Effects of a low oxygen environment on parental effort and filial cannibalism in the male sand goby, Pomatoschistus minutus

In fish, brood cycling parental males sometimes eat some orall of their eggs, a behavior termed filial cannibalism. Wetested predictions of filial cannibalism models related to thecost of parental care in the male sand goby, Pomatoschistusminutus, by increasing the parental effort (fanning expenditure)through reduced levels of dissolved oxygen to 39% in an experimentalgroup, whereas a control group had fully saturated water. Malesshowed both full-clutch cannibalism and partial-clutch cannibalismin both treatments. Giving the males one to three females tospawn with, we found that small clutches were completely eatenmore often than were larger ones, whereas partial-clutch cannibalismwas not affected by clutch size. Although treatment did notaffect filial cannibalism, it did affect a male's energy statesuch that males in the low oxygen treatment lost more body fat,indicating a greater fanning effort. This shows that males inthe low oxygen treatment allocated more energy to the presentbrood, potentially at the expense of future reproductive success.Our study strongly suggests that filial cannibalism in malesand gobies represents a strategic life-history decision asan investment in future reproductive success, and is not triggeredby a proximate need for food necessary for the male's own survival.Furthermore, males in the low oxygen treatment built nests withlarger entrances, and were less likely to rebuild their nestsafter destruction. Presumably, this makes fanning easier butthe nest more vulnerable to predators, suggesting a trade-offbetween fanning and nest defense.     

Egg cannibalism in Ascia monuste in the field; opportunistic, preferential and very frequent

Newly hatched caterpillars of the specialist herbivorous species Ascia monuste (Lepidoptera, Pieridae) feed on egg chorion before ingesting plant material. They also ingest their exoskeletons and perform egg cannibalism in both the field and the laboratory. Females lay eggs on plants which already have conspecifics (eggs and/or caterpillars) leading to the observation of cannibalism in the field. Two favorable conditions are required for such an occurrence: (1) eggs laid close together on the same leaf, and (2) the finding by older caterpillars of eggs on the same plant. Oviposition preference experiments in cages showed that females do not avoid laying on plants with eggs. However, they do avoid laying eggs on plants predated by caterpillars. All instars perform cannibalism in the field and, in addition to this, the ingested eggs are healthy and cannibalism occurs in the presence of abundant food. The ingestion of eggs from their own clutch was more intense when the clutch was large. Contact with eggs is also important for cannibalism. Caterpillars ingested healthy eggs during three successive days. According to the results, it is possible to state that egg cannibalism in A. monuste is opportunistic, preferential and highly frequent and that there is a tolerance to a diet based on animal protein in this species. Since A. monuste obtains nutrients from both animal and vegetable resources, it is probably better to consider it as opportunistic omnivorous.     

Egg sex ratio and paternal traits: using within-individual comparisons

Empirical studies of sex ratios in birds have been limited dueto difficulties in determining offspring sex. Since molecularsexing techniques removed this constraint, the last 5 yearshas seen a great increase in studies of clutch sex ratio manipulationby female birds. Typically these studies investigate variationin clutch sex ratios across individuals in relation to environmentalcharacteristics or parental traits, and often they find no relationships. In this study we also found that clutch sex ratiosdid not vary in relation to a number of biological and environmentalfactors for 238 great tit Parus major nests. However, interestingsex ratio biases were revealed when variation in clutch sexratios was analyzed within individual females breeding in successiveyears. There was a significant positive relationship betweenthe change in sex ratio of a female's clutch from one yearto the next and the relative body condition of her partner.Females mating with males of higher body condition in yearx + 1 produced relatively male-biased sex ratios, and the oppositewas true for females mated with lower condition males. Within-individualanalysis also allowed investigations of sex ratio in relationto partner change. There was no change in sex ratios of femalespairing with the same male; however, females pairing with anew male produced clutches significantly more female biased. Comparisons of clutch sex ratios within individuals may be apowerful method for detecting sex ratio variation, and perhapsfemale birds may indeed manipulate egg sex but require personalcontextual experience for such decisions.     

