Variation in group cohesion during the maternal social period of a subsocial spider

During the maternal social period, Amaurobius ferox spiderlings (Araneae: Amaurobiidae) show mutual tolerance, group cohesion and cooperation in prey capturing, which are recognized as the main characteristics in the evolution of spider sociality. Measuring spatial volume occupied by the spiderlings within the maternal web, this study investigated variation in group cohesion over the maternal social period, from emergence to dispersal. The results showed that the spatial volume of spiderlings varied greatly during the maternal social period and was associated with the development of the spiderlings. Strong group cohesion appeared to be related to maternal food provision, trophic egg laying and matriphagy. An increase of the spatial volume was obviously observed after matriphagy. The experiment revealed that group cohesion decreased after the second molt. A compact group of spiderlings should facilitate maternal food production at the prompted time, and reduced group cohesion after the second molt might lead to eventual dispersal of A. ferox spiderlings.     

Adaptive advantages of matriphagy in the foliage spider, Chiracanthium japonicum (Araneae: Clubionidae)

Young of the Japanese foliage spider, Chiracanthium japonicum, show matriphagy, whereby they consume their own mothers before dispersal. By removing mothers in the laboratory, I examined the importance of this sacrificial habit for offspring survival and dispersal behavior. Spiderlings that cannibalized their mothers gained weight more than threefold and dispersed from their breeding nests after molting into the third instar. The third-instar spiderlings had relatively longer legs than the previous instars and appeared to be more adapted to a solitary hunting life style. On the other hand, most spiderlings separated from their mothers could not molt into the third instar and dispersed significantly earlier than those with matriphagy. Furthermore, the lack of matriphagy decreased the survival rate of predispersal spiderlings. These results showed that matriphagy of C. japonicum has a great advantage in allowing offspring to disperse at a more developed and active instar. Received: October 23, 2000 / Accepted: March 15, 2001     

Trophic egg laying in the spider, Amaurobius ferox: mother-offspring interactions and functional value

Offspring of the spider Amaurobius ferox (Araneae, Amaurobiidae) were provided with trophic eggs of their mother the day after their emergence from the egg sac. This precisely timed egg laying followed after a series of mother-offspring interactions involving specific behaviors. Experiments showed that the trophic egg laying of the mother (providing she is in the appropriate reproductive condition) necessitated not only their presence, but also the stimulating behavior of the spiderlings. By stimulating their mother the spiderlings actually inhibited the normal maturation of the second generation of maternal eggs and prompted the release. Comparing to the trophic egg-deprived clutches, the clutches provided with the trophic eggs developed with higher body mass, earlier moulting and matriphagy. More offspring survived at the matriphagy with the mother normally provisioning the first clutch with trophic eggs rather than with the mother that did not produced the trophic eggs for her first clutch but for her second clutch. By turning her potential second generation into food, the mother increases her reproductive success.     

Adaptive advantages of maternal care and matriphagy in a foliage spider,Chiracanthium japonicum (Araneae: Coubionidae)

I examined the function of maternal care in a foliage spider,Chiracanthium japonicum. Females of this species make breeding nests with rolled-up grass leaves and provide themselves to spiderlings as food at the end of maternal care. By removing mothers from their offspring at 2 different times, the effects of maternal care on egg and spiderling survival rates were estimated separately. Mother attendance greatly improved survival and development of eggs as well as spiderlings. Detailed observations on the fate of immatures in breeding nests with and without their mothers showed lower hatching and spiderling emergence rates when mothers were removed. Furthermore, spiderlings that fed on their mother’s body showed accelerated growth and quickly molted into the 3rd instar with the delay of dispersal. This suggests that matriphagy, or eating the mother, enables spiderlings of this species to disperse at a later instar. Therefore, I conclude that the maternal care of this spider consists of guarding offspring, supporting offspring development and feeding spiderlings.     

