Cooperative colony founding alters the outcome of interspecific competition between Amazonian plant-ants

In myrmecophytes, plants with structures in which ants establish colonies, there is strong competition among ant queens for access to host plants. However, our knowledge of how queens of different partner species interact when attempting to colonize plants remains limited. The Amazonian myrmecophyte Maieta guianensis is colonized by queens of two ant species: Crematogaster laevis and Pheidole minutula. We elucidated the competitive ranking of queens of these species and tested the hypothesis that cooperative colony founding (pleometrosis) by P. minutula queens could alter this ranking. We found that C. laevis queens are behaviorally dominant to P. minutula when individual queens encounter each other. Despite being inferior in combat, however, P. minutula queens successfully colonized seedlings at similar rates whether they were placed alone or in concert with a C. laevis queen. This may have occurred because the smaller P. minutula queens frequently entered domatia before the more robust C. laevis queens. Although C. laevis queens can evict P. minutula queens that had previously colonized domatia, this was an infrequent phenomenon—perhaps because while not fatal, conflicts often resulted in serious injury. Furthermore, by colonizing the same plant cooperative P. minutula queens dramatically reduce the probability that C. laevis colonizes host-plants without reducing their own per capita rates of colonization success. To our knowledge, this is a novel benefit of pleometrosis, whose primary advantages have primarily been thought to occur after the critical stage of colony establishment. Given the decreased likelihood of colonization when faced with multiple P. minutula, it may be that C. laevis’ persistence at the landscape level is enhanced by such factors as priority effects, superior dispersal ability, or niche partitioning.     

Two new myrmecophytic associations from the Malay Peninsula: Ants of the genusCladomyrma (Formicidae,Camponotinae) as partners ofSaraca thaipingensis (Caesalpiniaceae) andCrypteronia griffithii (Crypteroniaceae)

Summary In Peninsular Malaysia the treesSaraca thaipingensis (Caesalpiniaceae) andCrypteronia griffithii (Crypteroniaceae) are inhabited by ants. In the vicinity of Gombak, near Kuala Lumpur, the hollow internodes of youngSaraca thaipingensis plants are colonized mainly by twoCladomyrma species. In larger trees aCrematogaster sp. is also found.Crypteronia griffithii is inhabited by a third species ofCladomyrma. None of these species is conspecific with any of the threeCladomyrma taxa so far described. The colonies are founded by single mated queens, which have a conspicuous, sphecid wasp-like behaviour when searching for host plants and nest sites. They chew holes into the plant internodes and hollow them out to provide nest sites. Coccids and pseudococcids are cultivated within the internodes. The homopterans are not carried by queens on their nuptial flights. They apparently find their way by themselves into the cavities or are perhaps carried there by the worker ants. TheCladomyrma ants onCrypteronia are not aggressive, in contrast to those onSaraca thaipingensis.The relationship ofCrypteronia with ants seems to be obligatory, whereasSaraca was only partly colonized byCladomyrma. The interaction ofSaraca withCrematogaster sp. is loose and facultative, since theCrematogaster sp. also lives on other three species. Our studies have now revealed fourCladomyrma spp. which are regularly associated with plants. The genus therefore seems to have an entirely myrmecophytic way of life.     

Asymmetric dispersal and colonization success of Amazonian plant-ants queens

Background

The dispersal ability of queens is central to understanding ant life-history evolution, and plays a fundamental role in ant population and community dynamics, the maintenance of genetic diversity, and the spread of invasive ants. In tropical ecosystems, species from over 40 genera of ants establish colonies in the stems, hollow thorns, or leaf pouches of specialized plants. However, little is known about the relative dispersal ability of queens competing for access to the same host plants.

Methodology/Principal Findings

We used empirical data and inverse modeling—a technique developed by plant ecologists to model seed dispersal—to quantify and compare the dispersal kernels of queens from three Amazonian ant species that compete for access to host-plants. We found that the modal colonization distance of queens varied 8-fold, with the generalist ant species (Crematogaster laevis) having a greater modal distance than two specialists (Pheidole minutula, Azteca sp.) that use the same host-plants. However, our results also suggest that queens of Azteca sp. have maximal distances that are four-sixteen times greater than those of its competitors.

