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.     

Physiological costs during the first maternal care in the wolf spider Pardosa saltans (Araneae,Lycosidae)

Many arachnids like other terrestrial arthropods, provide extensive maternal care. Few studies have quantified the underlying physiological costs of maternal care. We investigated how maternal care affects the free-moving wolf spider’s (Pardosa saltans) energy requirements. We described in detail their basic reproduction biology (i.e. carrying cocoon and young) and we evaluated the variation in the females’ energy reserves during maternal care. Our results show that mothers guard eggs until hatching and then guard their spiderlings for 27–30 more days. Laboratory observations indicated that spiderlings start leaving the maternal abdomen gradually 5–7 days after hatching. Females carry an egg sac (cocoon) that can weigh up to 77% of their post-reproduction weight and carry young that weigh 87–100% of their body mass. Females lost weight over time despite regular food intake, while carrying cocoon and young; but their weights increased gradually during the dispersal of young. The contributions of proteins, glucose and triglycerides to maintain females’ energy were calculated. Their energetic state varied during maternal care, in particular lipid levels declined, during the care of spiderlings when the females’ predatory behaviour was inhibited. Our results show that the maternal care provided by P. saltans females is particularly costly physiologically, during the 30 days following egg sac formation and development of spiderlings, even when food is available.     

Behaviour of colonial and solitary spiders of the theridiid species Anelosimus eximius

Most Anelosimus eximius live in colonies, but a few females emigrate short distances and establish small, individual webs. Field studies were conducted on one colony and nearby smaller webs in order to describe communal activities and division of labour, and to note costs and benefits of remaining in the parental colony and emigrating. Adult and juvenile females repaired the web and captured prey. Adult females, rather than juveniles, cared for egg sacs and fed young. When colonial egg sacs were abundant, females moved from sac to sac. Several colonial females regurgitated food to spiderlings which had recently emerged from a particular sac. In smaller webs inhabited by two or three females, the mother cared for the sac but all females fed the young by regurgitation. Males rarely participated in communal activities. Advantages of colony living include protection from predators, the availability of large prey and, in the event of a female's death, the care of her egg sac and feeding of her young. Cannibalism of the egg sac is a potential cost of coloniality, affecting about 10–20% of colonial sacs. Cannibalism was not observed in the smaller webs. However, the costs of emigrating are very high: most of the solitary females disappeared, leading to interspecific predation on their eggs and young.     

Spiderling sex ratio and maternal investment in the bolas spider Mastophora cornigera (Araneae,Araneidae)

Male Mastophora cornigera exit egg sacs as adults, which allowed us to determine spiderling sex ratios and patterns of maternal investment in this species. We collected 15 egg sacs produced by seven mothers, which yielded 1945 emergent spiderlings which were sexed, 1850 of which were weighed. Two emergent broods were significantly male and female biased and were unaffected by pre-emergence mortality. The weights of male and female spiderlings differed in eight broods, with males and females being heavier in four cases each. Five of these broods were derived from multiple egg sac sets produced by one mother, and in each case, the total mean male and female spiderling weights for all broods in a set were biased in the same direction as the biased brood(s) within that set. Mean emergent spiderling weight was independent of brood size and sex ratio for both males and females. Despite such independence, sex allocation in M. cornigera can favor sons, daughters, or both equally, and by numbers, by weight, or both at once. The proximate mechanisms and adaptive significance of such variability is unknown. We also review evidence for gender-biased allocations in arachnid offspring and suggested mechanisms for their applicability to M. cornigera.     

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.     

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.     

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.     

Protective function of maternal care against egg predators in the shield bug Sastragala esakii (Hemiptera: Acanthosomatidae)

Females of many acanthosomatid shield bugs attend eggs and young nymphs by covering them with their bodies. Although such form of maternal care has been suggested to have evolved independently in four genera, Elasmucha, Sastragala, Acanthosoma and Sinopla, previous studies exploring its adaptive function have solely focused on species of Elasmucha. This study pioneered an experimental examination of maternal care in the Japanese species Sastragala esakii. Field experiments demonstrated that unattended egg masses suffered intense predation, whereas egg masses attended by their mothers were rarely preyed upon. The ant Crematogaster sp. was the most common egg predator, while two other insect species were also observed to prey on eggs. The exclusion of ant workers and other walking predators from accessing egg masses through the utilization of a sticky trap resulted in a remarkably improved egg survival rate in orphaned egg masses, suggesting that, at least in our study site, maternal care of S. esakii primarily functions to protect eggs from walking predators. Laboratory observations revealed that egg protection against predators was achieved through at least two distinctive defensive behaviors shown by egg-tending females: (i) tilting their bodies when approached by predators; and (ii) fanning their wings when in contact with predators. However, females displayed a limited response to predators approaching from the posterior direction, suggesting that visual cues play a significant role in predator recognition. These results indicate a similarity in the maternal care functions between Elasmucha and Sastragala, and suggest the parallel evolution of female defensive behaviors.     

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.     