Maternal inheritance in a subsocial spider: web for collective prey capturing of the young

Construction of a safe refuge is the first step toward the evolution of social organization in spiders. In the case of the subsocial spider Amaurobius Jerox (Araneae, Amaurobiidae), the young remain in the natal nest after consuming their mother's body (matriphagy). The benefit that the young could obtain from the maternal web was investigated in their collective prey capture. The results demonstrated the adaptive value of the maternal web. The prey capturing activity was more effective when the young were allowed to stay on the maternal web than when they used the web constructed by themselves. Maternal web may also provide a better transmission of information on prey state and clutch mates' activities, because more spiderlings were recruited to contact and hold the prey in a shorter period of time than when they were on a spiderling-constructed web. The young's inheritance of the maternal nest is probably one of the crucial ways in which the mother influences the post-maternal social period of the young.     

Maternal Care and Infanticide by Males in Helvibis longicauda (Araneae: Theridiidae)

Maternal care in spiders varies from just the construction of a protective silken structure for the eggs and the selection of a safe site to place them, to a long period of association between the mother and spiderlings. Such extended care may involve the active protection from predators and parasitoids, food regurgitation, the production of trophic eggs and even matriphagy. In this study, we describe extended maternal care in Helvibis longicauda (Theridiidae) and evaluate the effectiveness of maternal protection against predators of eggs and spiderlings. We conducted experiments comparing the frequency of egg sac destruction and mortality of spiderlings in the presence and absence of mothers. We also observed the behaviour of the mother and spiderlings during prey capture events and interactions with possible predators. Helvibis longicauda females guard their egg sacs until the emergence of the young and guard the spiderlings for several instar stages, fighting possible predators, including conspecifics. We found that aggressive behaviour by females increased the survival of both eggs and spiderlings in our experiments. Intruder males were the main source of mortality in the absence of females. The benefits of maternal care for the young also include the acquisition of prey items that are captured, immobilized and pre‐digested by the mother. Effective maternal protection and the extended period of supplying food to juveniles probably contribute to the late dispersal of offspring in H. longicauda.     

Maturing and Coordination of Web-Building Activity

Experiments are reported in which 114 Zygiella x-notata Cl.spiderlings were taken out of the cocoon before the time atwhich they normally left (2 weeks after the second molt); 67were placed on wooden frames and 47 on wire frames. Both groupsbuilt partial, irregular thread-patterns at first, and, as theygrew older, constructed more and more regular geometric orb-webs:the transition can be traced through several stages. The shapeof the web, particularly the presence of the free-sector, wasdependent on the frame; webs built in a circular wire framelacked the free-sector. It is assumed that the accomplishmentof full, regular orb webs is essentially based on the progressivehistological differentiation of the central nervous system inthe spiderlings during the developmental phase under observation;the wire frame could influence the shape of the web throughmaking construction of a retreat difficult.     

Social Facilitation of Synchronized Molting Behavior in the Spider Amaurobius ferox (Araneae, Amaurobiidae)

During the social period, molting behavior of the young spider, Amaurobius ferox, is highly synchronized within the clutch. Result of the experimental study suggests that social facilitation among group members increased the synchronization. The duration of the molting period of grouped spiderlings was significantly shorter than that of individually isolated spiderlings. Involving the particular maternal strategy in food supply, this phenomenon might have adaptive values in the maintenance of mutual tolerance among the siblings by decreasing the interindividual difference in development and in the avoidance of cannibalism on molting individuals. This probably will also serve to make the peaceful collective behaviors of the spiderlings in matriphagy and cooperative prey capture during their social period.     

Reproductive state and care giving in Stegodyphus (Araneae: Eresidae) and the implications for the evolution of sociality

Cooperative brood care is a rare phenomenon in spiders and is restricted to a few social species, including three in the genus Stegodyphus. Brood care in Stegodyphus begins with regurgitation feeding followed by matriphagy: the young consume the body fluids of their mother causing her to die quickly. Whether such an extreme form of maternal care can become a communal task should depend on physiological or historical preconditions. I investigated whether femaleStegodyphus lineatus feed young or allow matriphagy according to their own reproductive state. Broods of young of two age classes (2 or 10 days after hatching) were isolated or fostered out to adult females that were unmated, had eggs or had young. Growth and survival of females and broods were followed over 21 days. The timing of matriphagy depended on the interaction between age of young and state of the foster mother. All broods that were fostered out to females with young grew and survived. Two-day-old young did not survive when isolated or fostered out to unmated females, but some survived and gained weight when placed with foster mothers that cared for egg sacs. Young of 10 days of age grew when fostered out to females with eggs but did not grow or lost weight when isolated or fostered out to unmated females. Survival among 10-day spiderlings was relatively high in all groups but differed significantly between treatments (young isolated or fostered out to unmated females or females with eggs) and control (left with the mother). The results show that these spiders will care for young from other females only when they are in the right developmental state. Such a constraint can have important consequences for the evolution of allomaternal care in social species: unless such a mechanism is overcome, nonreproductives cannot help in brood care. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Offspring recognition by mother crab spiders with extreme maternal care

