The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants

Abstract. The ant subfamily Pseudomyrmecinae comprises three genera of hyperoptic, arboreal ants, widely distributed in tropical and subtropical regions: Pseudomyrmex (∼200 species, New World), Myrcidris (two species, South America) and Tetraponera (∼100 species, Palaeotropics). The phylogenetic relationships among these ants were investigated using DNA sequence data (∼5.2 kb from 18S rDNA, 28S rDNA, wingless, abdominal-A, and long-wavelength rhodopsin genes) and 144 morphological characters, both separately and in combination. Data were gathered from a representative set of forty-nine pseudomyrmecine species, plus eighteen species from various outgroups. There was substantial agreement among the results obtained from different datasets, and from different methods of phylogenetic inference (parsimony, Bayesian inference). The monophyly of the following groups is strongly supported (100% bootstrap support and 1.00 posterior probability in the molecular dataset): Pseudomyrmecinae, Pseudomyrmex, and Pseudomyrmex + Myrcidris. The status of the genus Tetraponera is less clear: the DNA sequence data indicate that the genus is paraphyletic, but morphological features and a unique insertion in the 28S gene support the monophyly of this taxon. Seven of nine Pseudomyrmex species groups, established previously on the basis of morphology alone, are strongly upheld, but monophyly is rejected for the P. pallens group and the P. viduus group. In the latter case, molecular evidence indicates the existence of two independent clades, associated with the ant-plants Triplaris and Tachigali, respectively, whose convergent morphological features had caused them to be placed erroneously in the same species group. The present results confirm an earlier assertion that obligate associations with domatia-bearing plants have arisen at least twelve times in the subfamily. Molecular and morphological data support the hypothesis of a sister-group relationship between Pseudomyrmecinae and Myrmeciinae (84% parsimony bootstrap, combined dataset), which implies a Cretaceous origin of the stem-group pseudomyrmecines in the southern hemisphere. Pseudomyrmecines appear to have arisen in the Palaeotropics and later dispersed from Africa to South America, where they experienced a pronounced burst of diversification.     

Macroevolutionary assembly of ant/plant symbioses: Pseudomyrmex ants and their ant-housing plants in the Neotropics

Symbioses include some of the clearest cases of coevolution, but their origin, loss or reassembly with different partners can rarely be inferred. Here we use ant/plant symbioses involving three plant clades to investigate the evolution of symbioses. We generated phylogenies for the big-eyed arboreal ants (Pseudomyrmecinae), including 72% of their 286 species, as well as for five of their plant host groups, in each case sampling more than 61% of the species. We show that the ant-housing Vachellia (Mimosoideae) clade and its ants co-diversified for the past 5 Ma, with some species additionally colonized by younger plant-nesting ant species, some parasitic. An apparent co-radiation of ants and Tachigali (Caesalpinioideae) was followed by waves of colonization by the same ant clade, and subsequent occupation by a younger ant group. Wide crown and stem age differences between the ant-housing genus Triplaris (Polygonaceae) and its obligate ant inhabitants, and stochastic trait mapping, indicate that its domatium evolved earlier than the ants now occupying it, suggesting previous symbioses that dissolved. Parasitic ant species evolved from generalists, not from mutualists, and are younger than the mutualistic systems they parasitize. Our study illuminates the macroevolutionary assembly of ant/plant symbioses, which has been highly dynamic, even in very specialized systems.     

Ant colonization of Maieta guianensis seedlings,an Amazon ant-plant

The ants Pheidole minutula and Crematogaster sp. are obligate inhabitants of the ant-plant Maieta guianensis. They nest and reproduce exclusively in this and a few other Amazon melastome ant-plants. Experimental transplants of uncolonized M. guianensis seedlings to sites at different distances from established colonies of these two ant species, which are sources of founding queens, have shown that distance is an important factor affecting seedling colonization by ants. The proportion of colonized seedlings and the average number of colonizations per seedling, both by Crematogaster sp. and P. minutula, decreased as distance from established colonies increased. Seedling colonization was also affected by rainfall and fewer seedlings were colonized during the dry season, especially by P. minutula, than during the rainy season. P. minutula queens usually cooperate with each other during colony foundation, a behaviour not observed among founding queens of Crematogaster sp. Competition between these two ant species for host-plants appears to be strong as 63.6% of the seedlings colonized by Crematogaster sp. were also colonized by P. minutula during a 15-month observational period. However, there was generally an interval of a few months between colonizations, possibly giving the first species to colonize the plant a better chance of domination on this plant.     

