Female cannibalism and male courtship tactics in threespine sticklebacks

Summary Female threespine sticklebacks (Gasterosteus aculeatus) frequently raid male nests and eat all the eggs therein. We tested the hypothesis of Vickery et al. (1988) that females prefer to raid nests containing large numbers of eggs than ones with smaller numbers of eggs. This hypothesis is based on the finding that females spawning in nests containing many eggs will have reduced hatching success because of egg crowding. By consuming the male's eggs and forcing him to rebuild his nest, raiding females might obtain a new opportunity to spawn under better conditions. Our results were consistent with the first prediction of this hypothesis that females were more likely to spawn in nests containing fewer eggs than in nests with many eggs. However, this may be the result of males becoming less receptive to females as the number of eggs in their nests increases. Prediction 2 was that females should raid those nests containing the most eggs. Contrary to this prediction, males defending only one clutch were as likely to have their nests raided by groups of females as males defending several clutches of eggs. Female cannibalism is therefore unlikely to have evolved as a means of gaining access to a male defending a small number of eggs. We also examined the tactics used by males to counter female raids. Most raids occur when the male is courting, and nests are more vulnerable to shoals of females than to single females. Therefore, we hypothesized that males with eggs preferentially court a single female rather than large groups of females, and that males without eggs court both groups indiscriminately. We also predicted that males restrict the number of females they mate with when risk of having their nest raided is high. Our results indicate that: (1) both males with eggs and those without eggs minimize the risk of female cannibalism by courting solitary females rather than groups of females and (2) males limit the number of females that lay eggs in their nest when several potentially raiding females are present. Offprint requests to: G.J. FitzGerald     

Patterns of Predation Risk and Survival of Bird Nests in a Chilean Agricultural Landscape

Abstract: We used experimental nests baited with California Quail (  Callipepla californica ) eggs or clay eggs to examine relative risks of nest predation in an agricultural landscape and in two large forest preserves in a south-temperate rainforest in Chile. The most common predators, as identified by marks on clay eggs, were a caracara (   Milvago chimango ), a blackbird ( Curaeus curaeus ), and rodents. Nest losses from predation were similar in large and small forest patches and lower in patches than in extensive forest. In general, predation risk was higher (and nest survival therefore lower) on forest edges than in forest interior, in short-grass pasture than in tall-grass pasture, in narrow corridors than in wide corridors, and on visible nests than on concealed nests. High predation risks in pasture habitat tended to increase the risk of nest predation in adjacent forest edges. For open-cup nesters, the risk of nest predation was relatively high in the present agricultural landscape, indicating that much of the available wooded habitat (  forest edges, narrow corridors) offers poor nesting habitat, although it may be suitable for foraging and traveling. The numerous bird-plant mutualisms in this landscape may be at risk if nesting success of the principal mutualists is consistently low.     

Tactics of parasitic American coots: host choice and the pattern of egg dispersion among host nests

Summary I examined the tactics adopted by a conspecific brood parasite, the American coot (Fulica americana), and the degree to which these tactics reflect sources of mortality for parasitic eggs. Only 8% of parasitic eggs produced independent offspring, compared to a 35% success rate for non-parasitic eggs, and most mortality was due to egg-rejection by hosts or the consequences of laying eggs too late in the host's nesting cycle. Parasites usually laid parasitically before initiating their own nests and usually parasitized immediate neighbours. Parasites did not remove host eggs before laying their own egg, and egg disappearance in general was not more common at parasitized nests. I found no evidence for non-random host choice, either on the basis of stage of the host's nesting cycle or the host's brood size. The absence of adaptive host choice is likely a consequence of the fact that, due to host limitation, only a small proportion of parasites had meaningful variation among potential hosts to choose from. The pattern of egg dispersion among host nests by individual parasites appears to be a compromise between constraints imposed by host limitation and the increased success obtained from spreading eggs among nests. Most females laying fewer than five parasitic eggs laid them in a single host nest while females laying five or more eggs normally parasitized two or more hosts. An examination of egg rejection and survival rates showed that parasites would maximize success by laying a single egg per host nest, and the pattern of laying several eggs per host nest is likely a consequence of host limitation. However, no egg that was the fifth laid, or later, parasitic egg in a host nest was ever successful and this probably explains why most females laying five or more eggs parasitized more than one host.     

