Kin structure and the swarming behavior of the honey bee Apis mellifera

Summary Experimental hives obtained from cordovan queens that were instrumentally inseminated with semen from one cordovan and one Italian drone were set up and allowed to swarm. Cordovan provides a resessive genetic marker system (cuticle color) so that the workers from the cordovan and Italian male lines are distinguishable. Our results show that these patrilineal worker groups segregate non-randomly during colony fission and this segregation cannot be explained by observed age structure. Evidence of innate kin recognition in bees has been previously established. We argue that kin recognition could be responsible for the observed non-random grouping of kin during swarming.     

Chronic exposure effects of copper on growth,metamorphosis and thyroid gland,liver health in Chinese toad,Bufo gargarizans tadpoles

The impact of copper exposure on Chinese toad (Bufo gargarizans) tadpoles was investigated in this study. First, the 96h LC50 value of copper was 8.697 μM, by means of a 4 d acute toxicity test. Second, we studied the chronic effects of copper on B. gargarizans tadpoles at control, 0.025, 0.1 and 1.0 μM concentration. Survival, body length, body weight, developmental stage, incidence of metamorphic climax, and size at metamorphic climax were determined. In tadpoles developed to metamorphic climax (stage G42), liver and thyroid gland were assessed histologically. Copper at 0.1 and 1.0 μM could inhibit tadpole growth and prolong tadpole metamorphic progress relative to controls. Tadpole size (total length and weight) at stage G42 is also affected in the 0.1 and 1.0 μM treatments. In addition, histological examinations have revealed that 1.0 μM copper could cause significant pathological changes and hepatocytes degeneration in liver. Furthermore, histomorphological measurements indicated that copper at 0.1 and 1.0 μM reduced thyroid gland size, diameter and number of follicle. In conclusion, our study suggests that Cu could damage the liver and thyroid gland, so growth and metamorphosis of B. gargarizans tadpoles were inhibited resulted of disrupting liver metabolism and THs homeostasis.     

Kin recognition in the common lizard

The ability to recognize parents has never been reported in species in which parents do not provide care to their young; in such species, only sibling recognition has been found. However, there may be several advantages of parent recognition, even in the absence of parental care. We investigated the ability of neonates to recognize olfactory cues from both their mother and siblings in the common lizard, Lacerta vivipara, a species without parental care. Juveniles from 264 gravid females were reared for 2 days either with their mother, with another female, or separated from all other adults. Juveniles from some families were split into two or three groups so that each juvenile was unfamiliar with a subset of its siblings. After 2 days, we offered the juveniles a choice of two nocturnal shelters: one containing a lizard odor and the other without odor. The response to the odor of an unrelated and unfamiliar adult was influenced by both the sex of the adult and the sex of the juvenile. Juveniles of both sexes recognized the odor of their mother whether they were familiar with her or not (pre-natal determinism). Juveniles recognized familiar but not unfamiliar siblings (post-natal determinism). In the wild, spatial association with kin declines shortly after birth. Thus, recognition of the mother is likely to have biological relevance. Recognition of the mother may reduce competition and/or enhance juvenile establishment. Received: 15 May 1997 / Accepted after revision: 29 December 1997     

Kin conflict over caste determination in social Hymenoptera

We argue that caste determination, the process whereby females in the social Hymenoptera develop into either queens or workers, is subject to kin-selected conflict. Potential conflict arises because developing females are more closely related to their would-be offspring than to those of other females. Therefore, they may favour becoming queens contrary to the interests of other developing females and of existing queens and workers. We suggest two contexts leading to potential caste conflict. The first occurs when queens are reared in a reproductive phase following an ergonomic phase of worker production, while the second occurs when queens and workers are reared simultaneously. The first context assumes that workers' per capita contribution to colony survival and productivity falls with rising colony size. A critical feature influencing whether potential conflict is realized is the extent to which developing females can determine their own caste (“self-determination”). Self-determination is facilitated when female larvae control their own food intake and when queen-worker size dimorphism is low. We know of no strong evidence for actual conflict over caste fate arising in the first context. However, stingless bees and polygynous ants with excess queen-potential larvae that are either forced to develop as workers or are culled as adults demonstrate actual caste conflict in the second context. Caste conflict does not preclude caste regulation for “the good of the colony”, but such regulation is contingent on either the absence of potential conflict or on developing females losing control of their caste fate. Received: 22 March 1999 / Received in revised form: 15 June 1999 / Accepted: 19 June 1999     

