Aliens among us: nestmate recognition in the social huntsman spider, Delena cancerides

Unlike all other social spiders, the social huntsman spider, Delena cancerides, has been reported to rapidly respond to non-nestmates with lethal aggression, similar to the behavior of some eusocial insects. We tested for the presence of nestmate recognition in D. cancerides under laboratory conditions by introducing 105 unrelated alien conspecifics into foreign colonies and comparing their behavior to 60 control spiders removed and returned to their natal colony. Spiders demonstrated nestmate recognition by investigating alien spiders far more than nestmates and by resting closer to nestmates than to aliens. Serious attacks or deaths occurred in 23% of all trials; however, aggression was not directed significantly more toward aliens than to nestmates. Most notably, aggression was largely mediated by the adult females (resident or alien), who were most likely to attack or kill other subadult or mature individuals. Young individuals (resident or alien) were largely immune from serious aggression. Spiders recently collected from the field tended to be more aggressive than spiders born and raised in the laboratory, possibly due to blurring of recognition cues related to laboratory husbandry. Our findings support the prediction that nestmate recognition should evolve when there is a benefit to discriminating against non-kin, as in this social spider system where foraging individuals may enter a foreign colony and the colony retreat is a limited resource.     

Nest volatiles as modulators of nestmate recognition in the ant Camponotus fellah

When ants from alien colonies encounter each other the stereotypic reaction is usually one of aggressive behavior. It has been shown that factors such as queen-derived cues or nest-odors modulate this reaction. Here we examined whether nest volatiles affect nestmate recognition by observing the reaction of nestmates towards individual workers under one of four regimes: completely isolated; isolated but receiving a constant airflow from the mother colony; as previous but with the passage of nest volatiles towards the isolated ants blocked by adsorption on a SuperQ column; or reversed airflow direction-from the isolated ants to the nest interior. Ants that had been completely isolated for three weeks were subjected to aggressive behavior, but not those that had continued to receive airflow from the mother colony. Adsorbing the nest volatiles from the airflow by SuperQ abolished this difference, with these ants now also being subjected to aggression, indicating that nest volatiles can modulate nestmate recognition. Reverse airflow also reduced the level of aggression but to a lesser extent than airflow directed from the mother colony. In queenless colonies the overall aggression was reduced under all regimes, and there was no effect of flow, suggesting that the volatiles involved are queen-borne. The SuperQ adsorbed volatiles originated from Dufour's gland secretions of both workers and queen, implicating them in the process. Cuticular hydrocarbon profiles were not affected by exposure to nest volatiles, suggesting that the latter either constitute part of the recognition cues or affect worker behavior via a different, as yet elusive mechanism.     

Not one odour but two: A new model for nestmate recognition

Recognition systems play a key role in a range of biological processes, including mate choice, immune defence and altruistic behaviour. Social insects provide an excellent model for studying recognition systems because workers need to discriminate between nestmates and non-nestmates, enabling them to direct altruistic behaviour towards closer kin and to repel potential invaders. However, the level of aggression directed towards conspecific intruders can vary enormously, even among workers within the same colony. This is usually attributed to differences in the aggression thresholds of individuals or to workers having different roles within the colony. Recent evidence from the weaver ant Oecophylla smaragdina suggests that this does not tell the whole story. Here I propose a new model for nestmate recognition based on a vector template derived from both the individual’s innate odour and the shared colony odour. This model accounts for the recent findings concerning weaver ants, and also provides an alternative explanation for why the level of aggression expressed by a colony decreases as the diversity within the colony increases, even when odour is well-mixed. The model makes additional predictions that are easily tested, and represents a significant advance in our conceptualisation of recognition systems.     

