Filial cannibalism in a nest-guarding fish: females prefer to spawn in nests with few eggs over many

In fish, fecundity correlates with female body size and egg-tending males often eat small broods. Therefore, small females may prefer to spawn in nests that already contain many eggs, to ensure the brood is as large as possible. In contrast, large females may prefer nests with few eggs, if high egg number or density has a negative effect on egg survival, or if there are drawbacks of spawning last in a nest. To test the hypothesis that female body size affects nest (and male mate) choice, using the sand goby (Pomatoschistus minutus), we allowed small and large females to choose between two males that were matched in size — one guarding a small clutch and the other a large clutch, respectively. We recorded where females spawned (measure of female preference), the combined brood size, male courtship, egg care and nest building. We also quantified the effect of brood size and egg density on egg survival in a separate data set. Although the combined broods did not exceed the small brood sizes that are at risk of being eaten, both small and large females preferred to spawn in nests with smaller clutch sizes. This preference could not be explained by more courtship or male parental effort, nor by reduced survival of larger or denser broods. Instead, our result might be explained by females avoiding the danger of cannibalism of young eggs by males or the risk of reduced egg health associated with being near the nest periphery.     

The effects of nesting stage,sex, and type of predator on parental defense by killdeer (Charadrius vociferous): testing models of avian parental defense

Summary Two models predicting the temporal patterns of parental investment in offspring defense over the nesting cycle were tested. The first is based on offspring age, the other on the vulnerability of offspring to predation. Both models make very similar predictions for altricial species after eggs have hatched, i.e., increases in intensity of parental defense until fledging. For precocial species, however, the post-hatching predictions of each model are different: the offspring age model predicts a continued increase in defense intensity, while the vulnerability model predicts a decline. I examined the temporal patterns of parental defense of a precocial shorebird, the killdeer (Charadrius vociferus), and determined which model was supported. Killdeer responses to human and natural predators were observed. Killdeer were less willing to leave the nest, responded most intensely, and displayed closest to a potential predator around hatching. Defense intensity increased from early to late incubation as predicted by the offspring age model. However, after hatching killdeer parental defense declined for both males and females, thus supporting the vulnerability model for this stage. Males and females responded significantly differently to all types of predators. Males took greater risks, remained on the nest longer, defended offspring more intensely, and displayed closer to the predator than females at the approach of a potential predator. Responses to natural predators depended on the type of predator and the approach made by the predator; a greater range of defense behavior was used for predators approaching on the ground compared to aerial predators. In general, killdeer responses to humans were more intense and less variable than their responses to natural predators. This was most likely because the human intruder approached nests and chicks more directly and closer than natural predators.     

Why do males tolerate sneakers? Tests with the European bitterling, Rhodeus sericeus

In most species, males attack other males that attempt to gain fertilizations through sneak copulations. Here we report on a system where dominant males show a low level of aggression against sneakers at the initial stages of territory establishment. Females of the European bitterling, Rhodeus sericeus, lay their eggs in living mussels and males fertilize the eggs by releasing sperm over the mussels both before and after egg laying. When we allowed males to court females to a mussel containing no eggs at different male densities - one, two, four, or six males - the dominant male showed a low level of aggression against other males that released sperm. The dominant male became aggressive toward the other males only after eggs had been laid. This unusual pattern could be due to either some benefit of accepting sneakers or a high cost of aggression. We found support for both possibilities. The presence of several males decreased the time until a female spawned, whereas increased aggression by the dominant male against other males during a second female presentation, when the male was more territorial, interrupted courtship and increased the time until spawning. Females appeared to be attracted by both the presence of several males around a mussel and increased courtship under male competition. The bitterling mating system possibly differs from that of other species due to lack of investment in nest building and parental care, and high costs of defending the spawning site against sneakers.     