Polyphagy and Obligate Myrmecophily in the Butterfly Hallonympha paucipuncta (Lepidoptera: Riodinidae) in the Neotropical Cerrado Savanna

This study analyzes host plant use and obligate myrmecophily in Hallonympha paucipuncta, an endemic butterfly of the Cerrado. Larvae of H. paucipuncta are polyphagous, using at least 19 species of plants in 10 families. All larvae found were being tended by Crematogaster ants. Spatial distribution of larvae and tending ants were strongly aggregated, suggesting an influence of ants on oviposition and/or larval survival. Implications of obligate myrmecophily in the evolution of polyphagy in Riodinidae are discussed.     

Dietary specialization in mutualistic acacia‐ants affects relative abundance but not identity of host‐associated bacteria

Acacia‐ant mutualists in the genus Pseudomyrmex nest obligately in acacia plants and, as we show through stable isotope analysis, feed at a remarkably low trophic level. Insects with diets such as these sometimes depend on bacterial symbionts for nutritional enrichment. We, therefore, examine the bacterial communities associated with acacia‐ants in order to determine whether they host bacterial partners likely to contribute to their nutrition. Despite large differences in trophic position, acacia‐ants and related species with generalized diets do not host distinct bacterial taxa. However, we find that a small number of previously undescribed bacterial taxa do differ in relative abundance between acacia‐ants and generalists, including several Acetobacteraceae and Nocardiaceae lineages related to common insect associates. Comparisons with an herbivorous generalist, a parasite that feeds on acacias and a mutualistic species with a generalized diet show that trophic level is likely responsible for these small differences in bacterial community structure. While we did not experimentally test for a nutritional benefit to hosts of these bacterial lineages, metagenomic analysis reveals a Bartonella relative with an intact nitrogen‐recycling pathway widespread across Pseudomyrmex mutualists and generalists. This taxon may be contributing to nitrogen enrichment of its ant hosts through urease activity and, concordant with an obligately host‐associated lifestyle, appears to be experiencing genomewide relaxed selection. The lack of distinctiveness in bacterial communities across trophic level in this group of ants shows a remarkable ability to adjust to varied diets, possibly with assistance from these diverse ant‐specific bacterial lineages.     

Ecology of an Improbable Association: The Pseudomyrmecine Plant‐ant Tetraponera tessmanni and the Myrmecophytic Liana Vitex thyrsiflora (Lamiaceae) in Cameroon1

In young individuals of the obligate myrmecophytic liana Vitex thyrsiflora, several species of ants and other arthropods compete for resources offered by the plant. In mature individuals, the only inhabitant is the ant species Tetraponera tessmanni, which is completely restricted to Vitex lianas as its sole host. Established colonies of this ant provide effective defense against herbivores. The association between V. thyrsiflora and T. tessmanni is unusual in two respects. First, the climbing life form is rare among myrmecophytes. Secondly, it is surprising that a pseudomyrmecine should be the obligate associate of a liana. Pseudomyrmecine plant‐ants often prune vegetation contacting their host plant. This behavior functions in part to protect against invasion of the host by ecologically dominant ants. In contrast, T. tessmanni does not prune and is associated with a plant whose success, and thus that of its resident ant colony, depends on contacts with many other plants. Several traits of V. thyrsiflora and T. tessmanni combine to make the colonization of host plants by potential competitors very difficult. These include behavioral and morphological filters restricting entrance into the plant and exploitation of the resources it can supply; plant anatomical organization that enables T. tessmanni workers to carry out all activities, except leaf patrolling, within a single, branched private nesting space within which all food resources offered by the plant are produced; and polygyny, permitting the colony to monopolize a large, rapidly growing and long‐lived territory.     