Conclusions/Significance

We found large differences between ant species in both the modal and maximal distance ant queens disperse to find vacant seedlings used to found new colonies. These differences could result from interspecific differences in queen body size, and hence wing musculature, or because queens differ in their ability to identify potential host plants while in flight. Our results provide support for one of the necessary conditions underlying several of the hypothesized mechanisms promoting coexistence in tropical plant-ants. They also suggest that for some ant species limited dispersal capability could pose a significant barrier to the rescue of populations in isolated forest fragments. Finally, we demonstrate that inverse models parameterized with field data are an excellent means of quantifying the dispersal of ant queens.     

Relationship between Plant Size and Ant Associates in Two Amazonian Ant‐Plants1

A survey of two Amazonian melastome ant‐plants, Maieta guianensis and Tococa bullifera, revealed a significant difference in plant size according to the species of ant inhabiting the plant. Plants with Crematogaster laevis, on average, were smaller than those with Pheidole minutula (in M. guianensis) and those with Azteca sp. (in T. bullifera). There is no evidence that these patterns were due either to the deterministic replacement of C. laevis by another ant species during host‐plant ontogeny or to a habitat effect on plant growth rates coupled with colony survival. More likely, the smaller size of C. laevis plants can be explained by its effects on host‐plant performance. Plants with C. laevis lost their associated ant colonies more frequently than plants with P. minutula and Azteca sp. Plants that lost their C. laevis either died, or more commonly, were severely defoliated. Defoliated plants, once sprouted, tended to become recolonized, but such recolonizations were not deterministic so as to favor one species over another. Plants with C. laevis showed similar, or only slightly greater, standing levels of herbivory than plants with P. minutula or Azteca sp. This suggests that when C. laevis is present, it confers some degree of protection to its hosts. It was found that early in colony development, queens of C. laevis moved off their host plants to build satellite nests in dead twigs on the ground, a behavior not seen in the other two species and one that possibly renders colonies more vulnerable to mortality from predation, flooding, or nest decay. Comparable 8¹⁵N values in C. laevis and P. minutula indicate that the two species are equally dependent on food supplied by the host plant.     

The reproductive phenology of an Amazonian ant species reflects the seasonal availability of its nest sites

In saturated tropical ant assemblages, reproductive success depends on queens locating and competing for scarce nest sites. Little is known about how this process shapes the life histories of tropical ants. Here I investigate the relationship between nest site availability and an important life history trait, reproductive phenology, in the common Amazonian ant species Allomerus octoarticulatus. A. octoarticulatus is a plant-ant that nests in the hollow, swollen stem domatia on Cordia nodosa. I provide evidence that nest sites are limiting for A. octoarticulatus. Most queens produced by A. octoarticulatus colonies died before locating suitable host plants, and most queens that located hosts died before founding colonies, probably from intraspecific competition among queens for control of host plants. I further show that the reproductive phenology of A. octoarticulatus closely matches the seasonal availability of its nest sites, domatia-bearing C. nodosa saplings. Both the production and flight of A. octoarticulatus reproductives, and the number of C. nodosa saplings available for colonization by ants, peaked from March to May. There was correlative evidence that A. octoarticulatus colonies use temperature as a cue to synchronize their reproduction to the availability of C. nodosa saplings: both the production of reproductives by ant colonies and the number of C. nodosa saplings available for colonization were correlated with temperature, and not with rainfall. All of these results suggest that nest site limitation constrains the reproductive phenology of A. octoarticulatus.     

