Low relatedness among cooperatively breeding workers of the greenhead ant Rhytidoponera metallica

The greenhead ant Rhytidoponera metallica has long been recognized as posing a potential challenge to kin selection theory, because it has large queenless colonies where apparently many of the morphological workers are mated and reproducing. However, this species has never been studied genetically and important elements of its breeding system and kin structure remain uncertain. We used microsatellite markers to measure the relatedness among nestmates, unravel the fine‐scale population genetic structure, and infer the breeding system of R. metallica. The genetic relatedness among worker nestmates is very low but significantly greater than zero (r=0.082 ± 0.015), which demonstrates that nests contain many distantly related breeders. The inbreeding coefficient is very close to and not significantly different from zero, indicating random mating and lack of microgeographic genetic differentiation. On average, closely located nests are not more similar genetically than distant nests, which is surprising, as new colonies form by budding and female dispersal is restricted. Lack of inbreeding and absence of population viscosity indicates high gene flow mediated by males. Overall, the genetic pattern detected in R. metallica suggests that a high number of moderately related workers mate with unrelated males from distant nests. This breeding system results in the lowest relatedness among nestmates reported for social insect species where breeders and helpers are not morphologically differentiated.     

Nest site selection and longevity in the ponerine ant Rhytidoponera metallica (Hymenoptera, Formicidae)

Summary: The size of the nesting cavity and the nature of the nest-building material can constrain colony growth in eusocial insects. Rocks protect colony members against extreme temperatures as well as serve as a supplementary source of heat, thereby maintaining optimal conditions for brood development. The frequency distribution of available rocks and those occupied by colonies of Rhytidoponera metallica in the Avon River State Forest, Victoria, Australia, showed that colonies of R. metallica avoided nesting under small rocks (<200 cm²). A strong correlation between colony size and rock size indicates that nesting under a larger rock promotes colony growth. Laboratory choice experiments revealed that workers are capable of recognizing larger rocks from external physical characters (dimension). Field data indicates that colonies leave nests frequently, with nest abandonment peaking in the summer months.     

Regulated food recruitment through individual behavior of scouts in the ant,Myrmica sabuleti (Hymenoptera: Formicidae)

By presenting different kinds of food sources to colonies ofM. sabuleti, we have demonstrated that this species regulates its foraging activity by altering the proportion of scouts that return to the nest to recruit nestmates after discovering a food source and by varying the number of nestmates recruited by a scout. These two parameters are related to the kind of food discovered. Our behavioral experiments showed that the probability that a scout returned to the nest decreased with a decreasing quantity of sucrose solution. In contrast, the number of returned scouts that elicited recruitment from the nest and the mean number of nestmates recruited by one of these scouts were independent of the quantity of the sucrose solution. Recruitment even occurred toward a 1- or 0.25-µl droplet of sucrose solution. When a scout discovered a large dead prey, a large drop of prey juice, a cluster of 30 dead fruit flies, or 1 isolated fruit fly, it always went back to the nest, but it elicited recruitment only when the food source was a large dead prey or a large drop of prey juice. No recruitment occurred when the food source was a single fruit fly and recruitment occurred only once in 30 trials when a cluster of 30 fruit flies was discovered.     

Individual constancy of local search strategies in the giant tropical ant,Paraponera clavata (Hymenoptera: Formicidae)

Paraponera clavata workers engage in a period of local search after encountering a small amount of artificial nectar. Giving-up times from local search are not distributed normally; there is a strong skew to longer times. There is no statistically significant relationship between the amount of time required to collect the food and the subsequent search time. Giving-up time in response to the first reward presented to an ant is positively correlated with that ant's response to a second such reward. However, giving-up times diminish when an ant is presented with a series of rewards. Local search is a function of individual strategies, which remain relatively constant in the short term.     

Temporal polyethism, life expectancy, and entropy of workers of the ant Ectatomma vizottoi Almeida, 1987 (Formicidae: Ectatomminae)

We investigated the changes in the behavioral repertoire over the course of life and determined the life expectancy and entropy of workers of the ant Ectatomma vizottoi. Newly emerged ants were individually marked with model airplane paint for observation of behaviors and determination of the age and life expectancy. Ants were divided into two groups: young and old workers. The 36 behaviors observed were divided into eight categories. Workers exhibit a clear division of tasks throughout their lives, with young workers performing more tasks inside the colony and old workers, outside, unlike species that have small colonies. This species also exhibits an intermediate life expectancy compared to workers of other species that are also intermediary in size. This supports the hypothesis of a relationship between size and maximum life expectancy, but it also suggests that other factors may also be acting in concert. Entropy value shows a high mortality rate during the first life intervals.     

