Social wasp parasites affect the nestmate recognition abilities of their hosts (<Emphasis Type="Italic">Polistes atrimandibularis</Emphasis> and <Emphasis Type="Italic">P. biglumis</Emphasis>, Hymenoptera,Vespidae)

Summary. Queens of the parasitic social wasp, Polistes atrimandibularis, temporarily mimic the odor of their host species, Polistes biglumis, but their offspring have parasite-specific odors. As a consequence, in parasitized colonies individuals with different odors co-inhabit the colony and host workers, who are responsible for colony defense, accept wasps with different odors. In order to verify whether this particular condition causes a change in recognition abilities of hosts, we tested nestmate/non-nestmate discrimination in field colonies invaded by social parasites (and in non-parasitized colonies as controls). Results show that parasitized colonies distinguish between nestmates and non-nestmates, distinguish their parasite queen from those that usurped alien colonies, and accept their parasite's non-mimetic offspring but make more recognition errors than non-parasitized colonies. The optimal acceptance threshold model predicts that when the frequency of encountering non-kin increases, residents become less permissive towards intruders. However, my data show that parasitized colonies are more permissive towards non-nestmates with respect to non-parasitized colonies but they are also more aggressive towards nestmates, suggesting that host workers' learning abilities are impaired.     

Inhibition of Host Queen Reproductive Capacity by the Obligate Social Parasite Polistes atrimandibularis (Hymenoptera,Vespidae)

Polistes atrimandibularis is the obligate and permanent parasite of the social paper wasp Polistes biglumis bimaculatus. This parasite lives together with the host foundress for a considerable period on the invaded nest. Ovarian development was measured in females of the host species (foundresses and, when present, workers) from 18 parasitized and 14 non-parasitized colonies. The reproductive capacity of foundresses from parasitized nests decreased faster than that of foundresses from non-parasitized nests. These results indicate that the presence of the parasite lowers reproductive capacity of the host queen. Simultaneously, the fertility of the host worker is inhibited.     

Sulcopolistes atrimandibularis,Social Parasite and Predator of an Alpine Polistes (Hymenoptera,Vespidae)1

Sulcopolistes atrimandibularis Zimmermann is the obligate social parasite of Polistes biglumis bimaculatus Geoffry in Furcroy, a mountain species of paper wasp. Unlike all the other hymenopteran social parasites, the Sulcopolistes female obtains part of the food for her immature brood by plundering other nests of the host species. Parasite females can control more than one host nest: one of them (the nursery nest) she uses solely for reproduction purposes (from which the Sulcopolistes reproductives emerge) and others (supply nests) are used for exploiting the Polistes brood content. It is possible that this behaviour is an adaptation to the extremely short colonial cycle of the host.     

Parasites of the Antarctic rattail <Emphasis Type="Italic">Macrourus whitsoni</Emphasis> (Regan, 1913) (Macrouridae,Gadiformes)

A total of 386 Macrourus whitsoni from Antarctic waters were examined for ecto- and endoparasites. Sixty-five M. whitsoni collected near Halley Bay (Weddell Sea) and 321 specimens from the continental slope off King George Island (South Shetland Islands) were studied for sphyriid copepods directly after being caught. A subsample of 25 specimens from the Weddell Sea and of 9 specimens from King George Island were studied for the presence of other metazoan parasites. Twenty-two species were found, including one myxozoan, six digeneans, one monogenean, three cestodes, seven nematodes, one acanthocephalan and three crustacean species/taxa. While Auerbachia monstrosa and Capillaria sp. are reported for the first time from around the Antarctic, the other parasites have been recorded earlier in the Southern Ocean. Many parasite species found have a wide zoogeographical range and a low host-specificity. The parasite fauna of M. whitsoni revealed several similarities with its congeners M. carinatus and M. holotrachys from Antarctic and sub-Antarctic waters. This can be explained by a wide host range of many macrourid deep-sea parasites, together with an overlap in distribution patterns of their hosts. Other supporting factors are host migration and a close phylogenetic relationship between the hosts, which enable the parasites to infest all three macrourids. Eight new host and 14 new locality records are established.     

