Effects of local and landscape factors on the abundance of an endangered multivoltine butterfly at riverbanks

Plebejus argyrognomon is one of the grassland‐dwelling butterflies undergoing rapid decline in recent decades. Grassland habitats for butterflies are generally threatened by fragmentation and invasive species, hence are among the most vulnerable ecosystems. We studied the seasonal abundance of P. argyrognomon at habitat patches along the banks of the Kinugawa River in eastern Japan, to identify environmental factors suitable for population persistence of this species, including habitat patch connectivity. Results showed that the patch's host plant cover had a positive effect on abundance in all three seasons, while the shading of the host plants by surrounding non‐host plants and nearby forested area showed negative effects. Additionally, habitat patch connectivity and nectar richness could be considered as positive factors in autumn and summer, respectively. Analysis of habitat connectivity also showed that the Kinugawa River did not appear to act as a dispersal barrier for P. argyrognomon. Our findings emphasize the importance of understanding environmental factors that may vary among seasons, and such understanding could contribute to habitat management of multivoltine butterflies in fragmented landscapes.     

Comparative study of the morphology of stridulatory organs of the Iberian lycaenid butterfly pupae (Lepidoptera)

We compared the pupal stridulatory organs of 35 species and one subspecies of Iberian Lycaenidae using scanning electron microscopy. The studied species belong to the tribes Theclini, Eumaeini, Lycaenini, and Polyommatini. Nine species do not show stridulatory organs on the pupae but all other species possess them. Stridulatory organs are formed by a stridulatory plate (pars stridens) placed on the fifth abdominal segment and a file (plectron) in the sixth abdominal segment. The plate has tubercles in the Theclini and Lycaenini, tubercles, ridges, or undulations in the Eumaenini, and tubercles, teeth, or unspecialized structures in the Polyommatini. Morphological differences can be found in the files of the different tribes, regarding the number of teeth, their form and size. Cuticular formations of the organs were studied on a surface of 2,500 µm² and the average of ridges, tubercles, and teeth was measured searching for relevant taxonomic information. Stridulatory organs were thought to be an adaptation to myrmecophily but we show that they are present both in myrmecophilous and nonmyrmecophilous species; therefore, we suggest that this trait probably did not evolve in relation with myrmecophily, but may be used to enhance relationships with ants. J. Morphol. 275:414–430, 2014. © 2013 Wiley Periodicals, Inc.     

Reinstatement of a New Zealand copper butterfly,Lycaena rauparaha (Fereday, 1877)

Abstract

Previous confusion over the nomenclature and status of the New Zealand copper butterflies is reviewed. Lycaena rauparaha n. comb., which has been regarded as a variety of L. salustius in recent literature, is reinstated and adult, larval, and pupal characters are described. Its distinctions from L. salustius and L. feredayi are tabulated and a key to this species complex is given.     

Mutualism between Thisbe irenea butterflies and ants, and the role of ant ecology in the evolution of larval-ant associations

A facultative mutualism between the riodinid butterfly Thisbe irenea and the ponerine ant Ectatomma ruidum is described from Panama. Ants protect larvae against attacks of predatory wasps, but not against tachinid parasitoids. Several potential sources of ecological variation affecting the larval survival of Thisbe irenea are noted. A preliminary means of testing the ability of larvae to appease ants is described that may be applied to all butterfly-ant systems. Observations and literature records indicate that ant taxa which tend butterfly larvae are the same taxa that tend extrafloral nectaries and Homoptera. A general hypothesis for the evolution of myrmecophily among butterflies suggests that ant taxa which utilize secretions in their diet are major selective agents for the evolution of the larval ant-organs, and hence, ant-larval mutualisms. This idea is extended to suggest how appeasement of predaceous ant taxa through the use of larval ant-organs can influence an ant-larval relationship, eventually leading to mutualism.     

