toxic baits through efficient interference competition and monopolize bait resources. This has severe negative consequences for the survival of Argentine ants as colonies succumb to the toxic effects of the bait. In turn, T. sessile avoid areas occupied by Argentine ants, give up baits, and consequently suffer minimal mortality. Our results provide experimental evidence that highly efficient interference competition may have negative consequences for Argentine ants in areas where toxic baits are used and may provide a basis for designing innovative management programs for Argentine ants. Such programs would have the double benefit of selectively eliminating the invasive species while simultaneously protecting native ants from the toxic effects of baits.     

Allometry between leg and body length of insects: lack of support for the size–grain hypothesis

• 1 The size–grain hypothesis ( Kaspari & Weiser, 1999 ) states that (1) as organisms decrease in size, they perceive their environment as being more rugose; (2) long legs allow organisms to step over obstacles but hinder them from entering small gaps; and (3) as the size of an organism decreases, the benefits of long legs begin to be outweighed by the costs of construction. Natural selection should therefore favour proportionally longer legs in larger organisms, thereby leading to a positive allometry between leg and body length (scaling exponent b > 1).
• 2 Here we compare the scaling exponent of leg‐to‐body length relationships among insects that walk, walk and fly, and predominantly fly. We measured the lengths of the hind tibia, hind femur, and body length of each species.
• 3 The taxa varied considerably in the scaling exponent b. In seven out of ten groups (Formicidae, Isoptera, Carabidae, Pentatomidae, Apidae, Lepidoptera, Odonata adult), b was significantly greater than one. However, there was no gradual decrease in b from walking to walking/flying to flying insects.
• 4 The results of the present study provide no support for the size–grain hypothesis. We propose that leg length is not only affected by the rugosity of the environment, but also by (1) functional adaptations, (2) phylogeny, (3) lifestyle, (4) the type of insect development (hemimetabolism or holometabolism), and (5) constraints of gas exchange.

     

Preference of food particle size among several urban ant species

Appropriate particle size may be a critical characteristic for effective granular ant baits. We examined the particle size preference of six species of pest ants to an anchovy-based bait. We also examined head capsule widths of Argentine ants, Linepithema humile (Mayr) (mean = 0.54 mm), California harvester ants, Pogonomyrmex californicus (Buckley) (mean = 1.63 mm), red imported fire ants, Solenopsis invicta Buren (mean = 0.9 mm), and southern fire ants, Solenopsis xyloni McCook (mean = 0.76 mm) and compared them with the first and second most preferred particle size. There were differences between particle size of which the most mass was removed and of which there were more particles removed by ants. California Argentine ants, southern fire ants, and Alabama Argentine ants removed more 840 to 1,000-microm particle mass of the anchovy diet but had more visits to dishes containing 420 to 590 microm particles. California harvester ants and Allegheny mound ants, Formica spp., removed more >2,000 microm particle mass but visited dishes containing 1,000 to 2,000 microm particles more often. Red imported fire ants also removed more >2,000 microm particle mass but visited dishes with 590 to 840-microm particles most often. Pharaoh ants, Monomorium pharaonis (L.), removed and visited 420 to 590-microm particles more than any other size. A linear regression model determined that particle size preferred by each ant species relates to forager head width. The majority of particles of commercial ant bait, including Amdro, Ascend, Award, Bushwhacker, Max Force with fipronil, and old and new formulations of Max Force with hydramethylnon, were 1,000 to 2,000 microm, but the majority of Niban particles were <420 microm. Altering the size of particles of toxic ant baits to fit the particle size preference of each pest ant species may increase the efficacy of ant baits.     

Community structure and the habitat templet: ants in the tropical forest canopy and litter

The tropical forest canopy and litter differ in physical structure, resource availability, and abiotic conditions. We used standardized bait experiments in the canopy and litter of four neotropical tree species to explore how these differences shape the behavior, morphology, and diversity of ant assemblages. Ant activity (biomass at a bait after 32 min) was higher in the canopy, and higher on protein baits than carbohydrate baits. Aggressive bait defense occurred more frequently in the canopy (60%) than in the litter (32%), but was not associated with tree species or bait type in either habitat. The median size of workers of species in the canopy and litter was nearly identical, but body size distribution was unimodal in the canopy and bimodal in the litter. The colony size of the most aggressive species was an order of magnitude larger in the canopy. Species richness at a bait was relatively uniform across tree species and habitats. Litter and canopy shared no species, but overlap among tree species was three times higher in the litter assemblages. Litter assemblages showed less activity, less interference, less differentiation across the landscape, and different size distributions than canopy assemblages. The canopy and litter templets subsume a number of environmental gradients that combine to shape ant community structure.     