Species and Colony Components in the Recognition Odor of Young Social Wasps: Their Expression and Learning (Polistes biglumis and P. atrimandibularis; Hymenoptera: Vespidae)

In Polistes, nestmate recognition relies on the learning of recognition cues from the nest. When wasps recognize nestmates, they match the template learned with the odor of the encountered wasp. The social wasp Polistes biglumis use the homogeneous odor of their colony to recognize nestmates. When these colonies become host colonies of the social parasite P. atrimandibularis, colony odor is no longer homogeneous, as the parasite offspring have an odor that differs from that of their hosts. In trying to understand how the mechanism of nestmate recognition works in parasitized colonies and why parasite offspring are accepted by hosts, we tested the responses of resident Polistes biglumis wasps from parasitized and unparasitized colonies to newly emerged parasites and to nestmate and non-nestmate conspecifics. The experiments indicate that immediately upon eclosion both young parasites and young hosts lack a colony odor and that colony odor can be soon acquired from the accepting colony. In addition, while residents of nonparasitized colonies recognize only the odor of their species, resident hosts of parasitized colonies have learned a template that fits the odors of two species.     

Mammal density and patterns of ectoparasite species richness and abundance

Patterns of species richness, prevalence and abundance of ectoparasites have rarely been investigated at both the levels of populations and species of hosts. Here, we investigated the effects in changes in small mammal density on species richness, abundance and prevalence of ectoparasitic fleas. The comparative analyses were conducted for different small mammal species and among several populations during a long-term survey. We tested the hypothesis that an increase in host density should be linked with an increase in parasite species richness both among host species and among populations within host species, as predicted by epidemiological models. We also used host species density data from literature. We found that host density has a major influence on the species richness of ectoparasite communities of small mammals among host populations. We found no relationship between data of host density from the literature and parasite species richness. In contrast with epidemiological hypotheses, we found no relationships between abundance, or prevalence, and host density, either among host species or among host populations. Moreover, a decrease in abundance of fleas in relation with an increase in host density was observed for two mammal species (Apodemus agrarius and A. flavicollis). The decrease or the lack of increase in flea abundance in relation with an increase in host density suggests anti-parasitic behavioural activities such as grooming.     

Prevalence of the parasite Strepsiptera in <Emphasis Type="Italic">Polistes</Emphasis> as detected by dissection of immatures

Summary. hough the paper wasp genus, Polistes, is well studied, we know little of the incidence of parasitism in this group. Here we present details of 45 nest dissections for 4 species: P. dominulus (Christ), P. gallicus (L.), P. stabilinus Richards and P. carnifex (F.) to detail levels of parasitism of colony members by the obligate parasitic group of insects, the Strepsiptera. All 4 species showed evidence of parasitism among immature members. For 3 species, more than 50% of inspected nests were parasitized and the levels of parasitism among brood (larvae and pupae) was very high and did not differ significantly between parasitized nests. One species, P. stabilinus, suffered very low levels of parasitism, which may be related to its habitat choice. The number of parasites per host was positively related to the proportion of infected brood (parasite prevalence) and in some cases reached phenomenally high levels, which casts doubt on previ ously assumed mechanisms of infection for nest-making Hymenoptera, i.e. phoresy. We also document cases of egg parasitism and encapsulation in Polistes nests. Our data show that parasitism levels greatly varied among areas. Finally, the recent debate on the competitive advantage of P. dominulus in its introduced range, USA, has credited an absence of strepsipteran parasites of this species in facilitating its spread. For the first time, we document levels of parasitism for this species in its nature P range and this would appear to corroborate previous claims. We place our work in the context of other studies of parasitism of social insects and posit that the genus Polistes may have much to offer to this field.     

Coexistence of helminth species in Lysapsus limellus (Anura: Pseudidae) from an Argentinean subtropical area: influence of biotic and abiotic factors

The need for studies on helminth communities of South American amphibians was addressed by examining changes in composition and population dynamics of the helminth component and infracommunities in the frog Lysapsus limellus Cope, 1862, from 1994 to 1996. Two pond types were considered, one permanent and one semipermanent. The main goals of this study were (1) to investigate the relationships between pond type, season, study time, frog body size, and frog sex and the presence-absence and counts of helminth parasite species in the frog host, L. limellus, and (2) to examine the co-occurrence of the different parasite species in the frog host in terms of the structure, assembly, and dynamics of the helminth infracommunity. Parasite presence and absence were analyzed using logistic regression and parasite counts were analyzed using log-linear modeling. To examine the association between parasite species, a principal components analysis was carried out on the correlation matrix for the counts. The pattern of co-occurrences was also examined through a randomization test. The main results were as follows: (1) five parasite species were found in the infracommunity; (2) the pond type and the host size class were the principal factors related to the presence and absence of parasites for the three dominant species, while the year of study and the host sex were only important for one of the species; (3) for the parasite counts, many more factors were significant, with body size class and pond type always important factors for the three core species of the infracommunity, with season, year of study, and sex also sometimes important, and (4) strong associations were observed between some helminth species of the host from the permanent pond, but the same species did not co-occur in frogs in the semipermanent pond. The life histories of the five species can be considered as opportunistic or "r strategists."     

