Floral visitation patterns of two invasive ant species and their effects on other hymenopteran visitors

Abstract.  1. Floral nectar of the native Hawaiian 'ōhi'a tree, Metrosideros polymorpha , is an important food source for several native honeycreepers and yellow-faced bees, Hylaeus spp., but is also attractive to invasive ants.
2. I undertook this study to compare floral visitation patterns of two widespread invasive ants, the Argentine ant, Linepithema humile , and the big-headed ant, Pheidole megacephala , and to determine their effects on nectar volume and floral hymenopteran visitors.
3. In the first year of the study, Argentine ants visited inflorescences more frequently than big-headed ants at mid-day and in the afternoon, but did not occur in higher densities than big-headed ants at any time of day. In the following year, Argentine ants visited inflorescences both more frequently and in higher densities than big-headed ants. Argentine ant density had a stronger association with nectar concentration than big-headed ant density.
4. Nectar volume did not differ between ant-excluded and ant-visited inflorescences for either ant species. However, ant density was negatively associated with nectar volume for both species.
5.  Hylaeus spp. never visited inflorescences with big-headed ants, while non-native honeybees visited inflorescences with and without ants of either species in equal frequency.
6. Most studies of the effects of invasive ants on native arthropods have focused on interactions on the ground. Flowers should not be overlooked as microhabitats from which native arthropods may be displaced by invasive ants.     

Biogeography and morphological evolution in a Pacific island ant radiation

While insular radiations are documented for many terrestrial arthropods, few examples are known for eusocial insects. This study seeks to ascertain whether the spinescence observed among Fijian Pheidole ants was inherited from an oversea ancestor or is evidence of ecological release from interspecific competitors. We broaden our understanding of morphological convergence, insular radiation and Pacific biogeography by testing three hypotheses proposed previously for the Fijian Pheidole roosevelti group: (i) the group is monophyletic; (ii) spinescence is a plesiomorphic trait inherited from an overseas ancestor; and (iii) the group is closely related to spinescent New Guinean relatives. The analysis included the fragments of two mitochondrial genes (COI, cytb) and two nuclear genes (H3, EF1α-F2) from 66 taxa, including all members of the roosevelti group, representatives from the spinescent subgenus Pheidolacanthinus, Fijian congeners and widespread Pacific congeners. Our results yield new insights into the biogeographic history of Fiji, reveal a fascinating example of convergent evolution and serve as a novel example of ecological release occurring within an insular eusocial insect lineage. These findings recover the history of a presumably unremarkable ant species that colonized a remote oceanic archipelago in the Miocene (17-10 Ma) and radiated across the emerging islands into niche-space occupied elsewhere in the Pacific by distantly related spinescent congeners. We propose the radiation of Fijian Pheidole into spinescent morphotypes was the consequence of ecological opportunities afforded by the absence of competing ant lineages with conspicuous epigaeic foraging strategies.     

Induced biotic responses to herbivory and associated cues in the Amazonian ant-plant Maieta poeppigii

Ants inhabiting ant‐plants can respond to cues of herbivory, such as the presence of herbivores, leaf damage, and plant sap, but experimental attempts to quantify the dynamic nature of biotic defenses have been restricted to a few associations between plants and ants. We studied the relationship between certain features of the ant‐shrub Maieta poeppigii Cogn. (Melastomataceae) and the presence or absence of ant patrolling on the leaf surface in plants occupied by the ant Pheidole minutula Mayr (Hymenoptera: Formicidae). We also carried out field experiments to examine ant behavior following plant damage, and the potential cues that induce ant recruitment. These experiments included clipping of the leaf apex, as well as the presentation of a potential herbivore (live termite worker) and a foliar extract from Maieta on treatment leaves. The presence of ants patrolling the leaves of M. poeppigii is influenced by the number of domatia on the plant. Ant patrolling on the leaves of M. poeppigii was constant throughout a 24 h cycle, but the mean number of patrolling ants decreased from young to mature leaves, and from leaves with domatia to those without domatia. There was an overall increase in the number of ants on experimental leaves following all treatments, compared to control leaves. Visual and chemical cues associated with herbivory are involved in the induction of ant recruitment in the Maieta–Pheidole system. The continuous patrolling behavior of ants, associated with their ability to respond rapidly to foliar damage, may result in the detection and repellence/capture of most insect herbivores before they can inflict significant damage to the leaves.     