No reason to sneak: why males of all sizes can breed in the hole-nesting blenny, Aidablennius sphinx

Most blennioid fishes show a resource-based, promiscuous mating system with alternative 'sneaking' mating tactics. The hole-nesting species, Aidablennius sphinx, however, appears to be an exception; small males did not mimic females, most had their own nest and not once in 20 h of spawning observations did a second male enter a nest. In this field study, we ask if sneaker tactics are constrained and what factors favour independent nesting by small males. Larger males received more eggs sooner and attracted more females than smaller males. Larger males may be preferred because they released more sperm per ejaculation and fanned their eggs more often. In the long-term, however, larger males did not spawn more often than smaller males since: (1) the nests of large males filled up with eggs sooner than the nests of small males; (2) large males refused females more often than small males; and (3) a highly biased ratio of females to preferred large males meant many females had no option but to spawn with small males. Thus, small males nest independently, despite potential for female mimicry tactics (multiple females spawned simultaneously) and simultaneous parasitic spawning (males exited the nest during spawning)     

Why are larger convict cichlid (Cichlasoma nigrofasciatum) fry sometimes adopted into broods of smaller fry?

Females of Elasmucha grisea defend their eggs and small nymphs against invertebrate predators. Females sometimes guard their clutches side by side on the same birch leaf. We studied benefits of this joint guarding both in the field and in the laboratory. We found that adjacent females had significantly larger clutches than solitary females. In the laboratory, we studied the effectiveness of joint versus single defence against ant (Formica uralensis) predators. We established female pairs from initially singly guarding females by cutting off pieces of leaves with egg clutches and pasting them beside another female guarding her clutch. In the control group the females with their clutches were similarly cut off but these clutches were placed on another leaf without any female. The birch twigs where females guarded their clutches were placed in cages in close proximity to laboratory ant nests. In the experimental treatment, two females guarded their clutches together and at the same nest there was another birch twig without a female. In the control treatment two twigs with one female on each were placed close to another ant nest. Two females defended their clutches significantly more successfully, losing fewer eggs than did the single females. This primitive form of female sociality in parent bugs resembles colonial nesting in birds, where communal defence is also important. However, to our knowledge this is the first experiment where the benefit of joint guarding has been tested directly by manipulating the size of the breeding group rather than by measuring the risk of predation in groups of different size.     

Demographic plasticity in tropical reef fishes

We use age-based analyses to demonstrate consistent differences in growth, mortality, and longevity of coral reef fishes from similar habitats (exposed reef crests) 20 km apart. On outer-shelf reef crests of the northern Great Barrier Reef (GBR), size in four taxa of reef fishes (Chlorurus sordidus, Scarus frenatus, and S. niger and the acanthurid Acanthurus lineatus) was systematically and significantly smaller when compared with the same taxa on adjacent mid-shelf reef crests. Differences in size could be attributed to differences in growth between habitats (shelf positions). On outer reef crests the species examined had consistently lower size at age profiles and also reduced life spans compared with populations from mid-shelf reefs. To confirm this relationship, two of the most abundant species (C. sordidus and S. frenatus) were selected for more detailed spatial analysis of demographic patterns. Sampling adults of both taxa from reef crests on three mid- and three outer-shelf reefs revealed that most of the variation in growth was explained by shelf position, although C. sordidus also displayed differences in growth among mid-shelf reefs. We conclude that differences in body sizes across the continental shelf of the northern GBR are primarily determined by these trends in growth. Strong spatial patterns also existed in the mean ages of populations and longevity estimates for C. sordidus and S. frenatus between shelf positions. Both species on outer-shelf reefs displayed less variable cohort sizes, significantly reduced mean ages, and foreshortened longevity compared with populations on mid-shelf reefs. Furthermore, differences in these parameters were rare among replicate reefs within mid- and outer-continental-shelf positions. Age-based catch curves suggested that rates of S. frenatus natural mortality on the outer shelf were nearly twice as high as on the mid shelf. Visual surveys indicated that total scarid densities on outer-shelf reef crests are on average fourfold higher than for equivalent mid-shelf habitats. This fact, coupled with significantly reduced growth rates, reduced mean ages, and increased mortality rates, suggests that density-dependent processes may be responsible for observed differences among localities.     

