Nestmate recognition in the unicolonial ant Formica paralugubris

In unicolonial populations of ants, individuals can mix freelywithin large networks of nests that contain many queens. Ithas been proposed that the absence of aggression in unicolonialpopulations stems from a loss of nest mate recognition, butfew studies have tested this hypothesis. We investigated patternsof aggression and nest mate recognition in the unicolonial woodant, Formica paralugubris. Little aggression occurred, evenbetween workers from nests separated by up to 5 km. However,when aggression took place, it was directed toward non–nestmates rather than nest mates. Trophallaxis (exchange of liquidfood) occurred very frequently, and surprisingly, workers performedsignificantly more trophallaxis with non–nest mates thanwith nest mates (bias 2.4:1). Hence, workers are able to discriminatenest mates from non–nest mates. Higher rates of trophallaxisbetween non–nest mates may serve to homogenize the colonyodor or may be an appeasement mechanism. Trophallaxis rate andaggression level were not correlated with geographical distanceand did not differ within and between two populations separatedby several kilometers. Hence, these populations do not representdifferentiated supercolonies with clear-cut behavioral boundaries.Overall, the data demonstrate that unicoloniality can evolvedespite well-developed nest mate recognition. Reduced levelsof aggression might have been favored by the low rate of interactionswith foreign workers, high cost of erroneously rejecting nestmates, and low cost of accepting foreign workers.     

When supercolonies collide: territorial aggression in an invasive and unicolonial social insect

Some species of ants possess an unusual form of social organization in which aggression among nests is absent. This type of social organization, called unicoloniality, has been studied in only a handful of species and its evolutionary origins remain unclear. To date, no study has examined behavioural and genetic patterns at points of contact between the massive supercolonies that characterize unicoloniality. Since interactions at territory boundaries influence the costs of aggression and the likelihood of gene flow, such data may illuminate how supercolonies are formed and maintained. Here we provide field data on intraspecific territoriality for a widespread and invasive unicolonial social insect, the Argentine ant (Linepithema humile). We observed abrupt and well-defined behavioural boundaries at 16 contact zones between three different pairs of supercolonies. We visited nine of these zones weekly during a six-month period and observed consistent and intense intercolony aggression that resulted in variable, but often large, levels of worker mortality. Microsatellite variation along six transects across territory borders showed that F(ST) values were lower within supercolonies (0.08 +/- 0.01 (mean +/- SE)) than between supercolonies (0.29 +/- 0.01) and that this disparity was especially strong right at territory borders, despite direct and prolonged contact between the supercolonies. Matrix correspondence tests confirmed that levels of aggression and genetic differentiation were significantly correlated, but no relationship existed between geographic distance and either intraspecific aggression or genetic differentiation. Patterns of F(ST) variation indicated high levels of gene flow within supercolonies, but little to no gene flow between them. Overall, these findings are inconsistent with a model of relaxed ecological constraints leading to colony fusion and suggest that environmentally derived cues are not the prime determined of nestmate recognition in field populations of Argentine ants.     

Food availability and brood number do not influence intraspecific aggression in Argentine ants

Summary. Nestmate recognition cues can derive from both environmental and genetic factors, but can also be modulated in response to context-specific cues. Synchronous changes in nestmate recognition systems occur seasonally in some species of ants, however the mechanisms underlying these seasonal changes are often unknown. We studied two mechanisms, relative brood number and food availability, to determine if they generate temporal variation in intraspecific aggression in an introduced population of the Argentine ant, Linepithema humile. Using data from previous studies we found that seasonal increases in aggression levels correlate with seasonal increases in brood-to-worker ratios in the field. However, when we manipulated brood-to-worker ratios in paired experimental colonies, we found no direct evidence that relative brood numbers influenced aggression levels. To determine if food availability influenced aggression we conducted a second experiment in which we randomly assigned pairs of experimental colonies to starved or fed treatments and then measured aggression levels weekly for five weeks. We observed no difference in the level of aggression between these two treatment groups indicating that food availability also has no affect on aggression levels between hostile conspecific colonies.Received 24 August 2004; revised 15 September 2004; accepted 23 September 2004.     

