Within-group behavioral variation promotes biased task performance and the emergence of a defensive caste in a social spider

The social spider Anelosimus studiosus exhibits a behavioral polymorphism where colony members express either a passive, tolerant behavioral tendency (social) or an aggressive, intolerant behavioral tendency (asocial). Here we test whether asocial individuals act as colony defenders by deflecting the suite of foreign (i.e., heterospecific) spider species that commonly exploit multi-female colonies. We (1) determined whether the phenotypic composition of colonies is associated with foreign spider abundance, (2) tested whether heterospecific spider abundance and diversity affect colony survival in the field, and (3) performed staged encounters between groups of A. studiosus and their colony-level predator Agelenopsis emertoni (A. emertoni)to determine whether asocial females exhibit more defensive behavior. We found that larger colonies harbor more foreign spiders, and the number of asocial colony members was negatively associated with foreign spider abundance. Additionally, colony persistence was negatively associated with the abundance and diversity of foreign spiders within colonies. In encounters with a colony-level predator, asocial females were more likely to exhibit escalatory behavior, and this might explain the negative association between the frequency of asocial females and the presence of foreign spider associates. Together, our results indicate that foreign spiders are detrimental to colony survival, and that asocial females play a defensive role in multi-female colonies.     

The hydrozoan coral<Emphasis Type="Italic"> Millepora dichotoma</Emphasis>: speciation or phenotypic plasticity?

This article describes ecological and biological differences between two morphs of the Red Sea fire coral Millepora dichotoma. The species is divided into two main morphs: branching and encrusting, which were found to differ both in color and morphology. Each morph has two or more sub-morphs. A total of 372 M. dichotoma colonies were examined in a census at two study sites in the Gulf of Elat. Colony size and abundance of the two morphs were found to differ significantly between sites. Experimental examination of each morph's morphological plasticity revealed different growth rates and difference in growth plasticity between the branching and the encrusting morph. Most of the fragments from the branching colonies (94%) attached to experimentally placed Plexiglas substrate, compared with much less attachment by the encrusting fragments (11%). The growth form of the branching morph on the Plexiglas switched to encrusting, spreading over and covering the substrate. When the new encrusting colony reached the edge of this substrate, it started to produce tips, and returned to growth in the classic branching form. The encrusting morph did not change its growth form. Following attachment of the original fragments of the branching morph to the substrate, 8.1% of them produced new tips. When the original branches were removed, after converting to encrusting growth form, 19% of the fragments produced new tips. The capsule size of nematocysts of the two morphs was also significantly different (t-test, P<0.05). Molecular data (ITS region) clearly demonstrate that these two M. dichotoma morphs differ considerably. Molecular evidence (srRNA) from the symbiotic zooxanthellae also shows a different pattern of clades in the hosts. The ecological, biological and molecular data thus attest to the two morphs being distinguishable. Contrary to previous reports, we consequently suggest that the two morphs of M. dichotoma found in the Gulf of Elat are actually two distinct species.Communicated by O. Kinne, Oldendorf/Luhe     

Assessment of space utilisation in a subtidal temperate bryozoan community

On subtidal hard substrata, Bryozoa are often very abundant, both in terms of numbers of colonies and numbers of species. To investigate how so many species are able to co-exist, often in close proximity to each other, subtidal material was dredged from an area of bryozoan-rich coarse shelly sea-bed off the south-west of the Isle of Man (British Isles) in autumn 1984 and 1985. A survey was performed on the distribution of colonies of eight common, encrusting cheilostome bryozoan species across the surfaces of four types of empty, disarticulated bivalve shells. Each species was found to be significantly commoner on certain substratum types such as rough surfaces or inner concave surfaces. Fenestrulina malusii (Audouin) showed a further, more specific, tendency to be signficantly more abundant on the grooves of the corrugated inner surfaces of left valves of Pecten maximus (L.). In general however there was much overlap in distribution and it is proposed that such large numbers of species are able to co-exist because of the evolution of different life-history strategies ensuring that each species uses the available temporally and spatially heterogenous substrata in subtly different ways; some species, for example, occupying substrata that are optimal to bryozoan growth and other species, by reaching maturity earlier, being able to inhabit more ephemeral suboptimal substrata.     