Absence of measurable malaria-induced mortality in western fence lizards (Sceloporus occidentalis) in nature: a 4-year study of annual and over-winter mortality

Theoretical models of parasite virulence often quantify virulence by mortality. However, there is a lack of empirical studies of parasite-induced host mortality because it is often difficult to quantify in natural populations. I have estimated annual and over-winter mortality in a population of fence lizards (Sceloporus occidentalis) infected with a malaria parasite, Plasmodium mexicanum, in northern California. The duration of time a lizard was observed (an estimate of life-span) throughout the 4-year observation period, or following winter, was not related to either infection status or maximum parasitemia. In contrast to previous laboratory studies of this parasite-host system, I found no evidence of parasite-induced host mortality in nature.     

Larval niche selection and evening exposure enhance adoption of a predacious social parasite, Maculinea arion (large blue butterfly), by Myrmica ants

Larvae of the butterfly genus Maculinea feed briefly on a foodplant before being adopted as social parasites into Myrmica ant nests. Each Maculinea species typically survives only with a single Myrmica species, yet the eggs are laid across the overlapping territories of 3-5 Myrmica species and several other ants. The ability of Maculinea arion - a 'predatory' species of Maculinea - to influence its adoption into host Myrmica colonies was studied for the first time in the field. Some earlier reports, involving captive non-host ants, suggested that larvae of the predatory Maculinea follow ant trails into host nests or wander some distance from their foodplant before being discovered and (after a long interaction) carried away by Myrmica foragers. No dispersal from foodplants occurred in wild Maculinea arion larvae. Instead, they increased by >100-fold their probability of being discovered and adopted by Myrmica spp., rather than by ants of other genera, by exposing themselves in the micro-niche occupied by Myrmica foragers at their time of day of peak foraging. Despite a complex, hour-long interaction with Myrmica workers before being carried to the nest, Maculinea arion did not enhance its adoption by host species of Myrmica. Eggs were laid without bias in Myrmica sabuleti (host) and Myrmica scabrinodis (non-host) territories; larval survival on Thymus was the same in both ants' territories; larvae waited to be found beneath their foodplant rather than seek their host; Myrmica sabuleti and Myrmica scabrinodis foraged in the same vertical and temporal niches, and had the same probability of discovering larvae; both ants behaved identically after finding larvae and took the same time to adopt them; and the ratio of wild larvae taken into Myrmica sabuleti or Myrmica scabrinodis nests was the same as the distribution between these ants of Thymus, eggs and pre-adoption larvae.     

Prevalence and impact of a virulent parasite on a tripartite mutualism

The prevalence and impact of a specialized microfungal parasite (Escovopsis) that infects the fungus gardens of leaf-cutting ants was examined in the laboratory and in the field in Panama. Escovopsis is a common parasite of leaf-cutting ant colonies and is apparently more frequent in Acromyrmex spp. gardens than in gardens of the more phylogenetically derived genus Atta spp. In addition, larger colonies of Atta spp. appear to be less frequently infected with the parasite. In this study, the parasite Escovopsis had a major impact on the success of this mutualism among ants, fungi, and bacteria. Infected colonies had a significantly lower rate of fungus garden accumulation and produced substantially fewer workers. In addition, the extent of the reduction in colony growth rate depended on the isolate, with one isolate having a significantly larger impact than two others, suggesting that Escovopsis has different levels of virulence. Escovopsis is also spatially concentrated within parts of ant fungus gardens, with the younger regions having significantly lower rates of infection as compared to the older regions. The discovery that gardens of fungus-growing ants are host to a virulent pathogen that is not related to any of the three mutualists suggests that unrelated organisms may be important but primarily overlooked components of other mutualistic associations.     

