Indirect effects of granivory by harvester ants: plant species composition and reproductive increase near ant nests

Summary Of 36 plant species surveyed, 6 were significantly associated with nests of the desert seed-harvester ant Veromessor pergandei or Pogonomyrmex rugosus; two other plant species were significantly absent from ant nests. Seeds of two common desert annuals, Schismus arabicus and Plantago insularis, realize a 15.6 and 6.5 fold increase (respectively) in number of fruits or seeds produced per plant growing in ant nest refuse piles compared to nearby controls. Mass of individual S. arabicus seed produced by plants growing in refuse piles also increased significantly. Schismus arabicus, P. insularis and other plants associated with ant nests do not have seeds with obvious appendages attractive to ants. Dispersal and reproductive increase of such seeds may represent a relatively primitive form of ant-plant dispersal devoid of seed morphological specializations. Alternatively, evolution of specialized seed structures for dispersal may be precluded by the assemblage of North American seed-harvester ants whose workers are significantly larger than those ants normally associated with elaiosome-attached seed dispersal. Large worker size may permit consumption of elaiosome and seed.     

Nest mounds of red wood ants (Formicaaquilonia): hot spots for litter-dwelling earthworms

A previously undocumented association between earthworms and red wood ants (Formicaaquilonia Yarr.) was found during an investigation of the influence of wood ants on the distribution and abundance of soil animals in boreal forest soil. Ant nest mounds and the surrounding soil of the ant territories were sampled. The ant nest mound surface (the uppermost 5-cm layer) harboured a much more abundant earthworm community than the surrounding soil; the biomass of the earthworms was about 7 times higher in the nests than in the soil. Dendrodrilusrubidus dominated the earthworm community in the nests, while in soils Dendrobaenaoctaedra was more abundant. Favorable temperature, moisture and pH (Ca content), together with abundant food supply (microbes and decomposing litter) are likely to make a nest mound a preferred habitat for earthworms, provided that they are not preyed upon by the ants. We also conducted laboratory experiments to study antipredation mechanisms of earthworms against ants. The experiments showed that earthworms do not escape predation by avoiding contact with ants in their nests. The earthworm mucus repelled the ants, suggesting a chemical defence against predation. Earthworms probably prevent the nest mounds from becoming overgrown by moulds and fungi, indicating possible mutualistic relationships between the earthworms and the ants. Received: 21 November 1996 / Accepted: 3 April 1997     

Dietary similarity and foraging range of two seed-harvester ants during resource fluctuations

Summary Diets of desert seed-harvester ants Veromessor pergandei and Pogonomyrmex rugosus were studied for 3 years at two habitats where they are common and sympatric. Diets of the two species were similar, consisting mainly (87% of 23,913 seeds) of three annual plant species (Schismus arabicus, Plantago insularis, and Pectocarya recurvata). Diets converged following a drought in Winter/Spring 1984 which reduced seed production during this time. Foraging range of P. rugosus almost doubled following the drought while foraging range of V. pergandei remained constant. Neither ant species move their nests once established rendering them effectively sessile granivores. This limits the dietary options of a given colony and may explain interspecific dietary convergence.     

Negative effects of ant foraging on spiders in Douglas-fir canopies

 Spiders and ants are potential competitors and mutual predators. Indirect evidence from previous research has suggested that ant foraging may significantly lower the abundance of arboreal spiders in young Douglas-fir plantations in western Oregon. This study tested the effect of foraging by ants, dominated by Camponotus spp., on spider assemblages in Douglas-fir canopies in a 5-month ant-exclusion experiment. The biomass of potential prey organisms on foliage, dominated by Psocoptera, increased significantly by 1.9- to 2.4-fold following ant exclusion. The removal of ants did not affect the abundance of flying arthropods in the vicinity of tree canopies as indicated by sticky trap catches. The abundance of hunting spiders, the majority being Salticidae, increased significantly by 1.5- to 1.8-fold in trees without ants in the late summer; neither the abundance of web-building spiders nor the average body size of hunting and web-building spiders were significantly affected by ant removal. Spider diversity and community structure did not differ significantly between control and ant-removal trees. The majority of prey captured by ants were Aphidoidea (48.1%) and Psocoptera (12.5%); spiders represented only 1.4% of the ants’ diet. About 40% of observed ants were tending Cinara spp. aphids. Our observations suggest that the lower abundance of hunting spiders in control canopies with ants may be due to interference competition with ants resulting from ant foraging and aphid-tending activities. Direct predation of spiders by ants appeared to be of minor importance in this study system. This study did not provide sufficient evidence for exploitative competition for prey between ants and spiders. Received: 21 February 1996 / Accepted: 14 August 1996     