Spatial and behavioral interactions between a native and introduced salamander species

Behavioral interactions with native species may influence the invasiveness of introduced species. The salamanders Plethodon glutinosus and P. jordani in the eastern United States share many life history traits and demonstrate complex interspecific interactions that range geographically from competitive exclusion to sympatry. P. jordani was introduced to Mountain Lake Biological Station, Virginia, USA, between the years 1935 and 1945. We tested whether competition for space may influence the invasion of P. jordani into native P. glutinosus habitat by utilizing data from natural distributions, a field experiment, and controlled laboratory experiments. No environmental variables differed where P. glutinosus and P. jordani were collected in the field at the site of P. jordani introduction. In the field experiment, P. glutinosus was more fully exposed during foraging bouts in cages shared with heterospecifics as opposed to ones shared with conspecific salamanders. Condition (mass relative to body length) of salamanders at the end of the 3 months did not differ between conspecific and heterospecific treatments. In the laboratory, P. glutinosus most often attained the single burrow in the arena, but residency status had no effect. Species cohabited the burrow 50% of the time. Pair-wise encounters in the laboratory indicated that both species spend less than 20% of the time in aggressive behaviors as juveniles. Adults showed no behavior interpreted by us as aggression during pair-wise encounters. Received: 19 December 1999 / Accepted: 18 March 2000     

Genetic structure of local populations and divergence between growth forms in a clonal invertebrate,the Caribbean octocoral Briareum asbestinum

Although the genetic structure of many populations of marine organisms show little deviation from panmixia, in those marine species with limited larval dispersal, patterns of microgeographic genetic differentiation may be common. The octocoral Briareum asbestinum should show local population differentiation because colonies reproduce asexually by fragmentation, most matings occur between colonies in very close proximity, and the sexually produced larvae and sperm appear to disperse only short distances. Variability in secondary chemistry of individual B. asbestinum colonies from different populations in close proximity also suggests local population differentiation. We determined the genetic composition of local populations by surveying allozyme variation of three shallow and two deep populations within a 300 m² area at San Salvador Island, Bahamas and at a site 161 km away on Little San Salvador, Bahamas in July 1990. As B. asbestinum occurs as either an erect branching form or an encrusting mat often at the same sites, we sampled both morphs to examine the extent of genetic exchange between them. Five of 21 loci were polymorphic and most populations showed a deficit of heterozygotes. Allele frequencies differed significantly between morphs at each site where they occurred together. The mean genetic distance (D=0.065) between morphs is consistent with the interpretation that the two morphs are genetically isolated. Despite the close spatial proximity of the San Salvador populations, both the branching and encrusting morphs showed significant genetic heterogeneity among neighboring populations. Similarly, pooled allelic frequencies for samples collected from the islands of San Salvador and Little San Salvador differed significantly at 1 locus for the branching morph and at 3 out of 5 loci for the encrusting morph.     

Alloimmune responses between Acropora hemprichi conspecifics: nontransitive patterns of overgrowth and delayed cytotoxicity

Allogeneic interactions between Acropora hemprichi (Ehrenberg) colonies were followed in situ up to 18 mo between 1990 and 1992 in Eilat, Red Sea. We established all 21 possible branch pair combinations among seven large colonies in three to eight replicates each (n=92 pairs), in addition in 14 pairs of grafts within-colonies. All isografts resulted in fusion. Allogeneic interactions revealed a variety of effector mechanisms which involved the appearance of dark blue-purple color in the tissues of interacting branch tips (<1 cm), unilateral overgrowths (1.0 to 1.5 mm mo^-1 for 1 to 8 mo), rejections, healings, continuous overgrowths, reversals, repeated rejections and more. Direction of overgrowth was usually specific to a given combination of colonies. A hierarchial network of overgrowths was obtained by analyzing overgrowth directions of all pairwise combinations, which included 30 transitive and five nontransitive three-way combinations. These outcomes, similarly to responses of allogeneic encounters recorded from other coral species, show that colonies can respond selectively to different types of allogeneic challenges. It is suggested that corals may have the capacity to distinguish self from nonself by recognizing nonself attributes presented on conspecifics.     