Effects of familiarity and nestmate number on social interactions in two communal bees, Andrena scotica and Panurgus calcaratus (Hymenoptera, Andrenidae)

Summary: Communality is considered a widespread form of social organization in bees and wasps, one in which two or more females of the same generation share a common nest and amongst whom there is no reproductive division of labor. The evolution and maintenance of egalitarian communal societies poses theoretical difficulties where 'cheating' amongst nestmates is practised. Yet there is little knowledge of the intranidal social interactions among communal conspecifics that allow an understanding of the degree of cooperation versus exploitation amongst nestmates in such societies. Using a circle-tube arena, we staged interactions in the laboratory among pairs of female conspecifics using two communal andrenid species, Andrena scotica and Panurgus calcaratus, to score their cooperative and aggressive behaviors. For both species, females did not discriminate between familiar (nestmate) and unfamiliar (non-nestmate) conspecifics in their cooperative and aggressive behaviors, suggesting that there is universal acceptance of conspecifics. Levels of cooperation and aggression did not vary with the number of nestmates in the nest of origin for either species. In addition, frequencies of social interactions did not vary with body size, wing wear or ovarian development in P. calcaratus. Interspecific comparisons were made with published data on social interactions of other bee species at varying levels of social organization (solitary to eusocial) derived from analogous circle-tube experiments. Panurgus calcaratus displayed highly cooperative behavior, similar to that of a previously studied communal bee, Lasioglossum hemichalceum (Kukuk, 1992a). Andrena scotica displayed lower levels of cooperation, and was in this sense more similar to solitary species and to interactions amongst non-nestmate individuals of eusocial species. Both A. scotica and P. calcaratus displayed very low levels of aggressive behavior, again similar to L. hemichalceum. As a working hypothesis, we suggest the existence of a communal behavioral syndrome comprising high levels of intranidal cooperation and low aggression, both directed indiscriminately at conspecifics.     

Comparative morphology of Van der Vecht's organ in Polistes social parasites: host ecology and adaptation of the parasite

Camouflage strategies are common in insect social parasites. Being accepted into an alien colony as a dominant nestmate favours behavioural and morphological adaptations to mimic a specific odour. In Polistes social parasites, abdominal tegumental glands are involved in this camouflage strategy. These glands secreting cuticular hydrocarbons are connected with a modified cuticular area of the last gastral sternite of female wasps, named Van der Vecht's organ, whose secretion is involved in rank and dominance recognition. The size of this exocrine area has been demonstrated to be under selective pressure in Polistes, as a response to an efficient dominance recognition. Because chemical and behavioural integration differs between parasitic species, we carried out a comparison of Van der Vecht's organ size between the three Polistes social parasites and their respective hosts. The parasites Polistes sulcifer and Polistes semenowi, capable of a rapid chemical mimicry and specialized to exploit a lowland host, also show an enlarged Van der Vecht's organ. Conversely, the parasite Polistes atrimandibularis, specialized on a mountain species and showing a slow chemical integration, has a smaller organ. The time available for the parasite to tune up its chemical mimicry, before the emergence of workers to be accepted as a dominant nestmate, appears to be the most important selective pressure acting on the size of this abdominal organ. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013 , 109 , 313–319.     

Polyrhachis lama, a parasitic ant with an exceptional mode of social integration

Social parasitism is a common phenomenon amongst ants that occurs in manifold variations with differing levels of parasite–host integration. Particularly, high levels of social integration occur amongst closely related species (Emery’s rule), which form mixed colonies with their hosts and comprise the vast majority of social parasites. Considerable lower levels of integration are typically found amongst unrelated species that live in clearly separated colonies. The formicine ant Polyrhachis lama, however, parasitises a phylogenetically distant host species, Diacamma sp. of the subfamily Ponerinae, but lives spatially mixed with the host colonies. Studies on integration and communication have indicated that P. lama shows a high degree of host integration. However, the allocation of brood care behaviour, a central aspect of parasite integration, has not been studied. Because all known ant social parasites that are fully mixed with their host colonies are also true brood parasites, we investigated the integration of P. lama brood. Our results demonstrate that the parasite brood has a high degree of spatial integration, although it remains functionally separated regarding nutritive brood care. This can be attributed to behavioural and morphological differences between the phylogenetically distant species. The observed spatial confinement of parasite brood, however, is most likely due to an unusual method of chemical host integration. The parasite brood remains accepted in the Diacamma colonies only under the presence of adult parasites. Altogether, this suggests an active mechanism of chemical integration based on the acceptance allomones originating from P. lama workers. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Mutualistic interaction between a weevil and a rust fungus, two parasites of the weed Cirsium arvense