Maternal care is provided by several spider species, but there are no reports of mother spiders recognizing their young, which suggests that maternal care can be exploited by unrelated individuals. Diaea ergandros, a crab spider with extreme, sacrificial maternal care, does accept unrelated spiderlings (ca. 43.9% of spiderlings) into its nest in areas of high nest density. However, a field and a laboratory experiment with mother spiders and natural and adoptive spiderlings demonstrated that mothers did recognize their own offspring. Recognition was not expressed in survival as adopted (unrelated) spiderlings had similar survival rate to that of natural offspring. Instead it was displayed in growth; mother D. ergandros caught large prey items for their own offspring, but not for adopted spiderlings, and so natural offspring grew more than adopted spiderlings. Also, mothers produced trophic oocytes, which are important for the sacrificial care that influences spiderling survival, only when they lived with their own offspring.     

Effects of Early Contact with Maternal Parent on Locomotor Activity and Exploratory Behavior in Spiderlings of Hogna carolinensis (Araneae: Lycosidae)

In this study we investigated the effect of rearing conditions (stimulus complexity) on the locomotor activity and exploratory behavior of spiderlings of Hogna carolinensis in a novel open field arena. Experimental groups consisted of spiderlings that were allowed to remain with their maternal parent for 1 day (group 1), 3 days (group 2), and 5 days (group 3) after emergence from their egg sacs, and those who had no contact (group 4) with a maternal parent. Spiderlings that had physical contact with their mothers for 3 and 5 days required significantly less time to enter the arena, to reach the central grid area of the arena, and for an initial return to the start box than spiderlings in groups 1 and 4. In addition, spiderlings in groups 2 and 3 exhibited a significantly greater number of grid crossings as compared to groups 1 and 4, and spent more time in the arena. This is the first demonstration that environmental complexity, in the form of a social factor such as physical contact with a maternal parent, can affect the subsequent behavior of spiderlings.     

Agonistic behaviour of female Tegenaria atrica in the presence of different aged spiderlings

The agonistic behaviour of adult female T. atrica increases with increasing age of spiderlings until 80 days post-emergence, then decreases. In all cases, cannibalism of spiderlings increases markedly after completion of their pre-dispersal period with a correlation between the switch of behaviour, from tolerance to cannibalism, and a modification of individual’s cuticular compounds. Experiments that compare spiderlings weight and mobility indicate that female agonistic behaviour are always low with spiderlings in pre-dispersal period, and high with spiderlings in post-dispersal period. Female reproductive state and the weight/mobility of spiderlings are not the only factors controlling the adult female’s agonistic behaviour towards spiderlings on her web, as tactochemical information plays an important role in modulating agonistic behaviour after close contact between female spiders and spiderlings. The agonistic behaviour of females appears to correspond with a change in the increase of polar compound levels (methylesters and fatty acids) and the decrease of apolar compound levels (hydrocarbons) in spiderlings of different ages.     

Ontogenetic Changes in Web Design in Two Orb‐Web Spiders

The first orb web built by newly hatched spiders resembles the adult web in its overall form and structure. However, many details show ontogenetic changes. One possible explanation for these changes is that the tiny early‐instar spiders with their minute brains will make more mistakes and build less ‘perfect’ orb webs than older and larger juveniles and adults. To test this hypothesis, known as the size limitation hypothesis, I analysed orb webs from three developmental stages, spiderlings, juveniles and adult females, in two neotropical orb‐web spiders, the araneid Eustala illicita and the nephilid Nephila clavipes. Neither species showed clear signs of being behaviourally limited or more prone to committing errors as spiderlings than were older juveniles or adults. These findings therefore do not support the size limitation hypothesis in either species. Finally, I looked for evidence of the ‘biogenetic law’, which predicts that juveniles should build less derived orb webs than the adults. Evidence for this was found in E. illicita, but not in N. clavipes.     