Interpreting seawater temperature range using oxygen isotopes and zooid size variation in Pentapora foliacea (Bryozoa)

This is the first investigation of how two independent proxies for seawater temperature inference (zooid size variation and oxygen isotope ratios of skeletal carbonate) relate to the actual measured ranges of temperature experienced by cheilostome bryozoan colonies. Nine specimens of the bimineralic marine cheilostome bryozoan Pentapora foliacea (Ellis and Solander, 1786) were analysed, collected from ~18-m depth at two localities in Wales, UK—four from Skomer Island (51°42′510″N, 5°13′42.60″W) and five from Porth Ysgaden, Lleyn Peninsula (52°54′6.75″N, 4°38′47.34″W). The annual range of temperature implied by zooid size variability provides a good approximation of the actual range of temperature recorded by a datalogger. However, annual ranges of temperature reconstructed from skeletal oxygen isotope ratios were narrower, typically not showing the lowest temperatures experienced by the colonies. This can be explained by progressive thickening of zooid skeletal walls during the life of the colony that homogenises the temperature signal by time-averaging over the lifetime of the colonies. Our study provides evidence that a combined morphological isotope approach has great potential in the reconstruction of annual ranges in seawater temperatures from historical and fossil bryozoans, particularly for species that lack ontogenetic skeletal wall thickening and bimineralic skeletal composition. As cheilostome bryozoans have been common in benthic communities since the Late Cretaceous, they represent a valuable and underutilised resource for the interpretation of environmental regimes.     

Foraging and spatiotemporal territories in the honey ant Myrmecocystus mimicus wheeler (Hymenoptera: Formicidae)

Summary The honey ant Myrmecocystus mimicus is a scavenger, forages extensively on termites, collects floral nectar, and tends homoptera. Individual foragers of M. mimicus usually disperse in all directions when leaving the nest, but there are also groups of foragers that tend to swarm out of the nest primarily in one direction. Such massive departues are usually at irregular intervals, which may last several hours. The results of field and laboratory experiments suggest that these swarms of foragers are organized by a group recruitment process, during which recruiting scout ants lay chemical orientation trails with hindgut contents and simultaneously stimulate nestmates with a motor display and secretions from the poison gland. Usually these columns travel considerable distances (4–48 m) away from the nest, frequently interfering with the foraging activity of conspecific neighboring colonies.To prevent a neighboring colony from access to temporal food sources or to defend spatiotemporal borders, opposing colonies engage in elaborate display tournaments. Although hundreds of ants are often involved during these tournaments almost no physical fights occur. Instead, individual ants confront each other in highly sterotyped aggressive displays, during which they walk on stilt legs while raising the gaster and head. Some of the ants even seem to inflate their gasters so that the tergites are raised and the whole gaster appears to be larger. In addition, ants involved in tournament activities are on average larger than foragers.The dynamics of the tournament interactions were observed in several colonies over several weeks-mapping each day the locations of the tournaments, the major directions of worker routes away from the nest, and recording the general foraging activities of the colonies. The results indicate that a kind of dominance order can occur among neighboring colonies. On the other hand, often no aggressive interactions among neighboring colonies can be observed, even though the colonies are actively foraging. In those cases the masses of foragers of each colony depart in one major direction that does not bring them into conflict with the masses of foragers of a neighboring colony. This stability, however, can be disturbed by offering a new rich food source to be exploited by two neighboring colonies. This invariably leads to tournament interactions.When a colony is considerably stronger than the other, i.e., with a much larger worker force, the tournaments end quickly and the weaker colony is raided. The foreign workers invade the nest, the queen of the resident colony is killed or dirven off, while the larvae, pupae, callow workers, and honey pot workers are carried or dragged to the nest of the raiders. From these and other observations we conclude that young M. mimicus queens are unlikely to succeed in founding a colony within approximately 3 m of a mature M. mimicus colony because they are discovered and killed, or driven off by workers of the resident colony. Within approximately 3–15 m queens are more likely to start colonies, but these incipient groups run a high risk of being raided and exterminated by the mature colony.Although populations of M. mimicus and M. depilis tend to replace each other, there are areas where both species overlap marginally. Foraging areas and foraging habitats of both species also overlap broadly, but we never observed tournament interactions between M. mimicus and M. depilis.The adaptive significance of the spatiotemporal territories in M. mimicus is discussed.     