Mutualism between Maieta guianensis Aubl., a myrmecophytic melastome,and one of its ant inhabitants: ant protection against insect herbivores

Summary The hypothesis that ants (Pheidole minutula) associated with the myrmecophytic melastome Maieta guianensis defend their host-plant against herbivores was investigated in a site near Manaus, Amazonas, Brazil. M. guianensis is a small shrub that produces leaf pouches as ant domatia. Plants whose ants were experimentally removed suffered a significant increase in leaf damage compared with control plants (ants maintained). Ants patrol the young and mature leaves of Maieta with the same intensity, presumably since leaves of both ages are equally susceptible to herbivore attack. The elimination of the associated ant colony, and consequent increase in herbivory, resulted in reduced plant fitness. Fruit production was 45 times greater in plants with ants than in plants without ants 1 year after ant removal.     

Diversity of ant-plant interactions: protective efficacy in Macaranga species with different degrees of ant association

The pioneer tree Macaranga in SE Asia has developed manyfold associations with ants. The genus comprises all stages of interaction with ants, from facultative relationships to obligate myrmecophytes. Only myrmecophytic Macaranga offer nesting space for ants and are associated with a specific ant partner. The nonmyrmecophytic species are visited by a variety of different ant species which are attracted by extrafloral nectaries (EFN) and food bodies. Transitional Macaranga species like M. hosei are colonized later in their development due to their stem structure. Before the colonization by their specific Crematogaster partner the young plants are visited by different ant species attracted by EFN. These nectaries are reduced and food body production starts as soon as colonization becomes possible. We demonstrated earlier that obligate ant partners can protect their Macaranga plants against herbivore damage and vine cover. In this study we focused on nonspecific interactions and studied M. tanarius and M. hosei, representing a non-myrmecophyte and a transitional species respectively. In ant exclusion experiments both M. tanarius and M. hosei suffered significantly higher mean leaf damage than controls, 37% versus 6% in M. hosei, 16% versus 7% in M. tanarius. M. tanarius offers both EFN and food bodies so that tests for different effects of these two food rewards could be conducted. Plants with food bodies removed but with EFN remaining had the lowest mean increase of herbivore damage of all experimental groups. Main herbivores on M. hosei were mites and caterpillars. Many M. tanarius plants were infested by a shootborer. Both Macaranga species were visited by various ant species, Crematogaster spp. being the most abundant. We found no evidence for any specific relationships. The results of this study strongly support the hypothesis that non-specific, facultative associations with ants can be advantageous for Macaranga plants. Food bodies appear to have lower attractive value for opportunistic ants than EFN and may require a specific dietary adaptation. This is also indicated by the fact that food body production in the transitional M. hosei does not start before stem structure allows a colonization by the obligate Crematogaster species. M. hosei thus benefits from facultative association with a variety of ants until it produces its first domatia and can be colonized by its obligate mutualist.     

A shift in colony founding behaviour in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2

Summary. A shift in colony founding behaviour from single queen (haplometrosis) to multiple queens (pleometrosis) was observed locally in the obligate plant-ant Crematogaster (Decacrema) morphospecies 2, which is associated with Macaranga trees in Borneo. In addition, about a quarter of all mature colonies (27 of 95 trees examined) were found to be multiple queen colonies. They arose either directly from pleometrotic founding colonies or secondarily by adoption of additional queens. Using microsatellite markers, we showed that queens in colonies founded through pleometrosis are unrelated and each queen participates in producing worker offspring, albeit with significant skew in a third of the colonies. In mature polygynous colonies, all resident queens contributed to the production of workers and sexual offspring. Relatedness of queens in mature polygynous colonies was not significantly higher than in foundress associations. We hypothesize that increased nest site limitation in this specific interaction trigger the observed shift in colony founding behaviour. Crematogaster msp. 2 inhabits the light demanding pioneer plant species Macaranga pearsonii that is typical for early successional stages of secondary forests. Thus suitable host-plants for colonisation are abundant for only a short time in highly disturbed sites and become increasingly sparse when secondary forest matures.Received 6 September 2004; revised 29 November; accepted 10 December 2004.     