Residual vitellus and energetic state of wolf spiderlings Pardosa saltans after emergence from egg-sac until first predation

The aim of this study was to evaluate energetic source used by juveniles of a terrestrial oviparous invertebrate during the earliest periods of their life. Growth, behavioural activities and energy contents of Pardosa saltans spiderlings’ residual vitellus were monitored during 8 days after their emergence from their egg-sac until they disperse autonomously. The life-cycle of juvenile after emergence can be divided into three periods: a gregarious while juveniles are aggregated on their mother, dismounting off their mother’s back and dispersion. We present the first biochemical study of residual vitellus and energy expenditure during these three periods. At emergence, the mean weight of juveniles was 0.59 mg and energy stock from residual vitellus averaged 51 cal/g wet mass. During gregarious period, the weight of the juveniles aggregated on their mother did not vary significantly and juveniles utilized only 1 cal/day from their residual vitellus. During the period from dismounting until their first exogenous feed, juveniles lost weight and used 30% of their residual vitellus stock. Proteins from the residual vitellus contributed principally to their energy expenditure during this period: 1.5 µg protein/day. Juveniles’ first exogenous feeding was observed 7–8 days after emergence, when 70% of residual vitellus energy had been utilized. Juveniles dispersed after eating, reconstituting an energy stock comparable to that observed at emergence from egg-sac (50 cal/g wet mass). This new energy stock contains mainly lipids unlike the energy stock from the residual vitellus.

     

Involvement of the opioid system in the hypokinetic state induced in cockroaches by a parasitoid wasp

The parasitoid wasp Ampulex compressa stings and injects venom into the cockroach brain to induce a long-lasting hypokinetic state. This state is characterized by decreased responsiveness to aversive stimuli, suggesting the manipulation of a neuromodulatory system in the cockroach’s central nervous system. A likely candidate is the opioid system, which is known to affect responsiveness to stimuli in insects. To explore this possibility, we injected cockroaches with different opioid receptor agonists or antagonists before they were stung by a wasp and tested the escape behavior of these cockroaches to electric foot shocks. Antagonists significantly decreased the startle threshold in stung individuals, whereas agonists led to an increased startle threshold in controls. Yet, neither agonists nor antagonists had any effect on grooming. To further characterize the interaction between the venom and opioid receptors, we used an antenna-heart preparation. In un-stung individuals external application of crude venom completely inhibits antenna-heart contractions. In stung individuals the antenna-heart showed no contractions. Although acetylcholine restored contractions, the opioid receptor antagonist naloxone was unable to antagonize the venom inhibition. These results suggest that the venom of A. compressa might contribute to the manipulation of cockroach behavior by affecting the opioid system.     

Functional Value of Matriphagy in the Spider Amaurobius ferox

The adaptive value of matriphagy, the consumption of the mother by her offspring, in the sub-social spider Amaurobius ferox (Araneae, Amaurobiidae) was experimentally evaluated in terms of the benefits to the offspring and the costs to the mother. Matriphagy resulted in a 2.5-fold weight gain in the offspring over their initial weight, advancement of their moulting time, and larger body mass at dispersal in comparison with clutches deprived of matriphagy, but otherwise well provisioned with prey. Matriphagous offspring were also more successful at capturing large prey items, had a more extended social period, and a higher survival rate at dispersal. Mothers separated from their broods just prior to matriphagy were able to produce second clutches, effectively producing 33% more viable offspring (compared with the first clutches). However, the estimated reproductive outputs of the alternative maternal strategies (being devoted to the first clutch vs. deserting the first and producing a second one) suggest that mothers of A. ferox that are cannibalized by their broods enjoy greater reproductive success than those that escape cannibalism and produce second clutches.     