The influence of food supply on foraging behaviour in a desert spider

We tested the alternative hypotheses that foraging effort will increase (energy maximizer model) or decrease (due to increased costs or risks) when food supply increased, using a Namib desert burrowing spider, Seothyra henscheli (Eresidae), which feeds mainly on ants. The web of S. henscheli has a simple geometrical configuration, comprising a horizontal mat on the sand surface, with a variable number of lobes lined with sticky silk. The sticky silk is renewed daily after being covered by wind-blown sand. In a field experiment, we supplemented the spiders' natural prey with one ant on each day that spiders had active webs and determined the response to an increase in prey. We compared the foraging activity and web geometry of prey-supplemented spiders to non-supplemented controls. We compared the same parameters in fooddeprived and supplemented spiders in captivity. The results support the costs of foraging hypothesis. Supplemented spiders reduced their foraging activity and web dimensions. They moulted at least once and grew rapidly, more than doubling their mass in 6 weeks. By contrast, food-deprived spiders increased foraging effort by enlarging the diameter of the capture web. We suggest that digestive constraints prevented supplemented spiders from fully utilizing the available prey. By reducing foraging activities on the surface, spiders in a prey-rich habitat can reduce the risk of predation. However, early maturation resulting from a higher growth rate provides no advantage to S. henscheli owing to the fact that the timing of mating and dispersal are fixed by climatic factors (wind and temperature). Instead, large female body size will increase fitness by increasing the investiment in young during the period of extended maternal care.     

Aerial prey caching by solitary ground-nesting wasps: A test of the predator defense hypothesis

Many solitary ground-nesting wasps in the families Pompilidae and Sphecidae excavate nests after capturing prey for provisions. These wasps generally cache their immobilized prey temporarily during nest excavation, frequently by suspending the prey in a plant (aerial caching). Here I test the hypothesis that aerial caching by Ammophila spp. wasps (Sphecidae) functions to reduce prey theft by generalist predators, Formica spp. ants. Foraging ants removed baits placed on the ground more rapidly than baits suspended in plants; mean half-lives for ground and aerial baits were 14.5 and 145.7 min, respectively (mean values for experiments 1–3). Ant foraging activity decreased during the midday. Ant interference with nesting activities of Ammophila spp. also decreased during the midday, paralleling observed fluctuations in ant foraging activity.     

Kin recognition in a semi-natural context: Behaviour towards foreign conspecifics in the social waspRopalidia marginata (Lep.) (Hymenoptera: Vespidae)

Summary Female wasps of the tropical primitively eusocial speciesRopalidia marginata are known to discriminate unfamiliar nestmates from unfamiliar non-nestmates outside the context of their nests. Here, we show that when foreign conspecifics are introduced in the context of a nest in laboratory cages, genetic relatives among them are treated by nest inhabitants more tolerantly than non-relatives, but that no foreign conspecifics are accepted into the nests. However, some wasps may leave their nest and join the foreign relatives and non-relatives to found new colonies cooperatively. Very few of the introduced animals are severely attacked or killed; most are allowed to remain in parts of the cage away from the nest. These results suggest that factors other than genetic relatedness may be involved in regulating tolerance and acceptance of foreign conspecifics on a nest and its vicinity. Our results are different from those of similar experiments with ants, which have demonstrated that former nestmates that are removed as pupae and later introduced as adults are either accepted into the nest or attacked and killed. We attribute this difference to the fact that in a primitively eusocial species such asR. marginata, the rules governing tolerance and acceptance of foreign conspecifics must be quite different from those in highly eusocial species. We also attempt to test some predictions of the conspecific acceptance threshold models of Reeve (Am. Nat. 133:407–435, 1989). Our results uphold the predictions of his fitness consequence submodel but do not support those of his interaction frequency sub-model.     