Influence of atmospheric carbon dioxide enrichment on induced response and growth compensation after herbivore damage in Lotus corniculatus

Abstract 1. Plant growth and chemical defence compounds in four Lotus corniculatus genotypes exposed to factorial combinations of ambient and elevated carbon dioxide, and herbivory by caterpillars of Polyommatus icarus were measured to test the predictions of the carbon/nutrient balance hypothesis.
2. Shoot and root biomass, allocation to shoots versus roots, and carbon-based defence compounds were greater under elevated carbon dioxide. Pupal weight of P. icarus was greater and development time shorter under elevated carbon dioxide.
3. Herbivory decreased shoot growth relative to root growth and production of nitrogen-based defence (cyanide). Young leaves contained more defence compounds than old leaves, and this response depended on carbon dioxide and herbivory treatments (significant interactions).
4. Genotype-specific responses of plants to carbon dioxide and herbivory were found for the production of cyanide. Furthermore, maternal butterfly-specific responses of caterpillars to carbon dioxide were found for development time. This suggests the existence of genetic variation for important defence and life-history traits in plants and herbivores in response to rising carbon dioxide levels.     

Ants that associate with Lycaeninae butterfly larvae: diversity, ecology and biogeography

Abstract. Field data on associations of butterfly larvae with ants were collated for 435 species of the subfamily Lycaeninae to analyse patterns in the identity, diversity and ecology of attendant ants. Worldwide, members of at least fifty-three ant genera from six subfamilies associate with Lycaeninae larvae. All these ant genera also forage for other liquid carbohydrate food sources. Species of Crematogaster and Camponotus constitute by far the most important ant associates of Lycaeninae larvae. Diversity of butterfly–ant associations, as measured by Hurlbert rarefaction, is highest in the Oriental, Australian and Nearctic faunal regions, and lower in Africa and the Palaearctic. The Neotropical region could not be assessed separately due to paucity of data. Multiple regression analyses revealed that the number of Lycaeninae species associated with a particular ant genus in a facultatively mutualistic manner is largely determined by the ecological prevalence and geographical distribution, but does not correlate with species richness, of an ant genus. This opportunism is suggested to be the key variable supporting the geographically and taxonomically widespread occurrence of ant–caterpillar mutualism in the Lycaeninae. Obligate interactions are much more specific and almost exclusively involve ecologically dominant ants with long-lived colonies. Neither species richness nor geographical distribution of an ant genus are significant predictors. In such associations, the role of ants shifts from visitors to hosts, and patterns of host use and specificity resemble those known from inquiline ant guests.     

The significance of wing pattern diversity in the Lycaenidae: mate discrimination by two recently diverged species

Closely related species of lycaenid butterflies are determinable, in part, by subtle differences in wing pattern. We found that female wing patterns can act as an effective mate‐recognition signal in some populations of two recently diverged species. In field experiments, we observed that males from a Lycaeides idas population and an alpine population of L. melissa preferentially initiate courtship with conspecific females. A morphometric study indicated that at least two wing pattern elements were important for distinguishing the two species: hindwing spots and orange crescent‐shaped pattern elements called aurorae. We deceived male L. idas into initiating courtship with computer generated paper models of heterospecific females when these pattern elements were manipulated, indicating that the wing pattern elements that define the diversity of this group can be effective mate recognition signals.     

Effect of grazing intensity on the interaction between Shijimiaeoides divinus asonis (Matsumura, 1929) and its attendant ants

This study was carried out in the natural herbaceous grassland of Mt. Aso, which had been almost completely grazed, and which is subjected to routine grassland burning every spring (February or March) to conserve Shijimiaeoides divinus asonis (Matsumura, 1929). We clarify that ants protect the larvae of this butterfly and evaluate the effects of grazing intensity on the attendant ant population. The results obtained are summarized as follows: (i) Five species of attendant ant were identified, with the dominant species being Formica japonica (Motschulsky, 1866) and Camponotus japonicus (Mayr, 1866). (ii) The number of attendant ants decreased at night time; however, only one or two ants attended the larva until the following morning. (iii) Ten species of insect excluding ants and three species of spiders that approached the larva were recorded on the larval host‐plant. Formica japonica and C. japonicus fought off most newcomers of other insects including the natural enemies of these butterfly larvae. (iv) The number of ants in this butterfly habitat under regular grazing intensity was significantly higher than during low grazing intensity and non‐grazing periods. (v) A positive correlation was found between the number of attendant ants and the number of butterfly larvae on the host‐plant. We concluded that the interaction between this butterfly and attendant ants is one of facultative mutualism because the attendant ants protect the butterfly larva. Therefore, the numbers of this butterfly species may decrease if the number of attendant ants decreases due to the cessation of pasturage.