Ventilation or nest defense—parental care trade-offs in a fish with male care

Brood guarding animals face many critical trade-offs. Sand goby males (Pomatoschistus minutus) build nests with larger openings during low oxygen conditions, presumably to enhance ventilation. However, this may make the nest easier for egg predators to detect and harder for guarding males to defend. Manipulating oxygen level and predator presence (a small crab) for small and large males, we found support for a parental trade-off between fanning and nest defense. An increased fanning activity resulted in less time for guarding. Small males and males in low oxygen showed a higher fanning expenditure than large males and males in high oxygen, but surprisingly, filial cannibalism did not differ between these groups. Males built larger nest openings in low than high oxygen. However, males in both high and low oxygen treatments reduced their nest opening size in the presence of a predator, again indicating an important trade-off between ventilation and nest defense.     

Effects of costs and benefits of brood care on filial cannibalism in the sand goby

This study tested experimentally whether clutch size and the cost of care affect filial cannibalism in the sand goby, Pomatoschistus minutus. Evolutionary models of filial cannibalism suggest that egg eating has evolved as a way for the male parent to prolong his breeding season. These models assume that eggs function as an alternative energy source for the constrained parent. I manipulated clutch size by allowing males to mate with either one or two females, representing a small and a large clutch, respectively. The addition of a small male shore crab, a common egg predator, increased the cost of care. I quantified fat reserves as a measure of the condition of guarding males. Males who did not build nests had lower fat reserves than males who built nests, suggesting that males with low energy reserves do not start breeding. Males with small clutches lost their nest to the crab more often than males with large clutches. Neither filial cannibalism nor the amount of eggs eaten were affected by the treatments. Males who consumed eggs had a higher fat percentage than males who did not eat eggs. The result that males with small clutches lost their nests to the crabs supports the idea that eggs are defended only if the benefit from continued care will outweigh the cost and that males therefore are sensitive to the trade-off between present and future reproductive success. Received: 15 May 1997 / Accepted after revision: 15 November 1997     

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 overlooked side effect of nest-scattering behavior to decrease predation risk (Acari: Tetranychidae,Stigmaeidae)

The number of nests containing egg masses a female makes over her lifetime and the pattern of scattering nests vary among species in a genus of nest-weaving spider mites (Stigmaeopsis). We hypothesized that the scattered nests of small nest builders have a previously overlooked indirect effect in that the void nests created after predation take on a new role as hindering devices that effectively decrease predator searching efficiency. First, we demonstrated that the experimental design used in this study is a good reflection of the nest distribution pattern of Stigmaeopsis takahashii (an intermediate-sized nest builder) in the field. Using this species as a model, we tested how different nest-scattering patterns affect the predator to examine how scattering may indirectly provide an anti-predation strategy by increasing a predators searching time. Next, we observed how artificially arranged void nests disturb predatory behavior in both starved and fully fed predator females and showed that void nests have a strong hindering effect on predators. Thus, we concluded that the nesting behavior of this mite species not only has anti-predator effects but must also have a stabilizing effect on predator–prey interaction systems at the population level.     

Differences in Predators of Artificial and Real Songbird Nests: Evidence of Bias in Artificial Nest Studies

Abstract:  In the past two decades, many researchers have used artificial nests to measure relative rates of nest predation. Recent comparisons show that real and artificial nests may not be depredated at the same rates, but no one has examined the mechanisms underlying these patterns. We determined differences in predator-specific predation rates of real and artificial nests. We used video cameras to monitor artificial nests baited with quail and plasticine eggs and Field Sparrow ( Spizella pusilla ) and Indigo Bunting ( Passerina cyanea ) nests in field habitats in central Missouri (U.S.A.). Although daily predation estimates (all predators pooled) were similar between artificial and real nests, predators differed substantially in their depredation of artificial versus real nests. Snakes were the major predator at real nests, and raccoons ( Procyon lotor) were the major predator at artificial nests. We found strong support for models that distinguished predation between two or among three predator groups and between artificial and real nests. There was no snake predation of artificial nests, and the odds of predation of artificial nests was 115–551% (95% confidence interval) and 2–154% of the odds of predation of real nests by mammals and birds, respectively. Artificial nests with plasticine eggs could not be used reliably to identify predators. In several cases plasticine eggs were marked by mice, and raccoons were recorded on video removing the quail egg. Because biases for artificial nests were positive for some predators and negative for other predators (and could be compensating), and potentially existed for all predator groups, conclusions based on artificial nest studies should be suspect even when there is evidence that overall predation rates are similar among real and artificial nests.     

