Natural history of an ant–plant–butterfly interaction in a Neotropical savanna

Non-myrmecophilous lepidopteran larvae using plants bearing ant attractants such as extrafloral nectaries are good models for studying morphological and behavioural mechanisms against ant predation. Udranomia spitzi (Hesperiidae) is a butterfly whose larvae feed on leaves of Ouratea spectabilis (Ochnaceae), a plant with extrafloral nectaries. We described the early stages of U. spitzi, and used field observations and experiments to investigate the defensive strategies of caterpillars against predatory ants. Larvae pass through five instars and pupation occurs inside larval leaf shelters. Ant-exclusion experiments revealed that the presence of ants did not affect significantly caterpillar survival. Predation experiments showed that vulnerability to ant predation decreased with increase in larval size. The present study showed that predatory ants are not as relevant as demonstrated for other systems, and also illustrates how observational data and field experiments can contribute to a better understanding of the biology and ecology of a species of interest.     

Ant–herbivore interactions in an extrafloral nectaried plant: are ants good plant guards against curculionid beetles?

Extrafloral nectary drinking ants are known as effective plant guards, but some herbivores may circumvent ant attacks by foraging on different plant parts or presenting adaptations to avoid ant predation. Here we experimentally investigated the effect of Camponotus blandus on the florivory of the extrafloral nectaried shrub Banisteriopsis malifolia; and a possible spatial segregation between ants and herbivores (leaves and flowers). Flower buds are attacked by Anthonomus weevils. Results revealed no significant influence of C. blandus on the reduction of florivory. Adult Anthonomus are hard-bodied and were immune to ant bites; larvae are endophytic, so protected from ants. Ants and adult beetles were concentrated in different plant parts (leaves and flowers, respectively) so restraining the probability of encounters. Our results indicate that the system C. blandus–Anthonomus–B. malifolia is not stable, as ants receive extrafloral nectar, but are unable to protect the plant against weevils.     

Do caterpillars of Dryas iulia alcionea (Lepidoptera,Nymphalidae) show evidence of adaptive behaviour to avoid predation by ants?

Plants bearing extrafloral nectaries can attract wasps, bugs, and mainly ants. Ants can defend plants against herbivores by treating the plant as an extension of their own territory. We evaluated the efficiency of resting sites built by Dryas iulia alcionea larvae as protection against ant predation on their host plant Passiflora suberosa. The results showed that ants on P. suberosa have significant influence on the survival of D. iulia caterpillars. The use and construction of resting sites was recorded through the fourth larval stadium. A strong inverse relationship between the number of resting sites constructed per larva and the developmental stage of the larvae was observed. Additional results suggest that the efficacy of resting sites against cursorial predators decreases with larval development. The behaviour of constructing resting sites probably evolved by natural selection. It is likely that this behaviour was important for the ancestral lineage that gave origin to Heliconiinae genera Dryas, Dryadula and Phylaethria.     

The tortoise caterpillar: carnivory and armoured larval morphology of the metalmark butterfly Pachythone xanthe (Lepidoptera: Riodinidae)

ABSTRACT

The natural history and morphology of the later instars and pupa of the metalmark butterfly Pachythone xanthe H. Bates are described and illustrated for the first time. The caterpillars are myrmecophilous and carnivorous, feeding on scale insects (Hemiptera: Coccoidea) tended by Azteca cf. chartifex Forel (Formicidae: Dolichoderinae) ants and appear to be physically shielded from potential ant attacks by a carapace-like body that protects the head and appendages. In addition, the larvae present several types of specialised setae and ant-organs such as perforated cupola organs, glandular openings and functional tentacle nectary organs that may be related to appeasement of aggressive ant behaviour. Our results reveal the first detailed life cycle data for the Pachythonina subtribe and record a new independent origin of carnivory in Riodinidae.     

Within‐nest abundance of a tropical cockroach Pseudoanaplectinia yumotoi associated with Crematogaster ants inhabiting epiphytic fern domatia in a Bornean dipterocarp forest

Many epiphytic ferns in tropical forest canopies have domatia that provide habitat for arboreal arthropods such as ants. Two species of fern that were predominantly occupied by Crematogaster difformis ants were collected from the canopy of a forest in Borneo. In the fern domatia, two cockroach species, Pseudoanaplectinia yumotoi and Blatta sp., were found to live with C. difformis. Pseudoanaplectinia yumotoi were found in large numbers and accounted for approximately 20% of the total arthropod population, while Blatta sp. comprised only 1% of the arthropods living in the domatia. Behavioural experiments were conducted to evaluate the response of C. difformis workers to newly introduced cockroaches of each species and to allospecific ant workers. Crematogaster difformis workers were highly aggressive and quickly attacked allospecific workers. Both cockroach species could fend off ant attacks; however, dead Blatta sp. were often attacked by the ants while dead P. yumotoi were infrequently bothered.     