The presence of eggs in the nest and female choice in common freshwater gobies (Rhinogobius brunneus)

In many species of fishes with paternal care, females prefer to spawn with males who are already guarding eggs. We studied the effect of egg presence on female mate choice in common freshwater gobies, Rhinogobius brunneus sp. OR. In our tests, females did not prefer males with eggs, suggesting egg presence per se may not act as a cue to attract females. We also examined the effects of brood size on paternal care and offspring survival to look for possible benefits females could obtain when choosing males with eggs. Both fanning by egg-guarding males and egg survival increased with brood size. The presence of neighboring males did have a significantly negative effect on males' parental activity, which subsequently results in a lower level of egg survival than in solitary egg-guarding males. This result provides a partial explanation for the result of eggs not attracting females to mate.     

Maternal nest defense reduces egg cannibalism by conspecific females in the maritime earwig <Emphasis Type="Italic">Anisolabis maritima</Emphasis>

Nest predation imposes a major cost to reproductive females, who should therefore take measures to avoid encounters with predators. However, when predators are conspecifics, avoidance can be more difficult and may be a consequence of social or aggregative behaviors. In this study, we measured the consequences of conspecific egg cannibalism on hatching success in the maritime earwig (Anisolabis maritima), which occasionally form aggregations. We hypothesized that conspecific egg cannibalism is a byproduct of aggregation, and that cannibalism rates would increase with aggregation density; however, our results do not support this. We combined field data with a lab experiment to test the effectiveness of maternal nest defense in protecting nests from a conspecific. Nests with a guard had higher hatching success and lower rates of cannibalism than unattended nests in the presence of a conspecific. We also measured body and forcep size to see whether the outcome of contests was determined by relative size. Female guards who were larger relative to the invading conspecific maintained their nest and had higher hatching success than females who were relatively smaller, suggesting that the maritime earwig is under directional selection for larger body and/or forcep size.     

Does egg colour affect predation rate on open passerine nests?

The breeding success of many passerines is strongly reduced by egg predation. The adaptive significance of egg crypsis in open nesters is often taken for granted, but visually searching predators may first detect the nest or adult bird and not the eggs. Götmark predicted that selection should favour egg crypsis in the absence of conspicuous nests, whereas birds with conspicuous nests should have non-cryptic eggs. I compared the effect of egg colour treatment (white, blue, brown-spotted) on nest survival (1) among species characterized by different egg coloration, nest size and nest placement, and (2) between relatively well and poorly concealed nests within species. I used artificial nests (n=1,296) and eggs mimicking (except in egg colour) those of the yellowhammer (Emberiza citrinella), blackcap (Sylvia atricapilla) and song thrush (Turdus philomelos). Concurrently, I monitored survival of real nests (n=1,106). Nest survival differed among species, increased with nest concealment and throughout the breeding season, but was not significantly related to egg colour in any species. Nevertheless, the data for the yellowhammer suggest a trend in survival rates across the colour treatments. Brown eggs survived better than white eggs by 11% and 4% in 2 years, but this study had insufficient power to detect effects of this size. The results thus suggest that egg coloration in the song thrush and blackcap (shrub nesters) may be a neutral trait with regard to nest predation, whereas egg crypsis may be an anti-predation feature for the yellowhammer (ground/near-ground nester). The role of predation in the evolution of eggshell colour may vary not only between cavity and open nesters, but also across nest sites within the latter group.     

Effects of size and nuptial gifts on butterfly reproduction: can females compensate for a smaller size through male-derived nutrients?