Predation risk affects trade-off between nest guarding and foraging in Seychelles warblers

The fitness costs of egg loss for Seychelles warblers (Acrocephalus sechellensis)on Cousin Island are considerable because warblers have a single-eggclutch and no time to lay a successful replacement clutch. Onthe islands of Cousin and Cousine, with equal densities of Seychellesfodies (Foudia sechellarum), nearly 75% of artificial eggs placedin artificial nests were predated by fodies after 3 days. OnAride Island with no fodies present, loss of artificial eggswas not observed. Female warblers incubate the clutch, and malewarblers guard the clutch when females are absent. Deterrenceof fodies by male warblers is efficient: loss rate of eggs fromunattended warbler nests was seven times as high as from attendednests, and the more nest guarding, the lower the egg loss andthe higher the hatching success. Egg loss is independent ofthe amount of incubation by females. There is no trade-off betweenincubating and foraging by females. Nest guarding competes withforaging by males, and this trade-off has a more pronounced effecton egg loss when food availability is low. The transfer of breeding pairsfrom Cousin to either Cousine with egg-predating fodies or toAride without fodies allowed us to experimentally investigatethe presumed trade-off between nest guarding and foraging. OnCousine, individual males spent the same amount of time nestguarding and foraging as on Cousin, and egg loss was similarand inversely related to time spent nest guarding as on Cousin.Males that guarded their clutch on Cousin did not guard theclutch on Aride but allocated significantly more time to foragingand gained better body condition. Loss of warbler eggs on Aridewas not observed. Time allocation to incubating and foragingby individual females before and after both translocations remainedthe same.     

Clutch size and body size at first reproduction in Daphnia pulicaria at different levels of food and predation

Field data from seven alpine lakes in Serra da Estrela. Portugal.show that reproduction in Daphnia may be as efficiently controlledby fish predation and copepod predation on eggs in brood cavitiesas it is by food limitation. Body length and clutch size estimatesin Daphnia pulicaria revealed high inter- and intra-populationvariability in maturation size (body size at first reproduction).and in number of eggs per clutch. Daphnia at first maturationin lakes stocked with rainbow trout were half the size of thosefound in fishless lakes (body length of 0.86–0.95 and1.55–1.81 mm. respectively). The mean number of eggs perclutch was reduced to a similar degree by food limitation, predationby fish and copepod predation on eggs in brood cavities, butthe underlying mechanisms of this reduction were different.Food limitation caused smaller clutch sizes in all individuals,so variation remained the same. Fish predation caused the selectiveremoval of individuals with maximum clutches, so variation decreased.Copepod predation caused removal of eggs from brood cavitiesof randomly infested females, so that variation increased, particularlyat a high food level when non-infested females carried largeclutches of eggs.     

Reproduction in foundress associations of the social wasp, Polistes carolina: conventions, competition, and skew

Who reproduces in colonies of social insects is determined bysome combination of direct competition and more peaceful convention.We studied these two alternatives in foundresses of the paperwasp, Polistes carolina, by examining two different contexts:what determines who becomes the dominant reproductive and whatdetermines the amount of reproduction obtained by subordinates.The dominant queen on most nests was the foundress to arrivefirst, rather than the largest foundress, expected to be bestat fighting. This suggests that dominance is initially determinedby convention, although the persistence of some aggressiveconflict throughout the foundress period suggests that thisconvention is not absolute. Attempts to explain the divisionof reproduction using several skew theories were generallyunsuccessful. Skew was not correlated with relatedness, size differences, colony productivity, and challenges by the subordinate.P. carolina showed high constraints against solitary nesting,with a minority of females attempting to nest alone, and nonesucceeding. In this situation, most skew theories predict thatgroup stability will be independent of relatedness, yet nearlyall collected subordinates were full sisters to the queen.Reproductive partitioning in early P. carolina colonies may have more to do with enhancing worker production than with conflictover direct fitness.     

