Experimental tests of the mechanism for ant-enhanced growth in an ant-tended lycaenid butterfly

In a previous laboratory study, larvae of the ant-tended lycaenid butterfly Hemiargus isola developed into larger adults when reared with the ant Formica perpilosa than when reared without ants. Ants neither fed butterfly larvae nor significantly delayed developmental duration. We investigated two non-exclusive hypotheses for the mechanism of this effect: larvae tended by F. perpilosa (1) consume more food, and (2) digest the food they consume more efficiently, than those reared without ants. Larvae reared in the laboratory with F. perpilosa ants became significantly heavier adults but produced a significantly lighter fecal mass than their untended counterparts, suggesting that greater food consumption was not the primary mechanism for the higher growth rates of ant-tended larvae. Tended and untended larvae were equally proficient at digesting the contents of pollen (a major natural food source) throughout the tended portion of the life cycle. Taken together, the results suggest that neither greater consumption nor higher assimilation accounts for the larger size of F. perpilosa-tended larvae. We propose that tended larvae may expend less energy than their untended counterparts. Received: 3 January 1997 / Accepted: 18 June 1997     

Species-specific effects of tending ants on the development of lycaenid butterfly larvae

The transfer of nutrients between organisms is a common feature of mutualism. The production of these food rewards is often assumed to be costly. Estimation of the costs of producing food rewards is important for understanding the overall effects of the interaction on fitness. When food rewards are harvested by several species differing in foraging behavior, costs to the producer may differ. The larvae of many species in the butterfly family Lycaenidae produce secretions consumed by tending ants. Here I report that three North American ant species, Formica perpilosa, Dorymyrmex sp. (smithi complex), and Forelius foetida, had no negative effect on the duration of development and adult size of the lycaenid Hemiargus isola. Moreover, tending by the ant Formica perpilosa significantly enhanced larval growth, resulting in butterflies that were 20% heavier than their untended counterparts. Tending by the ants Dorymyrmex sp. (smithi complex) and Forelius foetida had no effect on butterfly weight. Tended, nonfeeding larvae lost 69% more weight than untended, nonfeeding larvae. Taken together, the results suggest that, although ant tending imposes a physiological cost, H. isola larvae use behavioral or physiological mechanisms to compensate or overcompensate for nutrients lost to ants.     

Wood ants prefer conifers to broadleaved trees in mixed temperate forests

1. Wood ants, species of the subgenus Formica s. str., are one of the most important groups of insects in forest ecosystems due to their high abundance and activity.
2. We assessed the foraging pressure of Formica polyctena and Formica rufa, two dominant wood ant species in the Białowieża Forest. In addition, we compared coniferous and mixed stands in these respects.
3. In mixed stands F. polyctena visited more trees per plot than F. rufa. F. polyctena might forage a larger area due to higher population sizes, which further confirms higher foraging pressure of F. polyctena in comparison with F. rufa.
4. In our study area, host trees of the ant–aphid mutualism were larger in diameter than nonhost trees. The ants also preferred trees situated closer to their nests. In both mixed and coniferous stands, the most visited tree species was Norway spruce. Thus, we can conclude that Norway spruce seems to be preferred by the ant. The decline of living spruce in the Białowieża Forest may endanger their food source and impact the vitality of ant colonies.

     

Stand type is more important than red wood ant abundance for the structure of ground‐dwelling arthropod assemblages in managed boreal forests

• 1 The relationships between red wood ants (Formica rufa group) and other ground‐dwelling arthropods were studied in young managed forests stands in Eastern Finland. The main objectives were: (i) to test the influence of stand type (dominant tree species; age: sapling versus pole stage) and numbers of red wood ants on the occurrence of other ground‐dwelling arthropods and (ii) to study the occurrence of red wood ants versus other arthropods on a distance gradient from ant mounds. We used pitfall traps set in 5–14‐year‐old sapling stands and 30–45‐year‐old pole‐stage stands of Scots pine (Pinus sylvestris L.) and birch (Betula spp.) forests.
• 2 Pitfall trap catches of red wood ants did not vary significantly between the forest stand types, although some groups of other arthropods showed clear responses to stand type (e.g. catches of other Formicinae and Gnaphosidae were higher in sapling stands than in pole‐stage stands). The number of red wood ants clearly explained less of the variation in assemblages of other ground‐dwelling arthropods than the forest stand type.
• 3 Red wood ant numbers decreased significantly with distance from the mounds, but the other ground‐dwelling arthropods were insensitive to this gradient or even showed a preference for proximity to ant mounds and high ant activity.
• 4 The results obtained in the present study suggest that wood ants do not have strong effects on several other ground‐dwelling arthropod groups in young managed forests other than in the immediate vicinity of their mounds.