Ecological barriers to early colony establishment in three coexisting acacia-ant species in Kenya

In black cotton uplands in East Africa, four symbiotic acacia-ant species compete for possession of a single swollen thorn tree species, Acacia drepanolobium, and yet coexist at fine spatial scales. Three of the four ant species produce independent foundress queens that establish colonies claustrally within swollen thorns, most often on small saplings. We conducted surveys of such saplings at two sites in 2001 and 2004, and examined foundresses and incipient colonies within their swollen thorns to determine what factors influence their success. Competition among foundresses for nest initiation sites was intense, with an average of over one founding attempt per swollen thorn in all samples, and with living and dead queens significantly hyper-dispersed among available thorns. Combat with other foundresses was the most common cause of death among claustral queens, especially for Tetraponera penzigi. In interspecific battles for nest initiation sites, T. penzigi was dominant over Crematogaster nigriceps and C. mimosae, and C. nigriceps won over 80% of its contests with C. mimosae foundresses. For singleton foundresses, brood parasitism by the braconid wasp Trigastrotheca laikipiensis typically results in the death of the entire ant brood. Host queens defend parasite larvae, pupae and eclosed adults, apparently unable to distinguish the wasps from legitimate offspring. Rates of brood parasitism were as high as 15–20% for incipient colonies of both Crematogaster species, but were extremely low for T. nigriceps in all samples. Although T. penzigi and C. nigriceps foundresses are always solitary, approximately 18% of claustral C. mimosae colonies contain cooperating pleometrotic queens. For unparasitized, claustral C. mimosae colonies, brood production per queen did not differ between solitary and cooperating foundresses. However, the per-capita risks associated with parastitism were reduced for pleometrotic queens. Received 8 March 2005; revised 23 May 2005; accepted 3 June 2005.     

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

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

A newly-discovered mode of colony founding among fire ants

Summary Queen ants start new colonies either unassisted by workers (independent founding), assisted by workers from their natal nest (dependent founding), or assisted by the workers of other species (dependent, socially parasitic). The monogyne form of the fire ant,Solenopsis invicta, founds independently in summer, but in the fall it also produces a few sexuals some of which overwinter, then fly and mate in early spring. These overwintered queens lack the nutritional reserves and behaviors for independent colony founding. Rather, they seek out unrelated, mature, orphaned colonies, enter them and exploit the worker force to found their own colony through intraspecific social parasitism. Success in entering orphaned colonies is higher when these lack overwintered female alates of their own. When such alates are present, orphaning causes some to dealate and become uninseminated replacement queens, usually preventing entry of unrelated, inseminated replacement queens. Such colonies produce large, all-male broods. Successful entry of a parasitic queen robs the host colony of this last chance at reproductive success. Only overwintered sexuals take part in this mode of founding.     

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.     

Pruning behavior and intercolony competition of Tetraponera (Pachysima) aethiops (Pseudomyrmecinae, Hymenoptera) in Barteria fistulosa in a tropical forest, Democratic Republic of Congo

The early stages of colonization by Tetraponera (= Pachysima) aethiops (Pseudomyrmecinae) to its obligate host plant, Barteria fistulosa (Passifloraceae), was studied in Kahuzi-Biega National Park, Democratic Republic of Congo (formerly Zaire). In our observations, as many as 36 queens colonized a seedling, and all of the established colonies had just a single queen each. The ants depend totally on the products of scale insects which inhabit the domatia. The ants damage the terminal meristems of the branches where other incipient colonies may inhabit. This tip-biting behavior may deteriorate the nutritional condition of scale insects and suppress the growth of the competing colonies. Ants pruned the plants near the edge of the trunk of the host tree most intensively, even the foliage of the lowest branch in shade. This observation supports the hypothesis that the main driving force which leads to pruning behavior is intercolony competition rather than improving the light condition of the host tree.     

