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.     

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.     

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.     

Ecology and field biology of two dominant Camponotus ants (Hymenoptera: Formicidae) in the Brazilian savannah

Camponotus renggeri and C. rufipes are very abundant in Brazilian cerrado savannah, where they feed extensively on liquid rewards and commonly associate with plants bearing extrafloral nectaries and honeydew-producing insects. Here, we provide a qualitative and quantitative field account on the natural history and ecology of these two ant species. The study was carried out in a cerrado reserve in south-eastern Brazil across a rainy/hot season (summer) and a dry/cold season (winter). The ants were found in two vegetation physiognomies: all nests of C. rufipes were located in the cerrado sensu stricto (scrub of shrubs and trees, 3–8 m tall), whereas C. renggeri occurred mostly in the cerradão (forest with more or less merging canopy, 10–12 m tall). Both species nested in fallen or erect dead trunks, as well as underground. In addition, C. rufipes built nests using dead plant material arranged or not around shrub bases. Colonies of C. rufipes were generally more populous than those of C. renggeri, and both species had colonies with more than one dealated queen. Both species were active mainly at night and foraged for resources near their nests, mainly extrafloral nectar and hemipteran honeydew (aphids and mealybugs). The average size of the home ranges of C. renggeri in cerrado sensu stricto and cerradão varied from ≈ 2.8 to 4.0 m² and apparently were not affected by season. In C. rufipes, however, foraging grounds in cerrado sensu stricto showed a twofold increase from dry/cold (≈ 4.5 m²) to rainy/hot season (≈ 9.8 m²). Our study highlights the importance of natural history data to understand the foraging ecology and role of these ants in cerrado savannah.     

Natural history and ecology of foraging of the Camponotus crassus Mayr, 1862 (Hymenoptera: Formicidae)

ABSTRACT

Ants are abundant in terrestrial ecosystems, especially in the Brazilian Cerrado, where they can play several roles at different levels of the food chain, including protection of plants against herbivores. Although there are many studies that evaluate the ant–plant interaction in the Cerrado, little is known about the natural history of most species of ants. Camponotus crassus Mayr, 1862, for example, is considered one of the main agents of plant biotic defence in Cerrado. But there are no studies specifically focused on this species, which hinders the understanding of how arthropod–plant interactions are structured in Cerrado. Here, we describe the natural history and ecology of the foraging of the C. crassus. We conducted the study from January 2013 to December 2014 in 10 quadrants of 40 m² to measure: the abundance, density and distribution of nests, location of the nests, the internal structure of the nests, the daily foraging of workers out of the nest, the food items they collect and the existence of territoriality and dominance of the workers on the soil and vegetation. We found 18 nests, 13 in the soil and 5 in hollow trunks on the ground with variable internal structures. The distribution of nests is aggregate, with density of 0.045 nests/m² and average distance of 3.73 m between nests. The foraging activity occurs on the daytime during the rainy and dry season. Extrafloral nectar and honeydew were the resources most collected, comprising 83.33% of the resources in the rainy period and 30% in the dry period. Camponotus crassus is a dominant species, especially on vegetation, although it also forages on the soil. This is the first study to evaluate in detail the natural history and foraging ecology of C. crassus, a diurnal, aggressive and territorial ant that mainly forage climbing onto the plants.     

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.     

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.     

Cubitermes subarquatus termitaries as shelters for soil fauna in African rainforests

Most soil fauna in tropical rainforests require a pre-existing shelter for at least part of their life cycles. Among termites and ants, the most frequently encountered taxa, the availability of appropriate nesting sites is a major factor controlling the multiplication of colonies. Work carried out in five forests in southern Cameroon indicates that termitaries of Cubitermes subarquatus, whether active or abandoned, provide shelter for large numbers of invasive invertebrates, particularly ants and other termite species, and may be a determining factor in the patchy distribution of many taxa. Dissections of 90 termitaries (65 active, 25 abandoned) produced a great diversity of invasive fauna, including 354 ant colonies representing 75 species in 31 genera. Ants occurred in all abandoned termitaries but 6·1% of active termitaries had no ants present. Nevertheless, in total active termitaries sheltered more ant colonies (273) than did abandoned ones (81). Every termitary examined also sheltered founding pairs or incipient colonies of alien termites, but active C. subarquatus termitaries yielded fewer mature colonies of alien termites (56·9%) than did abandoned ones (84%).     