马尾松-木荷不同比例混交林种内和种间竞争强度

分析不同比例组成的马尾松(Pinus massoniana)-木荷(Schima superba)混交林树木间的竞争关系,可为营造种间关系协调的马尾松混交林提供参考依据.以马尾松-木荷混交林为研究对象,根据其混交比例分别类型Ⅰ(对照,10马)、类型Ⅱ(8马2木)、类型Ⅲ(7马3木)、类型Ⅳ(6马4木)4种类型设置标准地...     

浙江天台山甜槠种内与种间竞争研究

采用Hegyi的单木竞争指数模型对天台山的甜槠种内、种间的竞争强度进行定量分析。结果表明 ,甜槠种内竞争强度随着林木径级的增大而逐渐减小 ,种内竞争较之与其伴生树种间的竞争剧烈。甜槠种内、种间竞争强度的顺序为 :甜楮 >木荷 >马尾松 >尾叶冬青 >虎皮楠 >短柄木包 >珍珠栗。竞争木对对象木的竞争强度与对象木的个体大小服从幂函数关系 ,竞争强度和对象木个体的大小呈极显著的负相关关系。当甜槠胸径达到 30cm后 ,竞争强度变化不明显 ,说明此时该生态系统已基本达到稳定状态。     

Argentine ant foraging activity and interspecific competition in complete vs. queenless and broodless colonies

Introduction experiments may prove useful in understanding the mechanisms underlying the successful establishment of invasive ant species into new areas. These manipulative introductions could be particularly helpful in exploring the interactions between invasive species and the local fauna and flora. However, the inherent risk of accidental establishment in such experiments poses unacceptable ethical concerns. Some of the worst invasive species are tramp ant species, which can adversely affect biodiversity and community structure after establishment. We conducted laboratory and field experiments investigating a safe methodology for carrying out introduction experiments using the sterile workers of the invasive Argentine ant, Linepithema humile, as a model. We found no difference in foraging rate between worker-only colonies of L. humile and complete colonies, containing queens, workers and brood. Worker-only L. humile colonies showed the same exploitative and interference ability as complete colonies in bait dominance trials with the odorous house ant, Tapinoma sessile, in both laboratory and field trials. We suggest that for those invasive ant species with sterile workers, worker-only colonies may be substituted for complete colonies in short-term field experiments in new areas. Received 18 January 2007; revised 19 June 2007; accepted 22 June 2007.     

Genetic and behavioural evidence for a city-wide supercolony of the invasive Argentine ant Linepithema humile (Mayr) (Hymenoptera: Formicidae) in southeastern Australia

The success of invasive ants is frequently attributed to genetic and behavioural shifts in colony structure during or after introduction. The Argentine ant ( Linepithema humile ), a global invader, differs in colony genetic structure and behaviour between native populations in South America and introduced populations in Europe, Japan, New Zealand and North America. However, little is known about its colony structure in Australia. We investigated the genetic structure and behaviour of L. humile across Melbourne, Victoria by quantifying variation at four microsatellite loci and assaying intraspecific aggression at neighbourhood (30–200 m), fine (1–3.3 km) and regional (5–82 km) spatial scales. Hierarchical analyses across these scales revealed that most genetic variation occurred among workers within nests (∼98%). However, although low genetic differentiation occurred among workers between nests at the fine and regional scales (∼2%), negligible differentiation was detected among workers from neighbouring nests. Spatial genetic autocorrelation analysis confirmed that neighbouring nests were genetically more similar to each other. Lack of aggression within and across these scales supported the view that L. humile is unicolonial and forms a large supercolony across Melbourne. Comparisons of genetic structure of L. humile among single nests sampled from Adelaide, Brisbane, Hobart and Perth with Melbourne showed no greater levels of genetic differentiation or dissimilar spatial structure, suggesting an Australia-wide supercolony.     

Population structure and intraspecific aggression in the invasive ant species Anoplolepis gracilipes in Malaysian Borneo