Coloration in New World orioles

Summary Although sexual dichromatism in birds is usually ascribed to sexual selection, some workers argue that avian coloration is better explained by predation-related selection pressures. Supporting evidence for these latter hypotheses comes primarily from broad interspecific comparisons, which can be biased by a variety of factors. This study examines the predation-based hypotheses of Baker and Parker (1979), particularly the Predator Deflection Hypothesis, with reference to two closely-related oriole species: Icterus parisorum, which is dichromatic, and I. gularis, in which both sexes are brightly coloured. To test the prediction that bright coloration serves to divert the attention of predators away from cryptic young, rates of predation on nests of conspicuous (2 years of age or older) and dull-coloured (first-year). I. parisorum males were compared. The results showed equal predation on the young of males in both age/plumage classes. The Predator Deflection Hypothesis also predicts that, once a predator has been detected, brightly-coloured birds should attempt to distract it, whereas cryptic individuals should not. Tests using models of avian predators showed that this was not the case: cryptic I. parisorum females responded as aggressively toward the model as did conspicuous conspecific males. The same was true for I. gularis. Overall, the results did not support the idea that bright coloration has evolved in response to predation pressure in these species.     

Hunted hunters? Effect of group size on predation risk and growth in the Australian subsocial crab spider Diaea ergandros

A reduced predation risk is considered to be a major adaptive advantage of sociality. While most studies are concerned with non-predatory prey species, group-living predators are likely to face similar threats from higher-order predators. We studied the relationship between group size and predation risk in the subsocial crab spider Diaea ergandros by testing predictions from theoretical models including attack abatement as well as the formation of protective retreats. In a field survey, we found predatory clubionid spiders in 35 % of the D. ergandros nests and as predicted, nest size did not correlate with predator presence. In a subsequent laboratory experiment, we observed survival probability, nest construction activity and feeding behaviour including weight development between groups of different sizes as well as in the absence or presence of a predator. Large groups had an advantage in terms of survival and growth compared to smaller groups or single individuals. They also built significantly larger nests than smaller groups, supporting the idea of protective retreat formation being an adaptive benefit to group living. Even though clubionids did attack D. ergandros, they did not significantly affect overall mortality of D. ergandros. The feeding experiment showed that spiders fed on a larger proportion of flies in the presence of a predator. However, these groups gained significantly less weight compared to the control groups, indicating that the potential predators not only act as predators but also as food competitors, constituting a twofold cost for D. ergandros.     

Behavioral resource depression and decaying perceived risk of predation in two species of coexisting gerbils

Summary Behavioral resource depression occurs when the behavior of prey individuals changes in response to the presence of a predator, resulting in a reduction of the encounter rate of the predator with its prey. Here I present experimental evidence on the response of two species of gerbils (Gerbillus allenbyi and G. pyramidum) to the presence of barn owls. I conducted the experiments in a large aviary. Both gerbils responded to the presence of barn owl predators by foraging in fewer resource patches (seed trays) and by quitting foraged resource patches at a higher resource harvest rate (giving-up density of resource; GUD). This reduced the amount of time gerbils were exposed to owl predation, and hence the encounter rate of owls with gerbils, i.e., behavioral resource depression. Thus, the presence of owls imposes a foraging cost on gerbils due to risk of predation, and also on the owls themselves due to resource depression. I then examined how resource depression relaxed over time following exposure to owls. In the days following an encounter with the predator, the reduction in foraging activity for both gerbil species eased. Increasing numbers of trays were foraged each day, and GUDs in seed trays declined. The two gerbils differed in their rate of recovery, with G. pyramidum returning to prepredator levels of foraging after 1 or 2 nights and G. allenbyi taking 5 nights or longer. Interspecific differences in recovery rates may be based on differences between the species in vulnerability to predation and/or ability to detect the presence of predators. The differences in recovery rates may be due to optimal memory windows or decay rates, where differences between species are based on risk of predation or on how perceived risk changes with time since a predator was last encountered. Finally, differences between or among competitors in recovery from resource depression may provide foraging opportunities in time for the species which recover most quickly and may have implications for species coexistence.     