Biological characterization of the obligate symbiosis between Acropyga sauteri Forel (Hymenoptera: Formicidae) and Eumyrmococcus smithii Silvestri (Hemiptera: Pseudococcidae: Rhizoecinae) on Okinawa Island,southern Japan

The ant Acropyga sauteri Forel has an obligate, mutualistic symbiosis with a mealybug, Eumyrmococcus smithii Silvestri, on Okinawa Island, southern Japan. The mealybugs live inside ant nests nearly all their lives, and the ants depend on them for food. Alate foundress queens carry mealybugs during their nuptial flights, using them to establish new colonies at new sites. However, important aspects of the symbiosis have not yet been elucidated. The present study characterizes the basic biology of the symbiosis and describes for the first time the morphologies of all growth stages of E. smithii. Our study suggests that E. smithii has only one nymphal stage, followed by a female pupal stage or male prepupal stage. Intensive sampling of ant nests across seasons showed that A. sauteri prefers nest sites 5–20 cm underground. Acropyga sauteri produced reproductive stages mainly in mid‐March or early April, and numbers of both ant workers and mealybugs increased from spring to summer. Experimental determination of colony identity with a method using nestmate recognition by ants suggested that each ant colony rarely has a perimeter greater than 30 cm, that the ants are monogynous, and that different ant colonies are densely aggregated along the root system of a plant, adjacent to each other but not interflowing. Both symbiotic partners were vulnerable to attacks by several common subaerial ant species following physical disturbance to their nests.     

Size,spines and crochets: defences of luna moth caterpillars against predation by brown anoles

Field-testing predator-prey interactions between invasive exotic brown anole lizards and sentinel larvae of the luna moth, Actias luna, suggests that late instars of this saturniid, and perhaps many other saturniids, are well defended by their spines and crochets against predation by lizards. However, Actias luna larvae remain vulnerable to anole predation until they reach about 3 cm in length. In contrast, even relatively small caterpillars that are chemically defended, such as those of the io moth, Automeris io, and the monarch butterfly, Danaus plexippus, were shown to be well defended against anoles by their toxins. There is an obvious advantage for caterpillars to be chemically defended. However, there is also a tradeoff: Danaus plexippus are restricted to feeding on a relatively small group of plants, which limits resources and requires specialisation; in polyphagous Automeris io, the constant production of elaborate spines and toxin leads to prolonged larval development, which is at least twice as long as that of Actias luna, and exposes the caterpillars to mortality factors such as parasitism for a longer duration.     

Ecology and behaviour of the weaver ant Camponotus (Myrmobrachys) senex

Abstract

Weaver ants use silk produced by their larvae to build their nests. This behaviour is one of the more notable instances of social cooperation in animals; however, there are few studies of Brazilian species. This study investigated the ecology, natural history and behaviour of the weaver ant Camponotus (Myrmobrachys) senex in Brazil and showed that the weaver larvae have a fundamental function in nest building. The nests were always arboreal (one nest/plant), with a round form, beige in colour, and with leaves and shoots adhered to the silk nest. The average size was 34.24 cm and the average weight was 163.87 g; nests contained up to 50,000 individuals and several queens. Fusion and fission of colony parts were observed for C. senex. Worker ants were frequently observed feeding on honeydew, fruits and insects; and defended their territory. We suggest that C. senex larvae could be considered an effective caste, valuable in nest construction.     

