Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. The mechanisms by which Argentine ants effectively compete against native ant species have been previously addressed in field studies that centered on interference and exploitation competition at baits and mainly examined the colony-level performance of Argentine ants. Detailed behavioral observations explaining the basis for the strong competitive ability of L. humile are comparatively rare. To gain a better understanding of the mechanisms by which Argentine ants displace native ants we examined the aggressive interactions between the Argentine ants and the odorous house ant, Tapinoma sessile in four different aggression assays: (1) worker dyad interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a resource competition assay which focused on competition for food and nesting space. Our results demonstrate a clear disparity between worker-level and colony-level fighting ability of Argentine ants and provide behavioral evidence to explain the superior interference ability of Argentine ants in group assays. Argentine ants experienced mixed success in fighting against odorous house ants in dyad interactions, but gradually gained a numerical advantage in symmetrical group interactions by active cooperation among nestmates. Results of the resource competition assay indicate that Argentine ants recruit rapidly, numerically dominate food and nesting sites, and aggressively displace T. sessile from baits. Taken together, the results of these assays allow us to pinpoint the behavioral mechanisms responsible for the remarkable competitive ability of Argentine ants.     

Patterns of DNase I sensitivity in the chromosomes of the ant Tapinoma nigerrimum (Hymenoptera,Formicidae)

We have analysed the patterns of DNase I/nick translation in the chromosomes of Tapinoma nigerrimum. The results show a non-uniform DNase I sensitivity in different chromosome domains. The hypersensitivity appears to be specially concentrated at both the NOR and the distal regions. The resemblance to and differences from the situation in other animal species, in which active genes are DNase I hypersensitive, are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Relationships between the ant Brachymyrmex obscurior (Hymenoptera, Formicidae) and Acacia pennatula (Fabaceae)

Summary. In central Mexico, the ant Brachymyrmex obscurior Forel feeds on nectar produced by extrafloral nectaries of Acacia pennatula (Schlecht. & Cham.) Benth. However, no studies have determined whether the ants visitation is related to plant nectar availability and whether ants protect A. pennatula from herbivory. The objectives of this 2-yr study (2000–2001) were to assess whether seasonal changes in ant visitation coincide with extrafloral nectar productivity in A. pennatula and to determine whether ants protect the plant. At the end of the dry season (April–June) B. obscurior was the only ant species on A. pennatula and extrafloral nectar production is limited to this period. Exclusion experiments, performed at the end of the dry season showed that A. pennatula did not receive a protective benefit when visited by ants. Branches with ants and branches where ants are excluded had similar numbers of the nonmyrmecophile leafhopper Sibovia sp. which was the only herbivore observed under natural conditions.Received 24 March 2004; revised 4 September 2004; accepted 8 September 2004.     

Seasonal polydomy in a polygynous supercolony of the odorous house ant,Tapinoma sessile

Abstract 1. The odorous house ant, Tapinoma sessile, is a native ant species common throughout North America. In its natural habitat, T. sessile is a low‐key species that consists of small colonies. In invaded urban areas, T. sessile exhibits extreme polygyny and polydomy and becomes a dominant invasive pest. 2. The current study examined: (i) the density, persistence, and the spatial distribution of nests in a large supercolony of T. sessile, (ii) trail abundance and overall colony connectivity as facilitated by the network of trails, (iii) the abundance and the spatial distribution of competing ant species, and (iv) the effect of environmental factors on the number and distribution of T. sessile nests. 3. A distinct pattern of seasonal polydomy was observed, whereby the colony undergoes an annual fission‐fusion cycle. The colony occupies one or a few nests during the winter, experiences rapid exponential growth in the spring to colonize available nesting sites, reaches maximum nest density in the summer, and again coalesces in the winter, returning to the same winter location year after year. The trails show spatio‐temporal variation as well, depending on the location of nesting and foraging sites. Furthermore, nest movements may be driven by soil microclimate and proximity to man‐made structures. 4. In total, 119 ant nests were discovered in a 3.15 ha plot, 90 (76%) of which belonged to T. sessile. Tapinoma sessile exhibited strong colony connectivity as 78/90 (87%) of nests were connected to at least one other nest by a trail. Mean persistence time for T. sessile nests was 133 ± 5 days. 5. Results indicate that T. sessile is a highly adaptable native ant species that exhibits a high degree of flexibility in its colony social structure. A high degree of polygyny and polydomy may contribute to its ecological dominance and pest status in urban environments.     