Wide home ranges for widely foraging lizards

Space usage by animals may be influenced by a range of factors. In this study we investigate whether foraging behaviour affects the home range size of lizards. Two distinct tactics of foraging have been recognized in predators: sit-and-wait foraging (SW) and active foraging (AF). Foraging activity level of a data set of lizard species, mainly compiled from literature, is compared with their home range sizes. Two opposite predictions can be made about foraging in connection with home range area: on the one hand, SW species may exhibit larger home ranges due to their mating system; on the other hand, AF species have higher metabolic energy and thus food requirements and can be expected to have larger home ranges that have to yield this food. This study shows that percentage of the time moving (as an index of foraging mode) correlates positively with home range, even after correcting for body mass, and these patterns remain when phylogenetic relationships are taken into account. We thus conclude that home range areas parallel activity levels in lizards.     

Cuticular hydrocarbon dynamics in young adult Polistes dominulus (Hymenoptera: Vespidae) and the role of linear hydrocarbons in nestmate recognition systems

In social insects, cuticular hydrocarbons (CHCs) play an important role in nestmate discrimination processes, but young individuals are usually not discriminated. We studied CHC changes in young workers of the social wasp Polistes dominulus. A quantitative estimation demonstrated that total quantities of CHCs increased after emergence, with branched alkanes increasing drastically when compared with other classes of hydrocarbons. The relative quantity of longer-chain compounds increased with respect to shorter ones; unsaturated compounds decreased. These changes might reduce the capacity of the cuticle to acquire compounds of environmental origin. We then tested whether individuals acquire hydrocarbons from the environment, and whether this capability equally characterises newly emerged and mature wasps. We exposed wasps of two age classes (adults younger or older than 24 h) to four linear hydrocarbons in turn, and observed how nestmates reacted to their re-introduction into the natal colony. Exposed young wasps elicited significantly more aggressive responses than control sisters; but treated wasps older than 24 h were generally accepted by nestmates. Chemical assays showed that exposed young wasps readily absorbed hydrocarbons; older ones did not incorporate hydrocarbons, suggesting that the chemical profiles of mature wasps are less prone to chemical shifts than those of newly emerged wasps.     

Intraspecific nestmate recognition in two parabiotic ant species: acquired recognition cues and low inter-colony discrimination

Parabiotic ants—ants that share their nest with another ant species—need to tolerate not only conspecific nestmates, but also nestmates of a foreign species. The parabiotic ants Camponotus rufifemur and Crematogaster modiglianii display high interspecific tolerance, which exceeds their respective partner colony and extends to alien colonies of the partner species. The tolerance appears to be related to unusual cuticular substances in both species. Both species possess hydrocarbons of unusually high chain lengths. In addition, Cr. modiglianii carries high quantities of hereto unknown compounds on its cuticle. These unusual features of the cuticular profiles may affect nestmate recognition within both respective species as well. In the present study, we therefore examined inter-colony discrimination within the two parabiotic species in relation to chemical differentiation. Cr. modiglianii was highly aggressive against workers from alien conspecific colonies in experimental confrontations. In spite of high inter-colony variation in the unknown compounds, however, Cr. modiglianii failed to differentiate between intracolonial and allocolonial unknown compounds. Instead, the cuticular hydrocarbons functioned as recognition cues despite low variation across colonies. Moreover, inter-colony aggression within Cr. modiglianii was significantly influenced by the presence of two methylbranched alkenes acquired from its Ca. rufifemur partner. Ca. rufifemur occurs in two varieties (‘red’ and ‘black’) with almost no overlap in their cuticular hydrocarbons. Workers of this species showed low aggression against conspecifics from foreign colonies of the same variety, but attacked workers from the respective other variety. The low inter-colony discrimination within a variety may be related to low chemical differentiation between the colonies. Ca. rufifemur majors elicited significantly more inter-colony aggression than medium-sized workers. This may be explained by the density of recognition cues: majors carried significantly higher quantities of cuticular hydrocarbons per body surface.     