Dominance and fertility in a functionally monogynous ant

Summary Colonies of some leptothoracine ants may contain several inseminated but sterile females in addition to a single, fertile queen (functional monogyny). We here report the first observations on the behavior of these supernumerary females in the nearctic ant Leptothorax sp.A, a species belonging to the L. muscorum complex.In four colonies, each with up to eight intermorphic females, ritualized or openly aggressive interactions between individuals were observed, similar to those among workers of some other leptothoracine ants. The responses of individual females during encounters with nestmates apparently reflect the existence of linear dominance hierarchies. In each colony, the highest ranking individual was fed and groomed significantly more often than other females, and was the only one to oviposit after hibernation and to become fully physogastric during the first weeks of spring. When these -females were removed from three colonies, several other females started to lay eggs. However, in each colony only the highest ranking individual remained fertile; the others either were pushed out of the nest and finally killed by the workers, or their ovaries degenerated again.Aggressive interactions among females may also play an important role in the foundation of new colonies either by inducing intermorphs to leave the maternal nest to found new colonies solitarily or by inducing colony fission. Offprint requests to: J. Heinze at his new address     

Offshore marine cheilostome Bryozoa from Fort Lauderdale,Florida

Cheilostome Bryozoa were collected upon marine fouling test panels exposed offshore from Fort Lauderdale, Florida, USA between 1961 and 1967. Panels were exposed at certain depths, ranging from the surface to the bottom, in several arrays extending to a distance of 4 miles offshore. Twenty-three species were identified from the 114 panels recovered and examined. Most species were found on panels in shallow water, or on panels near the bottom in deeper, offshore water. A notable exception was a large, arborescent colony of Biflustra savartii which engulfed a panel 50 feet (15 m) deep, 4 miles from shore, in the Florida current. The lack of cheilostome Bryozoa on shallow and mid-water panels at the offshore, deepwater stations was apparently due to the low number of larvae reaching them from the distant inshore colonies.     

Interspecific aggression in colonies of the slave-making ant Harpagoxenus sublaevis

Colonies of the slave-making ant, Harpagoxenus sublaevis, may simultaneously contain workers of several Leptothorax slave species. We observed aggressive interactions among slave-makers, between slavemakers and slaves, and among slaves in 11 mixed colonies. The first two types of aggression appear to be correlated with reproductive competition for the production of males. Aggressive interactions among slaves, however, occurred mainly between slaves belonging to different species. In two colonies, in which one slave species clearly outnumbered the other, the majority attacked and finally expelled all nestmates belonging to the minority species. Our observations thus suggest that in Harpagoxenus colonies a homogeneous colony odor is not always achieved and that heterospecific slaves may occasionally be mistaken for alien ants. Gas chromatographic analyses of ants from mixed colonies similarly show that cuticular hydrocarbon profiles may differ strongly between heterospecific nestmate slaves.     

Mutualist-induced morphological changes enhance growth and survival of corals

Species interactions can induce morphological changes in organisms that affect their subsequent growth and survival. In Moorea, French Polynesia, epibiotic gammaridean amphipods induce the formation of long, branch-like coral “fingers” on otherwise flat, encrusting, or plating Montipora coral colonies. The fingers form as corals encrust tubes built by the amphipods and lead to significant changes in colony morphology. This study examines the costs and benefits of this association to the amphipods and corals and demonstrates that the interaction is a mutualism. Amphipods gain protection from predators by living within corals, and corals benefit by enhanced growth and survival. Benefits to the coral arise through direct effects due to the amphipods’ presence as well as through benefits derived from the altered colony morphology. This study demonstrates that induced morphological plasticity can be a mechanism for facilitation, adding to our knowledge of the roles mutualism, and phenotypic plasticity play in ecology.     

Parasitoid learning during interactions with ants: how to deal with an aggressive antagonist

Pauesia picta, P. pinicollis and P. silvestris (Hymenoptera: Aphidiidae) are common parasitoids of the conifer aphid Cinara pinea, which is regularly attended by red wood ants (Formica spp.). In this study, I tested whether females of these parasitoid species learned during interactions with honeydew-collecting Formica polyctena workers that caution is not necessary if searching behaviour is adapted, and whether parasitoids benefit from being able to learn. When searching on Scots pines, naive females of P. picta and P. pinicollis generally retreated to a pine needle when making contact with a honeydew-collecting ant, did not approach ants from the side or from the front and kept a ”safe distance” from ants when sitting on needles. After some non- aggressive ant encounters, experienced female parasitoids changed their behaviour: they reduced their searching speed, approached ants from the side and even from the front, retreated less often in response to an approaching ant and reduced the ”safe distance”. These experienced females had a significantly higher rate of oviposition than naive females or females foraging for an unattended host. Thus, the ability of the parasitoid to learn during interactions with an antagonist led to a prolonged retention time and a higher oviposition rate. By contrast, there was no evidence of learning in P. silvestris. Females of this species showed no behavioural change in response to ant encounters, and there was no difference in the foraging success of naive and experienced female parasitoids. Received: 7 December 1999 / Revised: 23 September 2000 / Accepted: 10 March 2000     