We present a mutualism between a stem-boring weevil, Apion onopordi Kirby (Coleoptera: Apionidae), and a rust fungus, Puccinia punctiformis (Str.) Röhl. (Uredinales), both parasites of the creeping thistle, Cirsium arvense (L.) Scop. (Asteraceae). Females, but not males, of A. onopordi induced systemic rust infections of thistle shoots in the season after they were attacked by the weevil, indicating that insect oviposition is a crucial stage in pathogen transmission. Adult weevils emerged from systemically infected thistle shoots were heavier than weevils from healthy C. arvense shoots. Heavier females had a higher fecundity and laid larger eggs. The weevil preferred to deposit eggs in systemically rust-infected over healthy thistle shoots, which seemed to be a sub-optimal host. This is to our knowledge the first report of a mutualistic interaction between an herbivorous insect and a biotrophic plant pathogen. The mechanism responsible for the advantage of rust-infected shoots for A. onopordi causes a different outcome in other thistle herbivores, and therefore can not be explained by a general enhancement of nutritional quality in rust-infected tissue. This mutualism likely has evolved from a competitive relationship. Unlike other thistle herbivores A. onopordi seems to be better suited as mutualist for P. punctiformis because of its small impact on the host plant and its feeding niche on plant parts not directly associated with pathogen reproduction.     

Raid process, activity pattern and influence of abiotic conditions in the slave-making ant Rossomyrmex minuchae (Hymenoptera, Formicidae)

Summary: In the present work, we present the results of a study on the seasonal and daily activity, as well as on the raid process in Rossomyrmex minuchae. A comparative study of the annual and daily activity of the host species, Proformica longiseta, has also been made in parasitized and non-parasitized colonies. R. minuchae is an obligate slave-maker, in which the annual activity is strongly related to abiotic conditions, especially temperature and moisture of the soil. The raid process is comprised of three phases: scouting, assault of the host nest and robbing of the host brood. Recruitment of nestmates by mean of adult transport, concentration close to and invasion of the host nest, are the three tasks carried out by R. minuchae workers during the assault phase. Major differences between Rossomyrmex and Polyergus (the other obligate dulotic genus of the tribe Formicini) in terms of behaviour, both in nestmate recruitment and in the host-nest invasion, reflect different parasitic strategies in the two genera, supporting the assignment of these two genera to different phylogenetic groups.     

Variation in hostplant preference and performance by the milfoil weevil, Euhrychiopsis lecontei Dietz, exposed to native and exotic watermilfoils

Exotic plants often displace native plants and thus alter the availability of native hostplants for specialist herbivorous insects. The submersed aquatic weevil Euhrychiopsis lecontei Dietz is endemic to North America, but there are now source populations on the exotic Eurasian watermilfoil (Myriophyllum spicatum L.) as well as on the weevil's ancestral host, northern watermilfoil (Myriophyllum sibiricum Komarov). This provides an opportunity to examine a host range expansion in progress. To further define the host range of the weevil and to determine how population source and rearing plant influence host plant preference and performance, we conducted a series of preference and rearing experiments with weevils from two source populations reared on northern milfoil, on Eurasian milfoil, switched late in larval development from northern to Eurasian milfoil, and vice versa. We also included two rearing treatments with milfoils on which the weevil has not been documented: the native M. verticillatum L. and the exotic M. aquaticum Verd. Preference by weevils in the switched rearing treatments was similar to preferences exhibited by weevils reared solely on the second (later) milfoil species and an increase in preference for Eurasian milfoil was induced by adult exposure to Eurasian milfoil for 2 weeks. In contrast, sizes and development times of weevils in the switched rearing treatments were similar to sizes and development times exhibited by weevils reared solely on the first (early) milfoil species. These results indicate that preference by the milfoil weevil is determined late in larval development or later and Hopkins' host selection principle is not supported. However, size and development time were most affected by hostplant quality during larval development when larvae must acquire the resources needed for pupation. Oviposition preference in the milfoil weevil was a population attribute, not a fixed individual attribute and there was no significant variation in preference among individuals reared on northern milfoil, but significant variation in preference was detected among weevils reared on Eurasian milfoil. Weevils oviposited on all four milfoil species and completed development on three of them, but did not develop beyond the larval stage on M. aquaticum. Weevils reared on Eurasian milfoil developed faster and reached larger adult sizes than weevils in any other rearing treatment. The smallest sizes and longest development times were for weevils reared on the natives, northern milfoil and M. verticillatum. The milfoil weevil oviposits on an array of milfoil species and is unable to distinguish an unsuitable host (M. aquaticum) within this genus. The influence of rearing plant and adult exposure to Eurasian milfoil on hostplant preference suggests that host range expansion to novel congeners may occur more rapidly than predicted by models which assume that genetic variation is required. Significant variation among individuals in hostplant preference suggests the potential for a host shift to a plant for which E. lecontei appears pre-adapted.     

