Female defence polygyny in the bumblebee wolf,Philanthus bicinctus (Hymenoptera: Sphecidae)

Summary In the digger wasp Philanthus bicinctus the first adult males emerge about a week before the first females. The sex ratio of emerging wasps is near unity. Females appear to mate once and do so when they first start to nest, not when they first emerge.Males defend mating territories in a large area of female ground nests. Territoriality begins when females first start to nest and throughout the season coincides with the digging activities of females initiating rests. During the day territoriality occurs only when females are engaged in digging activities.Males compete for certain territory sites within the resting area; the most frequently occupied male territories are in areas where the density of nests is highest. About half of the males are excluded from nesting area territories. Some of these males defend areas among the flowers of snakeweed where females occasionally forage; this appears to be a loser tactic of males unable to obtain nesting area territories.Although there was no significant difference in the sizes of males on nesting area and snakeweed territories, within the nesting area larger males usually occupy territories with the higher nest densities.     

Nest defence by song sparrows: methodological and life history considerations

Summary Several aspects of nest defence behavior were investigated in song sparrows (Melospiza melodia) in eastern Ontario. Two independent tests were made of the hypothesis that the increase in nest defence observed through a nesting attempt is due to the birds becoming familiar with the nest threat, rather than because the nest contents increase in value to the parents. Neither test supported the hypothesis. As predicted by life history theory for species with age-independent mortality, males did not defend their nest more vigorously as they become older. Parents defended their nests less vigorously through the breeding season, contrary to the expected pattern of increased nest defence in response to declining renesting potential. This result may be attributable to a decline in offspring value through the breeding season. Nest defence behavior of mated individuals was positively correlated, independent of factors such as offspring age, renesting potential and brood size. From this result it is proposed that a source of variation in nest defence behavior may be individuals basing their own response on their mate's response in a positive feedback fashion. Males defended nests less vigorously than females, consistent with the expectation that males have lower certainty of parentage in the offspring. It is proposed that variation in paternal uncertainty could contribute to the unexplained variation reported in nest defence studies.     

An experimental study of competition for food between male and female nestlings of the red-winged blackbird

Summary Ten experimental broods of red-winged black-bird (Agelaius phoeniceus) nestlings, experimentally manipulated to contain two males and two females of similar age, were observed with the aid of video cameras to determine (a) which nestling characteristics were most important in influencing its chances of being fed and (b) if males and females differed with respect to these characteristics. Nestlings that could reach highest while begging were most successful at obtaining food from parents during individual feeding bouts. However, while there was a tendency for nestlings that begged earlier to be offered food first, a nestling's position in the nest during these bouts did not influence its feeding success. Males begged more than often females and were offered more food by parents. Males were also more likely to be fed when they begged. This was most likely because males, being larger than females, were able to reach higher while begging. Neither sex was more likely to beg sooner or occupy certain positions in the nest. However, while males that had been transferred from other nests received less food than natal males, there were no differences in food acquisition between transferred and natal females. Differences in the relative success of males and females under different conditions may help explain differences in fledgling sex ratios observed in many dimorphic species.     

Effects of nest site concealment on hatching success,reproductive success,and paternal behavior of the threespine stickleback,Gasterosteus aculeatus

Summary Male sticklebacks (Gasterosteus aculeatus) actively competed for a limited number of clay flowerpots as nest sites in circular wading pools. Males nesting in pots spawned earlier and more often, had higher mean and lower variance for hatching success (number of fry per gram of estimated clutch weight), and suffered fewer stolen fertilizations, nest raids, and territorial encounters than did males nesting outside pots, and in pools without pots (Table 2). Males nesting in pots had the more even temporal distribution of fanning bouts and interfanning intervals; the durations of fanning bouts and interfanning intervals were less variable for these males (Table 2).In replicate aquaria, one male nested on each side of a glass partition and both males spawned approximately simultaneously. Flowerpots were provided to one, both, or neither male. Males with pots had higher mean hatching success, fewer territorial encounters, and the more even temporal distribution of fanning bouts and interfanning intervals (Table 4).Thus it appears that nest site concealment, in the form of clay flowerpots, affected male hatching success, reproductive success, and parental behavior, even when physical contact between males was prevented. These data suggest that a causal relationship may exist between the paternal fanning regime and the subsequent hatching success. Furthermore, these data suggest that males which nest in high concealment increase their reproductive success, and that females which spawn with these males reduce the variance of their hatching success as well as increase their mean hatching success.     