Energetics of newly-mated queens and colony founding in the fungus-gardening ants Cyphomyrmex rimosus and Trachymyrmex septentrionalis (Hymenoptera: Formicidae)

Abstract.  The energetics of colony founding is investigated in the fungus gardening ants (Attini) Trachymyrmex septentrionalis and Cyphomyrmex rimosus . Similar to most ants, inseminated queens of these two species found nests independently unaccompanied by workers (haplometrosis). Whereas most ant founding queens seal themselves in a chamber and do not feed when producing a brood entirely from metabolic stores (claustral founding), the majority of fungus gardening ants must forage during the founding phase (semiclaustral founding). Laboratory-reared T. septentrionalis individuals comprise 84 dealate females collected after mating flights in June 2004. Twenty are immediately killed to obtain values for queen traits and another 20 after worker emergence for queen, fungus garden and worker traits. Cyphomyrmex rimosus comprise 22 dealate females collected in June 2005; ten of which are immediately killed and similarly prepared. Newly-mated T. septentrionalis queens have 25% of their dry weight as fat; whereas newly-mated C. rimosus queens contain 11% fat. These amounts are 50–75% less than most independently founding ant species. Trachymyrmex septentrionalis queens lose merely 5% of their energetic content during colony founding, whereas the total energetic content of their brood is more than three-fold the amount lost by the queen. Incipient T. septentrionalis colonies produce approximately half as much ant biomass per gram of fungus garden as do mature colonies. Similar to most ants, T. septentrionalis produces minim workers that are approximately 40% lighter than workers from mature colonies. Regardless of their size, T. septentrionalis workers contain much lower fat than do workers of claustral species. These data indicate that fungus gardening is adaptive because colonies can produce much cheaper offspring, making colony investment much lower.     

Habitat structure,dispersal strategies and queen number in two boreal Leptothorax ants

In two nearctic ants, Leptothorax canadensis and Leptothorax sp. A, young queens may either found their own nest solitarily after mating or seek adoption into an established colony. Whether a queen disperses or not is associated with genetically determined queen morphology in Leptothorax sp. A. Whereas a majority of winged queens attempt solitary colony founding after mating, most wingless, intermorphic queens return to their maternal nests and new colonies are founded by budding after hibernation. The latter strategy appears to be correlated with patchy, isolated habitats, whereas in extended boreal forests dispersal on the wing is probably more common. Alternative dispersal strategies strongly affect the average number of queens per colony and seasonal fluctuations of colony structure.     

The frequency of multi‐queen colonies increases with altitude in a Nearctic ant

1. Most ants in boreal and alpine habitats are facultatively polygynous, i.e. their colonies may contain one or several queens. It was investigated how the proportion of polygynous colonies varies along an elevation gradient from 60 to 2700 m in the Nearctic ant Temnothorax rugatulus (Emery). 2. Across all populations, the proportion of polygynous colonies was positively correlated with altitude. The correlation was considerably stronger when only populations in the narrow area of the Chiricahua Mts, Arizona, were compared. 3. The dominance of polygynous colonies at high altitudes may be associated with selection against solitary colony founding by young queens. In areas with short summers and long winters, hibernation mortality of solitarily founding queens may select for alternative reproductive tactics, such as polygyny and colony founding by budding. 4. Colony founding tactics need to be taken into account to more fully understand altitudinal and latitudinal patterns of ant faunas.     

Species specificity in settling-plant selection by foundress ant queens in <Emphasis Type="Italic">Macaranga</Emphasis>–<Emphasis Type="Italic">Crematogaster</Emphasis> myrmecophytism in a Bornean dipterocarp forest

Previous studies have demonstrated that the obligate myrmecophytism between Macaranga ant-plants and Crematogaster plant-ants is highly species specific, although multiple Macaranga species can coexist in a microhabitat. However, the species specificity has been described based on the study of trees with established plant-ant colonies. We studied how the process of settling into the partner Macaranga seedlings by single foundress Crematogaster queens contributes to species specificity. By sampling seedlings of three sympatric Macaranga myrmecophytes species in the field, we tested two hypotheses. The first is that foundresses correctly select their specific partner plant species when they settle into seedlings. The second hypothesis is that the seasons in which seedlings available for settling by foundresses appear are segregated among the Macaranga species, and the seasons in which foundress queens settle are synchronized to the appearance of seedlings of specific partner species; thus species specificity is consequently generated. Our results support the former hypothesis but not the latter: we always observed foundresses settling species-specific host plants, and seedlings suitable for settling were always available in each Macaranga species. Electronic Publication     