Complex display behaviour during the intraspecific interactions of myrmecomorphic jumping spiders (Araneae,Salticidae)

Jumping spiders (Salticidae) are known for their elaborate vision‐based display behaviour, and a sizeable minority of the species in this large family resemble ants. The display repertoire of two ant‐like salticid species from the Philippines is investigated. Myrmarachne assimilis is a specialist ant mimic, closely matching the appearance of the weaver ant Oecophylla smaragdina. Myrmarachne bakeri is a generalist mimic, which, although ant‐like, is not a close match to any sympatric ant species. Myrmarachne are sexually dimorphic, with males having enlarged chelicerae. The copulatory posture of Myrmarachne is unique among salticids, apparently as an adjustment for the male's elongated chelicerae. Details concerning the intraspecific behaviour of Myrmarachne appear to be adjustments to living in environments populated by numerous other animals with similar appearance (ants). However, the specialist and the generalist that were investigated have display repertoires that are broadly similar to each other and to those of more typical salticids. Generally, the display behaviour of Myrmarachne appears not to have necessitated dramatic deviation from typical salticid display and mating strategies.     

A preliminary study of nest structure and composition of the weaver ant Polyrhachis (Cyrtomyrma) delecta (Hymenoptera: Formicidae)

Polyrhachis weaver ants build their nests from vegetation bound together using silk produced by their larvae. Here we provide a pilot study of the composition and the physical structure of three arboreal silk nests of Polyrhachis (Cyrtomyrma) delecta based on examination of three colonies. We found broadly similar nest architecture and size of the nests with each containing six or seven identifiable chambers, and describe the distribution of ants of different castes and life stages between them. We also note the construction of silk ‘girder’ structures, which spanned larger chambers, and we hypothesize that these provide additional strength to the internal nest structure. This study highlights the importance of more detailed investigation of the internal nest structure and composition in Polyrhachis, and other weaver ant species, which will help to develop our understanding of this specialized form of nest construction and nesting habits in a diverse group of ants.     

Larval instars of the ant Strumigenys solifontis Brown (Hymenoptera: Formicidae): the fallacy of size distribution

Larval instars of the myrmicine ant Strumigenys solifontis Brown were determined by measuring the head width under a stereomicroscope and examining the morphology, especially chaetotaxy, by light microscopy and scanning electron microscopy. This study found that the larval stage consisted of four instars, but the first and second instars were not distinguishable by their head widths. However, the four instars could be discriminated clearly by a combination of size measurement and observations on the chaetotaxy. Within a population, there are few larvae in the first instar because of its short period (only 1–2 days before moulting into the second instar) and the low oviposition rate of the queen. Therefore, the first-instar larva needs to be collected separately from other instars so that it is not missed.     

Ants feeding on anal exudate from tortricid larvae: a new type of trophobiosis

A new type of trophobiosis between tortricid larvae of a hitherto unknown genus and species, and various ant species of the subfamilies Formicinae, Dolichoderinae and Myrmicinae, was found in Malaysia. The larvae live in silken shelters fixed to the leaves of bamboo. In response to mechanical stimuli from the ants, they discharge from the anus a liquid containing sugar and amino acids and/or protein. The tortricid is described as Semutophila saccharopa gen. nov., sp. nov.; a second species, S. susurra sp. nov., is described from New Guinea.     

Bionomics of the myrmecophilous oribatid mite Protoribates myrmecophilus (Acari: Oribatida) in the Oriental tropics

Behavioural interactions between the myrmecophilous oribatid mite Protoribates myrmecophilus and the host ant Myrmecina spp. are described and compared to those of Aribates javensis which shows a very specialized myrmecophily. The behaviour of P. myrmecophilous is similar to free-living oribatids: they can walk by themselves and can survive without ant attendance. In the ant nests, the mites are sometimes cared for by ants and they are brought by ants into a new nest site. Protoribates myrmecophilus is eaten by non-host species of Myrmecina. Protoribates myrmecophilus is a less specialized myrmecophilous oribatid species.     

Immature stages of Parrhasius polibetes (Lepidoptera: Lycaenidae): host plants,tending ants,natural enemies and morphology

Natural history and immature stage morphology of the facultative myrmecophilous butterfly Parrhasius polibetes (Stoll) (Lepidoptera: Lycaenidae) are described and illustrated through both light and scanning electron microscopy. Larvae underwent four instars. At the third instar, the dorsal nectar organ became present and functional, and larvae were facultatively tended by several ant species, those also tending plants bearing extrafloral nectaries and honeydew-producing hemipterans. Larvae were florivorous and polyphagous at the species level, using at least 44 species of plants in 19 families. Most host plants (～ 89%) had some type of ant-attractive features, such as extrafloral nectaries or ant-tended treehoppers. The host range of this butterfly species allows the use of floral resources throughout the year. Food sources that promote ant visitation, flower bud morphology and phenology appear to be related to the evolution and maintenance of polyphagy in this butterfly species.     