Invasive species are one of the main sources of the ongoing global loss of biodiversity. Invasive ants are known as particularly damaging invaders and their introductions are often accompanied by population-level behavioural and genetic changes that may contribute to their success. Anoplolepis gracilipes is an invasive ant that has just recently received increased attention due to its negative impact on native ecosystems. We examined the behaviour and population structure of A. gracilipes in Sabah, Malaysia. A total of 475 individuals from 24 colonies were genotyped with eight microsatellite markers. Intracolonial relatedness was high, ranging from 0.37 to 1 (mean +/- SD: 0.82 +/- 0.04), while intercolonial relatedness was low (0.0 +/- 0.02, range -0.5-0.76). We compared five distinct sampling regions in Sabah and Brunei. A three-level hierarchical F-analysis revealed high genetic differentiation among colonies within the same region, but low genetic differentiation within colonies or across regions. Overall levels of heterozygosity were unusually high (mean H(O) = 0.95, mean H(E) = 0.71) with two loci being entirely heterozygous, indicating an unusual reproductive system in this species. Bioassays revealed a negative correlation between relatedness and aggression, suggesting kinship as one factor facilitating supercolony formation in this species. Furthermore, we genotyped one individual per nest from Sabah (22 nests), Sarawak (one nest), Brunei (three nests) and the Philippines (two nests) using two mitochondrial DNA markers. We found six haplotypes, two of which included 82.1% of all sequences. Our study shows that the sampled area in Sabah consists of a mosaic of differently interrelated nests in different stages of colony establishment. While some of the sampled colonies may belong to large supercolonies, others are more likely to represent recently introduced or dispersed propagules that are just beginning to expand.     

Behavioral and Genetic Differentiation Between Native and Introduced Populations of the Argentine Ant

In this paper, we examine the hypothesis that reduced intraspecific aggression underlies the competitive prowess of Argentine ants in their introduced range. Specifically, we test three predictions of this hypothesis by comparing the genetic diversity, behavior, and ecology of Argentine ants in their native range to introduced populations. Differences between native and introduced populations of Argentine ants were consistent with our predictions. Introduced populations of the Argentine ant appear to have experienced a population bottleneck at the time of introduction, as evidenced by much reduced variation in polymorphic microsatellite DNA markers. Intraspecific aggression was rare in introduced populations but was common in native populations. Finally, in contrast to the Argentine ant's ecological dominance throughout its introduced range, it did not appear dominant in the native ant assemblages studied in Argentina. Together these results identify a possible mechanism for the widespread success of the Argentine ant in its introduced range.     

Kin recognition and the paradoxical patterns of aggression between colonies of a Mojave desert Pheidole ant

Populations of the desert seed-harvesting ant Pheidole xerophylla are often characterized by high nest density leading to competitive interactions between foragers from different nests. We investigated the inter-nest aggression, spatial distribution and genetic structure of a P. xerophylla population of the Mojave Desert in Southern California. Inter-nest aggression was quantified by standardized staged encounters in a neutral arena. Genetic relatedness within nests and relatedness between nests were calculated using allelic frequencies at four microsatellite-DNA loci. We found a bimodal distribution of inter-colony aggression levels with a first mode at low aggression levels and another mode at much higher aggression levels. Inter-colony aggression levels were largely non-transitive. No effect of geographical distance on inter-nest aggression levels was detected. Despite high amounts of variation in inter-colony relatedness ( − 0.24 to 0.37) this variable did not correlate with the level of aggression between nests. Intra-nest relatedness ranged from 0.40 to 0.75 and close inspection of worker genotypes within colonies revealed a high proportion of polygynous colonies or a mixture of polygyny and polyandry. Aggression levels among nests was found to decrease with increasing intra-nest relatedness. These results do not support the idea that aggression is modulated by a nestmate recognition mechanism based on overall genetic similarity. Instead, the absence of transitivity found in inter-colony aggression and bimodal distribution of aggression levels are compatible with a common label acceptance model of nestmate recognition and suggest that label diversity may be encoded by a limited number of loci. Received 29 March 2005; revised 8 September 2005; accepted 27 September 2005.     

Evidence that queens do not influence nestmate recognition in Argentine ants

Summary Introduced populations of many invasive ants exhibit low levels of intraspecific aggression. Argentine ants (Linepithema humile), for example, maintain expansive supercolonies in many parts of their introduced range. Recent studies demonstrate that variation in nestmate recognition in L. humile can derive from both environmental and genetic sources. In some ants, pheromones emitted by queens also influence nestmate-recognition behavior. To test if such a phenomenon occurs in Argentine ants, we examined whether levels of intraspecific aggression vary as a function of queen presence or absence in experimental lab colonies. For each of four known supercolonies from southwestern California, we set up a pair of experimental colonies and randomly assigned replicates within each pair to treatment (queen removal) and control (no queen removal) groups. Using two different behavioral assays, we then measured aggressive behavior for ten days, removed queens from colonies in the treatment group, and continued to monitor aggression in both experimental groups for an additional 65 days. Both assays yielded qualitatively similar results: intraspecific aggression remained high throughout the experiment in both experimental groups. These results suggest that L. humile queens fail to influence levels of intraspecific aggression in introduced populations.Received 2 June 2003; revised 1 September 2003; accepted 18 September 2003.     