Benefit of suites of defensive behavior induced by predator chemical cues on anuran tadpoles, Hyla japonica

When predator chemical cues are present, low activity of prey is a commonly seen defensive behavior. However, few studies have explored the functional implications of the defensive behaviors and, thus, elucidated the possible linkages between behavioral responses and its consequences. In this study, we experimentally investigated how behavioral responses of Hyla japonica tadpoles to predator chemical cues affect vulnerability to a dragonfly nymph Anax parthenope julius. The frequency of tadpoles attacked by dragonfly nymphs was lower with chemical cues of predator was present than without chemical cues, and most of attacks occurred when tadpoles were mobile. When tadpoles were exposed to chemical cues, on the other hand, their swimming speed was quicker and swimming distance was longer, respectively, and the rates of being approached of the swimming tadpoles by dragonfly nymph was lower than those not exposed to chemical cues. We found that the tadpoles are induced by predator chemical cues not only to generally lower activity but also to swim in bursts as additional behavior and that the suite of their behavioral responses reduce the vulnerability against dragonfly nymph. Tadpoles can receive information about the predation risks by chemical cues and adjust their defensive behavior accordingly.     

Vortex formation and foraging in polyphenic spadefoot toad tadpoles

Animal aggregations are widespread in nature and can exhibit complex emergent properties not found at an individual level. We investigate one such example here, collective vortex formation by congeneric spadefoot toad tadpoles: Spea bombifrons and Spea multiplicata. Tadpoles of these species develop into either an omnivorous or a carnivorous (cannibalistic) morph depending on diet. Previous studies show that S. multiplicata are more likely to develop into omnivores and feed on suspended organic matter in the water body. The omnivorous morph is frequently social, forming aggregates that move and forage together and form vortices in which they adopt a distinctive slowly rotating circular formation. This behaviour has been speculated to act as a means to agitate the substratum in ponds and thus could be a collective foraging strategy. Here we perform a quantitative investigation of the behaviour of tadpoles within aggregates. We found that only S. multiplicata groups exhibited vortex formation, suggesting that social interactions differ between species. The probability of collectively forming a vortex, in response to introduced food particles, increased for higher tadpole densities and when tadpoles were hungry. Individuals inside a vortex moved faster and exhibited higher (by approximately 27%) tailbeat frequencies than those outside the vortex, thus incurring a personal energetic cost. The resulting environmental modification, however, suggests that vortex behaviour may be an adaptation to actively create and exploit a resource patch within the environment.     

Herbicide exposure affects the chemical recognition of a non native predator in common toad tadpoles (<Emphasis Type="Italic">Bufo bufo</Emphasis>)