Thief workers and variation in nestmate recognition behavior in a ponerine ant, Ectatomma ruidum

Our findings give new insight into the relationship between nestmate recognition and cleptobiosis, intraspecific thievery of newly collected food items, in a neotropical ponerine ant, Ectatomma ruidum. The expression of discrimination of nestmates from non-nestmates varied among local aggregations of E. ruidum in a population at Barro Colorado Island, Panama. This result is due to differences in the behavior of the guard ants among aggregations, rather than differences in the expression of recognition cues by ants. Baiting experiments show that E. ruidum colonies in Panama have a greater tendency to have overlapping home ranges than a similar population in Costa Rica. The pattern of cleptobiosis, however, is strikingly similar between the Panamanian and Costa Rican populations.     

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.     

Can minimum convex polygon home ranges be used to draw biologically meaningful conclusions?

Many conclusions about mammalian ranging behaviour have been drawn based on minimum convex polygon (MCP) estimates of home range size, although several studies have revealed its unpredictable nature compared to that of the kernel density estimator. We investigated to what extent the choice of home range estimator affected the biological interpretation in comparative studies. We found no discrepancy when the question asked covered a wide range of taxa, as real and very large differences in range size were likely to have masked smaller differences due to the choice of home range estimator. However, when the question asked concerned within-species characteristics, the choice of home range estimator explained as much of the variation in range size as did the ecological variable in question. The implications for macro-ecological and intraspecific studies are discussed.     

Queen dimorphism and reproductive capacity in the ponerine ant, Ectatomma ruidum Roger

We report here the first case of queen dimorphism in a ponerine ant species. A total of 550 colonies of the Neotropical ponerine ant Ectatomma ruidum, from two natural populations in southeastern Mexico, were investigated for polymorphism and reproductive function within the queen caste. The distribution of different morphological traits (head and thorax widths, and scutum and alitrunk lengths) was shown to be bimodal. Thorax width and head width were significantly isometric for both macro- and microgynes and thorax proportions did not change with the category of the queen. Microgynes appear as an isometric reduction of the normal queens. On average, macrogynes were found to be approximately 20% larger in size and twice as heavy as microgynes (and up to three times heavier in terms of dry weight). Their wing surface was 72% greater. There was no difference between the two morphs in terms of their capacity to be inseminated and all dealate females present in natural colonies could be considered as true potentially reproductive queens whatever the class to which they belonged. Ovary size, number of ovarioles per ovary, and number of mature oocytes produced were significantly higher for macrogynes than for microgynes, but small queens were able to function as active egg-layers. However, the number of egg-laying individuals was significantly lower for this form, although egg-laying activity appeared to be independent of queen weight. Despite its markedly reduced reproductive capacity compared to macrogynes, the distribution of the microgyne form of E. ruidum in both studied populations was very broad, reaching one-third of all alate and dealate females and extending to one-third of all colonies. In 72.2% of the colonies where alate and/or dealate microgynes were present, both forms co-occurred. Fat content analysis results supported the idea that semi-claustral independent colony founding, typical for this species, would essentially be achieved by macrogynes, while microgynes would be adopted by established colonies. We argue that queen dimorphism in E. ruidum could represent an attractive alternative dispersal strategy for female sexuals. The small females may constitute an important reserve of potential reproductives at a very low energetic cost to the colony, the production of a macrogyne being about 7.5 times more costly than a microgyne. The significantly greater `wing surface/body weight' ratio of microgynes also suggests their greater capacity for dispersion. The combination of such a capacity for microgynes, along with their ability to contribute in the production of both female morphs, including the reproductively efficient macrogynes, would contribute to insuring genetic reassortment at the population level and could explain, in part, the ecological success of E. ruidum in Neotropical zones. Received: 4 January 1999 / Accepted: 12 May 1999     

A new ethological test to study nestmate recognition in adult ants

Summary An ethological test was designed to investigate nestmate recognition in adult ants. Contrary to classic dyadic tests, it consisted of a choice situation where a tested-worker was faced with nestmate and non-nestmate workers. These were kept alive, tied down, and thus immobilised in the neutral arena to record the tested-worker's reactions independently of the behaviour of the others. Such tests, applied to the ponerine ant,Ectatomma tuberculatum, enabled us to record various measures of discrimination and to limit aggressive behaviours between alien ants, which often leads to serious injury in dyadic tests. They also demonstrated the influence of age on discrimination behaviour in this species.     