Relative reproductive success of female moorhens using conditional strategies of brood parasitism and parental care

In a population of moorhens (Gallinula chloropus), at least27% of netting females laid one or more eggs in a neighbor'snest Females laid parasitically under three conditions: 56%of parasitic eggs were from nesting females that preceded layinga dutch in their own nest by a parasitic laying bout, 19% werefrom females whose nests were depredated before clutch completionand that laid the following egg parasiticaDy, and 25% were froma small number of females without territories, "non-nesting"parasites, that each laid a series of parasitic eggs. Clutchsizes varied greatly between females, but nesting females eachlaid a consistent clutch size both within and between seasonsfor a given mate and territory. Nesting females that employeda dual strategy of brood parasitism and parental care producedextra eggs that they laid in the nests of neighbors before layinga dutch in their own nests. Two out of ten females whose dutchesI experimentally removed during the laying period were successfullyinduced to lay their next egg in the nest of a neighbor. Nestingfemales that laid parasitically selected their hosts opportunisticallyfrom among the nests dosest to their territories. An experimentin which parasitic eggs were removed and hosts left to rearonly their own young showed that parasites did not choose hoststhat were better parents than pairs with contemporary neststhat were not parasitized. Females that only laid parasiticaDywithin a given season timed their parasitic laying bouts poorlyand achieved no reproductive success. Parasitic young rarelyfledged, and the mean seasonal reproductive success of nestingbrood parasites did not differ from that of nonparasitic females.However, the variance in reproductive success of nesting broodparasites was significantly higher than that of nonparasiticfemales.     

Clutch-size adjustments and skew models: effects on reproductive partitioning and group stability

Reproductive skew theory seeks to integrate social and ecologicalfactors thought to influence the division of reproduction amonggroup-living animals. However, most reproductive skew modelsonly examine interactions between individuals of the same sex.Here, we suggest that females can influence group stabilityand conflict among males by modifying their clutch size andmay do so if they benefit from the presence of subordinate malehelpers or from reduced conflict. We develop 3 models, basedon concessions-based, restraint, and tug-of-war models, in whichfemale clutch size is variable and ask when females will increasetheir clutch size above that which would be optimal in the absenceof male–male conflict. In concessions-based and restraintmodels, females should increase clutch size above their optimaif the benefits of staying for subordinate males are relativelylow. Relatedness between males has no effect on clutch size.When females do increase clutch size, the division of reproductionbetween males is not influenced by relatedness and does notdiffer between restraint and concessions-based models. Bothof these predictions are in sharp contrast to previous models.In tug-of-war models, clutch size is strongly influenced byrelatedness between males, with the largest clutches, but thefewest surviving offspring, produced when males are unrelated.These 3 models demonstrate the importance of considering third-partyinterests in the decisions of group-living organisms.     

Choosy females and indiscriminate males: mate choice in mixed populations of sexual and hybridogenetic water frogs (Rana lessonae, Rana esculenta)

For several decades, behavioral ecologists have studied theeffects of the environment on the behavior of individuals;but only fairly recently they have started to ask the reversequestion: how do the behavioral strategies of individuals affectthe composition and dynamics of populations and communities?Although intuitively obvious, this feedback from individualto higher levels is difficult to demonstrate, except in systemswith exceptionally fast and marked responses of the populationsto the behavior of its members. Such a system exists in sperm-dependentspecies. In European water frogs, for instance, successfulreproduction of a hybrid species (R. esculenta, genotype LR)requires mating with one of its parental species (R. lessonae,genotype LL), except in the rare cases where hybrids are triploid.The sexual host LL, however, should avoid matings with the sexual parasite LR, because the resulting LR offspring willeliminate the L genome from their germ line. In this studywe investigate how this conflict is solved. Since water froghybrids come in both sexes, rather than as females only likein other sperm-dependent systems, we performed the tests withboth females and males. One individual was given a choice betweentwo individuals of the opposite sex, one an LL and the otheran LR. In both species, females showed the predicted preferencefor LL males, whereas males did not discriminate between LLand LR females. On the individual level, we interpret the sexdifference in choosiness by the lower costs from mating withthe wrong species (LR) and the higher benefits from matingwith large individuals in males than in females. In "normal"species, male preference for large (i.e. more fecund) femalesis advantageous, but in this system such a choice can resultin mating with the larger LR females. With respect to the structureand dynamics of mixed populations, we discuss that the observed female preference is consistent with the higher mating successof LL males found in nature. Hence, mate female choice is astrong candidate for a mechanism promoting coexistence of thesperm-dependent hybrid and its sexual host. This confirms predictionsfrom previous theoretical models.     