Summary. In amphibians and fishes, evidence is increasing that chemical cues from injured conspecifics can play a role in the chemical labelling and learned recognition of unfamiliar predators. In this laboratory study, we tested the prediction that prior chemical exposure to a non-native predator feeding on conspecific tadpoles will subsequently allow tadpoles of the common toad (Bufo bufo) to recognize the chemical cues specifically released by this starved predator. Furthermore, we investigated the vulnerability of this chemically-mediated process to herbicide contamination. With these aims in view, groups of tadpoles were kept either unexposed or exposed for ten days to chemical cues from Turkish crayfish (Astacus leptodactylus) previously fed on tadpoles, both in uncontaminated water and in the presence of four sublethal concentrations of amitrole (0.01, 0.1, 1 and 10 mg.l⁻¹). We then assessed the effects of the six conditioning treatments on general activity and behavioural response to chemical cues from starved crayfish. Larval treatments did not affect the general activity of the tadpoles. By contrast, the treatments had significant effects on the behavioural response to the test solution prepared form starved crayfish. The only tadpoles to show an antipredator behavioural response to the chemical stimulation from starved crayfish belonged to the groups derived from chemical exposure to tadpole-fed crayfish in uncontaminated water and in contaminated water with the lowest concentration of amitrole (0.01 mg.l⁻¹). Conversely, this chemical stimulation produced no behavioural change in the control group or in the groups derived from exposure to tadpole-fed crayfish in contaminated water containing 0.1, 1 and 10 mg.l⁻¹ of amitrole. This study demonstrates that chemical cues released during the predator’s feeding activity can subsequently be used by common toad tadpoles in the recognition of an unfamiliar predator. In addition, our results show that the presence of sublethal amitrole concentrations can impair this recognition process. Such a pesticide effect might be especially detrimental for amphibian populations threatened by invasive predators.     

Kin and non-kin interventions in the aggressive disputes of vervet monkeys

Summary Interventions in aggressive disputes were investigated in a free-living troop of vervets (Cercopithecus aethiops sabaeus) in Barbados. Interventions on behalf of kin were more frequent than on behalf of non-kin. Both types of interventions were more likely when the intervening animal outranked the opponent; presumably because retaliation probability, and hence cost of intervening, is low against low ranking opponents. The number of interventions given on behalf of both kin and non-kin increased with the number of disputes in which they were involved. In contrast to kin interventions, the number of interventions given on behalf of non-kin was correlated with that received by non-kin, suggesting that reciprocation is a necessary component of non-kin interventions. Non-kin interventions were more likely when the recipient outranked the opponent, presumably because reciprocation probability is high. Pairs of non-kin form structured reciprocal relationships based on the proportion of interventions allocated to each other, and most non-kin interventions flowed through these relationships. Males intervened on behalf of non-kin more frequently than did females. The implications of the results for the evolution of kin and reciprocal altruism were discussed.     

Trichinella spiralis infection in voles alters female odor preference but not partner preference

Females may choose mates based on secondary sex traits that reflect disease resistance. Accordingly, females should be able to distinguish between unparasitized and parasitized males, and should prefer to mate with unparasitized individuals. Mate and odor preferences for uninfected males or males infected with the nematode, Trichinella spiralis, were examined among prairie voles (Microtus ochrogaster) and meadow voles (M. pennsylvanicus). In a 15-min odor preference test, only female meadow voles distinguished between bedding from parasitized and unparasitized conspecific males, and preferred to spend time with bedding from unparasitized males. Although T. spiralis infection influenced odor preference in female meadow voles, there was no effect of infection status on mate preference among either species. Testosterone and corticosterone concentrations were not different between parasitized and unparasitized males. However, among prairie voles, males that spent an increased amount of time with females during the mate preference test had elevated testosterone concentrations. Taken together, these data suggest that (1) female meadow voles can discriminate between unparasitized and parasitized males, (2) the effects of infection on steroid hormone concentrations may be masked by the effects of social interactions, and (3) parasites may represent a selective constraint on partner preference in voles; however, the life cycle of parasites may influence female preference and should be considered in studies of female preference. Received: 23 April 1998 / Accepted after revision: 25 October 1998     

Morphological and behavioral alterations induced by endocrine disrupters in amphibian tadpoles