Self-monitoring of reproductive success: nest switching in the redlip blenny (Pisces: Blenniidae)

Summary In the redlip blenny Ophioblennius atlanticus, commonly found on Caribbean reefs, eggs are laid in a male's nest and are guarded by the male until they hatch as planktonic larvae. Male reproductive success, defined as the number of egg batches hatched per reproductive period, was positively correlated with male size and with the inner surface area of the male's nest. Male blennies often switch nests between reproductive periods. Most nest switching (83%) occurred following periods of low reproductive success. Moreover, a male having low reproductive success was more likely to switch if the inner surface area of his nest was small. Most nest switching (72–80%) was to a nest of larger surface area than the previous nest. In 72% of all switches, males had higher reproductive success in the period following the switch than in the one preceding it. The results suggest that male redlip blennies monitor their current reproductive success and the characteristics of their current nests, switch to better nests if their reproductive success is low, and typically benefit from the decision to do so. These capabilities have seldom been demonstrated in any phylum.     

Female spawning patterns and male mating success in the sand goby Pomatoschistus minutus

In June 1989 in a study conducted near Träminne Zoological Station, Finland (60° N 23° E) I investigated whether or not male mating success could be explained by female choice for male size in sand gobies (Pomatoschistus minutus). Male mating success was constrained by nest size and increased markedly with increasing nest size. I also found a negative correlation between the length of spawning females and the fullness of the nest. As large females lay more eggs, they also need to find a nest with more space available for the eggs. The size of males without eggs was the same across nest size, whereas the size of males with eggs significantly increased with increasing nest size. This is interpreted as female discrimination against males as mates in nests that are often contested. There was no correlation between a male's size and his mating success when males with no eggs in their nests were excluded. A male removal experiment, however, showed that, in a specific nest, when male size increases so does mating success, whereas, if male size decreases, mating success also decreases. It is concluded that in the sand goby females prefer to mate with larger males, especially when the male possesses a high-quality nest that he most probably will have to defend against other males.     

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.     

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.     

Seasonal fluctuations in sand temperature: effects on the incubation period and mortality of loggerhead sea turtle (<Emphasis Type="Italic">Caretta</Emphasis><Emphasis Type="Italic">caretta</Emphasis>) pre-emergent hatchlings in Minabe,Japan

Incubation period, hatching success, and emergence percentage in loggerhead (Caretta caretta) nests were quantified during the 1993 and 1995 nesting seasons and following incubation seasons in Minabe, Wakayama, Japan. Sand and nest temperatures were also monitored. Over the seasons, daily mean sand temperature at nest depth fluctuated between 18.0°C and 33.3°C, with a steep increase in the second week of July and a peak in late August. Temperatures inside the nest chambers were a few degrees above those of the surrounding sand at the end of incubation. The incubation period ranged from 46 to 82 days. A significant negative correlation was found between mean sand temperature and incubation period. The relationship conformed to the day-degree concept. There was no significant seasonal trend in hatching success, but many pre-emergent hatchlings were found dead in most of the clutches during the warmest part of the season. Emergence percentage was correlated with mean sand temperature calculated for 4 days before emergence, suggesting that mortality may be due to heat. This heat-related mortality is considered to be a common phenomenon at our study site, because the peak in emergences coincides with the peak in high temperatures. These temperature effects on hatchling mortality must be taken into account in estimates of hatchling sex ratios. Because sand temperatures already exceed the optimal thermal range for incubation, this population is vulnerable to even small temperature increases resulting from global warming.     

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.     