Zooid size parallels contemporaneous oxygen isotopes in a large colony of<Emphasis Type="Italic"> Pentapora foliacea</Emphasis> (Bryozoa)

The relationship between seasonal changes in temperature and the size of zooids within cheilostome bryozoans is explored by comparing zooid size and contemporaneous carbonate mineralogy within a single colony of a perennial species that grew in a highly seasonal environment. Previously published oxygen isotope profile data from two fronds of a large colony of Pentapora foliacea from the Irish Sea record the cyclical patterns related to seasonal changes in temperature experienced by the colony over 3 years of growth. Zooid size data gathered from the same points at which the oxygen isotope data were taken reveal that zooid size and contemporaneous ¹⁸O values covary. These and previously published data show that zooid size profiling can be confidently used to easily estimate rates of growth and longevity of bryozoan colonies, and further illustrate the negative relationship between zooid size and ambient temperature. Some inconsistencies in timing between zooid size and ¹⁸O data may provide insights into the mechanisms behind temperature mediated body size changes in poikilothermic animals.Communicated by P. W. Sammarco, Chauvin     

The red and the black: habituation and the dear-enemy phenomenon in two desert Pheidole ants

Many species of territorial animals are more aggressive toward strangers than neighbors, a pattern of aggression referred to as the ’dear-enemy phenomenon.’ In many cases, the mechanism by which neighbors are discriminated from strangers and the function of neighbor-stranger discrimination remain controversial. We investigated the spatial patterns of inter-colony aggression within and between two Pheidole species of seed-harvesting ants in the Mojave Desert of California by quantifying aggression between colonies in standardized staged encounters. We also tested whether the level of fighting between workers of two colonies is affected by previous exposure to each other. We show that neighbors (i.e., colonies less that 2.6 m away) of either species are treated less aggressively than more distant colonies and that habituation may be a mechanism by which this discrimination is achieved. The variation in aggression among spatially distant colonies also suggests that additional genetic or environmental factors are involved in recognition. The function of the dear-enemy phenomenon in these ant species may be related to the greater risk to the resources of a colony presented by strange workers than workers from a neighboring colony. Received: 18 November 1999 / Received in revised form: 3 April 2000 / Accepted: 3 May 2000     

Colony activity integration in primitively eusocial wasps: the role of the queen (Polistes fuscatus,Hymenoptera: Vespidae)

Summary The queen's role in colony activity integration in small post-emergence colonies of Polistes fuscatus was investigated in the field. We continuously recorded the behaviors of all wasps in (1) undisturbed colonies, (2) colonies from which the queen had been removed, (3) colonies from which a single worker had been removed, (4) colonies with a cooled, relatively inactive queen, and (5) colonies with a cooled, relatively inactive worker (29 colonies; 148 h observation).The queen spends more time on the nest, spends more of its nest time active, participates in more interactions/h, and initiates more interactions/h than does the average worker. Overall, the queen is involved in more interactions than is any other colony member. Queen removal depresses worker activity level and causes episodes of worker activity to become less temporally coupled (less synchronized).The presence of a cooled, inactive queen on the nest produces an even greater reduction in worker activity level and also results in decreased synchrony in worker activity episodes. Removal or cooling of a single worker produces no systematic changes in the activities of the other colony members. We conclude that the P. fuscatus queen is a central pacemaker and coordinator of colony activity.     

Light-enhanced growth of the ascidian Didemnum molle/Prochloron sp. symbiosis

To obtain a direct measurement of the importance of Prochloron sp. to the ascidian Didemnum molle, ascidian colonies from Lizard Island Lagoon, Great Barrier Reef, were grown for 9 d at 0, 10, 40 and 100% sunlight in situ using unidirectional flow chambers. Growth (wet weight) was enhanced up to 40% of full sunlight, at which point growth appears to have been light-saturated. Colonies in 10 and 40% sunlight responded by (1) climbing up the sides of the growth chambers, and (2) flattening out to a more encrusting morphology; also (3) the chlorophyll content of three colonies in zero sunlight decreased by >80%, yet the ascidians remained healthy and did not lose weight. These data show that although the symbiosis may not be obligatory for D. molle, the ascidian's growth is enhanced by Prochloron sp., and the morphology of the ascidian colony is affected by its photobiology.Contribution No. 344 of the Australian Institute of Marine ScienceCommunicated by G. F. Humphrey, Sydney     