The importance of petiole structure on inhabitability by ants in Piper sect. Macrostachys (Piperaceae)

Several Central American species of Piper sect. Macrostachys have obligate associations with ants, in which the ant partner derives food and shelter from modified plant structures and, in turn, protects the plant against fungal infection and herbivory. In addition to these obligate ant-plants (i.e. myrmecophytes), several other species in Piper have resident ants only sometimes (facultative), and still other plant species never have resident ants. Sheathing petioles of sect. Macrostachys form the domatia in which ants nest. Myrmecophytes in sect. Macrostachys have tightly closed petiole sheaths with bases that clasp the stem. These sheathing petioles appear to be the single most important plant character in the association between ants and species of sect. Macrostachys . We examined the structure and variation of petioles in these species, and our results indicate that minor modifications in a small number of petiolar characters make the difference between petioles that are suitable for habitation by ants and those that are not.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 181–191.     

Molecular phylogeny and evolution of host-plant use in conifer seed chalcids in the genus Megastigmus (Hymenoptera: Torymidae)

Abstract. Phylogenetic relationships amongst Megastigmus species (Chalcidoidea: Torymidae) associated with conifer seeds were inferred from DNA sequence data. Twenty‐nine species of seed chalcids were analysed using two different genes, cytochrome b (mitochondrial DNA) and the D2 domain of the 28S ribosomal DNA. Maximum‐parsimony and maximum‐likelihood analyses showed that taxa formed two monophyletic groups, one clade comprising all species associated with Cupressaceae and Taxodiaceae hosts with the exception of Chamaecyparis, and the other clade composed of species associated with Pinaceae. Species infesting Cupressaceae and Taxodiaceae seemed to be specialized to particular host genera or even to be species specific, which was consistent with a taxonomic radiation following initial host adaptation. By contrast, Megastigmus species associated with Pinaceae appeared capable of shifting onto different congeneric species or even onto a new host genus, with their evolution apparently less constrained by plant association. We hypothesized that the Megastigmus group associated with Pinaceae may have a much higher invasive potential than that related to Cupressaceae. The study also confirmed the presence of invasive Nearctic species in the Palaearctic, and demonstrated the existence of a cryptic species complex.     

Slippery ant-plants and skilful climbers: selection and protection of specific ant partners by epicuticular wax blooms in Macaranga (Euphorbiaceae)

 In many ant-plant species of the genus Macaranga in South-East Asia, conspicuous blooms of epicuticular wax crystals cover the stem surface. We found that many ant species were unable to walk on these surfaces. Only the specific ant partners of glaucous Macaranga host plants were capable of moving on the slippery stems without difficulty. Therefore, the epicuticular coatings of Macaranga myrmecophytes appear to have a selective function and protect the associated ants against competitors. The epicuticular aggregates function as a physical barrier; no evidence of chemical repellence was found. The extent to which ”foreign” ant species are excluded from a tree strongly depends on inclination, diameter and length of the glaucous stem sections. The particular growth form of some glaucous Macaranga ant-plants enhances the influence of the wax barriers. The ant associates of glaucous and glossy Macaranga ant-plants (genera Crematogaster and Camponotus) differ strongly in their capacity to adhere to the glaucous stems. For this reason, the wax blooms in Macaranga can act as an ecological isolation mechanism for the sympiotic ants. Within the genus Macaranga, we find a high correspondence between the occurrence of glaucousness and obligatory ant association (50% in ant-plants; 6.7% in non-myrmecophytes). The genus Macaranga thus represents one of the few cases known so far where epicuticular wax crystals are likely to have evolved in relation to insects. Received: 2 January 1997 / Accepted: 9 June 1997     

Plant killing by Neotropical acacia ants: ecology,decision‐making,and head morphology

Mutualistic species often associate with several partners that vary in the benefits provided. In some protective ant–plant mutualisms, ants vary in the extent at which they kill neighboring vegetation. Particularly, in acacia ants (Pseudomyrmex), the area around the host tree that ants keep free from vegetation (“clearings”) vary depending on the species. This study assessed whether interspecific variation in clearing size corresponds to workers biting on plant tissue of different thickness. As expected, workers from species making the largest clearings bit more often on thicker plant tissues than workers from species making smaller clearings. Because head shape affects mandible force, I also assessed whether pruning on thick tissue in mutualistic ant species or being a predator in non‐mutualistic species correlated with broader heads, which yield stronger mandible force. The species with the broader heads were non‐mutualistic predators or mutualistic pruners of thick tissues, which suggest that pruning neighboring vegetation in non‐predatory species demands force even when the ants do not kill prey with their mandibles. The findings reveal that clearing size variation in mutualistic ant partners of plants can also be observed at the level of individual decision‐making processes among workers, and suggest that head morphology could be a trait under selection in protective ant–plant mutualisms. Abstract in Spanish is available with online material.     