Territorial behaviour in corn mice, <Emphasis Type="Italic">Calomys musculinus</Emphasis> (Muridae: Sigmodontinae), with regard to mating system

We studied territorial behaviour of Calomys musculinus, a promiscuous-polygynous species, in order to test the predictions that during the breeding period, home-range-owner females are more aggressive than home-range intruders and that males do not display territorial behaviour. At the core of the breeding season, we conducted 136 intrasexual and 66 intersexual encounters. The study was carried out in four 0.25-ha enclosures, each containing one independent population. We studied territorial behaviour utilising focal animal sampling to record behaviour. Encounters were conducted in a circular opaque arena (COA) near the home-range’s activity mathematical centre (AMC). Amicable behaviour was not observed in any of the 78 trials among females. The rates of aggressive and submissive behaviour per female–female encounters varied in relation to whether they were home-range owners or intruders, the former being aggressive, the latter submissive. So we could link these drives to the territorial behaviour of C. musculinus. This result fits well with the already known territoriality of C. musculinus during the breeding season. In contrast, the 58 intrasexual male encounters did not show direct aggressive interactions between them. The rates of the different behaviours between males did not vary in relation to whether they were home-range owners or intruders. In our study, which took place during the breeding period, C. musculinus males were nonterritorial, and they never exhibited aggressiveness towards females.     

Recognition of nestmate eggs in the ant Formicafusca is based on queen derived cues

Inclusive fitness benefits depend on recognizing the right individuals to interact with. Social insect nests protect themselves from non-kin intruders through nestmate recognition based on chemical cues. The recognition cues on adult individu- als are from a mixture of genetic and environmental sources, but the ontogeny and use of recognition cues on eggs has not been previously assessed. We studied recognition by workers of eggs that were either nestmates or non-nestmates, and the ontogeny of recognition cues on eggs in the ant Formica fusca, a species with precise egg recognition abilities. Workers were able to dis- criminate among freshly laid eggs with no nest derived cues on them, and the egg surface chemicals varied among nests in these eggs, suggesting that queen derived cues are used in nestmate recognition. The results are discussed in the light of their implica- tions on deceptive social parasite strategies and within colony conflicts     

Discrimination of nestmate workers and drones in honeybees

Summary Discrimination of nestmates from non-nestmates has mainly been investigated in female social insects. Little is known about discrimination of males. Here we show that under natural conditions at the nest entrance, honeybee workers can discriminate nestmate drones from non-nestmate drones as effectively as they can discriminate nestmate workers from non-nestmate workers.     

两种核域估算方法在野生藏狐家域研究中的比较

核域是野生动物重点利用、含有更多居所、隐蔽场所及依赖性食物资源的区域。关于核域的确定和划分一直存在争议,且很难找到客观的并能准确展示动物生物学信息的数理统计方法。2006 年4 月,2006 年9 ～10月和2007 年1 ～4 月,我们共记录青海省都兰县沟里乡3 只藏狐的199 个活动位点,通过固定核空间(Fixed kernel)和调和平均值(Harmonic mean)模型,结合独立区域法的计算原理,估算藏狐的核域,并对两种估算方法进行比较分析,以确定更理想的核域数学模型。研究结果表明:(1)两种估算方法中核域面积均受到家域总面积的影响,固定核空间法的核域面积受家域总面积影响较大,而且变量系数不稳定; (2) 随着样本量的增加,固定核空间法估算的核域面积逐渐减小,而调和平均值法则增大,前者从数理统计上更好地描述了藏狐对家域资源的利用分布,后者更符合客观实际;(3)调和平均值法在划定家域边界时容易包括不属于家域范围的区域,而在估算核域时可以在一定程度上克服该缺陷;(4)调和平均值法能够真实地反应动物的活动中心,估算的核域中包含有更多的活动位点。因此,尽管在估算藏狐核域时固定核空间法有稳定的计算结果等优点,调和平均值法为更理想的核域估算模型。     