Effects of sheltering fish on growth of their host corals

Stony corals are the foundation species of tropical reefs, and their structures can harbor a diverse range of mutualist taxa that can confer important benefits, including provision of nutrients. Prominent among the associates of branching coral in the genus Pocillopora are groups of zooplanktivorous damselfishes that take refuge in the coral to avoid their predators. In field and laboratory experiments, we explored the effects of colonies of resident damselfishes on growth of their host corals. Laboratory studies revealed a positive relationship between biomass of fish and output of ammonium. In the field, levels of ammonium were significantly elevated in the water surrounding the branches of Pocillopora occupied by colonies of damselfish, particularly in time periods following active feeding by the fish. Experimental manipulation of the presence of fish on host corals during a month-long field experiment revealed that corals hosting fish grew significantly more than those that lacked fish, and coral growth was positively correlated with the biomass of resident fish. The Pocillopora colonies in the field experiment varied in the degree of openness of their branching structure, and dye studies indicated that this affected their ability to retain waterborne nutrients. Together with biomass of resident fish, colony openness explained 76% of the variation in coral growth rate during the experiment. Corals can exhibit considerable morphological variability, and mutualistic fish respond to colony architecture during habitat selection, with some species preferring more open-branched forms. This makes it likely that corals may face tradeoffs in attracting resident fish and in retaining the nutrients they provide.     

Ammonium and free amino acid uptake by a deep-sea mussel (Bathymodiolus sp., undescribed) containing methanotrophic bacterial symbionts

The morphology of the lophophore and mouth was measured for freshly collected colonies of seven common bryozoan reef species at three sites along the central Caribbean coast of Panamá during the dry season, from the end of December 1988 through May 1989. Characters measured or scored include the number of tentacles, circumference of the lophophore, abanal and anal tentacle length, abanal and anal tentacle spacing, and perimeter of the mouth. Taxa studied include three species each of Stylophoma Levinsen, and Steginoporella (Smitt), and one species of Trematooecia Osburn. Individuals were collected from both continental and offshore fringing reefs from depths of 1 to>30 m. Terrestrial runoff from heavy tropical rains, sedimentation, and sea exposure vary greatly between sites. Congeneric species showed small but consistent differences in lophophore size, but little or no difference in spacing of tentacles or perimeter of the mouth. This suggests that closely related species feed on similarly sized particles despite their overall differences in zooidal size. Lopophore morphology also varied intraspecifically between reefs, particularly in the length of the tentacles and spacing between tentacles. Patterns of variation were similar for all species, which implies that species shift their diets in similar directions between the different reefs. The combination of these patterns suggests that niche diversification in feeding does not occur among congeneric species in the cheilostomes examined.     

Chemical camouflage by myrmecophilous beetles Zyras comes (Coleoptera: Staphylinidae) and Diaritiger fossulatus (Coleoptera: Pselaphidae) to be integrated into the nest of Lasius fuliginosus (Hymenoptera: Formicidae)

Summary. The myrmecophilous beetles, Zyras comes (Staphylinidae) and Diaritiger fossulatus (Pselaphidae) are guests of the black shining ant Lasius fuliginosus. Host worker ants never attacked these beetles, and often gave regurgitant to Z. comes following tactile communication with the beetle. By contrast, the workers from colonies without the myrmecophiles showed hostile responses towards Z. comes before tactile contact, but were not aware of D. fossulatus until contact. In L. fuliginosus, workers within a colony shared profiles, but the profiles differed among colonies. GC analyses showed that both Z. comes and D. fossulatus beetles had the same hydrocarbons as L. fuliginosus, and the profiles were more similar to those of the host colony workers than the foreign workers. Both Z. comes and D. fossulatus appear to imitate the hydrocarbon profile of their host workers, allowing integrating into the host nest. A Y-maze bioassay indicated that Z. comes can follow the trail pheromone of L. fuliginosus. This suggests that Z. comes may detect other chemical signals of L. fuliginosus to keep closer interactions with the workers. Received 22 June 2001; accepted 12 November 2001.