Interspecific predation as a mortality factor among overwintering spiders

Summary Results from field experiments indicate that predation occurs among spruce-living spiders during winter in SW Sweden. Field observations of natural activity showed that Philodromus spp and Pityohyphantes phrygianus together make up 80% of the spiders active on spruce in winter. They are therefore potential predators on other overwintering spiders. Laboratory experiments were performed at +4° C to assess the importance of such predation between spiders. Small spiders (length <2.5 mm) had a mean winter mortality of 58% when kept together with large spiders (2.5 mm) which had a mean mortality of 3% only. Among the small spiders the Erigninae spp seemed to be more vulnerable to predation than other taxonomic groups. Predation also occurred when large P. phrygianus were kept together, but such predation caused mortality of less importance to the spider populations than the mortality among small spiders. Differences in spider density and food availability did not change this pattern. Considerable weight increase occurred in subadult P. phrygianus when fed during winter. This suggests that winter foraging specimens increase their fitness. Interspecific predation among spiders is suggested to be an important mortality factor in natural populations at high spider densities in November and December, when the ambient temperature often is above 0° C and when the density of large spiders is not yet substantially reduced by bird predation.     

Nest raiding by the invasive Argentine ant on colonies of the harvester ant, Pogonomyrmex subnitidus

Invasive ants often displace native ants, and published studies that focus on these interactions usually emphasize interspecific competition for food resources as a key mechanism responsible for the demise of native ants. Although less well documented, nest raiding by invasive ants may also contribute to the extirpation of native ants. In coastal southern California, for example, invasive Argentine ants (Linepithema humile) commonly raid colonies of the harvester ant, Pogonomyrmex subnitidus. On a seasonal basis the frequency and intensity of raids vary, but raids occur only when abiotic conditions are suitable for both species. In the short term these organized attacks cause harvester ants to cease foraging and to plug their nest entrances. In unstaged, one-on-one interactions between P. subnitidus and L. humile workers, Argentine ants behaved aggressively in over two thirds of all pair-wise interactions, despite the much larger size of P. subnitidus. The short-term introduction of experimental Argentine ant colonies outside of P. subnitidus nest entrances stimulated behaviors similar to those observed in raids: P. subnitidus decreased its foraging activity and increased the number of nest entrance workers (many of which labored to plug their nest entrances). Raids are not likely to be the result of competition for food. As expected, P. subnitidus foraged primarily on plant material (85% of food items obtained from returning foragers), but also collected some dead insects (7% of food items). In buffet-style choice tests in which we offered Argentine ants food items obtained from P. subnitidus, L. humile only showed interest in dead insects. In other feeding trials L. humile consistently moved harvester ant brood into their nests (where they were presumably consumed) but showed little interest in freshly dead workers. The raiding behavior described here obscures the distinction between interspecific competition and predation, and may well play an important role in the displacement of native ants, especially those that are ecologically dissimilar to L. humile with respect to diet. Received 15 July 2005; revised 19 October 2005; accepted 26 October 2005.     