In butterflies and other insects, fecundity generally increases with female adult weight. Hence, most butterflies are essentially "capital breeders", because nutrients acquired during the larval stage are stored and subsequently used for egg production during the adult stage. However, in some species, males transfer a large nutritious ejaculate to the female at mating. These females can partly be characterized as "income breeders", and female mass can potentially be decoupled from fecundity to some extent. In the gift-giving green-veined white butterfly Pieris napi, it has been shown that female fecundity and longevity increase with number of matings and also that females mature at smaller size under poor food conditions compared to males. So it has been suggested that females can compensate for their smaller size through nuptial feeding. Here we test this hypothesis in P. napi by assessing female fecundity and longevity in relation to female mass and polyandry. The results showed no support for the hypothesis. Smaller females were not capable of increasing their mating rate to compensate for a low weight at eclosion. Instead, larger females remated sooner. Also, smaller females suffered from both a reduced daily and total fecundity compared to larger females and this decrease in fecundity was independent of female mating status, i.e. females allowed to mate only once and multiply mated females suffered to the same extent from their smaller size.     

Brood defense and filial cannibalism in the spottail darter (<Emphasis Type="Italic">Etheostoma squamiceps)</Emphasis>: the effects of parental status and prior experience

Male spottail darters (Etheostoma squamiceps) defend nest sites in which females deposit eggs over the course of several weeks. In laboratory experiments, I tested three predictions of the hypothesis that the presence of eggs increases the value of a nest site to male spottail darters: (1) males should preferentially defend nests with eggs over empty nests, and males that sired the eggs should (2) spend more time defending them and (3) consume less of their broods than should alloparent males. Parent males were tested using eggs they were guarding in the field; alloparents included males that were not defending eggs immediately prior to the experiment (inexperienced), and males that were defending eggs in the field when captured but were given eggs sired by a different male (experienced). In choice experiments in which males were offered nest sites differing only in the presence of eggs, males preferred to defend nest sites with eggs regardless of parental status. Parent males spent significantly more time defending eggs and consumed less of their broods than did inexperienced alloparent males. Experienced alloparent males were similar to parent males in brood defense but had levels of filial cannibalism similar to inexperienced males, suggesting that prior experience may influence brood defense but not egg consumption.     

Allopaternal care in the fathead minnow,Pimephales promelas: females prefer males with eggs

Summary Some species of fishes with exclusive male parental care exhibit the phenomenon of allopaternal care; that is, some males acquire and care for other males' eggs. We conducted a series of experiments to investigate the dynamics and evolution of allopaternal care in one such species, the fathead minnow (Pimephales promelas). In choosing a nest site, a newly reproductive male tended to take over the nest site of a parental male by evicting the resident male, rather than occupy a physically identical empty nest site. The new male generally cared for the old male's eggs, and in most cases, daily egg survival improved under the new male's care. When males were given a choice among unguarded nest sites, they preferred to occupy nest sites already containing eggs. When eggs were randomly assigned to nesting males, females preferred to spawn with males who had eggs in their nest sites. Thus, it appears that female preference for males with eggs led to the evolution of allopaternal care in the fathead minnow.     

Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).     

Histology of the gonads and observations on the social behavior of the Caribbean angelfish Holacanthus tricolor

Male and female specimens of Holacanthus tricolor were collected in May 1981 and September 1982 off the Island of St. Croix. U.S. Virgin Islands, and their gonads were examined histologically. All testes examined showed vasa deferentia which were characteristic of secondary males, suggesting protogynous sex change. A transitional individual supported this interpretation. Ovarian histology and size-specific egg counts revealed seasonal differences in spawning frequency and clutch size. Females appeared to spawn every day in May. The timing of final maturation and hydration of the oocytes indicated that spawning occurred in the late afternoon or early evening. Not all females spawned every day in September, and clutch sizes were smaller than those of May. Behavioral observations of H. tricolor in the field revealed that males defended harems of two to four smaller females. Within each harem, females of different sizes had overlapping home ranges, while the home ranges of females of similar size did not overlap. Besides size dimorphism between the sexes, H. tricolor exhibited a previously unreported sexual dichromatism.     

