Impact of the introduced yellow crazy ant Anoplolepis gracilipes on Bird Island, Seychelles

The introduced yellow crazy ant or long-legged ant Anoplolepis gracilipes was first reported in Seychelles in 1969 and now occurs on at least nine islands in the Central Seychelles. We describe the yellow crazy ant's effects on vegetation and invertebrate communities on one of these, Bird Island; in 2000, Anoplolepis (first reported in 1991) occurred there at densities at least 80 times higher than on other islands in the Central Seychelles. They were associated with high densities of coccid scale insects on foliage, especially of the native tree Pisonia grandis, in some instances causing tree death. Yellow crazy ants on Bird Island also significantly affected invertebrate communities on foliage and on the ground, both in terms of taxonomic composition and the density of specific taxa, apparently causing the local exclusion of some invertebrates.     

Prevalence and genetic diversity of three bacterial endosymbionts (Wolbachia, Arsenophonus, and Rhizobiales) associated with the invasive yellow crazy ant (Anoplolepis gracilipes)

Invasive species carry pathogens, parasites and mutualistic microorganisms into their new range. We examined the prevalence and genetic diversity of three bacterial endosymbionts (Wolbachia, Arsenophonus, and Rhizobiales) in four Anoplolepis gracilipes (Smith) populations in the Pacific region, four populations from mainland Australia and one population from Christmas Island in the Indian Ocean. Wolbachia was found in eight of the nine sampling sites, with between 31.8 and 100% of ants infected. These infection rates are substantially higher than those previously observed for other invasive ants such as Linepithema humile and Solenopsis invicta. All sequences of Wolbachia were genetically identical. Arsenophonus was observed in five of the nine sampling sites, with infection rates ranging between 4.5 and 50.8%. Like Wolbachia, Arsenophonus can modify the sex-ratio of its hosts via male-killing. Arsenophonus was found to co-occur with Wolbachia in the same ants in five of the nine sampling sites. Rhizobiales is a clade of symbiotic bacteria mostly found in herbivorous ants. These bacteria help provide nitrogen to their hosts and were found in only three of the nine sampling sites with an infection rate of 1.6–11.8%. It also co-occurred with the other bacteria. There was no genetic variation in Arsenophonus and Rhizobiales samples, with the exception of a sequence from one Arsenophonus sample in Samoa that differed by a single base pair. These bacterial endosymbionts may contribute to the population variability in A. gracilipes and may be manipulated for the purpose of pest management.     

Impact of the invasive crazy ant Anoplolepis gracilipes on Bird Island, Seychelles

The crazy ant (Anoplolepis gracilipes) invaded Bird island, Seychelles, in the 1980s. In 1997, its range expanded and population densities increased. The impacts of this change were studied in 2001 using a combination of arthropod collecting methods. The ant population excluded larger invertebrates (principally the large ant Odontomachus simillimus and the crabs, principally Ocypode spp.). Cockroaches, however, remained abundant in ant-infested areas and tree-nesting birds (Lesser Noddy Anous tenuirostris) appear to be able to breed successfully in the presence of the crazy ant. The ants are only abundant in areas of deep shade which provide cool nesting areas, yet enabling them to forage in the open when ground temperatures fall. The expansion of the ants was correlated with the regeneration of woodland on the island. Recommendations are made for the management of the woodland which may reduce the impacts of the crazy ant.     

Spatial dynamics of supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on Christmas Island, Indian Ocean

Key to the management of invasive species is an understanding of the scope of an invasion, the rate of proliferation and the rate at which invaded habitats become degraded. This study examines the spatial dynamics of high-density supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes , on Christmas Island, Indian Ocean, and the associated impacts at their boundaries. Since the early 1990s, A. gracilipes supercolonies have occupied over 30% of the 10,000 ha of rainforest on Christmas Island. Thirty-four discrete high-density supercolonies formed between 1989 and 2003, ranging in size across nearly three orders of magnitude from 0.9 to 787 ha. Supercolonies boundaries are diffuse, and ants were observed in low densities in some cases up to 200 m from the main high-density supercolony. The 13 boundaries examined were all dynamic over a 10–20 observation month period: nine boundaries expanded, and the maximum rate of spread was 0.5 m day⁻¹. Across boundary transition zones, between high-density supercolonies and intact rainforest, yellow crazy ants reduced other ant species richness, occupied red crab burrows and killed resident red crabs, which was the trigger for 'invasional meltdown' on Christmas Island. The highly variable and unpredictable nature of A. gracilipes boundaries poses a challenge for incorporation into a predictive framework, as well as for their management.     