Observation of group recruitment to prey in a primitive ponerine ant,Amblyopone sp. (reclinata group) (Hymenoptera: Formicidae)

Summary Group recruitment during foraging was observed in a primitive ponerine ant,Amblyopone sp. (reclinata group) under laboratory condition. Workers searched for prey singly; however, if a item of prey was stung by a worker, other ants joined the attack. After the prey became immobile, one of the workers laid a trail directly toward the nest. This scout worker recruited additional workers (between 3 and 33). They formed a single file procession to the point of prey capture, and cooperatively transported the prey. A scout worker could stimulate nest workers to leave the nest without direct contact, and the recruited workers could trace the trail without guidance by the scout worker. This is the first report of recruitment behavior during foraging in the primitive antAmblyopone.     

Dispersed central-place foraging in the polydomous odorous house ant, Tapinoma sessile as revealed by a protein marker

The odorous house ant, Tapinoma sessile, is a native ant species common throughout North America. In urban areas, this ant is classified a pest species and exhibits several attributes characteristic of invasive “tramp” ants (sensu Passera, 1994). These include: extreme polygyny, colony reproduction by budding, reduced internest aggression, generalist diet, and polydomy. Here we explore the organization of foraging and the pathways of food distribution in polydomous colonies of T. sessile in the laboratory and field using a novel marking technique (rabbit IgG protein) and enzyme linked immunosorbent assay (ELISA). Laboratory assays revealed patterns of food allocation from foragers to other castes and developmental stages. Foragers distributed the IgG- labelled sucrose to the majority of workers within 24 h, and workers retained significantly more sucrose than either queens or larvae. Approximately 50% of queens tested positive for the IgG marker and some queens received significantly more sucrose than others, indicating a possible reproductive dominance hierarchy. Larvae received little sucrose demonstrating their minor reliance on carbohydrates. The results of field experiments showed that odorous house ants are dispersed central-place foragers whereby ants from individual nests exhibit high foraging site fidelity, travel along well-established trails, and forage on a local scale. Dispersed central-place foraging most likely allows the odorous house ant to more efficiently secure both clumped and dispersed food sources and possibly increases its competitive ability. As a result, colonies become numerically large and ecologically dominant. The results of our study contribute to our understanding of the social behavior and colony organization in T. sessile. In addition, they provide a framework for designing more effective ant control programs based on liquid baits. Received 13 December 2005; revised 28 February 2006; accepted 3 March 2006.     

Mating flight, metrosis, and semi-claustrality in the seed-harvester ant Pogonomyrmex salinus (Hymenoptera, Formicidae)

Here we examine dispersal, metrosis, and claustrality in the seed-harvester ant Pogonomyrmex salinus at an unusually large mating aggregation. We found that mode of queen dispersal from the mating aggregation is not a function of queen mass and that wing damage among queens was relatively rare. P. salinus is haplometrotic in the field and foundress queens placed together in forced associations eventually fight to the death. While queens of Pogonomyrmex salinus can survive claustrally, producing a single minim from their body reserves in the laboratory, fed queens produce up to four significantly larger minims along with concurrent larvae and pupae during the same period. Since queens forage in the field, we interpret claustrality as a secondary reserve strategy when foraging fails, and suggest that foraging is obligate for P. salinus queens in an overdispersed and temperate environment. Thus, nest founding strategies employed by P. salinus may be environmentally determined and represent a continuum between fully claustral and obligate foraging. We discuss our results with reference to theories of pleometrosis and claustral colony founding. Received 12 November 2004; revised 12 April 2004; accepted 29 July 2005.     

The blind leading the blind in army ant raid patterns: Testing a model of self-organization (Hymenoptera: Formicidae)

We present field experiments and analyses that test both the assumptions and the predictions of a model that showed how the swarm raids of the army ant Eciton burchellimight be self-organizing, i.e., based on hundreds of thousands of interactions among the foraging workers rather than a central administration or hierarchical control. We use circular mill experiments to show that the running velocity of the ants is a sigmoidal function of the strength of their trail pheromones and provide evidence that the swarm raid is structured by the interaction between outbound and inbound forager traffic mediated by the pheromones produced by both of these sets of ants. Inbound traffic is also affected by the distribution of prey, and hence, sites of prey capture alter the geometry of the raid. By manipulating the prey distributions for E. burchelliswarms, we have made them raid in a form more typical of other army ant species. Such self-organization of raids based on an interaction between the ants and their environment has profound consequences for interpretations of the evolution of army ant species.     

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.     

