Two ways to be a myrmecophilous butterfly: natural history and comparative immature-stage morphology of two species of Theope (Lepidoptera: Riodinidae (original) (raw)
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Austral Ecology, 2018
Females of myrmecophilous butterflies tend to oviposit in plants visited by ant species that engage in stable associations with its larvae. In Banisteriopsis malifolia, caterpillars are attended by the same ants that feed on extrafloral nectaries. A conflict may arise when both the plant and caterpillars compete for ant attention, and ants are assumed to forage on the highest quality resource. By attending caterpillars, ants can be indirectly detrimental to plant fitness because florivorous larvae feed intensively until pupation. In this study, we specifically investigated (i) whether the occurrence of facultative myrmecophilous Synargis calyce (Riodinidae) caterpillars in B. malifolia was based on ant species (Camponotus blandus or Ectatomma tuberculatum) and abundance; (ii) the monopolization of ants by the butterfly larvae and (iii) the florivory rates incurred by the caterpillars on inflorescences. The abundance of S. calyce was six-fold greater in plants with C. blandus, compared to E. tuberculatum treatments. Caterpillars monopolized up to 50% of C. blandus on the plants, indicating that the resources offered by S. calyce were more attractive to ants than extrafloral nectaries. Florivory by riodinids incurred losses of almost 60% of flower buds. Myrmecophilous riodinids exploited an ant-plant mutualism by attracting aggressive ants that become larvae bodyguards. Thus, this ecological interaction is potentially detrimental to B. malifolia, since the ants, which can provide protection against herbivores, shift to provide defence for one of these herbivores.
Arthropod-Plant Interactions, 2017
Ant-related oviposition in facultatively myrmecophilous lycaenid butterflies is common, but not universal. In fact, our knowledge of ant-related oviposition in lycaenids is based on some common species (e.g., Rekoa marius, Allosmaitia strophius, Parrhasius polibetes), which limits generalizations about these systems. In this study, we experimentally investigated whether the oviposition pattern of the florivorous lycaenid Leptotes cassius was influenced by the presence of Camponotus ants and whether larvae were attended, rather than attacked, by ants. This might be evidence of myrmecophily. Both L. cassius and Camponotus ants occur on Bionia coriacea, an extrafloral nectaried legume shrub that grows in the Brazilian cerrado. Plants were randomly assigned to ant-present and ant-excluded treatments and were observed twice throughout the short reproductive season. Larvae of L. cassius were tended by ants, whose attendance was characterized by active antennation on the last body segments of the caterpillars. Therefore, Camponotus can be considered a partner of L. cassius. Lycaenid abundance was on average 1.9-and 1.21-fold higher in plants with ants in each sampling period, respectively, indicating a tendency of L. cassius to occur in plants with ants. Nonetheless, results were not statistically significant, suggesting that in this case ants are not a major cue for lycaenid oviposition. In many ant-lycaenid mutualisms, butterfly immatures benefit from reduced parasitism rates. However, no L. cassius immature, regardless of ant presence or absence, was parasitized. Furthermore, larvae may occur inside flower buds that may provide protection from natural enemies; thus, ants may not be required for immature protection.
Naturwissenschaften, 2014
Ovipositing adult females of myrmecophilous lycaenids are expected to select plants based on ant presence in order to maximize the survivorship of immature stages. Usually, larvae feed ants with honey-like solutions and, in turn, ants ward off parasitoids. Nonetheless, a rarely investigated approach is whether ant partners can also extend their protective behavior towards lycaenids eggs. Here, we investigated the ant-related oviposition pattern of Allosmaitia strophius and Rekoa marius; then, we compared egg parasitism according to the presence of ants. Lycaenid oviposition and egg parasitism (in percent) were experimentally compared in ant-present and ant-excluded treatments. The study plant, Heteropterys byrsonimifolia, is an extrafloral nectaried shrub which supports several ant species. We sampled 280 eggs, of which 39.65 % belonged to A. strophius and 60.35 % to R. marius. Both lycaenids eggs were significantly more abundant on branches with ants, especially those with Camponotus crassus and Camponotus blandus, two ant species known to attend to lycaenids. A. strophius and R. marius parasitism was 4.5-and 2.4-fold higher, respectively, in ant-present treatments, but the results were not statistically significant. Our study shows that ant-mediated host plant selection in lycaenids might be much more widespread than previously thought, and not restricted to obligate myrmecophilous species. Tending ants may be inefficient bodyguards of lycaenid eggs, because unlike larvae which release sugared liquids, eggs do not offer obvious rewards to ants. Ants can ward off parasitoids of larvae, as observed elsewhere, but our findings show that positive ant-lycaenid interactions are conditional and depend on immature ontogeny.
Biological Journal of the Linnean Society, 2007
Plant ants generally provide their host myrmecophytes (i.e. plants that shelter a limited number of ant species in hollow structures) protection from defoliating insects, but the exact nature of this protection is poorly known. It was with this in mind that we studied the association between Tetraponera aethiops F. Smith (Pseudomyrmecinae) and its specific host myrmecophyte Barteria fistulosa Mast. (Passifloraceae). Workers bore entrances into the horizontal hollow branches (domatia) of their host B. fistulosa, near the base of the petiole of the alternate horizontal leaves. They then ambush intruders from the domatia, close to these entrances. After perceiving the vibrations caused when an insect lands on a leaf, they rush to it and sting and generally spreadeagle the insect (only small caterpillars are mastered by single workers). Among the insects likely to defoliate B. fistulosa, adult leaf beetles and large katydids were taken as prey and cut up; single workers then retrieved some pieces, whereas other workers imbibed the prey's haemolymph. Other insects known to defoliate this plant, if unable to escape, were killed and discarded. Small Acrea zetes L. caterpillars were stung and then discarded by single workers; whereas locusts of different sizes were mastered by groups of workers that stung and spreadeagled them before discarding them (although a part of their haemolymph was imbibed). More workers were involved and more time was necessary to master insects taken as prey than those attacked and discarded. Consequently, the protection T. aethiops workers provide to their host B. fistulosa from defoliating insects results from predation, but more often from a type of aggressiveness wherein insects are killed and then discarded.