Thelytokous parthenogenesis in the exotic dacetine ant Strumigenys rogeri (Hymenoptera: Formicidae) in Taiwan

Only recently has it become clear that several species of eusocial hymenopterans regularly reproduce by thelytokous parthenogenesis, that is, the production of diploid female offspring by unmated females. This phenomenon suggests that parthenogenetic reproduction might be advantageous to organisms under certain environmental conditions. Here the occurrence of asexual reproduction is reported for the first time in the dacetine ant, Strumigenys rogeri, at least for the focal populations in Taiwan. Virgin queens of S. rogeri maintained with several workers produced both workers and young queens from unfertilized eggs under laboratory conditions in as short as 39 days, whereas workers were strictly sterile as no spermatheca was discovered after dissection. Combined with additional evidence (i.e. absence of males in field colonies), queen thelytoky is confirmed. Such a reproductive mode and short development time may jointly help explain the success of this tramp ant species in Taiwan and elsewhere.     

华南地区典型生境中红火蚁觅食行为及工蚁召集规律

观察研究了华南地区几种典型生境中红火蚁对不同类型食物的觅食行为和工蚁召集动态规律。研究结果表明红火蚁觅食行为存在搜寻、召集及搬运等主要过程。不同生境中红火蚁对食物的搜寻时间存在一定差异,荔枝园中搜寻时间明显长于其它生境;荔枝园、荒地、路边生境中红火蚁对蜂蜜的搜寻时间明显长于其它几种食物;同一生境中红火蚁对不同重量的同种食物搜寻时间无明显变化。发现食物后召集的工蚁数量随着时间延长呈现不断增加的趋势,对于较大的食物一般30min左右召集的工蚁数量达到最大,之后趋于稳定并逐渐减少,而对于可以直接搬动的食物发现后15min左右群体召集数量达到最大并很快将其搬走。召集工蚁数量与发现后时间的关系符合二次非线性方程,建立了火腿肠、花生油和蜂蜜等食物上红火蚁工蚁召集数量与发现时间的关系模型,分别为Na=4183.91e^-0.0327T-4231.48e^-0.0346T,Na=3253.78e^-0.0233T-3314.59e^-0.0271T,Na=117.97e^-0.0131T-163.93e^-0.0808T。对于不同食物红火蚁发现后召集的最大工蚁数量间有明显差异,其中花生油上最大,平均为176．3头,火腿肠上次之,为90．4头,蜂蜜上最少,为68．0头。对于不同重量的同种食物,工蚁的召集动态规律较为一致,发现食物后25-30min左右工蚁的召集数量均达到最大值,之后缓慢减少,发现食物后时间长度和食物上工蚁数量也符合房室模型函数Na=c1×exp（-c2T）-c3×exp（-c4T）。同种食物不同重量之间同一时间召集的工蚁数量存在较大差异,呈随食物重量增大而增大趋势。不同生境中对相同质量同种食物工蚁召集数量动态总体变化规律相近,但在发现食物后工蚁数量增长的速度和最大召集工蚁的数量存在明显差异,以路边工蚁数量增长最快、工蚁总数最多,苗圃次之,荔枝园、荒地中工蚁增长速度均较低。红火蚁对不同食物种类切割、搬运耗时存在很大差异,搬运完0．5g的火腿肠需要26．23h,而黄粉虫幼虫（重量约为0．1g）仅为15．6min。不同生境中红火蚁对相同重量同一种食物的搬运耗时也不同,荔枝园要长于其它生境。     

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.     

Dynamic matching of forager size to resources in the continuously polymorphic leaf-cutter ant, Atta colombica (Hymenoptera, Formicidae)

Abstract.  1. Ergonomic optimisation theory proposes that by increasing variation in worker morphology, social insect colonies may increase their dietary breadth; however, little is known about how this relationship operates at the colony level. This study examines the colony-level pattern of forager size allocation to resource sites in a natural setting.
2. Using a biologically relevant measure of toughness, it is shown that leaf-cutter ant colonies exploit a variety of plant resources that vary significantly in toughness at any given time.
3. Forager size is shown to be matched to the toughness of plant material, with larger ants harvesting tougher material.
4. Furthermore, outbound foragers travelling to a harvest site are matched in size to the toughness of plant material contained within the site and are not a random selection of available foragers. The match between forager size and plant toughness may reduce the number of wasted trips and ill-matched foragers.
5. The observed colony-level pattern of forager allocation could be the result of learning by individual foragers, or the result of information shared at the colony level.     