Foraging ecology and behaviour of the ponerine ant Ectatomma opaciventre Roger in a Brazilian savannah

This study provides a detailed account of the foraging behaviour of the ponerine ant Ectatomma opaciventre in a ‘cerrado’ savannah in south-east Brazil. Our observations suggest that this species has an exclusively diurnal foraging pattern. Feeding habits included both predation and scavenging, with termite workers and leaf-cutting ants as the most important food items. Contrary to all other Ectatomma species studied to date, no liquid food such as hemipteran honeydew or plant nectar was collected. Foragers showed clear individual foraging area fidelity.Workers of E. opaciventre employed a typical individual foraging strategy, i.e. there was no co-operation between foragers in the search for or retrieval of food, neither by tandem running nor by trail laying. Nest density was considerably lower than in other Ectatomma (0.015 nests per m²). The observed mean distance to the nearest neighbouring nest was 5.85 m, with a significant tendency toward over-dispersion. Nests were more frequently found in specific microhabitats, which may suggest active choice of nesting site by founding queens.     

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.     

Trophobiosis between ants and Eurystethus microlobatus Ruckes 1966 (Hemiptera: Heteroptera: Pentatomidae) a cryptic,gregarious and subsocial stinkbug

In this study we describe trophobiosis between ants and Eurystethus microlobatus (Hemiptera: Pentatomidae) at a highland quartzite rocky outcrop in southern Espinhaço Range, southeastern Brazil. We found stinkbugs exclusively on the mistletoe Psittacanthus robustus (Loranthaceae). The stinkbug species is gregarious, forming dense clusters, with females guarding eggs and young nymphs. In addition, this species presents cryptic colouration and most individuals were located on roots and shoots, camouflaged within the bark. Eurystethus microlobatus produced honeydew by day and night, and four ant species attended aggregations: Camponotus rufipes, Camponotus crassus, Camponotus melanoticus and Cephalotes pusillus. Camponotus rufipes was the most frequent and aggressive, and the only species to attend the stinkbugs by day and night. This is the first documented case of trophobiosis between true bugs and ants in the New World.     

Foraging ecology of the giant Amazonian ant Dinoponera gigantea (Hymenoptera,Formicidae, Ponerinae): activity schedule,diet and spatial foraging patterns

This study provides a detailed account of the natural history and foraging biology of the ponerine ant Dinoponera gigantea in a rainforest in north Brazil. The species nests on the ground and the colonies contain 70–96 workers. Ant activity is negatively correlated with temperature, and is more intense at dawn and dusk. Foragers leave the nest independently and search for food individually on the leaf litter, within ca 10 m around the nest. Workers are opportunistic feeders that collect seeds and fruits, and hunt for live prey as well as scavenge for dead animal matter. The dry weight of food items ranges from <10 mg (spiders, insect parts) to >400 mg (seeds, fruits). There is no nestmate recruitment during the search for or retrieval of food, irrespective of food type and size. Foragers have a high directional fidelity, and ants from neighbouring colonies may engage in ritualized territorial contests at the border of their foraging areas. The foraging ecology of D. gigantea is compared with other ponerine species living in tropical forests, as well as with other ant groups showing similar behavioural patterns.     

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.     