Costs and benefits of maternal behaviour in the green lynx spider (Oxyopidae,Peucetia viridans)

I examined the adaptiveness of maternal behaviour in the green lynx spider Peucetia viridans (Hentz), by measuring the costs and benefits to the female of egg-sac tending. P. viridans guards her egg sac for 6–8 weeks until the young have emerged and dispersed. I removed females from egg sacs in the field either immediately after oviposition or after they had guarded the egg sacs for 2 weeks, and compared the fate of these sacs with that of control sacs which had females present throughout. Survival rates of egg sacs were significantly improved by the presence of a female. The major sources of mortality were ant predation and sac dislodgement, both of which were significantly reduced by a guarding female, and mantispid brood parasitism, which was not affected by the female. To measure the spiders' lifetime reproductive output, I allowed caged females to construct egg sacs. Half had their first egg sac removed shortly after oviposition, while half were allowed to guard. Significantly more non-guarding than guarding females produced second egg sacs. Thus for the green lynx spider, egg sac guarding decreases lifetime egg production, but increases offspring survival. Under north Florida conditions, the benefits of guarding outweight the costs, and the females have a higher lifetime reproductive success than if they did not guard.     

Social behaviour of spitting spiders (Araneae: Scytodidae) from Singapore

The social organization of Scytodes sp., a spitting spider from Singapore, was investigated in the field and in the laboratory. The characteristics considered were colony structure, maternal care, prey capture, prey sharing, adults provisioning with prey and how feeding regime influences the level of tolerance expressed by juveniles that were living together. The characteristics of this species were compared with those of other Scytodes species that have been studied previously. Based on our findings, we conclude that in this species, social organization is 'periodic-social', with extended maternal care, this being a type of 'subsociality'. The females of Scytodes sp. wrapped their eggs in a silk egg sac and used their chelicerae to hold the eggs until they hatched. Spiderlings tended to remain in their mother's web until the fourth instar (i.e. after the third moult) but they were also highly aggressive and cannibalistic towards conspecifics after the third instar. Females carried prey to their spiderlings and either fed alongside the brood or left them to feed alone. Spiderlings also captured large prey cooperatively by spitting onto, biting and feeding on it together. The occurrence of cannibalism among her juveniles did not appear to be influenced by whether the adult female was present or not, nor did it appear to be influenced by prey abundance.     

Energy content of spider eggs

Summary The energy content of spider eggs was determined on samples from 12 species representing 6 families. These values ranged from 26.32 to 29.00 joules per mg ash-free dry mass with a mean of 27.30±SE of 0.27. The higher values were found in those species that overwinter as developmental stages within the egg sac. Rates of energy expenditure of developing eggs and spiderlings within the egg sac were only 7 to 19% of those of emerged spiderlings. The energy conserved by this reduction in rate of metabolism may facilitate survival without feeding during the potentially long periods of aerial dispersal by ballooning, a characteristic activity of most newly emerged spiderlings. The variation in mass-specific energy content is less than variation in clutch size and individual egg size on an intra-and interspecific basis. There was no correlation between energy content per unit egg mass and size of the female parent, egg size, or clutch size. Further analysis indicated that no single measure such as clutch size accurately represents the proportional amount of energy invested in reproduction in these animals.     

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.     

Egg sac damage and previous egg sac production influence truncated parental investment in the wolf spider,Pardosa milvina

Life history theory predicts that iteroparous animals adaptively partition reproductive effort between current and future reproduction. When rearing costs of current offspring exceed the potential benefits, parental care should be terminated and deferred toward future reproduction. We tested two related predictions that follow from life history theory: (a) parents should be sensitive to offspring viability and withhold parental care if offspring survival probability drops and future reproductive opportunities are likely, and (b) parents should be less sensitive to offspring survival probability when future reproduction is unlikely and maximize parental care late in life. The wolf spider, Pardosa milvina, demonstrates extensive parental care; however, they may also abandon or cannibalize their egg sacs. We tested the effects of egg sac damage and production of a previous egg sac on egg sac abandonment and cannibalism decisions. Among four egg sac groups (1st egg sac intact, 1st egg sac damaged, 2nd egg sac intact, 2nd egg sac damaged), we daily monitored egg sac abandonment and cannibalism and measured differences in egg sac searching, protection, and grooming among removed and damaged egg sacs (N = 116 with 1st egg sac and 88 with 2nd egg sac). Females with first egg sacs abandoned and cannibalized damaged egg sacs significantly more compared to unmanipulated egg sacs; however, females with second egg sacs were insensitive to egg sac damage. Females also spent significantly more time protecting second egg sacs compared to first egg sacs and groomed damaged egg sacs significantly more than undamaged. These results support the general predictions of life history theory that indicate that abandonment and cannibalism should decrease with diminished future reproductive potential and that parents should be less sensitive to indicators of offspring survival probability late in life.     