Ethology, 2010
Myrmecophilous lycaenid larvae possessing a dorsal nectar organ release strong recruitment behavior in ants attending them. Larvae lacking a functional dorsal nectar organ are unable to release such recruitment behavior.ยท Thus, the dorsal nectar organ with its carbohydrate secretions is responsible for the release of the recruitment behavior. The importance of the dorsal nectar organ for the evolution of mutualistic trophobiotic associations between myrmecophilous lycaenid larvae and their attending ants is discussed.
Assessing the quality of different ant species as partners of a myrmecophilous butterfly
Oecologia, 2001
We assessed the quality of different ant species as partners of the facultatively myrmecophilous lycaenid butterfly Glaucopsyche lygdamus. We compared disappearance and parasitism rates of G. lygdamus larvae in the field, and development of non-feeding prepupae in the laboratory, when individuals were untended or tended by one of four ant species. Formica podzolica was the only ant species to provide a clear benefit to G. lygdamus, in the form of reduced larval parasitism relative to untended larvae. F. 'neogagates' (F. neogagates + F. lasioides) and Tapinoma sessile were essentially neutral partners, providing no significant cost or benefit for any of the parameters measured. Relative to untended individuals, association with F. obscuripes significantly increased larval disappearance and significantly decreased pupal mass. Thus, F. obscuripes may act as a parasite of the general association between G. lygdamus and ants under certain conditions. Ant species also differed in their persistence as tenders of G. lygdamus larvae once an interaction was established. Over the lifetime of a larva, F. podzolica and F. obscuripes usually remained as the attendant ant species on plants over consecutive census dates, while F. 'neogagates' and T. sessile were frequently replaced, most commonly by F. obscuripes. It remains to be determined if disappearance and developmental outcomes reported here reflect true fitness costs (i.e. reduced survivorship and lower reproduc-tive success) for G. lygdamus. The potential and limitations for specialization in association between G. lygdamus and high quality ant partners are discussed.
Declassifying myrmecophily in the Coleoptera to promote the study of ant-beetle symbioses
Psyche
The symbiotic associations between beetles and ants have been observed in at least 35 beetle families. Among myrmecophiles, beetles exhibit the most diverse behavioral and morphological adaptations to a life with ants. These various associations have historically been grouped into discrete but overlapping behavioral categories, many of which are still used in the modern literature. While these behavioral classifications provide a rich foundation for the study of ant-beetle symbioses, the application of these systems in future studies may be less than effective. Since morphological characteristics often provide the only information of myrmecophilous beetles, they should be studied in a species-by-species fashion, as behavioral data are often limited or unavailable. Similarly, behavioral studies should focus on the target species at hand, avoiding discrete classification schemes. I formally propose the rejection of any classification scheme, in order to promote future studies of myrmecophily in both taxonomic and evolutionary studies.
Looking for the ants: selection of oviposition sites by two myrmecophilous butterfly species
Animal Biology, 2008
Obligate myrmecophilous butterfl y species, such as Maculinea teleius and M. nausithous that hibernate as caterpillar in nests of the ant species Myrmica scabrinodis and M. rubra respectively, have narrowly defi ned habitat requirements. One would expect that these butterfl ies are able to select for sites that meet all their requirements. Both butterfl y species occur in habitats where their initial larval resource, the host plant Sanguisorba offi cinalis , is abundant, while the ant nests are relatively less abundant. With ants in the proximity of a host plant, caterpillars have a chance of being found by their host ant species, while the host plant could become a sink as caterpillars will die when there are no ants close to the host plant. We tested whether females oviposit on host plants in the close proximity of host ants (ant-mediated oviposition) or whether there is random oviposition, using 587 1-m 2 plots in and around a nature reserve in Th e Netherlands where both species were reintroduced. We found that females of both species indeed more often deposit eggs on host plants nearby ants than on host plants without ants. Visits of adult butterfl ies to plots with the host plant but without the host ants only occurred in years with high butterfl y densities. We argue that understanding site selection for oviposition is important for these highly endangered butterfl y species with a specialised life cycle, especially after disturbances or reintroductions.
Adults of the South-East Asian lycaenid butterfly Allotinus unicolor search for trophobiotic homopterans associated with a particular ant species, Anoplolepis longipes. The butterflies settle down for feeding in such associations after a characteristic pre-alighting ("examination flight") and appeasement ("proboscis palpation") behaviour. Preimaginal instars of A. unicolor were always related with the same ant species as well. A. unicolor uses a wide range of homopterans for oviposition. The orientation mechanisms for specific ant recognition (probably visual and olfactory), the selective value of specific ant-associations for Miletinae butterflies, the consequences of such strategies for the larval food range, and the evolution of adult myrmecophily within the genus Allotinus are discussed.