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.     

An experimental study of paternal behavior in red-winged blackbirds

Summary The effect of brood size and female nesting status on male parental behavior was investigated in red-winged blackbirds Agelaius phoeniceus using brood size manipulation experiments. Male redwings allocated parental effort on the basis of brood size and nestling age. Males began assisting females only at nests with at least three offspring older than three days. Female nesting status had no singificant influence on male parental care. When females were unable to meet a brood's demand for food, males assisted females with nestling feeding. Females did not reduce the amount of food delivered to nestlings when males assisted. The amount of food brought to nestlings by the male was additional to the amount of food provided by the female. Male assistance increased fledgling success. When female provisioning was sufficient to meet a brood's demand for food males did not assist. The value of male parental care varied inversely with the ability of the female to meet nestling food demands. The ability of unassisted females to provide sufficient food and to raise a brood of nestlings successfully appeared to be influenced by resource abundance.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Nestling provisioning in polygynous great reed warblers (Acrocephalus arundinaceus): do males bring larger prey to compensate for fewer nest visits?

Most studies of social polygyny in birds have examined male provisioning on the basis of the number of feeding visits. This may be misleading if males compensate for infrequent visits by bringing larger prey at each visit. We investigated nestling provisioning in the socially polygynous great reed warbler, Acrocephalus arundinaceus, in south Central Sweden in 1996–1997. We collected data on rate of feeding visits, prey size and the amount of biomass delivered by males and females. Males had lower rates of feeding visits and provided smaller prey to nestlings in secondary than in monogamous and primary nests. Secondary females had higher rates of feeding visits and brought larger prey than monogamous and primary females. These results confirm that secondary females face a potential cost of polygyny through a lower rate of male feeding, and that this cost was reinforced by the significantly lower male provisioning rate (biomass h^–1) at secondary nests. Secondary females compensated for the lack of male assistance by increasing their rate of feeding and bringing larger prey. As a result, offspring in nests of secondary females received as much food as did those in nests of primary females. Prey load size increased with the parent’s proportion of feeding visits, suggesting that parents use different feeding strategies depending on their amount of responsibility for nestling provisioning. We suggest that parents which take the main responsibility for nestling feeding have to forage further away from the nest, and based on optimal-foraging theory, they should then on average bring larger prey to their nest. Received: 4 April 1999 / Received in revised form: 12 October 1999 / Accepted: 23 October 1999     

Cannibalize or care? The role of perceived paternity in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Parental care is a costly part of reproduction. Hence, natural selection should favor males which avoid caring for unrelated young. However, the decision to abandon or reduce care requires cues which are evaluated to give information on potential reproductive value of the offspring. The prediction that male sand gobies, Pomatoschistus minutus, care for foreign eggs as long as they were spawned in their own nest and at least some of such cues are fulfilled was tested. Egg-guarding males that had recently taken part in a spawning event were given a clutch of eggs that was sired either by themselves or another male, in either their own or another male’s aquarium. Males that had not taken part in a spawning event were used as controls and were given eggs sired by another male. We measured the amount of filial cannibalism and nest building. Control group males did not care for eggs and ate them all before rebuilding the nest. In the other treatments, there were no significant effects of paternity, though males moved to another male’s aquarium increased their clutch area threshold and completely consumed larger clutches than males that were not moved. There was no intermediate response in any treatment in the form of increased partial filial cannibalism or less well-constructed nests. Our results suggest that egg-guarding males cannot distinguish between eggs sired by themselves and those sired by other males but are able to react to cues indicating paternity state. Males do not adopt eggs to attract females in P. minutus.     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