Zooplankton growth rates: diel egg production in the copepods Oithona, Euterpina and Corycaeus from tropical waters

Clear diel periodicity in the timing of egg laying was observedin the tropical cyclopoid cope-pods Oithona plumifera, Oithonanana, Oithona simplex and Corycaeus amazomcus, and the harpacti-coidEuterpina acutifrons in waters surrounding Jamaica, West Indies.Individual females incubated under ambient food conditions formultiple clutches displayed remarkable constancy in their clutchdurations (= egg or embryonic development time). All speciesexhibited clutch durations of 20–23 h and cycle durations(= sum of clutch and inter-clutch durations) of 24 h, exceptfor the offshore species O.plumifera where the clutch durationwas 36 h and the cycle duration 48 or 72 h. Egg laying appearedto be synchronized by photoperiod, occurring around dawn formost species. In order to avoid biases, diel periodicity mustbe accounted for in estimates of egg production rates, and futureefforts should concentrate on the responses of clutch size andinterclutch durations to natural food climates.     

Brood parasitism,relatedness and sociality: a kinship role in female reproductive tactics

Conspecific brood parasitism (CBP) is a reproductive tactic in which parasitic females lay eggs in nests of other females of the same species that then raise the joint brood. Parasites benefit by increased reproduction, without costs of parental care for the parasitic eggs. CBP occurs in many egg‐laying animals, among birds most often in species with large clutches and self‐feeding young: two major factors facilitating successful parasitism. CBP is particularly common in waterfowl (Anatidae), a group with female‐biased natal philopatry and locally related females. Theory suggests that relatedness between host and parasite can lead to inclusive fitness benefits for both, but if host costs are high, parasites should instead target unrelated females. Pairwise relatedness (r) in host–parasite (h‐p) pairs of females has been estimated using molecular genetic methods in seven waterfowl (10 studies). In many h‐p pairs, the two females were unrelated (with low r, near the local population mean). However, close relatives (r = 0.5) were over‐represented in h‐p pairs, which in all 10 studies had higher mean relatedness than other females. In one species where this was studied, h‐p relatedness was higher than between nesting close neighbours, and hosts parasitized by non‐relatives aggressively rejected other females. In another species, birth nest‐mates (mother–daughters, sisters) associated in the breeding area as adults, and became h‐p pairs more often than expected by chance. These and other results point to recognition of birth nest‐mates and perhaps other close relatives. For small to medium host clutch sizes, addition of a few parasitic eggs need not reduce host offspring success. Estimates in two species suggest that hosts can then gain inclusive fitness if parasitized by relatives. Other evidence of female cooperation is incubation by old eider Somateria mollissima females of clutches laid by their relatives, and merging and joint care of broods of young. Merging females tended to be more closely related. Eiders associate with kin in many situations, and in some geese and swans, related females may associate over many years. Recent genetic evidence shows that also New World quails (Odontophoridae) have female‐biased natal philopatry, CBP and brood merging, inviting further study and comparison with waterfowl. Kin‐related parasitism also occurs in some insects, with revealing parallels and differences compared to birds. In hemipteran bugs, receiving extra eggs is beneficial for hosts by diluting offspring predation. In eggplant lace bugs Gargaphia solani, host and parasite are closely related, and kin selection favours egg donation to related females. Further studies of kinship in CBP, brood merging and other contexts can test if some of these species are socially more advanced than presently known.     

Gender-specific differences in cannibalism between a laboratory strain and a field strain of a predatory mite

Many phytoseiid species, including Phytoseiulus persimilis, are known to engage in cannibalism when food is scarce and when there is no possibility to disperse. In nature adult females of P. persimilis are known to disperse when prey is locally depleted. Males, in contrast, are expected to stay and wait for potential mates to mature. During this phase, males can obtain food by cannibalizing. Therefore, we hypothesize that male P. persimilis exhibit a higher tendency to cannibalize than females. Because rearing conditions in the laboratory usually prevent dispersal, prolonged culturing may also affect cannibalistic behavior. We hypothesize that this should especially affect cannibalism by females, because they consume far more food. We tested these hypotheses by comparing males and females from two strains, one of which had been in culture for over 20 years, whereas the other was recently collected from the field. It is known that this predator can discriminate between kin and non-kin and prefers cannibalizing the latter, hence to construct lines with high relatedness we created isofemale lines of these two original strains. We subsequently tested to what extent the adult females and males of the original strains and the isofemale lines cannibalized conspecific larvae from the same strain/line in a closed system. Relatedness with the victims did not affect cannibalistic behavior, but males engaged more often in cannibalism than females, and females of the laboratory strain engaged more in cannibalism than those of the field strain, both in agreement with our ideas. We hypothesize that the difference in cannibalism between the two genders will increase when they have the alternative to disperse.     