Predator-prey interactions between the digger wasp Clypeadon laticinctus and the harvester ant Pogonomyrmex occidentalis

Females of the digger wasp Clypeadon laticinctus prey exclusively on workers of the western harvester ant Pogonomyrmex occidentalis. There is considerable variability in the size of ant workers both within and between ant mounds, and wasps tend to select larger workers as prey. Worker ants were most likely to be outside the nest at surface temperatures of 25–35°C. Wasps were seen at ant mounds most frequently when surface temperatures were between 40 and 50°C. Wasps captured ants outside and inside ant nests. Predation attempts were equally likely to result in a successful capture in either location, but entering the ant nests was more time consuming. Approximately 63% of visits by wasps to ant mounds resulted in a successful prey capture. In general, ant mounds that were visually conspicuous or close to wasp nest aggregations were more likely to be visited by wasps. However, frequency of visitation and rates of predation at a given mound varied considerably from week to week. Agonistic interactions between female wasps involving chases and occasional fights were likely to occur whenever two wasps were simultaneously present at an ant mound, especially when the only access to prey was by entering the mound.     

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.     

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.     

The life cycle and natural history of the tropical aphid Cerataphis fransseni (Homoptera: Aphididae: Hormaphidinae), with reference to the evolution of host alternation in aphids

We describe the life cycle and general biology of the tropical cerataphidine aphid Cerataphis fransseni. We demonstrate that this aphid migrates between trees of Styrax benzoin and various species of palms; palm-feeding populations have previously been known as C. variabilis and C. palmae, which now become synonyms of C. fransseni. On S. benzoin the fundatrix induces a relatively simple gall which can contain >6000 aphids at maturity with a large number of reproductively sterile soldiers that protect the gall from predators. These galls are apparently produced throughout the year. Colonies on the secondary host plants, palms, are apparently obligately tended by ants whereas colonies within galls on Styrax are never tended by ants. We discuss the life cycle of this tropical aphid with respect to hypotheses for the evolution and maintenance of host alternation.     

The symbiosis between the weaver ant,Oecophylla smaragdina,and Anthene emolus,an obligate myrmecophilous lycaenid butterfly

Females of Anthene emolus use the presence of the weaver ant, Oecophylla smaragdina, as oviposition cue. Ovipositing females are not attacked by the ants. The first instar larvae are adopted by the ants and carried into their pavilions where the caterpillars feed. Outside the pavilions the larvae cannot survive. The older larvae leave the pavilions and are carried by the ants to their feeding places or back into the pavilions. The larvae are constantly attended by the ants. During the 3rd and 4th instar the caterpillars secrete high amounts of nutritive liquids representing an estimated energy content of 200 J per larva. Thus the larvae are important trophobionts of Oe. smaragdina and attract the ants by releasing food recruitment behaviour. The pupae are not attractive for ants, but are not attacked, either. The emerging adults are not protected from ant attacks and are sometimes killed by their host ants. The costs and benefits of this close obligate myrmecophilous relationship for both the ants and lycaenids as well as the evolution of ant-specific relationships of the lycaenids are discussed.     

Selection of ants by the American black bear ( Ursus americanus )

We examined selection of ants as food by black bears ( Ursus americanus ) in Utah by comparing the genera of ants we sampled from study plots in 5 vegetation types with those found in bear scats. In the plots we found 641 ant colonies representing 18 genera. Five genera comprised 85% of all colonies sampled: Formica, Lasius, Camponotus, Tapinoma, and Myrmica . Thirteen genera were found in scats, but 99% were composed of the same 5 listed above. Limited utilization of some ant genera by black bears may reflect body size, colony size, nest type, and presence or absence of a functional sting. Fifty-three percent of all ant colonies in sample plots were found under rocks. Energy values of ants ranged from 4.9 to 6.2 cal · g -1 dry weight. Pupae of Formica and Camponotus contained slightly more fat than respective workers, but the relationship was reversed for Lasius . Pupal cases were found in 33% of the scats, and bears may prefer ant brood because they are more digestible than adults and lack mechanisms of defense. Ants are a predictable and ubiquitous food source for black bears.     

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.