Discovery of independent‐founding solitary queens in the yellow crazy ant Anoplolepis gracilipes in East Java,Indonesia (Hymenoptera: Formicidae)

We collected four solitary queens of the invasive ant Anoplolepis gracilipes under stones in East Java, Indonesia. They produced nanitic workers by claustral colony foundation. This is the first report of independent colony foundation by queens in this species. The discovery may give an important insight into discussion on the origin of this invasive ant.     

Supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on an oceanic island: Forager activity patterns, density and biomass

Summary. Ants have the capacity to reach unusually high densities, mostly in their introduced ranges. Numerical dominance is often cited as key to the ability of exotic ants to displace native ant species, reduce the abundance of invertebrates and negatively impact upon bird, land crab and other vertebrate populations. On Christmas Island, Indian Ocean, the yellow crazy ant, Anoplolepis gracilipes (Jerdon), forms supercolonies, where extremely high densities of foraging ants have contributed to ‘invasional meltdown’ in rainforest areas. Densities of up to 2254 foraging ants per m² and a biomass of 1.85 g per m² were recorded, and nest densities reached 10.5 nest entrances per m². Populations of A. gracilipes can overcome and kill red endemic land crabs (Gecarcoidea natalis) over 100 times their own biomass. This is the highest recorded density of foraging ants, and adds another element to the definition of ‘supercolony’ of unicolonial ants. This paper documents one extreme in a continuum of densities of unicolonial, invasive ant species and highlights the need to incorporate forager densities into invasive ant research.Received 17 November 2004; revised 14 February 2005, accepted 21 February 2005.     

Characterization of microsatellite markers for the invasive ant species Anoplolepis gracilipes

We present primer sequences for eight polymorphic microsatellite loci for the formicine ant Anoplolepis gracilipes, a serious pest species in South‐East Asia and Pacific islands and still spreading on all continents. Microsatellite loci were isolated with a highly efficient method of enrichment. The number of alleles ranged from two to 19 with an observed heterozygosity ranging from 0.842 to 1.0. The markers were designed for a sociogenetic study as well as for population genetics.     

Colony structure, seasonality and food requirements of the crazy ant, Anoplolepis longipes (Jerd.), in the Seychelles

Abstract. 1. In Seychelles, A. longipes nests on the ground and in trees, underground nests do not occur, possibly because soils are too hard or unstable.
2. Nest density approached 700 ha^-1 in some areas and nests contained, on average, about 4000 ants. The total population (including foragers) in heavily infested areas may exceed 10 millions ha^-1.
3. On average, each nest contained about forty queens and fifty males, although only 50% of nests contained males. A few nests contained up to 300 queens, 1000 males, 36 000 workers and 23 000 brood.
4. Production of sexual adults and brood was related to the wet season, most being produced shortly before or after the heavy rains from November to March.
5. Natural spread is probably by budding of colonies which, during an expansive period, may advance 1 m day^-1.
6. Food collected by the colony included sugary substances from fruits, honey-dew-producing insects and plant exudates, and proteinaceous material, mainly in the form of insect prey and carcases. A forager collected, on average, 1.2 mg of liquid or 2.8 mg of solid material on each foraging trip.
7. Foraging occurred at fluctuating intensities throughout the day and night, but was limited by heavy rain, strong winds and substrate temperatures above 30°C. Maximum activity occurred at ground temperatures of 25–30°C and ceased about 44°C.
8. Availability of food and nest sites proably have the greatest influence on population size.     

Yellow crazy ant (<Emphasis Type="Italic">Anoplolepis gracilipes</Emphasis>) invasions within undisturbed mainland Australian habitats: no support for biotic resistance hypothesis

Ants are highly successful invaders, especially on islands, yet undisturbed mainland environments often do not contain invasive ants, and this observation is largely attributed to biotic resistance. An exception is the incursion of Yellow crazy ant Anoplolepis gracilipes within northeast Arnhem Land. The existence of A. gracilipes within this landscape’s intact environments containing highly competitive ant communities indicates that biotic resistance is not a terminally inhibitory factor mediating this ant’s distribution at the regional scale. We test whether biotic resistance may still operate at a more local scale by assessing whether ecological impacts are proportional to habitat suitability for A. gracilipes, as well as to the competitiveness of the invaded ant community. The abundance and species richness of native ants were consistently greater in uninfested than infested plots but the magnitude of the impacts did not differ between habitats. The abundance and ordinal richness of other macro-invertebrates were consistently lower in infested plots in all habitats. A significant negative relationship was found for native ant abundance and A. gracilipes abundance. No relationships were found between A. gracilipes abundance and any measure of other macro-invertebrates. The relative contribution of small ants (<2.5 mm) to total abundance and relative species richness was always greater in infested sites coinciding with a reduction of the contribution of the larger size classes. Differences in the relative abundance of ant functional groups between infested and uninfested sites reflected impacts according to ant size classes and ecology. The widespread scale of these incursions and non-differential level of impacts among the habitats, irrespective of native ant community competitiveness and abiotic suitability to A. gracilipes, does not support the biotic resistance hypothesis.     