Biparental defence in a spider mite (Acari: Tetranychidae) infesting Sasa bamboo

Summary Field observations on the nesting pattern of Schizotetranychus celarius (Banks) (Acari: Tetranychidae) revealed that many individuals live gregariously in a united nest. Behavioural experiments recorded on video tape recorder and 16 mm film showed that adult males and females can effectively defend their nest and offspring from a phytoseiid predator, larvae of Typhlodromus bambusae Ehara (Acari: Phytoseiidae), through counterattacks. The male spider mite often killed the predator's larva and succeeded in defending offspring and nest completely. Comparison of the behaviour patterns recorded between a nest owner parent S. celarius and two kinds of intruders (predators and conspecific males or females) revealed conspicuous differences. I suggest that there is a kind of biparental care in S. celarius, and that the life-pattern of this species therefore fits the definition of subsociality originally framed for insects.     

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)     

Fifteen-spined stickleback (Spinachia spinachia) females prefer males with more secretional threads in their nests: an honest-condition display by males

In the fifteen-spined stickleback (Spinachia spinachia), the male alone builds a nest and provides care for the eggs until they hatch. The nest is made of filamentous algae held together with shiny secretional threads of a glycoprotein, here called tangspiggin. Tangspiggin is produced by transformed kidney cells. I investigated how tangspiggin production was related to food intake by giving males low or high food rations. Males in the high-food group produced significantly more tangspiggin than those in the low-food group. Since low food rations did not lead to a significant drop in resting metabolic rate (oxygen consumption), tangspiggin production appears to be one of the first energy-demanding processes to be dropped during food deprivation. Among males given equal food rations, females preferred to mate with those with elevated levels of tangspiggin in their nests. Tangspiggin threads may signal safety for the offspring by keeping the eggs within the nest and protecting them from egg predators. Moreover, the threads become less shiny with age, due to particle deposition, and older nests that may house older (more valuable) eggs are thereby well camouflaged. Tangspiggin provides an honest signal of male condition, acting as an extra-bodily ornament, signaling direct benefits to females or possibly also indirect benefits of "good genes" from a male who is a successful forager.     

Male mating success and paternal care in Nicrophorus vespilloides (Coleoptera: Silphidae)

Summary If no female is present, male burying beetles Nicrophorus vespilloides (Coleoptera: Silphidae) co-operate in the burial of a corpse. Once a female has arrived, the males fight with one another. The defeated male stays near the corpse and to copulate with the female. Laboratory experiments using sterilised males showed that the defeated male was able sometimes to father some of the offspring raised on the corpse. Male N. vespilloides almost always participate in defence and feeding of the brood. This is not affected by the size of the male. Males quickly leave or are driven from 5 g corpses. Males feed the larvae as often as females do, and larvae raised by males alone are not significantly different in weight from larvae raised by females alone or by both parents. Males which cared for a succession of broods in the laboratory did not differ significantly in median lifespan from males which were removed from their corpses after eggs had been laid. Non-caring males weighed significantly more than caring males over a sequence of corpses, but the caring males did not differ significantly in weight from non-breeding controls.     

Chemosensory cues allow courting male garter snakes to assess body length and body condition of potential mates

When choosing between two potential mates, a male may benefit by picking a larger (longer and/or more heavy-bodied) female because she is likely to produce more or larger offspring. Males of many species use visual cues to evaluate the sizes of their mates, but in some situations (at night or in a crowded mating swarm), vision may be useless. Potentially, males may be able to use chemical cues that convey information about female body size. We manipulated cues available to free-ranging male garter snakes (Thamnophis sirtalis parietalis) in large courting aggregations near communal dens in Manitoba, Canada. Males not only directed disproportionate courtship to longer and heavier-bodied females, but also courted most vigorously in response to lipids extracted from the skins of such females. Our data show that with a flick of his tongue, a male garter snake can identify not only a female's body length, but also her body condition.     