Home ranges, movements, and activity of wolves (Canis lupus) in the Dalmatian part of Dinarids, Croatia

Home-range sizes, movements, and daily activity of wolves (Canis lupus L. 1758) were studied in Dalmatia, Croatia in 1998–2001. The total home ranges (100% MCP) of two packs were 160 km² and 141 km², mean=150.5 km². Core areas (50% kernel) were 26.2 km² and 3.3 km², respectively. Differences in core area sizes were influenced by human activity—hunting and sheep grazing. Compared with random locations, wolf locations were closer to the nearest water source (mean=937 m) and farther from houses (mean=653 m). Wolves were significantly more active during the night than during the day (activity indexes were 0.53 vs. 0.35), and night activity was higher during summer (0.58), and lower during winter (0.48). A correlation was found between distances traveled and activity index (r=0.58, p=0.003). Home range, seasonal variations in home-range size, habitat use, and activity of wolves in Dalmatia were oriented to make the compromise from danger of proximity to humans and also to benefit from human-related food sources.     

The role of cuticular hydrocarbons as chemical cues for nestmate recognition in the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Argentine ants (Linepithema humile) in their native South American range, like most other ant species, form spatially restricted colonies that display high levels of aggression toward other such colonies. In their introduced range, Argentine ants are unicolonial and form massive supercolonies composed of numerous nests among which territorial boundaries are absent. Here we examine the role of cuticular hydrocarbons (CHCs) in nestmate recognition of this highly damaging invasive ant using three supercolonies from its introduced range. We conducted behavioral assays to test the response of Argentine ants to workers treated with colonymate or non-colonymate CHCs. Additionally, we quantified the amount of hydrocarbons transferred to individual ants and performed gas chromatography-mass spectrometry (GC/MS) to qualitatively characterize our manipulation of CHC profiles. The GC/MS data revealed marked differences in the hydrocarbon profiles across supercolonies and indicated that our treatment effectively masked the original chemical profile of the treated ants with the profile belonging to the foreign individuals. We found that individual workers treated with foreign CHCs were aggressively rejected by their colonymates and this behavior appears to be concentration-dependent: larger quantities of foreign CHCs triggered higher levels of aggression. Moreover, this response was not simply due to an increase in the amount of CHCs applied to the cuticle since treatment with high concentrations of nestmate CHCs did not trigger aggression.The results of this study bolster the findings of previous studies on social insects that have implicated CHCs as nestmate recognition cues and provide insight into the mechanisms of nestmate recognition in the invasive Argentine ant. Received 6 February 2007; revised 31 May and 27 July 2007; accepted 16 August 2007.     

小兴安岭南坡野猪家域分析

采用笼式活捕野猪,利用无线电遥测技术、R2V、Arcview 和SPSS 软件技术,于2004 年7 月15 日至2006年1 月19 日,对小兴安岭南坡野猪家域进行测定、计算和分析,以了解该地区野猪的家域变化规律。研究结果表明:季节变化影响野猪家域面积,在春、秋季,野猪的家域面积显著大于冬季,而春、夏季间无显著差异;不同性别及年龄野猪家域大小也不同,成体雄性秋季发情期和亚成体春季分窝期家域面积显著增加;在冬、春季家族群野猪的家域面积显著大于独体野猪家域面积,说明家族群对家域面积有影响;亚成体家域大小主要受家族群家域的影响,家族群野猪面积大,相应地家族野猪中亚成体家域面积也大。