The influence of forager number and colony size on food distribution in the odorous house ant, Tapinoma sessile

Stomodeal trophallaxis plays a major role in ant colony nutrition and communication. While the rate of food distribution at the individual level (worker to worker) is rapid, factors affecting the rate of food distribution at the colony level remain poorly understood. We used the odorous house ant, Tapinoma sessile (Say), as a model species to investigate the factors affecting the rate of spread of liquid carbohydrate food throughout a colony. To track the movement of the food we used protein marking and double antibody sandwich enzyme-linked immunosorbent assay, DAS-ELISA. Increasing colony size while keeping the number of donor workers constant significantly decreased the number of individuals testing positive for the marker. After 8 h of trophallactic interactions with ten donors, 92 ± 5% of recipient workers tested positive in a colony of 125 and 38 ± 5% tested positive in a colony of 1,000. Interestingly, as colony size increased and the percentage of workers testing positive decreased, the proportion of workers actually receiving food increased. Food originating from a single donor fed approximately 12 individuals in colonies comprised of 125 recipients and approximately 38 individuals in colonies comprised of 1,000 recipients. Thus, the per capita consumption of food decreased as colony size increased, most likely because the amount of food reaching the colony was limited. Increasing the number of donors while keeping colony size constant significantly increased the number of recipient ants testing positive for the marker. As the number of donor workers doubled, the percentage of recipients testing positive more than doubled suggesting that the number of individuals receiving food increases with increasing colony size, while the per capita amount of food decreases. When food was available ad libitum and in close proximity to the nest, numerous workers fed directly at the food source. This dramatically increased the rate and the extent of food distribution to both the workers and the queens and colony size had no significant effect on the spread of the marker in the workers or the queens. The rate and the extent of food distribution at the colony level may depend on a number of factors including the number of successful foragers, the size and density of the recipient colony, and the recipient caste.     

黑头酸臭蚁微卫星标记的分离与开发(英文)

【目的】本研究旨在分离黑头酸臭蚁Tapinoma melanocephalum基因组微卫星标记,确定这些微卫星位点的多态性。【方法】使用454 GS-FLX焦磷酸测序技术开发来自中国华南陆地和岛屿的11个黑头酸臭蚁地理种群基因组微卫星位点。从随机设计的100对微卫星引物中筛选出10对引物,用于确定黑头酸臭蚁4个地理种群［东澳岛(DAD)、荷包岛(HBD)、梅州(MZ)和山咀(SJ)］10个微卫星位点的多态性,分析种群遗传多样性和种群分化。【结果】从11个黑头酸臭蚁地理种群基因组中成功开发和分离10对微卫星引物。在DAD, HBD, MZ和SJ 4个地理种群中,10个微卫星位点中7个有高多态性,这10个位点均显著偏离Hardy-Weinberg平衡;每个位点的等位基因数量(A)是3.50~9.00个,每个地理种群每个位点等位基因丰富度(A_R)在1.992~12.938之间。岛屿地理种群(DAD和HBD)的AR和预期杂合度(H_E)与大陆地理种群(MZ和SJ)的相比差异不显著。4个地理种群均显示高水平遗传分化(F_ST=0.15969);HBD和MZ种群与其他配对地理种群相比,遗传分化较高(F_ST=0.185),基因流较低,说明这两个种群基因流被限制。此外,遗传变异来自种群内个体之间。【结论】筛选新的微卫星位点能够为研究黑头酸臭蚁种群结构和繁殖结构提供有效工具,以深入了解其传播机制。     

Interactions between the ant Tapinoma nigerrimum (Hymenoptera: Formicidae) and the main natural enemies of the vine and citrus mealybugs (Hemiptera: Pseudococcidae)

Honeydew-excreting hemipterans, such as mealybug pests, can be protected from their natural enemies by tending ants in return for honeydew, thereby compromising the aims of biological control. In this respect, antagonistic interactions between the ant Tapinoma nigerrimum, native to the Mediterranean basin, and the main natural enemies of both the vine mealybug (VMB), Planococcus ficus, and the citrus mealybug (CM), Planococcus citri, were assessed in laboratory conditions. Parasitism of vine and CMs by their respective parasitoids, Anagyrus sp. nr. pseudococci and Leptomastix dactylopii, was negatively affected by the ant T. nigerrimum. Similarly, T. nigerrimum was shown to significantly disrupt the predatory potential of ladybird larvae, Cryptolaemus montrouzieri, when foraging on host CMs. By contrast, the presence of the ant did not negatively influence the predatory activity of C. montrouzieri adults when feeding on CMs. Consequently, the encyrtid parasitoids A. sp. nr. pseudococci and L. dactylopii and the larval stage of the predator C. montrouzieri may be considered as T. nigerrimum-sensitive, whereas the adults of C. montrouzieri may be regarded as T. nigerrimum-resistant predators. Accordingly, the ant T. nigerrimum constitutes a threat to the biological control of mealybugs by either the encyrtids A. sp. nr. pseudococci and L. dactylopii or the larval stage of the ladybird C. montrouzieri. Hence, adequate control of T. nigerrimum is highly recommended before any release of these mealybugs' natural enemies.     