The influence of ants on patterns of colonization and establishment within a set of coexisting lycaenid butterflies in a south-east Asian tropical rain forest

In Peninsular Malaysia ten species of lycaenid butterflies use leaf flushes or inflorescences of the legume tree Saraca thaipingensis as larval hostplant. Resource partitioning among these species is regulated by a complex mixture of patterns of interaction with ants. Females of obligately myrmecophilous species lay their eggs exclusively on trees colonized by their specific host ants. On trees colonized by weaver ants, only specialist mutualists adapted to these territorial ants are able to survive, while larvae of other species are killed. The formicine ant Cladomyrma petalae, which inhabits hollow twigs of the myrmecophytic hostplant, likewise precludes oviposition by female butterflies. Lycaenid larvae confronted with this ant species never survive, but one concealed feeding species (Jamides caeruleus) escapes removal due to the cryptic life-habits of the larvae. Two facultative myrmecophiles associate in a mutualistic way with a wide and largely overlapping range of ant genera which forage at the extrafloral nectaries of leaf flushes. One species (Cheritra freja) is not myrmecophilous, but is tolerated by all but the most territorial ants. Ant-dependent hostplant selection and egg-clustering characterize the obligate mutualists, whereas facultative myrmecophiles and the non-myrmecophile distribute their eggs singly over appropriate hostplants. Signals mediating caterpillar-ant communication are highly specialized in one obligate myrmecophile (Drupadia theda), but rather unspecific in four other species tested. Altogether our observations indicate that colonization and establishment of lycaenid butterflies on S. thaipingensis trees are governed by specializations as well as opportunistic use of resources (ants and hostplant parts). Therefore, the diversity of this species assemblage is maintained by deterministic as well as stochastic factors.     

Wedged between bottom-up and top-down processes: aphids on tansy

Abstract. 1. Many species of aphids exploit a single host‐plant species and have to cope with changing environmental conditions. They often vary greatly in abundance even when feeding on the same host. In a field experiment, the bottom‐up (plant quality/patch type frequency) and top‐down (ant attendance/predation) effects on the abundance of four species of aphids feeding on tansy (Tanacetum vulgare) were tested using a full factorial design. In addition, a model was used to examine these patch characteristics for their relative effects on the population dynamics and abundance of different aphid species. 2. Aphid numbers changed significantly depending on the quality of the host plant and the presence/absence of attending ants. The obligate myrmecophile, Metopeurum fuscoviride, was abundant on high‐quality plants, while on poor quality plants or on plants without attending ants these aphids did not survive until the end of the experiment. The facultative myrmecophiles, Aphis fabae and Brachycaudus cardui, and the unattended aphid species, Macrosiphoniella tanacetaria, all reached similar peak population densities, but M. tanacetaria did best in poor quality patches. 3. Natural enemies reduced aphid numbers, but those species feeding on high‐quality plants survived longer than those on poor‐quality plants, which existed only for a short period of time, especially when associated with ants. Losses due to migration of winged morphs and mortality caused by parasitoids were insignificant. 4. Varying the frequency of different patch types in a model indicates that different degrees of associations with ants are favoured in different environments. If the proportion of high‐quality patches in a habitat is large, obligate myrmecophiles do best. On increasing the number of poor‐quality patches, unattended species become more abundant. 5. The results suggest that, in spite of large species specific differences in growth rates, degree of myrmecophily or life cycle features, the temporal and spatial variability in top‐down and bottom‐up forces differentially affects aphid species and allows the simultaneous exploitation of a shared host‐plant species.     

Herbivory in a spider through exploitation of an ant–plant mutualism

Spiders are thought to be strict predators [1]. We describe a novel exception: Bagheera kiplingi, a Neotropical jumping spider (Salticidae) that exploits a well-studied ant–plant mutualism, is predominantly herbivorous. From behavioral field observations and stable-isotope analyses, we show that the main diet of this host-specific spider comprises specialized leaf tips (Beltian food bodies; Figure 1A) from Vachellia spp. ant-acacias (formerly Acacia spp.), structures traded for protection in the plant's coevolved mutualism with Pseudomyrmex spp. ants that inhabit its hollow thorns [2]. This is the first report of a spider that feeds primarily and deliberately on plants.     