Temporal variation of vitellogenin synthesis in Ectatomma tuberculatum (Formicidae: Ectatomminae) workers

Workers of the ant species Ectatomma tuberculatum (Ectatomminae) have active ovaries and lay eggs that are eaten by the queen and larvae (trophic eggs). Vitellogenins are the main proteins found in the eggs of insects and are a source of nutrients. The aim of this study was to characterize the period of vitellogenin production in workers of E. tuberculatum. The vitellogenin was identified from queen and worker eggs by SDS-PAGE. Anti-vitellogenin antibodies were obtained and used to detect this protein in the fat body and haemolymph of workers at different ages. Vitellogenin from E. tuberculatum consists of two polypeptides of 31 and 156 kDa. In the eggs of queens, the 156 kDa polypeptide is cleaved into two subunits of 36 and 123 kDa. The analysis of the haemolymph of workers showed that the secretion of vitellogenin varies with age. The secretion is initiated around the fifth day after emergence, with peak production from days 20 to 60, and stops around day 100. The variation in production is related to the different activities performed by the workers within the colony, suggesting that vitellogenin may have an important role in maintaining age polyethism.     

黑翅土白蚁巢群觅食群体大小与工蚁体长呈正相关

【目的】明确黑翅土白蚁Odontotermes formosanus巢群的觅食群体大小与工蚁体长之间的关系,为进一步研究其生态学特点以及更好地开展防治提供依据。【方法】在杭州植物园内选择5处有黑翅土白蚁活动区域作试验点,诱捕并以中性红(neutral red)标记捕获的白蚁工蚁,通过“捕获-标记-释放-重捕”试验测定黑翅土白蚁的觅食群体大小。【结果】5个试验点中有4个成功实施了“捕获-标记-释放-重捕”试验,4个试验点黑翅土白蚁巢群的觅食群体个体数量分别为443 133±45 469, 495 360±67 429, 674 345±101 703和1 224 662±93 112头。黑翅土白蚁觅食群体大小(Y)与觅食工蚁的体长(X)呈正相关,拟合的指数函数方程式为：Y=1.8389e^0.7185X(R²=0.7834)。【结论】本研究明确了黑翅土白蚁的觅食群体大小以及工蚁体长之间的函数关系,为开展区域防治提供了依据。     

Worker laying in leafcutter ant Acromyrmex subterraneus brunneus （Formicidae, Attini）

We studied the process of offspring production in queenless colonies of Acromyrmex subterraneus brunneus, and particularly evaluated the ovary development of workers as a function of their age. For this, subcolonies were set up and evaluated at different periods of isolation from the queen （2, 4 and 6 months）, besides individually labeled age groups. The subcolonies were assessed according to offspring production and ovaries containing oocytes or not. The evaluations showed worker oviposition and development of males originating from worker-laid eggs. At 2 months＇ absence of the queen, eggs and larvae were found, with eggs in a higher proportion than larvae. After 4 months, the proportion of eggs had reduced while larvae had increased, and a pupa was found in one subcolony. At 6 months, besides a higher share of larvae, one pupa and one adult male were found. Dissection of workers revealed ovaries containing oocytes during the periods of evaluation. Only a group of medium-sized and large workers, 23.3%, 20.9% and 37.5% of the population from each period assessed in queenless subcolonies respectively, presented developed oocytes in the ovary. The same was observed in colonies with a queen, with 17.6%, 19.6% and 7.8% of the group of dissected workers from each time period, respectively. With respect to worker age, we observed by dissection of the ovary, that the greatest percentage of individuals with ovarioles containing oocytes occurred at 45 days （6 weeks） up to 90 days （12 weeks）. These results probably are associated with the workers reproduction and the laying of trophic and reproductive eggs in colonies with and without a queen; these eggs have distinct functions in each situation.     

Raiding and Foraging Behavior of the Blood-Red Ant,Formica sanguinea Latr. (Hymenoptera,Formicidae)

In this paper we report the results of a detailed study on the behavioral ecology of slave raiding and foraging activity in the European blood-red ant, Formica sanguinea Latr. The field study was conducted over an unbroken period of 78 days, during which the activity of two dulotic colonies of this facultative slave-maker was observed for 10 h each day. It was possible to observe 26 raids distributed over 23 days, among which 18 were followed by the sacking of nests belonging to the species F. cunicularia, F. fusca, and Lasius emarginatus, whereas 8 failed. Simple, continuous, and simultaneous raids occurred. We recorded the timing, frequency, distance, and direction of slave raids, including the number of participants and the type of booty. Particular attention was devoted to the scouting behavior and raiding organization. Moreover, every day, we observed foraging and predatory behavior, during which adult insects (mainly ants), seeds, and berries were retrieved to the dulotic colonies. On the basis of our observations F. sanguinea seems to be a very efficient slave-maker and predatory species of the Raptiformica subgenus. Moreover, its dulotic behavior may be regarded as a continuation and an expansion of its foraging and predatory behavior, as predicted by Darwin's hypothesis for the origin and evolution of slavery in ants.     