     

The effect of red wood ant abundance on feeding damage by the pine weevil Hylobius abietis

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The fate of an intentional introduction of Formica lugubris to North America from Europe

Red wood ants (Formica s.str.) are not prevalent in the forests of North America, but commonly occur in conifer and mixed conifer forests in northern Europe and Asia. In 1971, a European red wood ant species, Formica lugubris, was intentionally established in a 35‐year‐old predominantly mixed conifer plantation approximately 30 km north of QC, Canada. The purpose of its introduction was to evaluate the potential of this species as a biological control agent against conifer‐defoliating Lepidoptera species. This red wood ant introduction was monitored periodically for about 5 years after establishment, but its long‐term fate has not been reported. We visited this field site in 2005 and found that this species was well established, and we could locate some of the nests that resulted from the original release. We mapped and measured over 100 nests around the site of original release, which ranged from 5 cm in height to over 1 m. We estimated the population of introduced ants to have grown to over 8 million in the last 34 years. Significant clustering of nests suggests that these nests may be one supercolony. F. lugubris has become a dominant understory arthropod in this mixed forest, and is likely to have ecological impacts, including effects at the community and ecosystem level.     

First report of seed dispersal by ants in Dicentra peregrina (Papaveraceae), an alpine plant in the Japanese Alps

Seed dispersal by ants, known as myrmecochory, is commonly observed among various plant taxa. The seeds of these plants have an elaiosome to attract ants. In Japan, myrmecochory has been well studied in several lowland plant species, but not in highland plant species that grow above the tree line. We investigated whether the seeds of Dicentra peregrina, known as the “queen of the alpine plants” in Japan, are carried by Formica gagatoides ants at 2510 m a.s.l. on Mt Norikura, Kita‐Alps, Japan. We observed F. gagatoides workers picking up D. peregrina seeds by grasping the elaiosome and carrying the seeds into their nests. We inferred from the observed ant behavior and the seed morphology that D. peregrina is a myrmecochorous species.     

Interference competition,not predation,explains the negative association between wood ants (Formica rufa) and abundance of ground beetles (Coleoptera: Carabidae)

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The role of an attractive brood pheromone in the obligatory,slavemaking ant,Polyergus breviceps (hymenoptera: Formicidae)

Freeliving workers of Formica occulta, an ant species enslaved by the obligatory slavemaking ant Polyergus breviceps, retrieve and nurse Polyergus pupaejust as well as conspecific pupae in a choice test. No such attraction was found toward pupae of the facultative slavemaker; Formica wheeleri,which also enslaves F. occulta. Formica neogagates,a sympatric species which is not parasitized by either slavemaker, preferentially retrieves and tends conspecific brood over that of Polyergusand F. wheeleri.It is proposed that brood of obligatory slavemaking species must possess an attractive pheromone for slavemaker colony foundation to be successful, since slavemaker brood must be nursed by adult slave workers with no prior exposure to slavemaker brood. An attractive pheromone is not necessary in the brood of facultative slavemakers, since this brood is cared for by newly eclosed slave workers who imprint on the slavemaker brood.     

An orb‐weaver spider eludes plant‐defending acacia ants by hiding in plain sight

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Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.     

A nectar-secreting gall wasp and ant mutualism: selection and counter-selection shaping gall wasp phenology, fecundity and persistence

Abstract.

• 1 The natural history of a gall wasp including interactions with inquilines, parasites, and a mutualistic ant are examined. The stability of the system is described from the perspective of influences on gall wasp life history characteristics.
• 2 An exclusion experiment demonstrated that the nectar-secreting gall of Disholcaspis perniciosa mediates a mutualism with the tending ant, Formica obscuripes. Survivorship increased from 0% in the absence of ants to 25.3% in their presence, largely due to the exclusion of inquilines.
• 3 Specialized parasites, Eudecatoma spp., attacked before the ant-gall interaction began, when the developing gall was still beneath the host plant (Quercus gambellii) epidermis and ants were not in attendance. They may select for later developing gall wasps, which benefit by having fewer individuals parasitized. However, counter-selection for earlier development may result from decreased gall wasp size, decreased fecundity, and an increase in gall failures resulting from late development.
• 4 Local persistence of the gall wasp population despite increased pressure from inquilines and parasites was attributed to gall wasp escape in time due to polymorphic emergence resulting from diapause. Most individuals emerge at the end of the summer, but approximately 15% remain in the galls as prepupae for 1–5 years.