Genetic variation,multiple paternity,and measures of reproductive success in the critically endangered hawksbill turtle (Eretmochelys imbricata)

The Yucatán Peninsula in Mexico contains some of the largest breeding groups of the globally distributed and critically endangered hawksbill turtle (Eretmochelys imbricata). An improved understanding of the breeding system of this species and how its genetic variation is structured among nesting areas is required before the threats to its survival can be properly evaluated. Here, we genotype 1195 hatchlings and 41 nesting females at 12 microsatellite loci to assess levels of multiple paternity, genetic variation and whether individual levels of homozygosity are associated with reproductive success. Of the 50 clutches analyzed, only 6% have multiple paternity. The distribution of pairwise relatedness among nesting localities (rookeries) was not random with elevated within‐rookery relatedness, and declining relatedness with geographic distance indicating some natal philopatry. Although there was no strong evidence that particular rookeries had lost allelic variation via drift, younger turtles had significantly lower levels of genetic variation than older turtles, suggesting some loss of genetic variation. At present there is no indication that levels of genetic variation are associated with measures of reproductive success such as clutch size, hatching success, and frequency of infertile eggs.     

Colony genetic structure in a facultatively eusocial hover wasp

The degree of genetic heterogeneity among the individuals inan animal society depends on the society's genetic structure.Genetic heterogeneity, in turn, means that group members willdiffer in their reproductive objectives and conflicts over reproductionmay arise. The resolution of these conflicts may be reflectedin the way that reproduction is partitioned between potentialreproductives. We used 5 microsatellite loci to investigategenetic structure and reproductive skew in 17 nests of the Malaysianhover wasp, Parischnogaster alternata. Parischnogaster alternatacolonies are small (1–10 females), and all adult colonymembers are capable of mating and producing offspring. We foundthat colonies tended to consist of closely related individualsand that at any one time the production of both female and maleoffspring was nearly always monopolized by a single dominantfemale, despite considerable variation between nests in parameterspredicted to affect skew. Subordinate females that remainedin their natal colonies obtained indirect fitness benefits byhelping to raise offspring to which they were related. Subordinatefemales also appeared to be positioned within an age-based queuefor inheritance of the dominant egg-laying position. We suggestthat the high skew in P. alternata may result from strong ecologicalconstraints on solitary nesting, high relatedness, and a relativelyhigh probability that subordinates will eventually inherit theposition of dominance.     

Simultaneous spawning by female stream goby Rhinogobius sp. and the association with brood cannibalism by nesting males

A laboratory experiment was conducted by varying the undersurface area of nesting substratum and the number of females in an experimental tank to elucidate the determinants of the mating pattern in the stream goby, Rhinogobius sp. cross‐band type. Males with larger nests tended to attract two or more females to their nest in a tank. Moreover, males spawned simultaneously with multiple females and entire brood cannibalism by males was rarely observed under a female‐biased sex ratio. When males spawned with a single female with low fecundity, however, entire brood cannibalism occurred at a high frequency, suggesting that a male guarding a nest with fewer eggs consumes the brood. Therefore, spawning behaviour of females that leads to a large egg mass would decrease the risk of entire brood cannibalism. In this species, simultaneous spawning by multiple females in a nest serves as a female counter‐measure against entire brood cannibalism. These results suggest that a conflict of interest between the sexes through brood cannibalism is a major determinant of simultaneous spawning.     