Endophytic fungi alter foraging and dispersal by desert seed-harvesting ants

Endophytic fungi are thought to interact mutualistically with host plants by producing alkaloid metabolites that deter herbivory. Since such fungi are transmitted via seed in some grasses, the presence of endophytes may also protect plants from seed predators. We conducted seed choice experiments for two dominant seed harvesting ants, Pogonomyrmex rugosus in the Sonoran desert and Pogonomyrmex occidentalis at a higher elevation, riparian zone in Arizona, USA. Non-infected fescue (Festuca arundinacea) seeds and seeds infected with the endophytic fungus, Acremonium coenophialum, were presented to ant colonies in three different populations. Infected seeds were harvested less frequently than non-infected seed for the two populations of Pogonomyrmex rugosus but not for the population of Pogonomyrmex occidentalis. We also a conducted seed dispersal experiment for one population of Pogonomyrmex rugosus. Of the seeds that were harvested, most of the colonies discarded more infected seeds into refuse piles than expected by chance. Seeds discarded into refuse piles have greater germination success than surrounding areas. The most important interaction of endophytes and grasses may be deterrence of seed predation and enhancing the probability of germinating in favorable sites, since these processes directly increase plant fitness.     

Complex Nesting Behavior by Two Neotropical Species of the Ant Genus Stenamma (Hymenoptera: Formicidae)1

Two Neotropical ant species, Stenamma expolitum Smith and S. alas new species (described here), exhibit three unusual nesting behaviors: (1) they build architecturally sophisticated nest entrances that elevate the nest opening away from the surface; (2) they maintain multiple identical nests but occupy only one of them; and (3) they keep a round “door pebble” at the nest entrance, with which they plug the opening in response to army ants. Adaptive hypotheses for these behaviors are discussed, including the possibility that there are multiple lines of defense against army ant predation.     

Chemical mimicry of the ant Oecophylla smaragdina by the myrmecophilous spider Cosmophasis bitaeniata: Is it colony-specific?

Workers of most social insects can distinguish between nestmates and non-nestmates, and actively attack the latter if they attempt to intrude into the nest or surrounding territory. Nevertheless, there are many records of heterospecific organisms living within the nests of social insects, and they are thought to gain access through chemical mimicry. The salticid spider Cosmophasis bitaeniata lives within the leaf nests of the ant Oecophylla smaragdina, where it preys on the ant larvae. We investigated, using behavioural bioassays and chemical analyses, whether the previously reported resemblance of the cuticular hydrocarbons of ant and spider was colony-specific. Behavioural experiments revealed that the spiders can distinguish between nestmate and non-nestmate major workers and are less inclined to escape when confined with ants that are nestmates. More significantly, C. bitaeniata were more likely to capture ant larvae from nestmate minor workers than non-nestmate minor workers. The chemical analyses revealed that the cuticular hydrocarbon profiles of the spiders and the major workers of the ant colonies were colony-specific. However, the hydrocarbon profiles of C. bitaeniata do not match those of the major workers of O. smaragdina from the same colony. Perhaps the colony-specific cuticular hydrocarbon profiles of C. bitaeniata function to obtain prey from the minor workers rather than avoid eliciting aggression from the major workers.     

Colony cycle of an Australian swarm-founding paper wasp,Ropalidia romandi (Hymenoptera: Vespidae)

Summary Colonies of an Australian swarm founding polistine wasp,Ropalidia romandi, on the Atherton Tableland, Queensland, manage perennial nests by practicing seasonal colony activity. In summer nests, most cells were occupied by immatures of all developmental stages. The proportion of cells that did not contain immatures increased towards winter, when most nest cells did not contain immatures and half of these winter nests had rather large amounts of nectar deposited in empty cells. While the visits to tree flowers byR. romandi were not frequent in summer, nearly 60 % of insects collected on tree flowers in winter wereR. romandi. With such seasonal switching in the modes of colony reproductive activity and of nectar foraging behavior,R. romandi responds to its physical and biotic environment. In this way,R. romandi may be able to continue colonies throughout the year, with much reduced level of colony activity in winter, in areas with seasons that are unfavorable for reproductive activity of a colony.     