Studies on the relationship between the ant ectoparasite Antennophorus grandis (Acarina: Antennophoridae) and its host Lasius flavus (Hymenoptera: Formicidae)

The mite Antennophorus grandis (Berlese, 1903) is a large obligate ectoparasite of workers of the ant Lasius flavus(Fabricius). It rides under the head of the ants and uses its long front legs to communicate with its hosts and stimulate them to give it food. We present allometric and morphometric data showing that L. flavus workers can occur in two size classes. We also present the first quantitative ethograms of mite-bearing and mite-free L. flavus workers of the two size categories. The mites tend to occur on the smaller nurse workers and receive food from them at an extremely high frequency. Antennophorus grandis also frequently gain food when one ant is donating food to the one they are riding upon. The mites seem to inhibit the ability of their host worker to show most social behaviours such as tending ant larvae. The mites frequently move from one host worker to another. For these reasons the mites may have a larger impact upon their host colony than their relative rarity first suggests. The ants do not seem to have any specific defence against these parasites. The mites live in small populations and show female-biased sex ratios consistent with local mate competition. Preliminary evidence suggests that the mites have haplodiploid sex determination which may explain how they are able to adapt their sex allocation to their population size.     

Arboreal substrates influence foraging in tropical ants

1. Physically complex substrates impart significant costs on cursorial central‐place foragers in terms of time spent outside the nest and total distance travelled. Ants foraging in trees navigate varied surfaces to access patchy resources, thus providing an appropriate model system for examining interactions between foraging efficiency and substrates. 2. We expected that the speed of recruitment, body size distribution and species richness of foraging arboreal ants would differ predictably among common substrate types occurring on tropical tree trunks. We measured changes in ant abundance and species composition over time at baits placed on bare tree bark, moss‐covered bark, and vine‐like vegetation appressed to bark. We also measured average body size and body size frequency on the three substrate types. Ants discovered baits sooner and accumulated at baits relatively faster when using vine substrates as the primary foraging trail. Average body size was smaller on vine substrates than on bark. Experimental removal of vine and moss substrates nullified these differences. Contrary to our predictions, species richness and body size distributions did not differ among the three substrate types, due in part to the frequent presence of a few common ground‐nesting species at baits on bare bark. 3. Our results collectively indicate that linear substrates facilitate access of foraging ants to patchy resources. Ant use of vine‐like substrates appears to be opportunistic; vine use is not confined to certain species nor constrained by body size.     

More food,less habitat: how necromass and leaf litter decomposition combine to regulate a litter ant community

1. In brown food webs of the forest floor, necromass (e.g. insect carcasses and frass) falling from the canopy feeds both microbes and ants, with the former decomposing the homes of the latter. In a tropical litter ant community, we added necromass to 1 m² plots, testing if it added as a net food (increasing ant colony growth and recruitment) or destroyer of habitat (by decomposing leaf litter). 2. Maximum, but not mean, colony growth rates were higher on +food plots. However, neither average colony size, nor density was higher on +food plots. In contrast, +food plots saw diminished availability of leaf litter and higher microbial decomposition of cellulose, a main component of the organic substrate that comprises litter habitat. 3. Furthermore, necromass acted as a limiting resource to the ant community only when nest sites were supplemented on +food plots in a second experiment. Many of these +food +nest plots were colonised by the weedy species Wasmannia auropunctata. 4. Combined, these results support the more food–less habitat hypothesis and highlight the importance of embedding studies of litter ant ecology within broader decomposer food web dynamics.     

Importance of large colony size for successful invasion by Argentine ants (Hymenoptera: Formicidae): Evidence for biotic resistance by native ants

Abstract The Argentine ant, Linepithema humile (Mayr), is a widespread invasive ant species that has been associated with losses of native ant species and other invertebrates from its introduced range. To date, various abiotic conditions have been associated with limitations to the spread of Argentine ants, however, competitive interactions with native ant fauna may also affect the spread of Argentine ants. Here, we experimentally manipulated colony sizes of Argentine ants in the laboratory to assess whether Argentine ants were able to survive and compete for resources with a widespread, dominant native ant, Iridomyrmex‘rufoniger’. The results showed that over 24 h, the proportions of Argentine ants that were alive, at baits, and at sugar water decreased significantly in the presence of Iridomyrmex. In addition, Argentine ant mortality increased over time, however, the proportion of the colony that was dead decreased with the largest colony size. Argentine ants were only able to overcome Iridomyrmex when their colony sizes were 5–10 times greater than those of the native ants. We also conducted trials in which colonies of Argentine ants of varying sizes were introduced to artificial baits occupied by Iridomyrmex in the field. The results showed that larger Argentine ant colonies significantly affected the foraging success of Iridomyrmex after the initial introduction (5 min). However, over the first 20 min, when the Argentine ants were present at the baits, and over the entire 50 min experimental period, the numbers of Iridomyrmex at baits did not differ significantly with the size of the Argentine ant colony. This is the first experimental study to investigate the role of colony size in the invasion biology of Argentine ants in Australia, and the results suggest that Iridomyrmex may reduce the spread of Argentine ants, and that Argentine ants may need to attain large colony sizes in order to survive in the presence of Iridomyrmex. We address the implications of these findings for the invasion success of Argentine ants in Australia, and discuss the ability of Argentine ants to attain large colony sizes in introduced areas.     