Reproductive skew, costs, and benefits of cooperative breeding in female wood mice (Apodemus sylvaticus)

Two current models seek to explain reproduction of subordinatesin social groups: incentives given by dominants for peacefullyremaining in the group (reproductive skew model) or incompletecontrol by dominants. These models make different predictionsconcerning genetic relatedness between individuals for thedistribution of reproduction and the stability of cooperativebreeding associations. To test these models and to furtherexplore the relationships between reproductive skew, geneticrelatedness, and investment of each participant, we performedbehavioral observations of female wood mice (Apodemus sylvaticus)raising pups communally. Our results do not support previousmodels. Differences in lifetime reproductive success were significantlygreater within mother—daughter pairs than within pairsof sisters or unrelated females. Subordinate females of eitherbreeding unit did not differ in their direct reproduction.Calculations of inclusive fitness based on our results leadto the following predictions: (1) Communal nests should occuronly when ecological circumstances prevent solitary breeding.(2) Subordinate females gain the highest inclusive fitnessjoining their mothers; they also show the highest nursing investment.(3) Mothers should accept daughters, who have no opportunityfor solitary breeding. (4) Dominant sisters and unrelated femalesshould reject subordinate females because cooperative breedingreduces their reproductive success. However, breeding unitsof dominant sisters and unrelated females nevertheless occurand can be explained by our finding that such females significantlyreduce nursing time, which may help them save energy for futurebreeding cycles. Our data demonstrate that both genetic relatednessand investment skew are important in the complex evolutionof reproductive skew in cooperative breeding.     

Kin recognition and incest avoidance in a group-living insect

Mate choice theories predict that animals evolved strategiesto mate with optimally genetically dissimilar partners, providingfitness benefits. In group-living species, when adults do notdisperse, assessment of relatedness between conspecifics canbe a key factor for choosing mates. Here, we report for thefirst time, kin recognition abilities and their implicationin mate choice in the gregarious cockroach, Blattella germanica(L.). Binary choice tests showed that females mated preferentiallywith nonsibling rather than with sibling males, thus avoidingincest. In addition, inbreeding induced an important decreaseof their reproductive success. Contrary to what could be expectedwhen females had the choice between a nonsibling strain memberand a nonstrain member, they did not avoid mating with distantlyrelated nonstrain members, and extreme outbreeding induced anincrease of their reproductive success. Furthermore, our matechoice experiments disentangled the influences of familiarityfrom those of relatedness and evidenced that kin discriminationwas based on genetic cues independently of familiarity. Phenotypematching was a plausible mechanism for kin recognition. Contraryto many insect species, body size was not a salient criterionfor mate choice and had no consequences on reproductive success.     

Safer sex with feeding females: sexual conflict in a cannibalistic spider

Mating strategies are to a large degree shaped by conflictsbetween the sexes, causing a rapid antagonistic coevolutionof traits involved in reproduction. The view that sexual cannibalismrepresents a form of sexual conflict leads to the predictionof male traits that facilitate escape from cannibalistic females.A variety of traits have been suggested to serve this functionin spiders, where sexual cannibalism is comparatively common.Empirical evidence, however, is virtually absent. Here we showexperimentally that opportunistic mating with feeding females,which has been reported from several species of orb-weavingspiders, greatly reduces the risk of cannibalism and injuryfor males in the spider Nephila fenestrata. This has directconsequences for a male's fertilization success because survivingmales can reduce the female's remating probability by guardingher against rivals. Although copulation with previously matedfemales sometimes appears to be mechanically impossible, secondmales that do copulate can expect to fertilize on average 64%of a female's eggs. Our results support the view that opportunisticmating may have evolved as a male tactic in a context of sexualconflict over sexual cannibalism.     

Clutch size variation in the Nazca booby: a test of the egg quality hypothesis

In obligately siblicidal bird species, aggressive behavior bya dominant chick results in a fixed brood size of one, yetthese species usually show clutch size variation between individuals.Simmons proposed that variation in clutch size in obligatelysiblicidal species is related to a trade-off between egg qualityand egg quantity: some individuals produce a single highly hatchable egg, while others produce two small, lower qualityeggs. We tested the egg quality hypothesis as an explanationfor observed clutch size variation in the Nazca booby (Sulagranti), an obligately siblicidal seabird. We tested the assumptionthat egg volume is positively correlated with hatchabilityand the prediction that eggs from one-egg clutches are largerthan eggs from two-egg clutches. We did not find a positive relationship between egg volume and hatchability in this species.Eggs from two-egg clutches were either equivalent in volumeor larger than eggs from one-egg clutches. Thus, the egg qualityhypothesis was rejected as an explanation for clutch size variationin the Nazca booby. Instead, two-egg clutches appear to befavored because of the insurance value of the second-laid egg,while one-egg clutches result from food limitation.     

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.