Developmental toxicity effects of endocrine disrupter chemicals, acephate and cypermethrin were studied in Bufo melanostictus tadpoles. Thirty developing eggs of B. melanostictus were exposed to each concentration (0.01, 0.05, 0.1, 0.5, and 1?µg?L^?1) of acephate or cypermethrin in the laboratory (temperature: 23?±?1°C; photoperiod: 11.5–12.5?h). Eggs maintained in conditioned water alone served as controls. After hatching, larvae were fed on boiled spinach until the completion of metamorphosis. In control group, larvae that hatched on 3rd day were heavily pigmented, voracious feeders, and active swimmers; in these tadpoles, hind limb and forelimb-buds emerged on 16th and 24th day and metamorphosis was complete on 32nd day. Eggs exposed to acephate also hatched on 3rd day but larvae exhibited deformities such as, (i) tail distortions, (ii) laterally crooked trunk, (iii) decreased pigmentation, (iv) inactivity, (v) peeling of the skin, and (vi) delay in emergence of limbs and completion of metamorphosis. Cypermethrin-exposed eggs exhibited a delay (4–8 days) in hatching, there was no mortality, deformities in tail, trunk and head region, delay in the emergence of limbs, and completion of metamorphosis were evident. The demonstrated data indicate that these pesticides interfere with amphibian development when present in the aquatic system.     

Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997     

Kin recognition in rainbowfish (Melanotaenia eachamensis): sex, sibs and shoaling

Living with relatives can be beneficial to individuals via the evolution of kin-directed altruism, but this is tempered by the increased risk of inbreeding. Therefore, in social species, the ability to recognise relatives can be highly advantageous. This study focuses on kin discrimination in the Lake Eacham rainbowfish, Melanotaenia eachamensis, an endangered freshwater species from north-east Queensland, Australia. First, I examined kin recognition abilities when a combination of both chemical and visual recognition cues was available. When given a choice of shoaling with same-sex groups, females spent significantly longer with full-sibs rather than half-sibs, full-sibs rather than non-relatives and half-sibs rather than non-relatives. Males spent significantly longer shoaling with full-brothers versus half-brothers, but showed no other shoalmate preferences. Second, in the presence of only chemical cues, females did not discriminate among groups of different levels of relatedness, but males showed a non-significant tendency to associate with full-sibs rather than non-relatives. Male shoaling behaviour seemed to be more influenced by factors other than relatedness, e.g. intra-sexual competition. Finally, I found that the shoaling preferences of females changed when exposed to groups of males. Females preferred to associate with non-relatives rather than half-brothers and non-relatives rather than full-brothers. There was no significant difference in the time spent with half-brothers versus full-brothers. Taken together, my results suggest that females have very good kin recognition abilities. They prefer to shoal with female relatives but avoid male relatives, and so are able to balance the benefits of nepotism and the costs of incest. Received: 2 May 2000 / Revised: 26 July 2000 / Accepted: 30 July 2000     

Predator and prey space use: dragonflies and tadpoles in an interactive game

Predator and prey spatial distributions have important population and community level consequences. However, little is known either theoretically or empirically about behavioral mechanisms that underlie the spatial patterns that emerge when predators and prey freely interact. We examined the joint space use and behavioral rules governing movement of freely interacting groups of odonate (dragonfly) predators and two size classes of anuran (tadpole) prey in arenas containing two patches with different levels of the prey's resource. Predator and prey movement and space use was quantified both when they were apart and together. When apart from predators, large tadpoles strongly preferred the high resource patch. When apart from prey, dragonflies weakly preferred the high resource patch. When together, large prey shifted to a uniform distribution, while predators strongly preferred the high resource patch. These patterns qualitatively fit the predictions of several three trophic level, ideal free distribution models. In contrast, the space use of small prey and predators did not deviate from uniform. Three measures of joint space use (spatial correlations, overlap, and co-occurrence) concurred in suggesting that prey avoidance of predators was more important than predator attraction to prey in determining overall spatial patterns. To gain additional insight into behavioral mechanisms, we used a model selection approach to identify behavioral movement rules that can potentially explain the observed, emergent patterns of space use. Prey were more likely to leave patches with more predators and more conspecific competitors; resources had relatively weak effects on prey movements. In contrast, predators were more likely to leave patches with low resources (that they do not consume) and more competing predators; prey had relatively little effect on predator movements. These results highlight the importance of investigating freely interacting predators and prey, the potential for simple game theory models to predict joint spatial distributions, and the utility of using model choice methods to identify potential key factors that govern movement.     