Trail Communication During Foraging and Recruitment in the Subterranean Termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)

The search for food in the French subterranean termite Reticulitermes santonensis De Feytaud is organized in part by chemical trails laid with the secretion of their abdominal sternal gland. Trail-laying and -following behavior of R. santonensis was investigated in bioassays. During foraging for food termites walk slowly (on average, 2.3 mm/s) and lay a dotted trail by dabbing the abdomen at intervals on the ground. When food is discovered they return at a quick pace (on average, 8.9 mm/s) to the nest, laying a trail for recruiting nestmates to the food source. While laying this recruitment trail the workers drag the abdomen continuously on the ground. The recruitment trail is highly attractive: it is followed within a few seconds, by more nestmates, and at a quicker pace (on average, 6.4 mm/s) than foraging trails (on average, 2.9 mm/s). The difference between foraging and recruitment trails in R. santonensis could be attributed to different quantities of trail pheromone. A caste-specific difference in trail pheromone thresholds, with workers of R. santonensis being more sensitive to trails than soldiers, was also documented: soldiers respond only to trails with a high concentration of trail pheromone.     

Macronutrient Regulation in the Tropical Terrestrial Ant Ectatomma ruidum (Formicidae): A Field Study in Costa Rica

Using the geometric framework for the first time in a field study of ant nutritional biology, we show that Ectatomma ruidum colonies actively regulate protein–carbohydrate intake, self-selecting a carbohydrate-biased diet. These data, combined with quantitative recruitment data, provide a more comprehensive understanding of colony-level nutritional requirements of ants. Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.     

Biological constraints and colony founding in the polygynous invasive ant Lasius neglectus (Hymenoptera, Formicidae)

Summary. The polygynous invasive ant Lasius neglectus was described from Budapest, Hungary, as an unicolonial species, with no apparent colony barriers, and inferred intra-nidal mating without a nuptial flight. Here we analyze additional morphological characteristics of gynes, their physiological condition at emergence and at the time of mating and we describe the productivity of different types of colony founding in the laboratory. A low increase in dry weight and in fat content from emergence to mating indicates that gynes can only succeed in dependent colony founding. However, the queen-worker thorax volume ratio is typical of a species with independent colony founding and we were able to demonstrate independent colony founding in the laboratory (both haplo- and pleometrotic). Brood development in independent founding is rapid and the number of nanitics higher than in other Lasius species. Both colony budding and dependent colony founding could also be demonstrated in the laboratory. Worker oviposition was absent. The carbohydrate content of newly mated queens is consistent with the observed loss of mating flight of this species. However the relative wing area clearly indicates that L. neglectus queens should be able to fly. Therefore, both queen morphology and physiological state at maturity show that L. neglectus is intermediate between a monogynous, free-living, non pest ant and a multiqueened (polygynic) invasive tramp ant. This neglected ant thus offers interesting opportunities to study the origin of unicoloniality and the spread of this species in northeast Spain.     

Resource Assessment, Recruitment Behavior, and Organization of Cooperative Prey Retrieval in the Ant Formica schaufussi (Hymenoptera: Formicidae)

Foragers of the ant Formica schaufussi recruit nestmates to large anthropod prey and cooperatively transport the prey to the nest. The size of the group of ants retrieving prey is significantly correlated with the prey mass at the point at which the retrieval group reaches the nest entrance. To understand the mechanism involved in this size matching process, the regulation of retrieval group size was investigated by examining the modulatory role of trail pheromones in recruitment communication and the behavioral processes that might adjust retrieval group size to prey mass. Laboratory studies of hindgut, poison, and Dufour's gland extracts showed that the contents of the hindgut, which was determined to be the source of trail pheromone, induced recruitment and orientation behavior in ants and regulated the recruitment response of ants in the absence of any other communication signal. However, chemical mass communication alone did not appear to explain the regulation of retrieval group size. Scout ants assess whether to collect prey individually or recruit nestmates to group-retrieve 100-, 200-, or 400-mg prey but did not vary group size in relation to either the prey mass or the presence of interspecific competitors once the decision to initiate group retrieval was made. The number of recruits leaving the nest was independent of these factors and first matched prey mass during prey transport, possibly through a process of differential individual response to immobile versus mobile prey items. Unpredictable factors such as prey resistance to movement and rapidly changing degrees of interspecific competition may preclude scouts from fine-tuning the retrieval group size before it reaches the prey.