Sexual selection for male parental care in the sand goby, Pomatoschistus minutus

Male parental care is typically thought to come at a cost to mate attraction and future mating success. However, it has also been hypothesized that paternal care may be under sexual, as well as natural, selection, such that good fathers actually attract more mates. Here we show experimentally that in the sand goby, Pomatoschistus minutus, females prefer to mate with males that provide higher levels of parental care. We manipulated male behavior using (1) different nest sizes and (2) an application of low-O₂ water in the nests, and found that females consistently preferred males with elevated levels of care in dichotomous mate choice tests. This complements our earlier study in which we showed that males increase the amount and quality of care they provide in the presence of females. Our results demonstrate that male care may have evolved as a result of sexual selection rather than natural selection alone, and furthermore, that male care may not necessarily be in conflict with mate attraction.     

Nest-size variation reflecting anti-predator strategies in social spider mites of<Emphasis Type="Italic"> Stigmaeopsis</Emphasis> (Acari: Tetranychidae)

The social spider mites (Acari: Tetranychidae) of Stigmaeopsis weave dense nests on the underside of host leaves. Four species occur on the leaves of bamboo in Japan: Stigmaeopsis longus, S. celarius, S. takahashii and S. saharai. We initially reconfirmed the occurrence of distinct variation in nest size among the species. Based on the hypothesis that this variation plays a role in protecting the spider mites from predators, we looked at the behavior of the natural enemies that occur on the host plants along with members of Stigmaeopsis. We found considerable variation in the ability of nests to protect the spider-mite eggs. The smallest nests protected the eggs against three predators, whereas the largest nests protected the eggs against only one predator species. So, decreases in nest size increased egg defense. Thus we concluded that nest-size variation reflects a strategy for reducing predation.Communicated by D. Gwynne     

Non-consumptive effects of predator presence on copepod reproduction: insights from a mesocosm experiment

Reproduction in planktonic animals depends on numerous biotic and abiotic factors. One of them is predation pressure, which can have both direct consumptive effects on population density and sex ratio, and non-consumptive effects, for example on mating and migration behaviour. In copepods, predator vulnerability depends on their sex, motility pattern and mating behaviour. Therefore, copepods can be affected at multiple stages during the mating process. We investigated the reproductive dynamics of the estuarine copepod Eurytemora affinis in the presence and absence of its predator the mysid Neomysis integer in a mesocosm experiment. We found that the proportion of ovigerous females decreased in the presence of predators. This shift was not caused by differential predation as the absolute number of females was unaffected by mysid presence. Presence of predators reduced the ratio of males to non-ovigerous females, but not by predation of males. Our combined results suggest that the shift from ovigerous to non-ovigerous females under the presence of predators was caused by either actively delayed egg production or by shedding of egg sacs. Nauplii production was initially suppressed in the predation treatment, but increased towards the end of the experiment. The proportion of fertilized females was similar in both treatments, but constantly fell behind model predictions using a random mating model. Our results highlight the importance of non-consumptive effects of predators on copepod reproduction and hence on population dynamics.     

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.     