Raiding behavior of the dulotic antChalepoxenus muellerianus (Finzi) in the field (Hymenoptera: Formicidae,Myrmicinae)

Summary Little information is available regarding the raiding behavior in nature of dulotic ants belonging to the tribeLeptothoracini. Between July 17 and 24, 1991, several raids ofChalepoxenus muellerianus (Finzi) were observed in nature near Tignale/Lago di Garda, Italy. Apparently, this species raids frequently during its summer raiding season (5 raids were observed during 7 days spent observing 8 colonies). A singleChalepoxenus colony sometimes raids more than one host colony more or less simultaneously. Observations during which oneChalepoxenus colony raided another and captured slavemaker brood indicate that intraspecific raids can occur either accidentally, or as a result of competition or territoriality when there is a sufficient dense slavemaker population.     

Host colony usurpation by the queen of the Japanese pirate ant, Polyergus samurai (hymenoptera: formicidae)

Colony usurpations by newly mated queens of Polyergus samurai were observed under artificial conditions. Newly mated queens of P. samurai were introduced into three kinds of Formica japonica host colonies: queenright, queenless (artificially orphaned), and workerless (only a queen remaining) colonies. In the queenright condition, the P. samurai queen intruded into the host nest and killed the host queen, and was subsequently adopted by the host workers. In all queenright and queenless host colonies, seven of 13 queens of P. samurai succeeded in colony usurpation, although the starting time of grooming, a nestmate behavior, by host workers in the queenright condition occurred earlier than in the queenless condition. In workerless conditions, four of five P. samurai queens ignored the F. japonica queen. The results suggest that while host-queen killing is not necessary, it is important to win acceptance by host workers.     

Trail-following responses ofTapinoma simrothi (Formicidae: Dolichoderinae) to pygidial gland extracts

Summary The pygidial (anal) gland was found to be the source of trail pheromone in the antTapinoma simrothi. Bioassays conducted with fractionated pygidial gland secretion indicated that the fraction containing iridodials and iridomyrmecin is responsible for the trail pheromone activity. Thus workers ofT. simrothi may utilize the same glandular exudate for alarm and trail following. At high emission rates from a point source, the ants responded in alarm, e.g., rushed to the source with open mandibles and raised abdomen. When concentrations were low and drawn as a line, the ants followed the secretion calmly. Trails ofT. simrothi are long-lived, having a biological half-life of 10 to 19 days. Quantitative studies of the evaporation rates of the iridodials by gas chromatography resulted in a half-life of 11 days, agreeing with the biological data. The implications of the use of the same glandular secretion for alarm and food recruitments are discussed.     

Patterns in foraging and nesting ecology in the neotropical ant, <Emphasis Type="Italic">Gnamptogenys moelleri</Emphasis> (Formicidae,Ponerinae)

Summary This study provides quantitative field data on the natural history and foraging behaviour of the Neotropical bromeliad-nesting ant Gnamptogenys moelleri (Ponerinae) in a sandy plain forest in Southeast Brazil. The ant nested on different bromeliad species and the nests were more frequently found in bigger bromeliads. The species used a wide array of invertebrates in its diet, hunting for live prey and scavenging the majority of the items from dead animals. The food items varied greatly in size (1 to 26 mm). Hunting was always performed by solitary workers. Retrieving was performed by solitary workers (small items), or by a group of 3 to 12 workers recruited to the food source (large items). Almost all G. moelleri foraging activity was restricted to the nest bromeliad. In the warm period more ants left the nest to forage, and foraging trips achieved greater distances compared to the cool season. Trap data revealed that overall availability of arthropod prey is higher in the summer than in the winter. The opportunism in nest site use and in foraging behaviour, the small foraging area, as well as the seasonal differences in foraging activity are discussed and compared with other tropical ants.Received 30 May 2003; revised 22 September 2003; accepted 3 October 2003.     

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.     

Communication in the primitive cryptobiotic ant Prionopelta amabilis (Hymenoptera: Formicidae)

Summary The Neotropical ant Prionopelta amabilis, a cryptobiotic species in the phylogenetically primitive tribe Amblyoponini, lives in subterranean habitats, where it preys preferentially on campodeid diplurans and other small arthropods. Here we report that the species employs chemical recruitment and orientation trails during foraging and nest emigrations. The trail pheromone originates in a hitherto unknown basitarsal gland located in the basitarsus of the hindlegs. Trails are laid by a special foot dragging behavior. During the recruitment process the chemical trail signal is complemented by body shaking on the part of the recruiting ant. Foragers frequently wipe the basitarsal gland opening in the hindlegs with the grooming apparatus of the front legs. This latter structure is equipped with unusual glands evidently specialized for this purpose.