Intraspecific nest parasitism in the swallow Hirundo rustica: the importance of neighbors

Summary Intraspecific nest parasitism in the swallow Hirundo rustica involves several parasite and anti-parasite tactics. Neighboring swallows breed asynchronously, perhaps because neighboring nests with overlapping egg-laying periods have higher frequency of intraspecific nest parasitims than neighboring nests with no such overlap. Since nest guarding prevents nest parasitism, the effect of nest guarding was removed by putting up old swallow nests. Such experimental nests were parasitized by swallows more often when the nests contained eggs, than when empty, and when the nests were far away from, rather than close to, an active neighboring nest. Close experimental nests with eggs were, however, parasitized less frequently if asynchronous (non-overlapping egg-laying periods), rather than synchronous, to active neighboring nests. Aggressive anti-parasite behavior of nest owners was determined from responses towards female intruders. Responses were more often aggressive before and especially during egg laying than during incubation. Nest owners also behaved aggressively more often when the intruder approached close to as compared to at some distance from their nest. Focal pairs behaved aggressively more often towards closely approaching intruders during the incubation period than towards distantly approaching intruders during the egg-laying period. Swallows reduce the frequency of intraspecific nest parasitism by nesting asynchronously with and close to a neighboring nest. Aggression by neighbors may reduce the success of potential parasitic swallows.     

Wave action: the environmental trigger for hatching in the California grunion Leuresthes tenuis (Teleostei: Atherinopsidae)

 California grunion Leuresthes tenuis (Teleostei: Atherinopsidae) emerge from the ocean to spawn on beaches of southern California. Grunion eggs do not hatch at a set developmental age, but remain in the sand until turbulent surf at high tide washes them out to sea. In previously studied fishes and amphibians that lay eggs terrestrially, low oxygen is the trigger for hatching in water, and high oxygen tensions inhibit hatching. For the grunion, however, eggs placed in air, seawater and oil did not hatch at any at oxygen tension until they were agitated in fluid. Following agitation in seawater, all eggs hatched within several minutes. Grunion eggs in normoxic or hyperoxic water hatched significantly faster than eggs agitated at the lowest oxygen tensions. Mechanical agitation, not hypoxia, is the environmental trigger for hatching in California grunion eggs. Received: 15 May 1999 / Accepted: 5 April 2000     

Aggregation and dispersal based on social cues as a nest-site selection strategy in a resource-defence polygynandry mating system

Many animals must choose a nest site in order to reproduce. However, it is unclear how nest-site selection strategies vary across different mating systems. We must therefore explore nest-site selection strategies in a range of mating systems, including the interaction between resource-defence polygyny and polyandry (i.e. polygynandry). In this study, we imposed a re-settlement event in the terrestrial toadlet Pseudophryne bibronii and measured the influence of the spatial position of each male’s nest site with respect to rival males on the likelihood that it would be abandoned or receive eggs. We captured every calling male in a population, measured their breeding success and released them back into the breeding area. We then recorded the establishment and abandonment of nest sites by males over 26 consecutive nights. Spatial positioning did not have any significant effects on male-breeding success, supporting claims that females show less discrimination between nest sites when they are polyandrous and spread their eggs amongst multiple male nests. However, we found that males consistently selected nest sites according to a site’s spatial position, which suggests that fitness benefits unrelated to male breeding success (e.g. reduced mortality risk) might influence male nesting decisions. Overall, our study provides new evidence that the mating system adopted by a population can influence the cues that individuals respond to when selecting nest sites.     

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     

Variation in nesting patterns affecting nest temperatures in two populations of painted turtles (<Emphasis Type="Italic">Chrysemys picta</Emphasis>) with temperature-dependent sex determination

Mechanisms maintaining sex ratios in populations with temperature-dependent sex determination (TSD) remain elusive. Although geographic variation in embryonic sex determination (i.e., pivotal temperature) has been widely investigated in reptiles exhibiting TSD, no previous studies have directly addressed geographic variation in maternal behavior affecting nest thermal conditions. I evaluated patterns of nest-site selection and its effects on thermal and hydric nest conditions for a population of painted turtles ( Chrysemys picta bellii) exhibiting TSD in New Mexico. These results are compared to data collected from a well-studied, conspecific population experiencing relatively cooler climatic conditions in Illinois. Since canopy vegetation cover reduces nest temperatures in Illinois, I expected females in New Mexico to nest under high amounts of canopy vegetation cover. However, females from New Mexico placed nests under significantly less canopy vegetation cover, but closer to standing water, than did females from Illinois. Experimental nests in New Mexico demonstrated that increased canopy vegetation cover and soil moisture reduced nest temperatures. By nesting close to standing water rather than under canopy vegetation cover, females in New Mexico nested in habitats more closely associated with maximizing moisture around nests rather than reducing nest temperatures through shading. Mean July nest temperatures were similar for both populations. Since nest hydric conditions affect hatching success and hatchling size in C. picta, nesting patterns in New Mexico may primarily reflect selection for microhabitats affecting offspring survivorship or size.     