     

Immigration and formation of mixed colonies in red wood ants (Hymenoptera,Formicidae)

Long-term field studies of the composition and spatial structure of settlements of ants of the Formica rufa group were carried out in two regions of Russia (Moscow and Arkhangelsk provinces). Fragmentation of damaged nests followed by reintegration of the fragments is the main way of formation of mixed colonies of ants from different nests (including different species). The principal factor of nest fragmentation is their damage by wild boars, bears, and in some localities, by poachers. The formation of mixed nests and nest complexes with participation of different Formica species was observed. They are formed by joining the ants from several damaged nests or by a colony from a destroyed nest immigrating into an intact one. Regular damage of many nests leads to the formation of broad zones of mixed colonies. The mixed colonies including 2–3 species of wood ants have recently become common. The phenomenon of mixed colonies raises a question as to the relative importance of two basic principles (sociality and specific identity) in the life of ant societies and demonstrates the priority of the social principle.     

Size matters: nest colonization patterns for twig‐nesting ants

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Ants protect conifer seedlings from feeding damage by the pine weevil Hylobius abietis

• 1 Ants that protect food resources on plants may prey on (or deter) herbivores and thereby reduce damage. Red wood ants (of the Formica rufa group) are dominant ants in boreal forests of Eurasia and affect the local abundance of several herbivorous species.
• 2 The pine weevil Hylobius abietis (L.) is a herbivore that causes severe damage by feeding on the bark of coniferous seedlings within areas of forest regeneration.
• 3 We investigated whether ants can protect conifer seedlings from pine weevil feeding. In a manipulative experiment, ants were attracted to sugar baits attached to spruce seedlings and the damage caused by pine weevils was compared with control seedlings without ant‐baits.
• 4 The feeding‐scar area was approximately one‐third lower on the seedlings with ant‐baits compared with the controls. Besides red wood ants, Myrmica ants were also attracted in high numbers to the ant baits and the relative effects of these species are discussed.
• 5 The results obtained in the present study support the trophic cascade hypothesis (i.e. damage to herbivores is suppressed in the presence of predators). The decreased pine weevil feeding on the baited seedlings was probably a result of nonconsumptive interactions [i.e. the presence of (or harassment by) ants distracting pine weevils from feeding].
• 6 Understanding the role of ants may have important implications for future strategies aiming to control pine weevil damage. For example, maintaining suitable conditions for ants after harvesting stands may be an environmentally friendly but currently unexploited method of for decreasing weevil damage.

     

Temporal and spatial patterns of foraging in two syntopic ant species,Manica yessensis and Formica lemani,on Mount Fuji,Japan

Spatiotemporal characteristics of terrestrial foraging were studied in two ant species, Manica yessensis and Formica lemani, in a volcanic desert on the southeast slope of Mount Fuji, Gotenba, Japan. Both ants are common in this habitat, and they construct underground nests in this dry area with sparse vegetation. Nests of M. yessensis have multiple nest-openings on the surface, whereas nests of F. lemani have very few openings, but their nesting and foraging areas overlap completely. A “mark-and-observe” method applied to M. yessensis demonstrated that worker ants of this species move between openings more than 3 m away. A study plot (6 m × 12 m quadrat) was set up, in which all nest-openings of both species were mapped. Day-long observations on numbers of foragers in this plot revealed that foraging M. yessensis are active in morning and evening, while F. lemani continues foraging all day, but both species cease activity at night. Associations between locations of foragers and nest-openings differed significantly between the two species, that is, surface foraging of M. yessensis workers was largely confined to the vicinity of their nest-openings, whereas foragers of F. lemani travelled far from their nest-openings. The function of multiple nest-openings in M. yessensis is discussed.     

Influence of native ants on arthropod communities in a vineyard

• 1 Ants can have a range of effects on arthropods in crops, including suppressing herbivores such as caterpillars. However, ants can also increase hemipteran densities while reducing natural enemy numbers. In vineyard ecosystem, the effects of native ants and their interactions with other arthropods are poorly understood.
• 2 An ant‐exclusion experiment was designed to test the impact of native ants on both canopy and ground arthropods concurrently. The potential influence of ants on predation and parasitism of light brown apple moth (LBAM) eggs, a grape pest, was also examined. Adult grapevine scale insects and earwigs under bark were counted after a season of ant‐exclusion.
• 3 Among 23 ground ant species collected, six were found to forage in the canopy, with two Iridomyrmex species being the most commonly encountered.
• 4 There was no difference in the abundance of most arthropod orders and feeding groups between ant‐excluded and control vines, although ground spiders were more abundant under ant‐excluded vines, despite increased ground ant foraging pressure. LBAM egg parasitism and predation were low and probably affected by weather and other arthropods. Ant exclusion did not reduce survival of scale insects, although the distribution and abundance of scale insects were negatively associated with earwigs.
• 5 In conclusion, native ants did not consistently suppress arthropod assemblages, including natural enemies, and they did not promote the survival of scale insects. Interactions among native ant species within a vineyard might minimize their effects on other arthropods, although this needs further study.