Relationship between Plant Size and Ant Associates in Two Amazonian Ant‐Plants1

A survey of two Amazonian melastome ant‐plants, Maieta guianensis and Tococa bullifera, revealed a significant difference in plant size according to the species of ant inhabiting the plant. Plants with Crematogaster laevis, on average, were smaller than those with Pheidole minutula (in M. guianensis) and those with Azteca sp. (in T. bullifera). There is no evidence that these patterns were due either to the deterministic replacement of C. laevis by another ant species during host‐plant ontogeny or to a habitat effect on plant growth rates coupled with colony survival. More likely, the smaller size of C. laevis plants can be explained by its effects on host‐plant performance. Plants with C. laevis lost their associated ant colonies more frequently than plants with P. minutula and Azteca sp. Plants that lost their C. laevis either died, or more commonly, were severely defoliated. Defoliated plants, once sprouted, tended to become recolonized, but such recolonizations were not deterministic so as to favor one species over another. Plants with C. laevis showed similar, or only slightly greater, standing levels of herbivory than plants with P. minutula or Azteca sp. This suggests that when C. laevis is present, it confers some degree of protection to its hosts. It was found that early in colony development, queens of C. laevis moved off their host plants to build satellite nests in dead twigs on the ground, a behavior not seen in the other two species and one that possibly renders colonies more vulnerable to mortality from predation, flooding, or nest decay. Comparable 8¹⁵N values in C. laevis and P. minutula indicate that the two species are equally dependent on food supplied by the host plant.     

Detrimental effects of two widespread invasive ant species on weight and survival of colonial nesting seabirds in the Hawaiian Islands

Invasive ants are a significant conservation concern and can have far-reaching effects in ecosystems they invade. We used the experimental control of ant numbers on two pairs of small (<5 ha) offshore islets dominated by either the big-headed ant, Pheidole megacephala or the tropical fire ant, Solenopsis geminata to investigate the influence of these species on seabird hatching success, fledging success and weight. Limited unpublished observations of both ant species attacking nesting seabirds exist, but the frequency of attacks or how they affect seabird growth and survival are unknown. Island-wide treatments with hydramethylnon resulted in the eradication of P. megacephala and the temporary reduction of S. geminata densities. No difference in hatching success, growth, or fledging success of Wedge-tailed Shearwaters (Puffinus pacificus), a common colonial nesting seabird in the Hawaiian Islands was observed on the pair of islets dominated by P. megacephala. On islets dominated by S. geminata, ant control resulted in a temporary increase in fledging success. Injury frequency increased dramatically on the untreated islet (8.3–100%) while remaining the same on the treated islet (27–38%). Severely injured chicks (i.e., chicks that lost >20% of tissue on their feet) weighed significantly less than uninjured chicks and did not fledge. It is unclear if the chicks were being preyed upon or stung in defense of nearby ant colonies. Radical changes in invasive ant populations have been noted, and booming ant populations could cause short-term, but widespread damage to seabird colonies. The negative effects of invasive ants on seabirds may be difficult to detect, and therefore unknown or underestimated throughout the world where the two groups overlap.     

Internest sex-ratio variation and male brood survival in the ant Pheidole pallidula

Sex allocation in social insects has become a general modelin tests of inclusive fitness theory, sex-ratio theory, andparent-offspring conflict. Several studies have shown that colonysex ratios are often bimodally distributed, with some coloniesproducing mainly females and others mainly males. Sex specializationmay result from workers assessing their relatedness to malebrood versus female brood, relative to the average worker-relatednessasymmetry in other colonies of their population. Workers thenadjust the sex ratio in their own interest This hypothesis assumesthat workers can recognize the sex of the brood in their colonyand selectively eliminate males. We compared the primary sexratio (at the egg stage) and secondary sex ratio (reproductivepupae and adults) of colonies in the ant Pheidole pallidula.There was a strong bimodal distribution of secondary sex ratios,with most colonies producing mainly reproductives of one sex.In contrast, there was no evidence of a bimodal distributionof primary sex ratios. The proportion of haploid eggs producedby queens was 0.35 in early spring and decreased to about 0.1in summer. Male eggs also were present in virtually all fieldcolonies sampled in July, although eggs laid at this time ofyear never give rise to males. All male brood is, therefore,selectively eliminated beginning in July and continue to beeliminated through the rest of the year. Finally, the populationsex-ratio investment was female-biased. Together, these resultsare consistent with the hypothesis that workers control thesecondary sex ratio by selectively eliminating male brood inabout half the colonies, perhaps those with high relatednessasymmetry.[Behav Ecol 7: 292–298 (1996)]     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