Run,robber, run: parasitic acacia ants use speed and evasion to steal food from ant‐defended trees

Ants that are obligate plant associates protect their host against herbivores and aggressively defend the resources offered by the plant. Workers of Pseudomyrmex nigropilosus Emery (Hymenoptera: Formicidae), an acacia ant that parasitizes the mutualism by not defending the tree, are seen stealing food from other ant‐defended acacia trees. In the present study, hypotheses of evasion, chemical crypsis, chemical repellence and temporal activity patterns are tested in the field aiming to determine how P. nigropilosus enters other acacia trees, successfully circumventing the defence of the resident ants. When parasitic ants are stealing, resident ants are evaded by stopping walking, changing their walking direction or walking faster. Resident and parasitic workers have similar temporal activity patterns. Parasitic workers can walk 2.6‐fold faster compared with any of the three species of acacia‐ants from which they usually steal food. Behavioural assays suggest that P. nigropilosus do not have chemical repellence but that chemical crypsis may be involved in the evasion strategy. This last hypothesis needs to be explored further by chemical and olfactory analyses. The combination of speed and evasive reactions allows parasitic ants to access well‐defended acacia trees.     

Sperm ultrastructure of five species of the Neotropical ant genus Pseudomyrmex (Hymenoptera: Formicidae)

The seminal vesicles of adult males of five species of Pseudomyrmex were prepared for light and transmission electron microscopy. The Pseudomyrmex spermatozoa are long and slender with similar morphology. The head region has an acrosome and a nucleus. In all the studied species, two morphologically distinct types of acrosomal vesicles were observed, a long structure, as observed in all known ants, and a pear‐shaped one, never before observed in ants. The nucleus is elongated and both condensed and loose chromatin are present. The flagellum has an axoneme, a centriolar adjunct, two mitochondrial derivatives and two accessory bodies. The centriolar, the mitochondrial derivatives and the accessory bodies are similar to observations in most ant species that have been studied. The axoneme presents an uncommon 9 + 9 + 1 microtubule arrangement and the central microtubule has 13 protofilaments. The acrosomal dimorphism and the different levels of chromatin organization are exclusive characteristics of Pseudomyrmex. Furthermore, the 9 + 9 + 1 microtubule arrangement is different from all Hymenoptera, as well as from most insects, which present a 9 + 9 + 2 arrangement. These new morphological characters that are specific to Pseudomyrmex, are valuable synapomorphies of the genus and can be used in taxonomic characterization of the Pseudomyrmecinae subfamily and in phylogenetic analyses in Formicidae family.     

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.     

Two ways to be a myrmecophilous butterfly: natural history and comparative immature‐stage morphology of two species of Theope (Lepidoptera: Riodinidae)

Symbiotic interactions between butterfly larvae and ants, termed myrmecophily, require a range of behavioural and morphological adaptations (ant‐organs). Here, using light and scanning electron microscopy, we describe the complete life cycle of two species of Theope (Lepidoptera: Riodinidae) that have contrasting ways of life. Theope thestias larvae are facultatively tended by several ant species, whereas Theope pieridoides have obligate symbiotic interactions with Azteca ants that inhabit a myrmecophytic tree. Morphological differences associated with their different degrees of intimacy with tending ants are visible from hatching. In T. thestias, the untended first‐instar larva has elongated bifurcated setae and a few tiny perforated cupola organs (PCOs), whereas in T. pieridoides, the ant‐tended first instar has short dendritic setae, larger and more numerous PCOs, and functional tentacle nectary organs (TNOs). Throughout ontogeny, T. pieridoides always shows more conspicuous ant‐organs than T. thestias, with the exception of balloon setae, which are larger and more numerous in T. thestias. In addition, mature T. pieridoides larvae have an anterior set of ant‐organs, including a new type, here described and termed anterior glandular openings (AGOs). Based on the behavioural responses of ants in contact with these structures, a new interpretation for the mechanism whereby Theope larvae can manipulate the behaviour of their tending ants is proposed. Until now, three ecological syndromes can be defined among Theope species: (1) oligophagous larvae with facultative myrmecophily; (2) monophagous larvae with obligate myrmecophily; and (3) polyphagous larvae with obligate myrmecophily. These results suggest that differences in the degree of specificity in the ant–plant interactions may have an important role in the evolution of host‐plant use in Theope. © 2013 The Linnean Society of London     