Potential latitudinal variation in egg size and number of a geographically widespread reef fish,revealed by common-environment experiments

Variation in maternal reproductive traits was examined in field and reared populations of a geographically widespread reef fish, Pomacentrus coelestis (Pomacentridae), drawn from three different latitudes in Japan. Size-specific clutch size and clutch weight of wild fish increased with increasing latitude. Conversely, latitudinal variation in egg size of wild fish was obscure in same-season comparisons, probably because of the temperature effect on egg size. Common-environment experiments conducted at three temperatures showed that egg size decreased with increasing temperature in all populations. In the experiments, egg size, clutch size and clutch weight differed among populations at all temperatures, showing clear latitudinal clines. Females from low latitude spawned larger eggs at every experimental temperature. Size-specific clutch size and weight were greater in females from high latitude. Thus, the northern fish had a larger reproductive output per spawning and a larger number of smaller eggs in a spawning. Such interpopualtion variation in this fish is likely to be partially genetically based, although environmental effects on the variation cannot be entirely ruled out. This study provides evidence of potential latitudinal variation in the egg size and number in a coastal fish, by common-environment experiments. The close correspondence between latitudes and these maternal reproductive traits may be a consequence of local adaptation.     

Food and predators affect egg production in song sparrows

Although the possibility that food and predators may interact in limiting avian populations has long been recognized, there have been few attempts to test this experimentally in the field. We conducted a manipulative food addition experiment on the demography of Song Sparrows (Melospiza melodia) across sites that varied in predator abundance, near Victoria, British Columbia, Canada, over three consecutive breeding seasons. We previously showed that food and predators had interactive effects on annual reproductive success (young fledged per female). Here, we report the effects on egg production. Our results show that food limits the total number of eggs laid over the breeding season ("total egg production") and that interactive food and predator effects, including food effects on nest predation, determine how those eggs are "parceled out" into different nests. Food addition alone significantly affected total egg production, and there was no significant interannual variability in this result. At the same time, both food and predators affected the two determinants of total egg production: "clutch number" (total number of clutches laid) and average clutch size. Both clutch number and size were affected by a food x predator x year interaction. Clutch number was lower at low-predator locations because there was less nest predation and thus less renesting. Food addition also significantly reduced nest predation, but there was significant interannual variation in this effect. This interannual variation was responsible for the food x predator x year interactions because the larger the effect of food on nest predation in a given year, the smaller was the effect of food on clutch number; and the smaller the effect of food on clutch number, the larger was the effect of food on clutch size. Potential predator and year effects on total egg production were thus cancelled out by an inverse relationship between clutch number and clutch size. We suggest that combined food and predator effects on demography could be the norm in both birds and mammals.     

Egg adoption can explain joint egg-laying in common eiders

Hypotheses regarding the evolution and maintenance of intraspecific nest parasitism were tested with data collected during a 3-year study of common eiders (Somateria mollissima) breeding near Churchill, Manitoba. The nest parasitism rate was highest (42.4% of nests) during the year with the highest nest density and the best environmental conditions, and lowest (20.2% of nests) in the year with the lowest nest density and the poorest environmental conditions. Over the nesting season, parasitic eggs were laid at the same time as normally laid eggs. Most parasitic eggs (>75%) were laid before the host female laid her third egg. The majority of the parasitic eggs were the first or second egg produced by the parasitic female. When a parasitic egg was laid before or on the same day as the host female initiated her clutch, the probability of her first egg being depredated before incubation was significantly lowered. First- and second-laid eggs suffered a high rate of predation probably because nesting females do not attend their clutch until their second or third egg is laid. Hypotheses that some females use intraspecific nest parasitism to parasitize the parental care of other females were inconsistent with these data. Egg adoption is a likely explanation for the prevalence of females incubating parasitic eggs in this population. Received: 30 September 1997 / Accepted after revision: 6 May 1998     

Nest-maintenance effort and health status in chinstrap penguins, Pygoscelis antarctica: the functional significance of stone-provisioning behaviour