The nest architecture of the ant, Pheidole oxyops Forel, 1908 （Hymenoptera： Formicidae）

Pheidole oxyops builds subterranean nests, with an external architecture that is distinctive and easily recognizable by its wide and specific entrance hole, measuring up to 12.2 cm in diameter, denoting a pitfall-trap. In order to study the nests＇ internal architecture, seven nests were excavated; four were identified with neutral talc, while the others were cast in cement and then excavated. Measurements were made in order to gain a better understand- ing of their structures, and a photographic documentation was obtained as well. The excavations revealed that the nests are perpendicular relative to the ground, beginning with a cylindrical channel with a mean length of 13.5 cm, containing irregular formations, and whose diameter becomes progressively narrower until the first chamber is formed. As the channel continues, dish-like chambers appear, interconnected by channels that become progressively narrower and longer, while the chambers are arranged at greater distances from each other as nest depth increases. Both channels and chambers are located on the vertical projection of the entrance hole. Nests may reach a depth of up to 5.09 m, with a number of chambers ranging between 4 and 14.     

Survey of the natural enemies of Dysmicoccus mealybugs on pineapple in Hawaii

Surveys for mealybugs, associated natural enemies and ants were conducted in abandoned pineapple fields on the Hawaiian islands of Oahu and Maui from July 1992 to November 1993. Whole plant samples were taken, and mealybugs and ants found were identified. Mealybug-infested plant parts were isolated and held until natural enemies emerged from parasitized host material. At sample sites where only Dysmicoccus brevipes was present, its densities ranged from a mean of 23 to 157 mealybugs per plant, while in areas with mixed populations of this mealybug and Dysmicoccus neobrevipes Beardsley, densities ranged from a mean of 23 to 118 mealybugs per plant. Ants were present at all sample sites and on all dates. Pheidole megacephala (F.) was the most common ant species found. Anagyrus ananatis Gahan was the most common parasitoid reared. However, it attacked only D. brevipes, the dominant mealybug in the pineapple fields surveyed. Percent parasitization of D. brevipes by A. ananatis in the presence of ants ranged from 0.3 to 9.9%. Percent parasitization of D. brevipes and D. neobrevipes per plant by Euryrhopalus propinquus Kerrich ranged from 0.05 to 2.2%. Mean densities of the predators Lobodiplosis pseudococci (Felt), Nephus bilucernarius Mulsant and Sticholotis ruficeps Weise ranged from 0.05 to 5.75, 0.1 to 1.8, and 0.05 to 0.2 individuals per plant, respectively.     

Response of an open‐forest ant community to invasion by the introduced ant,Pheidole megacephala

The introduced tramp ant, Pheidole megacephala, is a well‐known pest of urban areas and coastal dune ecosystems in eastern Australia. Until recently, establishment and spread of P. megacephala colonies has been regarded as likely only in disturbed areas. Here we describe the extent of an established colony of P. megacephala in a long undisturbed open forest near Maryborough in southeast Queensland and compare ant community structure with those of nearby uninfested sites. Tuna baiting revealed three distinct zones: (i) a zone totally dominated by P. megacephala (at least 10 ha) where few other ant ant species were detected; (ii) a zone where P. megacephala was absent and many other ant species were found; and (iii) a zone where opportunists (species of Ochetellus and Paratrechina) competed with P. megacephala at baits. Pitfall trapping over a 9‐month period resulted in 12 species being recorded at the infested site, compared with a mean of 25 species recorded at adjoining uninfested forest. Over 94% of ants recorded in pitfalls at the infested site were P. megacephala. Most notably, P. megacephala had completely displaced dominant Dolichoderines (species of Iridomyrmex), subordinate Camponotini (species of Camponotus, Opisthopsis and Polyrhachis) and other species of Pheidole which are common at forest sites.