Origin and evolution of the dependent lineages in the genetic caste determination system of Pogonomyrmex ants

Hybridizing harvester ants of the Pogonomyrmex barbatus/rugosus complex have an exceptional genetic caste determination (GCD) mechanism. We combined computer simulations, population genomics, and linkage mapping using >1000 nuclear AFLP markers and a partial mtDNA sequence to explore the genetic architecture and origin of the dependent lineages. Our samples included two pairs of hybridizing lineages, and the mitochondrial and nuclear data showed contradicting affinities between them. Clustering of individual genotypes based on nuclear markers indicated some exceptions to the general GCD system, that is, interlineage hybrid genes as well as some pure-line workers. A genetic linkage map of P. rugosus showed one of the highest recombination rates ever measured in insects (14.0 cM/Mb), supporting the view that social insects are characterized by high recombination rates. The population data had 165 markers in which sibling pairs showed a significant genetic difference depending on the caste. The differences were scattered in the genome; 13 linkage groups had loci with F(ST)>0.9 between the hybridizing lineages J1 and J2.The mapping results and the population data indicate that the dependent lineages have been initially formed through hybridization at different points in time but the role of introgression has been insignificant in their later evolution.     

Wing disc development during caste differentiation in the ant Pheidole megacephala (Hymenoptera: Formicidae)

The genus Pheidole has three distinct castes in females: queen, major, and minor workers. It has been believed that the larvae of major workers have prominent mesothoracic wing discs, although the minor worker larvae lack them. Here we conducted histological examinations of wing discs during larval development in P. megacephala. We show that all three castes have mesothoracic wing discs, at least in their early stage of the final larval instar, and that the wings degenerate differently in the dimorphic worker castes. The minute wing discs of minor workers neither grow nor metamorphose but disappear during the prepupal stage. On the contrary, the wing discs of major workers evaginate at the onset of the prepupal stage but subsequently degenerate by apoptotic cell death. This apoptotic wing degeneration in the prepupal stage was contradistinguished from wing degeneration in some lepidopteran insects, in which apoptosis occurs in the pupal wing buds. Our results suggest that each worker caste shows a different degeneration process to express the wingless character and that apoptotic degeneration has been adopted in association with the evolution of worker dimorphism.     

The microsporidian pathogen Myrmecomorba nylanderiae: Intracolony transmission and impact upon tawny crazy ant (Nylanderia fulva) colonies

The microsporidian pathogen Myrmecomorba nylanderiae Plowes et al. infects introduced tawny crazy ants (Nylanderia fulva (Mayr)), and constitutes one of the first natural enemies known to attack this invasive ant. We assess how infection is transmitted within colonies and how infection impacts N. fulva colony fragment growth under carbohydrate‐deficient and carbohydrate‐sufficient dietary conditions. Carbohydrate scarcity is a common source of stress for ant colonies. Infected workers efficiently pass infection to developing larvae but all other potential pathways for within colony transmission are rare or non‐viable. For unknown reasons, queens within infected colony fragments are generally uninfected, limiting the role of transovarial transmission in intracolony transmission. In the laboratory, infection by M. nylanderiae primarily impacts the growth of N. fulva colonies by reducing pupal production. Colony growth showed a substantially greater impact under carbohydrate‐deficient conditions implying that the effect of the pathogen may depend on seasonally variable carbohydrate availability. In the colony growth assay, worker mortality did not differ with infection status under either nutrient regime. However, in a longer, direct test of survivorship, infected worker survivorship was significantly lower. Recently, some established N. fulva populations with high prevalence of M. nylanderiae infection have declined precipitously, though it is unknown if M. nylanderiae is a causative agent in these declines. The combination of chronic impacts, presence in North America, and potential association with population declines makes M. nylanderiae a promising prospect for the biological control of N. fulva.     

Molecular genetic markers provide no evidence for reproductive isolation among retreat building phenotypes of the net-spinning caddisfly Macrostemum carolina

Larvae of the stream-dwelling, filter-feeding caddisfly Macrostemum carolina construct silken catchnets within protective retreats. In the Savannah River, M. carolina individuals make three different retreats, each with a distinct water entrance hole: (i) at the end of a silken tube; (ii) with a approximately 180 degrees silken backstop; and (iii) flush with the top of the retreat. To resolve whether these different retreats represent alternative behavioural phenotypes within a single panmictic population or fixed phenotypes within three genetically distinct populations or species, we compared the allele frequencies at three polymorphic nuclear loci (allozyme electrophoresis for Gpi, Mpi and Pgm) and the mitochondrial DNA (mtDNA) haplotype frequencies among individuals displaying the three retreat morphs. We also calculated pairwise exact tests of population differentiation using the allozyme and mtDNA allele frequencies. No significant genetic differentiation was detected among caddisflies exhibiting the different retreat morphs. Therefore, these morphs apparently represent a single panmictic population in the Savannah River. Consequently, additional study is required to assess whether this retreat polymorphism is a phenotypically plastic trait under conditional control, or is mediated by alternative alleles at a Mendelian gene or genes (or a combination of the two).     