Does the great spotted cuckoo choose magpie hosts according to their parenting ability?

When brood parasites are about to lay an egg, they have to decide which nest to parasitize. The best nest in which to lay will depend on the parenting ability of the host. We have studied selection of magpie (Pica pica) hosts by great spotted cuckoos (Clamator glandarius). Great spotted cuckoos preferentially parasitize large host nests. Nest volume in magpies is a good indicator of territory quality, since there is a negative relationship between magpie nest size and breeding date, and timing of breeding in magpies is known to be positively related to territory quality. Moreover, magpies occupying high-quality territories have high breeding success. Therefore, nest size is positively related to the quality of magpies. Parasitized magpie nests were of greater volume than the nearest neighbouring nest not parasitized by the great spotted cuckoo. In order to test whether the great spotted cuckoos might select high-quality magpie hosts, we manipulated pairs of parasitized and non-parasitized nests with identical laying dates and habitats, introducing into each of the nests the same number of parasitic and non-parasitic eggs. The number of fledglings reared (magpie plus great spotted cuckoo chicks) in naturally parasitized nests was higher than in experimentally parasitized nests. Thus, the probability of survival of the parasite chicks increased if cuckoo eggs were laid in the nests of high-quality hosts originally chosen by the parasite.     

Sexual dimorphism and male mating success in the tentacled blenny, <Emphasis Type="Italic">Parablennius tentacularis</Emphasis> (Teleostei: Blenniidae)

Although external sexually dimorphic traits are commonly found in males of combtooth blenny species, little is known about the benefit they can convey to male mating success. Indeed, while female preferences for large males have been demonstrated in some species, the possible role played by dimorphic ornaments has been neglected. We now report on the tentacled blenny, Parablennius tentacularis, a species where males are characterized by bulb glands on the anal fin and both sexes exhibit a dark spot on the dorsal fin and orbital tentacles. Males are territorial, make nests in empty bivalve shells, and provide solitary parental care for the eggs. Using morphometric analysis and field collected data on male and female external features, nest characteristics and number of eggs in the nests, we have assessed the development of dimorphic traits in both sexes and male mating success. The results reveal that orbital tentacles of males are more developed and more variable in size than those of females. Larger males exhibit longer orbital tentacles and larger anal glands but do not necessarily occupy larger nests. Male mating success is significantly correlated with the inner nest surface area and with orbital tentacle size but not with body size. These results provide support for a primary role of male ornaments in enhancing blenny male mating success and are discussed in the context of mate choice for direct and indirect benefits.     

Influence of emergence success on the annual reproductive output of leatherback turtles

Reproductive output of leatherback turtles (Dermochelys coriacea) is affected by the stochastic nature of emergence success. Average emergence success of nests at Playa Grande, Costa Rica was 0.38 ± 0.27. Incubation temperature affected development of leatherback turtle eggs and emergence of hatchlings from the nest. We found that high temperatures reduced hatching success and emergence rate and increased embryonic mortality both early and late during incubation at Playa Grande. There was a temporal effect on emergence success that resulted in more hatchlings being produced at the beginning of the season, because of higher emergence success, than toward the end. Likewise, production of hatchlings varied from year to year. The average annual reproductive output was 252 ± 141 hatchlings per female. The 2005–2006 nesting season had the highest emergence success and produced the greatest number of hatchlings per female compared to the 2004–2005 (+120%) and 2006–2007 (+41%) seasons. However, average clutch size (62 ± 10) and clutch frequency (9.45 ± 1.63), were not different among years. Turtles that had nested a high number of years exhibited greater clutch frequency and arrived earlier to nest than turtles that had nested in fewer numbers of years. Nesting when environmental conditions favor high developmental success and emergence rate may constitute an advantageous reproductive strategy.     