     

Antipredatory Activity of the Weevil Oxyops vitiosa: A Biological Control Agent of Melaleuca quinquenervia

The larvae of the leaf-feeding weevil Oxyops vitiosa, a biological control agent of Melaleuca quinquenervia, are covered with a viscous orange coating that is thought to protect against generalist predators. This coating is gradually lost as the larvae drop to the ground and pupate in subterranean pupal cells. To test the antipredator activity of this species, four immature life stages (early instars, late instars, prepupae, pupae) were exposed to a common generalist predator, the red imported fire ant Solenopsis invicta. Choice tests were conducted by placing an O. vitiosa individual and a control larva of the weevil Neochetina eichhorniae into an arena containing a S. invicta colony and observing subsequent ant behaviors. S. invicta workers contacted O. vitiosa early instars, late instars, and prepupae less frequently than control N. eichhorniae larvae, and upon contact S. invicta was less likely to behave aggressively toward these O. vitiosa life stages than toward N. eichhorniae larvae. However, S. invicta contacted, attacked, and consumed naked (nonencased) O. vitiosa pupae and N. eichhorniae larvae with equal frequency. Encased O. vitiosa pupae buried in sand were not attacked compared to susceptible encased pupae on the sand surface. By shifting from a chemical defense during the larval stages to a physical defense during the pupal stage, O. vitiosa reduces the risk of attack by this generalist predator.     

Nesting preferences of the threatened wood ant <Emphasis Type="Italic">Formica exsecta</Emphasis> (Hymenoptera: Formicidae); implications for conservation in Scotland

Wood ants are a dominant and ecologically important component of northern coniferous forests with interactions at many trophic levels. Each species exhibits specific habitat preferences which need to be understood if conservation measures are to be successful. In Britain, the rare narrow-headed ant Formica exsecta has disappeared from much of its former range and is now largely restricted to the highlands of Scotland where it is found in open canopy woodland and along forest edges. Nest locale of one small and vulnerable population at the edge of its current range, were compared with those of random locations within the same habitat and with actual nests within the stronghold of the Abernethy-Glenmore complex in Strathspey. After data exploration with Redundancy Analysis, stepwise multiple regression was used to create a model which best estimated the variance in nest location using a parsimonious selection of vegetation and environmental variables. The input variables included were light, soil moisture, altitude, tree stature and distribution, vegetation structure and composition, and ground characteristics. F. exsecta clearly exhibited preferences for the position of nest mounds in relation to light, vegetation and tree cover. Forest location was also important in determining which variables nests were affected by. This study highlights the importance of maintaining a dynamic mosaic of different-aged woodland enabling early successional species such as F. exsecta, with suitable areas to move to as conditions change and allowing the co-existence of all wood ant species.     

Species‐specific levels of ant attendance mediate performance costs in a facultative myrmecophilous butterfly

Trophobiont butterfly larvae offer caloric rewards to ants through specialised glands and, in return, gain ant‐derived protection from natural enemies. Thus, from the larva's perspective, the major cost of myrmecophily comprises the reward production. Larvae of the butterfly Parrhasius polibetes (Stoll) (Lycaenidae) are facultatively tended by several ant species, which might differ in the intensity of tending behaviour. The performance costs (development time, survival, pupal mass and adult dry mass) of P. polibetes are examined when tended by two ant species differing in size and foraging strategies (Camponotus melanoticus Emery and Camponotus crassus Mayr), along with the corresponding intensity of tending behaviour towards late instars. Untended larvae serve as controls. Larvae tended by C. melanoticus take longer to pupate compared with both C. crassus and control larvae. By contrast, pupae whose larvae are tended by C. crassus are lighter than control larvae but do not differ from those tended by C. melanoticus. No differences are found in the adult stage, indicating compensation in all cases. Both at short‐ and long‐term scales, C. melanoticus tends larvae of P. polibetes more intensely than C. crassus. The increase in tending activity of C. melanoticus presumably delays the development time of larvae tended by this ant species. The results of the present study show that tending intensity varies depending on the ant species, and that P. polibetes has compensatory mechanisms to minimise myrmecophily costs, regardless of tending intensity. To the authors' knowledge, this is the first experimental evidence that intensity of ant‐tending behaviour is species‐specific and affects performance in a trophobiont insect.