Sucrose phosphate synthase in leaves of mistletoe: its regulation in relation to host (<Emphasis Type="Italic">Abies alba</Emphasis>) and season

Sucrose-phosphate synthase (SPS; E.C. 2.4.1.14) was studied in 1-year-old leaves of the xylem-parasitic mistletoe (Viscum album L.), growing on Abies alba. Glucose-6-phosphate served as an allosteric activator of mistletoe SPS, increasing the affinity for both substrates, fructose-6-phosphate and UDP-glucose. The activation state of SPS, i.e. the ratio of substrate limited versus non-limited activity, showed two clear peaks between February and July which coincided with increased rates of net photosynthesis of the parasite. Periods of decreased SPS activity were accompanied by a transient accumulation of sucrose and starch. In samples exhibiting a high activation state, activity was decreased by incubation of the extract with ATP; however, ATP did not affect SPS activity in samples exhibiting a low activation state of SPS. In parallel to the first increase of the activation state in March, pool sizes of the positive effector glucose-6-phosphate were high, whereas pool sizes of fructose-2,6-bisphosphate, an inhibitor of sucroneogenesis, were low. The decline in the activation state in April occurred in parallel with increased rates of transpiration of the parasite. This could have increased the availability of host-derived sugars, although the xylem sap of A. alba showed rather consistent concentrations of total soluble sugars throughout the vegetation period (1.1-3.9 mM). We thus speculate that sugar availability in the host xylem controls carbohydrate metabolism in the parasite.     

Burning bridges: priority effects and the persistence of a competitively subordinate acacia-ant in Laikipia, Kenya

Theoretical models suggest that subordinate competitors may rely on strong colonization ability and/or high persistence (e.g., the ability to resist invasion) as a strategy to coexist with competitively dominant species. While strong colonization ability among subordinate competitors has been widely documented, we know less about the role of persistence in facilitating species coexistence. In upland East Africa, four species of acacia-ants (Crematogaster sjostedti, C. mimosae, C. nigriceps, Tetraponera penzigi) compete for possession of Acacia drepanolobium host trees. Despite a strong dominance hierarchy, the four acacia-ant species coexist at fine spatial scales. Here we present evidence that T. penzigi, the least aggressive competitor, modifies host trees in two ways that reduce the probability of aggressive takeover by neighboring colonies. First, T. penzigi workers destroy virtually all leaf nectaries on their host trees. Second, T. penzigi workers create and maintain entryways into their swollen thorn domiciles that are too small to allow entry by their Crematogaster competitors. In a 2Ƕ factorial experiment, we manipulated nectar availability and swollen-thorn entryway size to determine the influence of these factors on the probability of aggressive displacement by a dominant competitor (C. mimosae) in staged conflicts. Addition of artificial nectaries and enlargement of swollen-thorn entryways on T. penzigi-occupied trees increased the probability of aggressive displacement of T. penzigi by C. mimosae from swollen thorns 14-fold and 8-fold, respectively. Further, empty saplings with nectaries destroyed by T. penzigi workers were colonized by half as many C. mimosae workers as saplings where nectaries were left intact. Our results demonstrate that T. penzigi's unusual strategy of nectary destruction and the maintenance of small entryways in swollen thorns produce priority effects, effectively reducing the probability that T. penzigi colonies will be displaced from host trees by more dominant competitors.