Heavy metals and pesticides in anuran spawn and tadpoles,water, and sediment

High concentrations of metals (cadmium, copper, lead, and zinc) and of lindane were measured in the sediment and in the spawn and tadpoles of Bufo bufo, Rana dalmatina, and Rana ridibunda from two Austrian locations. Residues in spawn suggest maternal transfer. Increase of the metal concentrations from spawn to advanced tadpole stage is explained by the larval microphaguous feeding habits which cause high exposition of the tadpoles to substances concentrating in sediments and suspended particles. Metal and pesticide concentrations in spawn and tadpoles are toxic to various other aquatic organisms and are furthermore considered to be potentially hazardous to the anurans themselves. The results are conform with world‐wide observations of contamination of anuran larvae with heavy metals, and prove the significance of non‐point source chemical exposure due to allochthonous toxicant input by wind load and precipitation.     

Kin associations and direct vs indirect fitness benefits in colonial cooperatively breeding sociable weavers Philetairus socius

Indirect fitness benefits are believed to be an important force behind the evolution of cooperative breeding. However, helpers may associate with their relatives as a result of delayed dispersal, hence, kin associations might be a consequence of demographic viscosity rather than active choice. In addition, recent studies showed that helpers may have access to reproduction therefore direct benefits might also play an important role. Here, we investigate the possible roles of direct genetic benefits and kin associations on helping behavior in the sociable weaver Philetairus socius, a colonial and cooperatively breeding passerine. We used a microsatellite-based genotyping method to describe the genetic structure within nests and colonies. Within a colony, we found considerable genetic structure between males but not females. Sociable weaver colonies have several nests that are simultaneously active, giving individuals a choice of associating with a range of first-order kin to unrelated individuals. Helpers were significantly more related to the young in the helped nests than in other nests of the colony, suggesting an active choice for associating with kin. The helpers were generally offspring or first-order relatives of one (50%) or both (43%) breeders, although more infrequently, seemingly unrelated individuals also helped (7%). We found no supporting evidence of extrapair parentage and hence no direct genetic gains from helping in our population. This strong reproductive skew is contrary to theoretical models predicting conflicts over reproduction in stepfamilies. We discuss whether female decisions and/or other direct benefits of remaining in kin associations or helping might explain the high skew observed.     

Recency preference of odour memory retrieval in honeybees

Free-flying honeybees (Apis mellifera) were trained on tasks in which they had to choose one of three odours for a reward of sugar water. In acquisition, the bees learned this task in about five trials of training. Unrewarded retention tests showed that the odour memory was retained after 24-h delay. These findings are unsurprising. Integration experiments were then performed in which the bees had to learn two successive tasks of odour discrimination with conflicting demands. In task 1 (20 trials), one of three odours provided sugar water while the other two provided tap water. In task 2 (ten trials), which followed task 1 immediately, a different odour provided the reward. The bees were given unrewarded tests immediately after training on task 2 and then tested again after 10 min, 22 h or 24 h. The 22-h delay coincided with the circadian time for the start of task 1 training, while the 24 h coincided with the circadian time for the end of the task 2 training. Bees strongly preferred the rewarded odour for task 2 on immediate testing and after a 10-min delay. After delays of 22 and 24 h, they still preferred the rewarded odour for task 2. We conclude that the most recently acquired odour memory dominates behaviour in honeybees. The close association between floral odour and reward availability under natural circumstances may predispose honeybees to rely more on the most recently rewarded odour cue rather than on circadian time.