Global-scale analyses of chemical ecology and population genetics in the invasive Argentine ant

Ants are some of the most abundant and ecologically successful terrestrial organisms, and invasive ants rank among the most damaging invasive species. The Argentine ant is a particularly well-studied invader, in part because of the extreme social structure of introduced populations, known as unicoloniality. Unicolonial ants form geographically vast supercolonies, within which territorial behaviour and intraspecific aggression are absent. Because the extreme social structure of introduced populations arises from the widespread acceptance of conspecifics, understanding how this colonymate recognition occurs is key to explaining their success as invaders. Here, we present analyses of Argentine ant recognition cues (cuticular hydrocarbons) and population genetic characteristics from 25 sites across four continents and the Hawaiian Islands. By examining both hydrocarbon profiles and microsatellite genotypes in the same individual ants, we show that native and introduced populations differ in several respects. Both individual workers and groups of nestmates in the introduced range possess less diverse chemical profiles than ants in the native range. As previous studies have reported, we also find that introduced populations possess much lower levels of genetic diversity than populations in the native range. Interestingly, the largest supercolonies on several continents are strikingly similar to each other, suggesting that they arose from a shared introduction pathway. This high similarity suggests that these geographically far-flung ants may still recognize and accept each other as colonymates, thus representing distant nodes of a single, widely distributed supercolony. These findings shed light on the behaviour and sociality of these unicolonial invaders, and pose new questions about the history and origins of introduced populations.     

Resource-based models of competitive interactions. I. Intraspecific competition in Ratibida columnifera ( Asteraceae)

We constructed and tested a series of regression models of intraspecific competition in Ratibida columnifera (Asteraceae), based on the growth and water use of individual plants. Models were constructed from a set of plants ("model") grown without competition under three watering regimes. Each model was then tested on another set of plants ("test") grown, singly or in pairs, under two watering regimes, one of them different than those of the "model" plants. Both sets of plants were grown simultaneously. Models that used only estimates of plant dry mass (ISON [interval by interval size only], and SON [size only]) were outperformed (i.e., the difference between predicted values of final dry mass and their true values were larger on average) by models that incorporated direct measures of water uptake and usage (ISAW [interval by interval size and water], WON [water only], SAW [size and water]). Harvest biomass predictions given by these three last models deviated from values of true biomass by an average of only 6.1%.     

Kin structure and queen execution in the Argentine ant Linepithema humile

Every spring, workers of the Argentine Ant Linepithema humile kill a large proportion of queens within their nests. Although this behaviour inflicts a high energetic cost on the colonies, its biological significance has remained elusive so far. An earlier study showed that the probability of a queen being executed is not related to her weight, fecundity, or age. Here we test the hypothesis that workers collectively eliminate queens to which they are less related, thereby increasing their inclusive fitness. We found no evidence for this hypothesis. Workers of a nest were on average not significantly less related to executed queens than to surviving ones. Moreover, a population genetic analysis revealed that workers were not genetically differentiated between nests. This means that workers of a given nest are equally related to any queen in the population and that there can be no increase in average worker–queen relatedness by selective elimination of queens. Finally, our genetic analyses also showed that, in contrast to workers, queens were significantly genetically differentiated between nests and that there was significant isolation by distance for queens.     