A latitudinal gradient in intensity of applying ant-repellent substance to the nest petiole in paper wasps (Hymenoptera: Vespidae)

Summary Observations on pre-emergence, single-foundress colonies of Japanese paper wasps (Polistes) revealed that there was a latitudinal gradient in intensity of application of an ant-repellent substance (secreted by the metasomal sternum VI glands) to the nest petiole. Thus the lower the latitude, the more frequently a foundress rubbed ant repellent onto the nest petiole. Estimation of potential ant predation on wasp brood using bait traps showed that there was a positive correlation between the frequency of rubbing and potential predation pressure from ants which are guided predominantly by substrate cues for foraging. There was also a latitudinal gradient in the degree of temporal association of rubbing behavior with foraging: the lower the latitude, the more closely foundress departure from the nest was associated with rubbing. Enlargement of the nest petiole by applying oral secretion potentially obliterated previous coats of ant repellent; however, this behavior was not always followed by rubbing behavior. The ant-repellent chemical barrier around the nest petiole may have evolved in tropical regions of the world as a defense against ant predation on wasp brood. I argue that as ant predation pressure diminishes towards the cooler regions, so does selection maintaining the behavioral sequence where foundress departure from the nest is preceded by rubbing behavior.     

Temporal patterns of seed use and availability in a guild of desert ants

ABSTRACT.

• 1 Temporal patterns of seed use were studied from late winter to autumn in three species of seed-harvesting ants in the Sonoran Desert. Measures of effective foraging activity, dietary niche breadth and dietary niche overlaps were obtained each month and were tested for correlation with estimates of the available seed resource.
• 2 Seeds were the only numerically important type of food in the diets of all species.
• 3 The ants partitioned the resource according to both seed species and seed size, although there was considerable overlap.
• 4 Pheidole xerophila had the smallest forager body size and is a specialist on small seeds because it harvested them in greater proportion than their rank in the soils and expanded its diet to larger seeds only when the abundance of small seeds declined.
• 5 When the abundance of the small seeds of Bouteloua barbata decreased, the middle-sized ant, Veromessor pergandei, showed a decrease in foraging activity, increase in niche breadth, and a decrease in overlap with P.xerophila.
• 6 Seed size preferences of V.pergandei did not vary seasonally, except that during the month of highest seed abundance, V.pergandei showed no size preference.
• 7 Pogonomyrmex rugosus was the largest ant; it preferred larger seeds and was inactive when small seeds were most abundant. Seasonal foraging activity and niche parameters were random in relation to seed abundance.
• 8 We suggest that nocturnal foraging by P.rugosus during the summer months was a response to interference with diurnal foraging by either predation frorn horned lizards or competition from V.pergandei.
• 9 Seasonal abundance of small seeds explains most of the seasonal foraging patterns of P.xerophila and V.pergandei. The summertime abundance of larger seeds during years of adequate precipitation may account for the seasonal activity patterns of P.rugosus.

     

Foraging,recruitment and predation by Decamorium uelense (Sanstchi) (Formicidae: Myrmicinae) on termites in Southern Guinea Savanna,Nigeria

Summary The myrmicine ant Decamorium uelense is a predator of small termites which feed inside their food sources of roots and plant debris. The ant uses a specialized recruitment regime to secure its prey. A scout ant searches for foraging termites and returns to the nest where it recruits a column of 10–30 ants which attack and immobilize the termites. A mass recruitment phase is then instigated, with larger numbers of ants retrieving the prey. The major prey item is Microtermes (consumed at an annual rate of 632 termites m^-2), with other small termites accounting for 0.5% of the total predation. The annual predation of Microtermes in primary savanna woodland by D. uelense removes 74% of the standing population.     