Individual differences in nest defense in the colonial breeding Black-tailed Gulls

Often in colonial seabirds, all colony members are believed to defend against nest predators and experience equal nest predation risk. However, the variation of defense behavior among members and its reproductive consequences are largely unknown. We investigated (1) individual variation in the nest defense of breeding Black-tailed Gulls Larus crassirostris against a natural egg predator, the Jungle Crow Corvus macrorhynchos and (2) how this behavioral variation affects an individual’s own nest predation risk and that of their neighbors. Results were compared between 2 years where crow attack levels were manipulated to average 5 and 22 times normal rates (“low” and “high” predation risk years, respectively) by the placement of varying numbers of artificial nests containing unguarded eggs at the perimeter of the gull colony. In both years, 23–38% of parents, mostly males, showed “aggressive” defense behavior (strikes or chases) against crows and decoys. Other “non-aggressive” gulls showed no defense. In the year of low predation risk, intrusion rates by crows (landing within 0.5 m of an individual gull’s nest) were similar for aggressive and non-aggressive gulls. In the year of high predation risk, however, the rates of intrusion for aggressive gulls (4%) and for non-aggressive gulls with an aggressive neighbor (37%) were significantly lower than for non-aggressive gulls without an aggressive neighbor (76%). These results indicate that aggressive individuals reduce nest predation risk for themselves and conspecific neighbors in a colonially breeding species.     

Parental care and the cost of reproduction in a Mediterranean fish

Summary Symphodus tinca is a common near-shore Mediterranean labrid fish in which females may sometimes spawn their eggs over hundreds of square meters, or alternatively spawn into well-defined algal nests. Eggs spawned in either manner are fertilized, but widely scattered eggs receive no parental care, whereas eggs spawned into nests are usually guarded by the male until they hatch. Here, I report weight changes of individual marked fish that engaged in a variety of different reproductive behaviors during three breeding seasons. Males gained weight at 0.15% per day outside the spawning season, and added 29–78% to their overall body weight between reproductive seasons, even following substantive weight losses in a spawning season (up to 20% among nesting males). Nesting and nest-guarding males lost an average of 0.32% and 0.41% of their body weight per day in 1986 and 1987. This cost is four times greater than reproduction for nonnesting males, which registered a 0.03% daily weight gain. Actively spawning females lost 0.06% of their body weight daily during the spawning season. While long-term growth rates did not appear to be substantially affected by reproduction in either sex or by parental care in males, present work does not exclude the possibility that such long-term effects may exist.     

Female sand gobies gain direct benefits by choosing males with eggs in their nests

In some fish species with paternal care, females prefer to spawn with males whose nests already contain eggs. Several hypotheses have been put forward to explain this behaviour, such as reduced risk of predation or cannibalism (the dilution effect), increased parental investment, and mate copying. This experimental study focuses on female mate choice in the sand goby, Pomatoschistus minutus. Females were found to choose males with eggs in their nests. In addition, hatching success increased with clutch size, mainly because males with larger clutches showed less filial cannibalism. Increased egg survival in large clutches may thus be explained by a combination of the dilution effect and higher parental investment. In another experiment, females did not seem to copy the observed mate choice of other females. In conclusion, female preference for males with eggs in their nests is adaptive, and can be explained by direct benefits, as more surviving offspring are produced. Received: 23 December 1995/Accepted after revision: 11 May 1996     

Characterizing temperate rocky shores from the perspective of an early juvenile snail: the main threats to survival of newly hatched Nucella emarginata

Mortality factors most likely to constitute substantial selective pressures for early juvenile gastropods on temperate rocky shores were identified by examining the vulnerability of hatchlings of an intertidal snail, Nucella emarginata, to heat stress, desiccation, and predation in 1992 and 1993. The highest temperature of substrata measured at tidal heights colonized by N. emarginata in Barkley Sound, British Columbia, Canada, was 28.5°C. This temperature was not lethal to hatchlings in laboratory tests. In laboratory and field desiccation experiments, all hatchlings died within 6 h of emersion. Early juveniles could not survive direct exposure to even moderate drying conditions for the duration of a low tide. Hence, intertidal microhabitats which dry up even for short periods during low tides would prove lethal. Of 45 intertidal animal species to which hatchlings were exposed in the laboratory, small decapod crustaceans were the only organisms to cause substantial hatchling mortality. Of these, Pagurus hirsutiusculus and Hemigrapsus nudus were by far the most abundant in the field, and are probably the only important predators of early juvenile N. emarginata at most sites. Total predator densities in the field were as high as 438 individuals m^–2, suggesting that predation pressure may be intense. Desiccation and predation by decapod crustanceans appear to be the most significant threats to early juvenile N. emarginata. These factors commonly occur on most temperate rocky shores and undoubtedly constitute major selective agent influencing population parameters and shaping life-history strategies and early juvenile traits of intertidal invertebrates.     