Studies in European ant-decapitating flies (Diptera: Phoridae): ant alarm pheromone as host finding cue in Pseudacteon brevicauda,a parasite of Myrmica rubra (Formicidae: Myrmicinae)

Pseudacteon brevicauda Schmitz parasitizes Myrmica rubra in the Middle Rhine region in Germany. Female flies are attracted to disturbed ant colonies, where they attack Myrmica workers for oviposition. We experimented with different body parts of the ants, with dissected mandibular glands, and with synthetic gland constituents to locate the source of the fly attractant and the chemical basis of the signal. The attracting kairomone originates from the mandibular glands, which are used by Myrmica ants in alarm communication. Among the gland constituents, the ketones 3-octanone and 3-nonanone attracted and alerted the parasitic flies. The alcohol 3-octanol did not attract flies over long distance but it released an alerting response over short distance. A comparison of the chemical compositions of mandibular glands between ant species, focusing on the bioactive compounds, can explain both host specificity of P. brevicauda and the extension of the host spectrum to the (chemically similar) Myrmica scabrinodis.     

Comparison of natural histories and karyotypes of two closely related ant‐eating spiders,Zodarion hamatum and Z. italicum (Araneae,Zodariidae)

Based on morphological characters, two species of the genus Zodarion, Z. hamatum and Z. italicum, were placed into the italicum group. The aim of this study was to evaluate similarity of these two species in respect of various aspects of their natural history and karyotypes. Both species had similar phenological patterns, being univoltine, but some specimens reached adulthood the same year before winter. Both representatives had nocturnal activity and were able to capture various ant species. Zodarion hamatum seems to be specialized on Myrmicinae ants, and Z. italicum on Formicinae ants. Zodarion hamatum showed more agile predatory behaviour than Z. italicum presumably due to specialization on Myrmicinae ants, which have lower population density. Both species are generalized mimics of bicoloured (orange‐dark brown) ants, such as Lasius emarginatus; they performed similar courtship and mating. The two study species exhibited contrasting reproductive strategies. Zodarion hamatum spiders had higher fecundity than Z. italicum as a result of larger body size but Z. italicum had larger eggs than Z. hamatum. This is presumably an adaptation to different availability of prey ants. Both species had the same diploid chromosome number, sex chromosome system and pattern of X chromosome heteropycnosis in testes. Results support a very close relationship between these two Zodarion species.     

New trophobiotic symbioses of ants with South East Asian Bugs

A trophobiotic relationship between two species of phloem-feeding plataspid bugs and an ant, Meranoplus mucronatus, was discovered on tree trunks in Malaysia. Similar relationships were found between coreid bugs and Crematogaster sp. and Anoplolepis longipes, on bamboo in the same area. The ants recruit to groups of the bugs and feed on the liquid, sugar-rich faeces of the larvae, stimulating release of the honeydew by tactile signals. They protect all stages of the bugs from disturbance by biting and by the use of defensive scretions. Phloem-feeding bugs in the families Plataspidae and Coreidae need long stylets to pierce the thick bark of their host tree. The different methods of accommodating the resting stylets in these two families are described. The plataspids are described as Tropidotylus servus sp. nov. and T. minister sp. nov. A coreid previously reported in association with M. mucronatus in Malaya is described as Hygia cliens sp. nov. The coreids on bamboo were determined as Cloresmus spp. and Notobitus affinis.     

Carton nest building and trophobiont manipulation in the south-east Asian ant Dolichoderus sulcaticeps (Mayr 1870) (Hymenoptera: Formicidae)

Dolichoderus sulcaticeps, a rare canopy living dolichoderine ant, was studied in the rainforest of Malaysia. It forms large polycalyc colonies comprising partial nests on trees and climbers. The three dimensional multichambered partial nests are situated on the leaf underside. Fibrous nest material is gathered from the surface of the nesting plants and soaked with water from the plant surface. Building activity, therefore, is strictly dependent on the presence of rainfall or dew. Within the cell with plant leaf surface contact, as well as on free feeding sites, non specific coccid and membracid throphobionts, are tended. Free feeding sites are permanently protected. Mealybugs are transported between existing and new free feeding sites, as well as to and from nest pavilions, especially during their construction or destruction. Prey hunting has not been observed in D. sulcaticeps. The workers, however, recruit to offered dead prey, and scavenged. During the day they were permanently on the surface of plants around their nests and feeding sites.