Intraspecific brood parasitism as a conditional reproductive tactic in the treehopper<Emphasis Type="Italic"> Publilia concava</Emphasis>

In species exhibiting egg guarding as well as communal egg laying, females may adopt the strategy of laying eggs in the nests of conspecifics and leaving without providing care (termed intraspecific brood parasitism). This study is the first to describe such a behavior in the insect Publilia concava (Hemiptera: Membracidae) through field studies that followed 849 marked females across 1,828 brood associations. While brood parasitism increased the total number of eggs in a host brood, it did not reduce the overall hatching success of host broods. Solitary females exhibited a range of guarding durations while parasitic females rarely remained to guard eggs. Females exhibiting the parasitic tactic increased their lifetime number of clutches without decreasing the number of solitary clutches that they were able to initiate. Estimates of egg number for these individual broods revealed that females adopting the parasitic tactic (in addition to solitary breeding) had higher lifetime fecundity relative to females that did not parasitize. In females that exhibited both tactics (solitary and parasitic), the parasitic tactic yielded a higher rate of oviposition. The major component of oviposition rate was the time to find hosts and this time decreased with increasing host availability across 222 replicate groups. Females exhibited a shift toward the parasitic tactic when host broods were more abundant (i.e. in larger groups and later in the season). However, the time to find hosts increased as the frequency of the parasitic tactic increased, suggesting that this tactic may be maintained through negative frequency dependence. The results of this study suggest that brood parasitism may be the preferred tactic, as part of a conditional strategy, when hosts are readily available with solitary breeding being the preferred tactic when hosts are in short supply.     

Incubation feeding in snow buntings: female manipulation or indirect male parental care?

Summary Male snow buntings regularly feed their mates on the nest during the incubation period. We removed males from 7 females at the start of incubation (Early Widows) and from 7 others when the eggs hatched (Late Widows) to experimentally assess the effects of incubation feeding on the behaviour of females and the reproductive success of both parents. Early Widows spent significantly more time off their nests than Late Widows and Controls. As a consequence, Early Widows had significantly longer incubation periods and a significantly higher proportion of them lost two or more eggs during development. There was no difference between Early and Late Widows in any index of reproductive success measured during the nestling period although significantly earlier brood reduction suggests that Early Widows were in poorer condition than Late Widows. Since both parents benefitted from incubation feeding by increased hatching success and shorter incubation periods, we conclude that this behaviour is an adaptive form of indirect parental care by males and is not the result of female manipulation.     

Behaviorally mediated spatial and temporal refuges from a cleptoparasite,Argochrysis armilla (Hymenoptera: Chrysididae), attacking a ground-nesting wasp,Ammophila dysmica (Hymenoptera: Sphecidae)

Summary It has been suggested that parasite pressure favors the evolution of sociality within the Hymenoptera. I analyzed the impact of a chrysidid nest parasite, Argochrysis armilla, on its host, the solitary ground-nesting wasp, Ammophila dysmica, to assess the role of parasitism in favoring two steps towards sociality: aggregated nesting and delayed nest provisioning. The foraging strategy of Argochrysis armilla involves discovering host nests during excavation, learning the locations of discovered nest, and returning to oviposit in nests during nest provisioning; I therefore assessed the influence of host behavior on (1) parasite discovery of nests and (2) parasite oviposition in nests. Significantly fewer parasites discovered host nests that were excavated during the early morning and late afternoon, due to partial asynchrony of host and parasite activity. Nests excavated in areas of low nest density were also less likely to be discovered; use of low density nest sites increased during periods of high parasite activity. Due to a rapidly decelerating rate of parasite recruitment to nests under excavation, the duration of nest digging had only a limited influence on nest discovery by parasites. The probability of parasite oviposition in a host nest was determined by the number of parasites discovering the nest during excavation and by the time between nest excavation and provisioning; delaying nest provisioning reduced the risk of parasite oviposition. Delayed provisionings primarily appeared to be a result of the stochastic process of hunting and prey encounter. The number of provisions placed in a nest (one vs two) had no effect on the probability of nest parasitism. Spatial patterns of parasitism were directly density dependent in 1984 and density independent in 1986. In this system parasite pressure acts against the formation of nesting aggregations and in favor of delayed nest provisioning. The dependence of these results on species-specific aspects of the parasite's foraging strategy and the host's defensive strategy suggests, however, that different parasite species may generate qualitatively different selection pressures, potentially contributing to the diversity of nesting behavior in the Hymenoptera.