Predation risk affects courtship and attractiveness of competing threespine stickleback males

The effect of predation risk and male-male competition on male courtship behaviour and attractiveness to females was studied in the threespine stickleback (Gasterosteus aculeatus) by presenting dummy or live females to solitary and competing males under different predation risks. In the presence of a predator, males decreased courtship activity. Different courtship components were, however, adjusted to different extents and in opposing directions to predation risk, probably because the single components may have varied in riskiness. The presence of a competing male decreased overall courtship activity, but increased the frequency of zigzags, suggesting zigzagging to be a competitive strategy against other males. In the presence of a predator male courtship activity was not affected by a competitor. Female mate choice correlated with the males' previous frequency of zigzags towards a dummy female. However, when a live female paid attention to a male, the male decreased zigzagging and instead increased leading and fanning behaviours, probably trying to attract the female to the nest to mate. Predation risk affected the attractiveness of males as females reduced their attention to a male when he faced a predator and reduced his courtship activity. As females instead increased their attention to a competing male that had increased his courtship activity, due to decreased competition, males clearly are balancing mating opportunities against predator avoidance. When males vary in their susceptibility to predators, predation risk may thus affect mating success of competing males. Received: 31 January 1997 / Accepted after revision: 15 April 1997     

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.     

Kill before being killed: an experimental approach supports the predator-removal hypothesis as a determinant of intraguild predation in top predators

Intraguild predation (IGP) has been explained in terms of competitor-removal, food-stress and predator-removal hypotheses. Only the first two hypotheses have been fairly well studied. To test the predator-removal hypothesis as a force determining IGP in avian predators, we performed a field experiment to simulate the presence of an IG predator (eagle owl Bubo bubo dummy) in the surrounding of the nests of four potential IG prey (black kite Milvus migrans, red kite Milvus milvus, booted eagle Aquila pennata and common buzzard Buteo buteo). To discard the possibility that an aggressive reaction towards the eagle owl was not related to the presence of the IG predator, we also presented a stuffed tawny owl Strix aluco, which is a potential competitor but cannot be considered an IG predator of the studied diurnal raptors considered in the experiment. While almost always ignoring the tawny owl, raptors chiefly showed an interspecific aggressive behaviour towards their IG predator. Our results seem to support the predator-removal hypothesis, as the IG prey may take advantage of the diurnal inactivity of the IG predator to remove it from their territory. However, the recorded behaviour may be also considered as a special variety of mobbing (i.e. a prey’s counter-strategy against its predator), where the mobber is sufficiently powerful to escalate predator harassment into deliberate killing attempts. In their turn, eagle owls can respond with an IG predatory behaviour aimed at removing IG prey species which are highly aggressive mobbers.     

Blue eggs do not reduce nest predation in the song thrush,Turdus philomelos

Summary Many passerine birds with open cup-shaped nests lay blue or blue-green eggs. In thrushes, blue eggs may be cryptic and provide camouflage by imitating spots of light on green leaves. Alternatively, egg coloration may be selectively neutral because nest predators detect nests and not eggs, or it may be maladaptive because organisms are not always well adapted to their present environment. I evaluated these hypotheses by studying predation on artificial song thrush (Turdus philomelos) nests with quail eggs, painted either white, blue, or spotted (cryptic to a human eye). Corvids were the major nest predators. For concealed as well as exposed nests, I found no differences in the predation rates of nests with white, blue, or spotted eggs. Predators apparently detected the nests, and not the eggs, first. In a second experiment, I placed egg groups without nests in trees to study the effect of color per se. The predation rate of the spotted egg groups was significantly lower than that of the white and blue egg groups, for concealed as well as exposed egg groups. These results suggest that blue eggs in the song thrush are not cryptic but may be selectively neutral or even maladaptive with regard to nest predation.