Chemical disguise as particular caste of host ants in the ant inquiline parasite Niphanda fusca (Lepidoptera: Lycaenidae)

The exploitation of parental care is common in avian and insect 'cuckoos' and these species engage in a coevolutionary arms race. Caterpillars of the lycaenid butterfly Niphanda fusca develop as parasites inside the nests of host ants (Camponotus japonicus) where they grow by feeding on the worker trophallaxis. We hypothesized that N. fusca caterpillars chemically mimic host larvae, or some particular castes of the host ant, so that the caterpillars are accepted and cared for by the host workers. Behaviourally, it was observed that the host workers enthusiastically tended glass dummies coated with the cuticular chemicals of larvae or males and those of N. fusca caterpillars living together. Cuticular chemical analyses revealed that N. fusca caterpillars grown in a host ant nest acquired a colony-specific blend of cuticular hydrocarbons (CHCs). Furthermore, the CHC profiles of the N. fusca caterpillars were particularly close to those of the males rather than those of the host larvae and the others. We suggest that N. fusca caterpillars exploit worker care by matching their cuticular profile to that of the host males, since the males are fed by trophallaxis with workers in their natal nests for approximately ten months.     

Turkish honeybees: genetic variation and evidence for a fourth lineage of Apis mellifera mtDNA

The mtDNA of bees from 84 colonies of Turkish honeybees (Apis mellifera) was surveyed for variation at four diagnostic restriction sites and the sequence of a noncoding intergenic region. These colonies came from 16 locations, ranging from European Turkey and the western Mediterranean coast to the Caucasus Mountains along the Georgian border, the eastern Lake Van region, and the extreme south. Combined restriction site and sequence data revealed four haplotypes. Three haplotypes belonged to the eastern Mediterranean mtDNA lineage. The fourth haplotype, which had a novel restriction site pattern and noncoding sequence, was found in samples from the extreme south, near the Syrian border. We found two different noncoding sequences among the eastern Mediterranean haplotypes. The "Caucasian" sequence matches that described from A. m. caucasica, and the "Anatolian" sequence matches that of A. m. carnica. The frequency of the "Caucasian" sequence was highest (98-100%) in sites near the Georgian border and decreased steeply to the south and west. Elsewhere the Anatolian sequence was found. In European Turkey (Thrace) a restriction site polymorphism previously reported from A. m. carnica in Austria and the Balkans was present at high frequency. A novel mtDNA haplotype with a unique restriction site pattern and noncoding sequence was found among bees from Hatay, in the extreme south near the Syrian border. This haplotype differed from the three previously known lineages of honeybee mtDNA--African, western European, and eastern Mediterranean-and may represent a fourth mitochondrial lineage.     

Queen size dimorphism in the ant Tetramorium moravicum (Hymenoptera, Formicidae): Morphometric, molecular genetic and experimental evidence

Summary. By combining different methods we evaluate whether the ant Tetramorium rhenanum is specifically separated from T. moravicum or whether it is a conspecific microgyne form. High-precision morphometry shows a clear difference in queen size. Sequence comparison of 1031 bp of COI reveals that T. rhenanum falls into a clade with T. moravicum, which is significantly separated from T. forte and T. chefketi. T. rhenanum shares at least two haplotypes with T. moravicum and is considered as a junior synonym. Sexual behaviour and colony foundation experiments corroborate conspecificity. The queen dimorphism is discussed in the context of social parameters such as queen number and reproductive strategy.Received 10 June 2004; revised 28 October 2004; accepted 12 November 2004.     

Molecular features of a defined genetic marker for the determination of the Porphyra tenera lineage

Nucleotide sequences of the nuclear SSU rDNA and ITS1 are presented as a defined genetic marker for Porphyra tenera as a species. Exon nucleotide sequences were identical within all the P. tenera specimens. Intron nucleotide sequences varied between populations. The introns and ITS1 variations are presented as defined genetic markers to establish the Porphyra tenera strains. Wild-collected thalli identified by morphological systematics, from five populations of Porphyra tenera throughout Japan, were discriminated by comparing sequences of the various regions utilizing the results of this and previous studies.