Stone provisioning is a nest-maintenance activity accomplished by pygoscelid penguins after reliefs during the incubation/brooding period. The functional significance of this behaviour has been mainly explained as a parental strategy preventing nest flooding under detrimental weather conditions. In addition, and in the light of recent studies, this behaviour could also fit into the sexual selection process. In this study, we tested the first idea, that is, whether stone provisioning is a nest-maintenance behaviour to increase egg/nestling survival by lowering the risk of nest flooding, and can thus be considered a form of parental care. Additionally, we investigated if the effort invested by parents in nest maintenance is constrained by physiologically limiting resources. The effort of stone collection and the perceived risk of nest flooding were experimentally manipulated during the incubation and early brooding phases in a chinstrap penguin, Pygoscelis antarctica, colony. Three groups of nests were established. After weighing, control nests were left unmanipulated. In a second group of nests (reduced group), only one-half of the initial weight of stones was returned to the nests. In a third group of nests (snow-added group), we both reduced nest weight by a half and added snow outside the nest bowl over 6 consecutive days. Ten days after manipulation, the difference in nest weight between initial and final conditions was significantly related to treatment: penguins increased stone provisioning in the reduced group (44% of half-reduced nests), but drastically more in the reduced and snow-added group (123% of half-reduced nests), while the weight of control nests was unchanged compared to premanipulation conditions. The intensity of stone provisioning was affected by nest date, peaking about hatching time and shortly after, and declining with advancing chick age. These results suggest that stone provisioning is a mechanism that has evolved to prevent egg or chick mortality by nest flooding. The haematocrit, but not leukocyte numbers as expressed by the buffy coat layer, varied with the experimental conditions. Penguins investing more time in nest maintenance had a lower haematocrit, suggesting a physiological trade-off probably mediated by competition between the time devoted to nest maintenance versus foraging activities. The amount of stones collected and the haematocrit were positively related to the number of neighbour nests, so those individuals surrounded by more nests seemed to obtain benefits in the availability of nest material and energy savings. This study indicates that stone-provisioning behaviour is a nest-maintenance activity evolved to improve thermal nest characteristics potentially increasing offspring survival, and competing in time and energy with other reproductive activities. Stone provisioning in penguins should therefore be regarded as a form of parental care and an important part of individual reproductive effort in species breeding in harsh environments. Furthermore, nest size and nest-maintenance effort should be considered reproductive traits indicative of parental quality and thus could also be involved in the post-mating sexual selection process.     

Egg mimicry and allopaternal care: two mate-attracting tactics by which nesting striped darter (Etheostoma virgatum) males enhance reproductive success

In a variety of fish species with paternal care of offspring, females prefer to spawn in nests that already contain eggs. This female preference has been hypothesized to explain egg thievery in male sticklebacks, allopaternal care of eggs in minnows, and the evolution of egg-mimicking body features in male cichlids and darters. Here we employ microsatellite-based parentage analyses to evaluate the reproductive success of striped darter (Etheostoma virgatum) males that appear to utilize two of these functionally related tactics to entice females to spawn in their nests. In an isolated population (Clear Creek, Ky.), we observed that breeding males develop conspicuous white spots on their pectoral fins. If these spots are egg mimics, as we suspect, then this represents the fourth independent evolutionary origin of egg mimicry documented to date in darters, the first based on pigmentation (as opposed to physical structures), and the first in which the egg mimics vary greatly in number among males. From direct counts of microsatellite genotypes in clutches of embryos, at least 3.8 females contributed to the progeny within a typical nest, and females tended to spawn preferentially with males that were larger and displayed more egg-mimic spots. In another population (Hurricane Creek, Tenn.) without egg mimics, the multi-locus genetic data document that allopaternal care is common, especially among the smallest males who sometimes tend nests containing their own as well as an earlier sire's offspring. Thus, these foster males had adopted egg-containing nests and then successfully spawned with subsequent females. Overall, the genetic data on paternity and maternity, in conjunction with field observations, suggest that egg mimicry and allopaternal care are two mate-attracting reproductive tactics employed by striped darter males to exploit female preferences for spawning in nests with 'eggs'. Electronic Publication     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.