Dynamics of the association between a long-lived understory myrmecophyte and its specific associated ants

Myrmecophytic symbioses are widespread in tropical ecosystems and their diversity makes them useful tools for understanding the origin and evolution of mutualisms. Obligate ant–plants, or myrmecophytes, provide a nesting place, and, often, food to a limited number of plant–ant species. In exchange, plant–ants protect their host plants from herbivores, competitors and pathogens, and can provide them with nutrients. Although most studies to date have highlighted a similar global pattern of interactions in these systems, little is known about the temporal structuring and dynamics of most of these associations. In this study we focused on the association between the understory myrmecophyte Hirtella physophora (Chrysobalanaceae) and its obligate ant partner Allomerus decemarticulatus (Myrmicinae). An examination of the life histories and growth rates of both partners demonstrated that this plant species has a much longer lifespan (up to about 350 years) than its associated ant colonies (up to about 21 years). The size of the ant colonies and their reproductive success were strongly limited by the available nesting space provided by the host plants. Moreover, the resident ants positively affected the vegetative growth of their host plant, but had a negative effect on its reproduction by reducing the number of flowers and fruits by more than 50%. Altogether our results are important to understanding the evolutionary dynamics of ant–plant symbioses. The highly specialized interaction between long-lived plants and ants with a shorter lifespan produces an asymmetry in the evolutionary rates of the interaction which, in return, can affect the degree to which the interests of the two partners converge.     

Phylogeny and floral hosts of a predominantly pollen generalist group of mason bees (Megachilidae: Osmiini)

Within the genus Osmia, the three subgenera Osmia, Monosmia, and Orientosmia form a closely‐related group of predominantly pollen generalist (‘polylectic’) mason bees. Despite the great scientific and economic interest in several species of this clade, which are promoted commercially for orchard pollination, their phylogenetic relationships remain poorly understood. We inferred the phylogeny of 21 Osmia species belonging to this clade by applying Bayesian and maximum likelihood methods based on five genes and morphology. Because our results revealed paraphyly of the largest subgenus (Osmia s.s.), we synonymized Monosmia and Orientosmia under Osmia s.s. Microscopical analysis of female pollen loads revealed that five species are specialized (‘oligolectic’) on Fabaceae or Boraginaceae, whereas the remaining species are polylectic, harvesting pollen from up to 19 plant families. Polylecty appears to be the ancestral state, with oligolectic lineages having evolved twice independently. Among the polylectic species, several intriguing patterns of host plant use were found, suggesting that host plant choice of these bees is constrained to different degrees and governed by flower morphology, pollen chemistry or nectar availability, thus supporting previous findings on predominantly oligolectic clades of bees. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 78–91.     

Differential host use in two highly specialized ant-plant associations: evidence from stable isotopes

Carbon and nitrogen stable isotopes were used to examine variation in ant use of plant resources in the Cecropia obtusifolia / Azteca spp. association in Costa Rica. Tissue of ants, host plants and symbiotic pseudococcids were collected along three elevation transects on the Pacific slope of Costa Ricas Cordillera Central, and were analyzed for carbon and nitrogen isotopic composition. Worker carbon and nitrogen signatures were found to vary with elevation and ant colony size, and between Azteca species groups. Ants in the A. constructor species group appear to be opportunistic foragers at low elevations, but rely more heavily on their host plants at high elevations, whereas ants in the A. alfari species group consume a more consistent diet across their distribution. Further, isotope values indicate that both ant species groups acquire more nitrogen from higher trophic levels at low elevation and when ant colonies are small. Provisioning by the host is a substantial ecological cost to the interaction, and it may vary, even in a highly specialized association. Nonetheless, not all specialized interactions are equivalent; where interaction with one ant species group appears conditional upon the environment, the other is not. Differential host use within the Cecropia-Azteca association suggests that the ecological and evolutionary benefits and costs of association may vary among species pairs.