Intraspecific competition through food robbing in the harvester ant,Messor aciculatus (Fr. Smith), and its consequences on colony survival

Summary Intraspecific interference competition in the harvester ant,Messor aciculatus, was studied. Colonies of this species were found not to have territories. Some nests were located very close to each other, and the foraging areas of the neighbors usually overlapped. Even though the frequency with which alien and resident ants met was very high in the vicinity of the nest entrances, aggressive interactions between them rarely occurred. However, when hostile workers encountered each other, they exhibited a kind of ritualized combat and the winner ejected, but did not injure the loser. If any aliens entered the nest, some of them were pulled out, mainly by the residents.Aliens roaming near a neighbor's nest entrance ferociously attacked the residents carrying seeds in their mandibles and robbed them. On other occasions, aliens entered the nest and stole the collected seed. Although seed robbing and stealing occurred among neighboring colonies, there were remarkable differences in the frequency of their occurrence. The results of field observations and experiments suggest the existence of a dominance order among the neighbors. In one instance, extermination of an inferior colony by its neighbor was observed. The raider colony transferred the stored seeds from the nest of the inferior colony to its own and deposited the larvae and workers some distance away from the nest.The influence of ritualized combat and food robbing on colony activities, and the ecological significance of this interference behavior in terms of spatial distribution and temporal persistence of the nest sites, is discussed.     

Variation in the level of aggression, chemical and genetic distance among three supercolonies of the Argentine ant in Europe

In their invasive ranges, Argentine ant populations often form one geographically vast supercolony, genetically and chemically uniform within which there is no intraspecific aggression. Here we present regional patterns of intraspecific aggression, cuticular hydrocarbons (CHCs) and population genetics of 18 nesting sites across Corsica and the French mainland. Aggression tests confirm the presence of a third European supercolony, the Corsican supercolony, which exhibits moderate to high levels of aggression, depending on nesting sites, with the Main supercolony, and invariably high levels of aggression with the Catalonian supercolony. The chemical analyses corroborated the behavioural data, with workers of the Corsican supercolony showing moderate differences in CHCs compared to workers of the European Main supercolony and strong differences compared to workers of the Catalonian supercolony. Interestingly, there were also clear genetic differences between workers of the Catalonian supercolony and the two other supercolonies at both nuclear and mitochondrial markers, but only very weak genetic differentiation between nesting sites of the Corsican and Main supercolonies (F(ST) = 0.06). A detailed comparison of the genetic composition of supercolonies also revealed that, if one of the last two supercolonies derived from the other, it is the Main supercolony that derived from the Corsican supercolony rather than the reverse. Overall, these findings highlight the importance of conducting more qualitative and quantitative analyses of the level of aggression between supercolonies, which has to be correlated with genetic and chemical data.     

Absence of aggression but not nestmate recognition in an Australian population of the Argentine ant Linepithema humile

Intraspecific aggression is rare within introduced populations of the Argentine ant Linepithema humile, and colonies exhibit a structure known as unicoloniality, in which aggression among nests is atypical. We document a similar form of colony structure in an introduced population of Argentine ants in Victoria, Australia, in which aggression is extremely rare among nests ranging over hundreds of kilometres. However, using a highly sensitive behavioural bioassay we found that workers display subtle differences in their behaviour towards non-nestmates and nestmates. In particular, non-nestmates consistently engage in antennating behaviour with greater frequency than nestmates, perhaps providing a mechanism for homogenization of nest odour. Further, we found that non-nestmates at seaport sites (where populations may derive from multiple introductions) antennate each other with greater frequency than their counterparts from non-seaport sites. These data suggest that the Victorian population of L. humile may comprise multiple independent introductions. Received 4 July 2007; revised 15 January and 4 March 2008; accepted 4 March 2008.     

Experimental Evidence of Large-Scale Unicoloniality in the Tramp Ant Wasmannia auropunctata (Roger)

We examined intraspecific colonial aggressiveness in Wasmannia auropunctata (Roger), a tramp species originating from the neotropics. By observing the results of one-on-one confrontations, we compared the behavioral responses of workers originating from six New Caledonian locations (introduced range) and four Brazilian cocoa plantations (original range). We recorded interindividual aggressive behavior on four levels ranging from physical contact, with no aggressive response, to prolonged aggressiveness, including stinging by one or both ants. In Brazil, we often observed high intraspecific aggressiveness between populations originating from distant locations, indicating that W. auropunctata may behave as a multicolonial species in its native range. In New Caledonia, paired encounters resulted in low agonistic behavior, as shown by the absence of full attacks (which include stinging by one or both opponents). Our results suggest that W. auropunctata behaves as a single supercolony throughout New Caledonia and that the scale of its unicoloniality (widespread colonies with interconnected nests without aggressiveness between workers originating from distant areas) is different in introduced and native populations. According to the present study, it seems likely that differences in intraspecific aggressiveness between native and introduced populations of W. auropunctata contribute to its invasive success.