寡居种悦目金蛛若蛛群居机制初步研究

对寡居种悦目金蛛Argiope amoena若蛛群居生活对其结网的影响、温度和种群大小对若蛛存活率的影响、若蛛对限制性空间的利用及其扩散方式进行了观察和研究.结果表明,若蛛从卵袋出蛰后不经历群居扩散即具备结网能力,可结完整网,也具备在扩散前不进食水存活的能力;在变温条件下若蛛存活率要远远高于室温条件下的存活率,而在室温条件下若蛛开始死亡和半数死亡的时间比其聚居期长得多;若蛛主要通过群体空中扩散的方式进行扩散,可以减小能耗与敌害的威胁;不同种群大小的若蛛在限制性空间的分布均从第5d左右开始发生较大变化的,这与在空间不受限的条件下若蛛在第5d开始扩散的结果是一致的.据此我们推断:个体扩散代价最小化(包括御敌和能耗)和维持种群较高的生存率可能是寡居种悦目金蛛若蛛群居生活的主要原因.     

Ionic analyses and effects of ions on contractions of sponge tissues

Summary Contractions of oscular membranes of several marine sponges in response to mechanical stimuli were recorded in various ionic media. Previously, failure to record action potentials and to stimulate electrically was reported.Contractions persisted when sodium was replaced by lithium or potassium but not when the replacement was by choline or sucrose. Contractions were maintained in sea water in which the only univalent cation was potassium; they also persisted in Na-sea water without potassium.Contractions stopped when either calcium or magnesium was omitted, but addition of an equivalent extra amount of the other divalent cation then restored contractions. Contractions were also maintained when strontium was substituted for calcium and magnesium.Analyses of contractile membranes yielded average values for Naples sponges of Na 375 mM/kg H₂O, K 29.8 mM/kg H₂O and inulin space 48.9% of H₂O; for Woods Hole Microciona in mM/kg H₂O: Na 300, K 66, Ca 8.4, Cl 225, inulin space 40% H₂O. The calculated intracellular concentrations for Microciona in mM/kg H₂O were: Na 216, K 104.5, Ca 7.8, Cl 45 yielding i/o ratios of Na 0.5, K 11.6, Ca 0.84, Cl 0.09.It is concluded that, despite relatively normal ionic ratios, mechanical stimuli activate the contractile system in sponges more directly than in muscle.Dedicated to Prof. H. Autrum on the occasion of his 60th birthday.This work was done in part while the author held a Guggenheim Fellowship. Support from NSF grant G 8795 is acknowledged.Assistance of Donald Job, particularly in making all calcium and magnesium measurements, is gratefully acknowledged.     

Araneophagic behavior of Gardena brevicollis Stål (Heteroptera: Reduviidae): Foraging on pre‐dispersal spiderlings in the spider's web

Araneophagic behavior of an Emesinae assassin bug, Gardena brevicollis Stål (Hemiptera: Reduviidae) was observed in the field. The bug invaded a web of Acusilas coccineus Simon (Araneae: Araneidae) spiderlings and hunted five juvenile spiders. The bug showed stalking tactics to prey on spiders, whereas luring tactics, which have been reported in other Emesinae bugs, were not observed. To the best of my knowledge, the present study is the first report of araneophagy in the genus Gardena and the first report of araneophagic behavior in East Asian assassin bugs.     

Costs and Benefits of Maternal Care in a Subsocial Spider,Coelotes terrestris

The maternal social spider Coelotes terrestris demonstrates extended care towards its progeny: the mother guards its egg sac for 3–4 weeks, then stays with its young from the time of their emergence until their dispersal about 1 month later. The present investigation evaluates the adaptiveness of these maternal behaviours by comparing the fitness of females performing them with that of females separated from the egg sac or the spiderlings. By protecting their egg sacs from predation and parasites, and by pursuing this task while supplying the young with food, mothers enhance the survival rate and the development of many of their spiderlings. The costs linked with these activities, estimated by the ability to produce another clutch, appear variable according to the stage in the reproductive cycle. In such terms, the egg sac guarding appears to have a low cost in relation to the care given to the spiderlings.