Interspecific competition and coexistence between ants and land hermit crabs on small Bahamian islands

Numerous studies have demonstrated the existence of intra- and interspecific competition among ants, but few have investigated direct competitive interactions between ants and other taxa. In this paper, I present the first evidence of direct competitive interactions between ants and crabs. Evidence of competition for food between ants and the land hermit crab, Coenobita clypeatus (Herbst), was derived from observations and experiments in an archipelago of small islands in the central Exumas, Bahamas. Correlational evidence of competition for food based on occurrences at baits was found between ants and hermit crabs in multiple years. Observations at baits over time revealed species turnover occurred due to aggressive interactions. C. clypeatus discovered food items rapidly, but lost control of food over time, particularly to the ant Brachymyrmex obscurior Forel, which took longer to find food items but recruited large numbers of workers that drove off hermit crabs. A second ant species, Dorymyrmex pyramicus Roger, discovered baits quickly but did not recruit to baits in large numbers, and was not a superior competitor to either C. clypeatus or B. obscurior. Competition between ants and land hermit crabs was not intense enough to cause complementary distributions, and mechanisms of coexistence apparently include temporal variation in foraging activity and complementary foraging strategies when ants and crabs are active at the same time. Because of the widespread distributions and generalist scavenger diets of many ants and crabs, such competitive interactions are likely to be a common facet of many tropical and subtropical insular and coastal communities.     

Limited effects of dominant ants on assemblage species richness in three Amazon forests

1. Ants are highly interactive organisms and dominant species are considered to be able to control the species richness of other ants via competitive exclusion. However, depending on the scale studied, inter‐specific competition may or may not structure biological assemblages. To date, ant dominance–richness relationships have only been studied in small sample units, where a few dominant colonies could plausibly control most of the sample unit. 2. We conducted a comprehensive survey of terrestrial ant assemblages using bait, pitfall, and litter‐sorting methods in three sites in Brazilian Amazonia. Using a spatially structured rarefaction approach, based on sampling units with linear dimensions ranging from 25 to 250 m, the mesoscale patterns of ant dominance–richness relationships (sampling units covering hundreds of meters separated by kilometers) were investigated. 3. Interference–competition models (parabolic or negative linear relationships between species richness and the abundance of dominant ants) tended to be more frequent in smaller sample units or in assemblages sampled with interactive methods, such as baits. Using more inclusive sampling methods, the relationship was generally asymptotic rather than parabolic, with no reduction in species diversity because of the presence of dominants. Random co‐occurrence patterns of species within sites support the interpretation of a limited role for present‐day competition in structuring these assemblages. 4. Competition from dominant species may reduce species richness in small areas, especially when artificial baits are used, but appears to be less important than environmental constraints in determining ant species richness across scales of hectares and greater in these Amazon forests.     

Ants protect conifer seedlings from feeding damage by the pine weevil Hylobius abietis

• 1 Ants that protect food resources on plants may prey on (or deter) herbivores and thereby reduce damage. Red wood ants (of the Formica rufa group) are dominant ants in boreal forests of Eurasia and affect the local abundance of several herbivorous species.
• 2 The pine weevil Hylobius abietis (L.) is a herbivore that causes severe damage by feeding on the bark of coniferous seedlings within areas of forest regeneration.
• 3 We investigated whether ants can protect conifer seedlings from pine weevil feeding. In a manipulative experiment, ants were attracted to sugar baits attached to spruce seedlings and the damage caused by pine weevils was compared with control seedlings without ant‐baits.
• 4 The feeding‐scar area was approximately one‐third lower on the seedlings with ant‐baits compared with the controls. Besides red wood ants, Myrmica ants were also attracted in high numbers to the ant baits and the relative effects of these species are discussed.
• 5 The results obtained in the present study support the trophic cascade hypothesis (i.e. damage to herbivores is suppressed in the presence of predators). The decreased pine weevil feeding on the baited seedlings was probably a result of nonconsumptive interactions [i.e. the presence of (or harassment by) ants distracting pine weevils from feeding].
• 6 Understanding the role of ants may have important implications for future strategies aiming to control pine weevil damage. For example, maintaining suitable conditions for ants after harvesting stands may be an environmentally friendly but currently unexploited method of for decreasing weevil damage.

     

The size–grain hypothesis: do macroarthropods see a fractal world?

Abstract.  1. In the size–grain hypothesis (a) long legs allow walking organisms to step over gaps and pores in substrate but prohibit them from entering those gaps; (b) the world is more rugose for small organisms; and (c) the relative cost of long legs increases as organisms grow smaller. The hypothesis predicts a positive allometry of leg length ( = mass ^b where b > 0.33 of isometry), a pattern that robustly holds for ants.
2. Toward testing for leg length allometries in other taxa, arthropods were extracted from the Panama leaf litter and measured. Three common taxa (spiders, diplopods, Coleoptera) yielded b s that exceeded 0.33 while three others (Acarina, Pseudoscorpiones, and Collembola) did not. The exponent b tended to increase ( P = 0.06, n = 7) with an arthropod taxon's average body mass.
3. Since leg length in cursorial organisms tends toward isometry in very small and very large taxa (i.e. mammals) this suggests that the size–grain hypothesis may best apply at a transition zone of intermediate body mass: the macroarthropods.
4. Body length was a robust predictor of mass in all groups despite variation in shape.