Haltung und Zucht der Geißelspinne (Damon variegatus) (Phrynichidae) (Chelicerata: Amblypygi)

In the temporary carboniferous hall of „GONDWANA – Das Prähistorium“ in Germany, whip spiders (Damon variegatus) were kept and bred. 3,3 were purchased from a dealer in summer 2007. The night active spiders were kept together in a terrarium. No aggresions were observed. The animals were fed little locusts, young hissing cockroaches, and crickets. In 2008 courtship display and mating could be observed. The biggest male was the only active one. After 7 days after the intake of the spermatophores all females carried eggsacs under the ophistosoma. Each egg sack contained approximately 25–30 eggs. During the carrying phase only one female feasted on a cricket. 98 days after the egg sack was produced the young spiderlings hatched. The coloration of the spiderlings was completely different from the coloration of the adults. The spiderlings were carried by the female for the next 9 days. Then, the spiderlings moulted and left the female immediately. The adults were now separated from their young. The three groups of spiderlings stayed together in a smaller terrarium. However, each group from each female stayed separate from the others for weeks. The second moulting took place 41 days after the first one. No cannibalism was observed. Obviously Damon variegatus seems to be a spider with a social touch. If the terrarium and the climate conditions inside are adapted to the needs of the whip spiders keeping and breeding is possible. The species is highly attractive to visitors.     

Haltung und Zucht der Geißelspinne (Damon variegatus) (Phrynichidae) (Chelicerata: Amblypygi)

In the temporary carboniferous hall of „GONDWANA - Das Prähistorium“ in Germany, whip spiders (Damon variegatus) were kept and bred. 3,3 were purchased from a dealer in summer 2007. The night active spiders were kept together in a terrarium. No aggresions were observed. The animals were fed little locusts, young hissing cockroaches, and crickets. In 2008 courtship display and mating could be observed. The biggest male was the only active one. After 7 days after the intake of the spermatophores all females carried eggsacs under the ophistosoma. Each egg sack contained approximately 25-30 eggs. During the carrying phase only one female feasted on a cricket. 98 days after the egg sack was produced the young spiderlings hatched. The coloration of the spiderlings was completely different from the coloration of the adults. The spiderlings were carried by the female for the next 9 days. Then, the spiderlings moulted and left the female immediately. The adults were now separated from their young. The three groups of spiderlings stayed together in a smaller terrarium. However, each group from each female stayed separate from the others for weeks. The second moulting took place 41 days after the first one. No cannibalism was observed. Obviously Damon variegatus seems to be a spider with a social touch. If the terrarium and the climate conditions inside are adapted to the needs of the whip spiders keeping and breeding is possible. The species is highly attractive to visitors.     

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.     

Subsociality and female reproductive success in a mycophagous thrips: An observational and experimental analysis

Oviparous females of the haplodiploid, facultatively viviparous thrips Elaphrothrips tuberculatus(Thysanoptera: Phlaeothripidae) guard their eggs against female conspecifics and other egg predators. The intensity of maternal defense increases with clutch size. Field and laboratory observations indicate that cannibalism by females is an important selective pressure favoring maternal care. Experimental removals of guarding females showed that egg guarding substantially increases egg survivorship and that the survivorship of undefended eggs is higher in the absence of nonguarding female conspecifics than in their presence. The fecundity of viviparous females increases with the number of eggs cannibalized. The reproductive success of oviparous females increases with body size and local food density and decreases with local density of breeding females. Social behavior may not have advanced beyond maternal care in Elaphrothrips tuberculatusbecause, relative to Hymenoptera, capabilities for helping relatives are few or nonexistent, and the causes of variation in female reproductive success are not influenced easily by cooperation among females.     

Proximate Factors Regulating Maternal Options in the Eggplant Lace Bug, Gargaphia solani (Heteroptera: Tingidae)

The eggplant lace bug, Gargaphia solani, was used to investigate the proximate factors regulating maternal care and a noncaring, condition dependent strategy called egg dumping. We hypothesize that the act of delaying oviposition while searching for a dumping opportunity suppresses oogenesis and triggers guarding behavior. We examined several predictions of this hypothesis by measuring: (1) whether females do delay oviposition in the absence of dumping stimuli, (2) whether females in transition between egg dumping and egg guarding are capable of expressing either reproductive option, (3) the effect of nymphal interactions and antennal ablation on the duration of maternal care, and (4) oogenesis in guarding and dumping females. We found that females without a dumping opportunity wait, on average, 30 h longer to oviposit than females exposed to a dump mass. Females that had initiated their own egg mass could resume eggdumping if they had laid less than half of their eggs but were unlikely to abandon their eggs when most had been laid. Maternal care in G. solani can be prolonged if interactions with nymphs are artificially prolonged. Females require antennae to maintain maternalcare. Presumably antennae transduce cues from eggs and nymphs. Dissections of dumping and guarding females 72 h after their first oviposition demonstrated that dumpers continue to produce primary oocytes after a dumping event but guarders terminate oogenesis whilecaring for their first brood. We interpret all of these results within the context of the hypothesis that juvenile hormone titers regulate the expression of both egg dumping and egg guarding.