Effects of Mesopredators on Nest Survival of Shrub-Nesting Songbirds

ABSTRACT Although nest predation is often the single largest source of mortality in avian populations, manipulative studies to determine predator impacts on nest survival are rare, particularly studies that examine impacts of mid-size mammalian predators (hereafter, mesopredators) on nest survival of shrub-nesting birds. We quantified nest survival and identified nest predators of shrub-nesting songbirds within 4 large (approx. 40-ha) exclosures and 4 control sites within a longleaf pine (Pinus palustris) ecosystem. During 2003–2006, we located and monitored 535 shrub nests (222 with videography) for 4,804 nest-days to quantify daily nest survival and document predation events. We found no support for a treatment effect, suggesting mesopredators had little impact on daily nest survival (0.9303 in controls and 0.9260 in exclosures) of shrub-nesting songbirds. For the 5 most commonly monitored species, daily nest survival within species was constant. Our analysis suggested that shrub nests were most vulnerable during the nestling stage and presence of cameras on nests increased survival with the increase in survival being more pronounced during the incubation stage. We filmed 107 nest predation events, identifying predators at 88 nests. Of these 88 nests, snakes caused 33%, red imported fire ants (hereafter fire ants, Solenopsis invicta) 28%, raptors 17%, corvids 8%, mesopredators 6%, and small mammals 8% of nest predations. Cause-specific nest predation in controls and exclosures did not differ from expectation, providing evidence that compensatory predation did not occur. Nest predators differed from expectation with regard to nest stage; fire ants and raptors only depredated nests during the nestling stage. Presence of cameras had no effect on nest abandonment. Fire ants were the most prevalent nest predator, and nest predation by fire ants was only observed on nestlings, potentially reducing likelihood of renesting. Magnitude and timing of fire ant predation suggests that fire ants may be the most influential nest predator of shrub-nesting birds within the longleaf pine ecosystem. Our data suggest that controlling mesopredators will have no effect on nest success of shrub-nesting birds within longleaf pine forests.     

Seed dispersal curve of a Mediterranean myrmecochore: Influence of ant size and the distance to nests

Euphorbia characias is a Mediterranean spurge with a diplochorous dispersal system: after a ballistic dispersal that scatters the seeds, some ant species find and retrieve the seeds to their nest (myrmecochorous dispersal). The seed dispersal curve generated by ants in an abandoned field was described and partitioned according to ant size and to the distance to nest entrance from where seeds fell after ballistic dispersal. Both variables (ant size, distance to nest) affected dispersal distance. The seed dispersal curve showed a peak at short distance (median=1m) and a tail extending to 9.4m. The peak and the tail are explained differently. Short distances are usually generated by small ants (Pheidole pallidula and Tapinoma nigerrimum; 0.56±0.41m [n=48]) both from the nearest or farther nest entrances. The tail of the curve is generated disproportionately by big ants (Aphaenogaster senilis and Messor barbarus; 2.09±1.71m [n=61]) from farther nests. Seeds have a much greater probability (P=0.734) of being transported to nests which are not the nearest. This effect is largely due to transportation by big ants.     

Nest-plugging: interference competition in desert ants (Novomessor cockerelli and Pogonomyrmex barbatus)

Summary Pogonomyrmex barbatus and Novomessor cockerelli, sympatric species of harvester ants in the Lower Sonoran desert, compete for seed resources. This study reports on a method of interference competition. Early in the morning, before P. barbatus' activity period, N. cockerelli fills the nest entrances of P. barbatus with sand. This delays the beginning of the P. barbatus activity period for 1–3 h. P. barbatus colonies near N. cockerelli nests were more likely to be plugged. Nest-plugging shifts the typical daily sequence of P. barbatus activities, including the onset of foraging, forward towards midday, when high temperatures force the colony back inside the nest. P. barbatus colonies do not compensate for late emergence or events impeding foraging by increasing foraging rate. Thus nest-plugging by N. cockerelli decreases the foraging capacity of P. barbatus colonies.     

Social behaviour ofRopalidia fasciata (Hymenoptera: Vespidae) females on satellite nests and on a nest with multiple combs

The social behaviour ofRopalidia fasciata females on 3 satellite nests and on a nest with multiple, independent combs was observed. Many females often visited satellite nests, and even dominant foundresses, who usually spent more of their time on the nest, became active when satellite nests were being constructed. Three foundresses laid eggs on a satellite nest and eggs laid by 2 of these (who performed foraging) survived. On another satellite nest, only the dominant foundress laid an egg which survived. On a nest consisting of 2 independent combs, females frequently shifted between the 2 combs even after the emergence of many adults. Constructions of satellite nests and of multiple independent combs are considered to be adaptive for this species living in Okinawa, where most of nests are destroyed by frequent typhoons or abandoned after predation by ants.