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.     

Solitary nesting as an alternative breeding tactic in colonial nesting bluegill sunfish (Lepomis macrochirus)

Colonial breeding can evolve in response to benefits afforded by clumped individuals, such as reduced predation and increased ease of assessing potential mates. However, colonial breeding can also impose costs such as increased disease transmission or increased cuckoldry. Here, we investigate solitary nesting as a potential alternative breeding tactic in colonial breeding bluegill sunfish (Lepomis macrochirus). Most male bluegill, termed parentals, compete for nesting sites in colonies and then court and spawn with females and provide sole care of the eggs. Although nesting in a colony results in reduced predation and fungal infection of broods, it comes at a cost of increased parasitism by specialized cuckolder males that do not nest. We found that 4.5% of parentals forgo spawning in a colony and instead construct nests solitarily. Solitary males were of similar size and age to colonial males, but were in significantly better condition. Solitary males obtained as many eggs as males nesting in the center of colonies, and significantly more than males nesting on the periphery of colonies. Thus, females do not appear to discriminate against solitary males. Solitary males had smaller ear tabs, a presumed sexually selected character used by parental males in intrasexual competition, than colonial males. Tracking data revealed consistency in nesting tactic (but not nest position within the colony) between spawning attempts. We suggest that solitary nesting represents either a facultative decision made by parental males in top condition at the onset of breeding, or a life history decision to forgo spawning in colonies.Communicated by K. Lindström     

Mate guarding in the swallowHirundo rustica

Summary The importance of mate guarding by males in the monogamous swallowHirundo rustica was studied by temporarily detaining the males. Mate guarding reduced the frequency of extra-pair copulations and of sexual chases involving female mates. Males participated in sexual chases more frequently if they had a non-fertile female. Neighbouring males of ‘widowed’ females increased their own mate guarding presumably in response to the experimentally increased rate of sexual chases. Neighbouring males with a fertile female increased their mate guarding more than did males with a non-fertile female. Addition of eggs to swallow nests in the post-fledging period of the first brood induced mate guarding by male nest owners. These males also copulated more frequently with their mates than did control males. Neighbouring male swallows responded to the increased mate guarding by showing sexual interest in the guarded females. removal of eggs from swallow nests during the laying period, leaving only one egg in the nest, resulted in reduced nest attendance by females. Male mates responded by increasing their mate guarding intensity as compared to controls, and neighbouring males showed an increased sexual interest in these females.     

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.     

The effect of maternal body size on embryo survivorship in the broods of pregnant male pipefish

The occurrence of male pregnancy in the family Syngnathidae (seahorses, pipefishes, and sea dragons) provides an exceptionally fertile system in which to investigate issues related to the evolution of parental care. Here, we take advantage of this unique reproductive system to study the influence of maternal body size on embryo survivorship in the brood pouches of pregnant males of the broad-nosed pipefish, Syngnathus typhle. Males were mated with either two large females, two small females, a large then a small female, or a small then a large female. Our results show that offspring survivorship depends on an interaction between female body size and the number of eggs transferred by the female. Eggs of larger females deposited in large numbers are more likely to result in viable offspring than eggs of smaller females laid in large numbers. However, when females deposited smaller numbers of eggs, the eggs from smaller females were more likely to produce viable offspring compared to those from larger females. We found no evidence that this result was based on mating order, the relative sizes of competing females, or egg characteristics such as dry weight of eggs. Additionally, male body size did not significantly influence the survivorship of offspring during brooding. Our results suggest that the factors underlying offspring survivorship in pipefish may be more complex than previously believed, with multiple factors interacting to determine the fitness of individual offspring within the broods of pregnant males.