Polygynous supercolonies of the acacia‐ant Pseudomyrmex peperi,an inferior colony founder

In ant–plant protection mutualisms, plants provide nesting space and nutrition to defending ants. Several plant–ants are polygynous. Possessing more than one queen per colony can reduce nestmate relatedness and consequently the inclusive fitness of workers. Here, we investigated the colony structure of the obligate acacia‐ant Pseudomyrmex peperi, which competes for nesting space with several congeneric and sympatric species. Pseudomyrmex peperi had a lower colony founding success than its congeners and thus, appears to be competitively inferior during the early stages of colony development. Aggression assays showed that P. peperi establishes distinct, but highly polygynous supercolonies, which can inhabit large clusters of host trees. Analysing queens, workers, males and virgin queens from two supercolonies with eight polymorphic microsatellite markers revealed a maximum of three alleles per locus within a colony and, thus, high relatedness among nestmates. Colonies had probably been founded by one singly mated queen and supercolonies resulted from intranidal mating among colony‐derived males and daughter queens. This strategy allows colonies to grow by budding and to occupy individual plant clusters for time spans that are longer than an individual queen’s life. Ancestral states reconstruction indicated that polygyny represents the derived state within obligate acacia‐ants. We suggest that the extreme polygyny of Pseudomyrmex peperi, which is achieved by intranidal mating and thereby maintains high nestmate relatedness, might play an important role for species coexistence in a dynamic and competitive habitat.     

Roads Alter the Colonization Dynamics of a Keystone Herbivore in Neotropical Savannas1

Roads can facilitate the establishment and spread of both native and exotic species. Nevertheless, the precise mechanisms facilitating this expansion are rarely known. We tested the hypothesis that dirt roads are favorable landing and nest initiation sites for founding‐queens of the leaf‐cutter ant Atta laevigata. For 2 yr, we compared the number of attempts to found new nests (colonization attempts) in dirt roads and the adjacent vegetation in a reserve of cerrado (tree‐dominated savanna) in southeastern Brazil. The number of colonization attempts in roads was 5 to 10 times greater than in the adjacent vegetation. Experimental transplants indicate that founding‐queens are more likely to establish a nest on bare soil than on soil covered with leaf‐litter, but the amount of litter covering the ground did not fully explain the preference of queens for dirt roads. Queens that landed on roads were at higher risk of predation by beetles and ants than those that landed in the adjacent vegetation. Nevertheless, greater predation in roads was not sufficient to offset the greater number of colonization attempts in this habitat. As a consequence, significantly more new colonies were established in roads than in the adjacent vegetation. Our results suggest that disturbance caused by the opening of roads could result in an increased Atta abundance in protected areas of the Brazilian Cerrado.     

Three queen morphs with alternative nest-founding behaviors in the ant, Temnothorax longispinosus

In many ant species, multiple modes of founding new colonies occur in the same population. These modes include dependent founding, independent founding by haplometrosis (single queen), and independent founding by pleometrosis (multiple queens). In several cases, a dimorphism in queen size has been found, such that each morph specializes in a particular nest-founding behavior. I investigated queen size in the ant Temnothorax longispinosus in several southern Wisconsin populations and found three distinct queen morphs: small queens with very low fat content and short wings, large queens with low fat and long wings, and large queens with high fat and very long wings. Several traits associated with founding behavior correlated with these queen sizes. Small queens were produced in lower numbers, were more common in polygynous nests, and returned to the nest in higher proportions than both large queen morphs. The size ranges and fat levels of each queen morph were similar to those of other species that specialize in either haplometrosis (very large, high fat), pleometrosis (large, low fat), or dependent founding (small and low fat). However, there was extensive overlap in several of the founding behaviors, suggesting that the morphs in these populations have some flexibility in founding behavior. The queen morphs in these populations of T. longispinosus may resemble early stages in the evolution of more specialized dispersal polymorphisms found in other ant species. Received 11 January 2006; revised 15 September 2006; accepted 18 September 2006.