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1.
The egg deposition behavior of the turnip sawfly, Athalia rosae (Hymenoptera: Symphyta), is described. Both unmated and mated females lay eggs individually inside of fresh young leaves of cruciferous plants. During an oviposition event, females exhibit a distinct pause in abdominal contractions just before the actual egg deposition act. Unmated females show a longer pause (11.31 s on average) than mated females (4.38 s on overall average). By employing an eye color mutation, the sex of the eggs laid by females was ascertained. Females mated once lay mostly fertilized (diploid female) eggs initially but begin to lay a considerable number of unfertilized (haploid male) eggs later in life. The laying of an unfertilized egg is associated with a longer pause (6.98 s on average) than the laying of a fertilized egg (3.76 s on average). These results are in contrast to previous reports on apocritan Hymenoptera, where the presence of a pause or a longer pause during oviposition was associated with the deposition of fertilized eggs rather than unfertilized eggs. The possibility that mated Athalia rosae females control fertilization and its implications for sex allocation strategies are discussed.  相似文献   

2.
The number of mature eggs remaining in the ovaries and the time left for oviposition determine the reproductive decisions of the hyperdiverse guild of insects that require discrete and potentially limiting resources for oviposition (such as seeds, fruits or other insects). A female may run out of eggs before all available oviposition sites are used (egg limitation), or die before using all of her eggs (time limitation). Females are predicted to change clutch size depending on whether eggs or time is the limiting resource. We extend this framework and ask whether the same constraints influence a strategy in which females modify eggs into protective shields. In response to egg parasitism cues, female seed beetles (Mimosestes amicus) lay eggs in vertical groups of 2–4, modifying the top 1–3 eggs into shields in order to protect the bottom egg from attack by parasitoids. We made contrasting predictions of how egg and time limitation would influence egg size and the incidence and level of egg protection. By varying access to seed pods, we manipulated the number of remaining eggs a female had at the time she received a parasitism cue. Although egg size was not affected, our results confirm that egg‐limited females protected fewer eggs and time‐limited females protected more eggs. Female body size explained the number of eggs in a stack rather than host deprivation or the timing of parasitoid exposure. Our results clearly show that host availability relative to female age influences the incidence of egg protection in M. amicus. Furthermore, our study represents a novel use of life history theory to explain patterns in an unusual but compelling defensive behaviour.  相似文献   

3.
Many bird species lay eggs speckled with protoporphyrin‐based spots, however, for most of them the function of eggshell spotting is unknown. A plausible hypothesis is that protoporphyrin might have a structural function in strengthening the eggshell and is therefore deposited when calcium is scarce. In this study, we experimentally provided Great Tit Parus major females with supplemental calcium to examine its effect on the protoporphyrin‐based maculation of their eggs. In addition, we studied variation in eggshell pigmentation patterns in relation to other egg parameters and laying order. Calcium‐supplemented females laid larger eggs but shell thickness was not significantly affected by the treatment. Calcium supplementation may reduce the time and energy females devote to searching for calcium‐rich material, so that they can collect more nutrients and so lay larger eggs. Furthermore, pigment darkness was associated with egg volume and shape, which suggests that female quality and environmental food availability may also influence the shell pigmentation pattern. Within clutches, later‐laid eggs had larger and darker spots that were distributed more unevenly on the shell surface. This within‐clutch pattern could be explained by the increase in egg volume and egg shape and a decline in shell thickness with egg‐laying order, which characteristics were all related to shell‐spotting pattern. Eggs with a coronal ring had thinner shells, but pigment intensity and spot size were not related to shell thickness. Thus, our results suggest that concentrated spotting distribution may have a mechanical function, supporting the structural‐function hypothesis.  相似文献   

4.
Oviparous, facultative egg retention enables Drosophila females to withhold fertilized eggs in their reproductive tracts until circumstances favor oviposition. The propensity to retain fertilized eggs varies greatly between species, and is correlated with other reproductive traits, such as egg size and ovariole number. While previous studies have described the phenomenon, no study to date has characterized within‐species variation or the genetic basis of the trait. Here, we develop a novel microscope‐based method for measuring egg retention in Drosophila females and determine the range of phenotypic variation in mated female egg retention in a subset of 91 Drosophila Genetic Reference Panel (DGRP) lines. We inferred the genetic basis of egg retention using a genome‐wide association study (GWAS). Further, the scoring of more than 95,000 stained, staged eggs enabled estimates of fertilization success for each line. We found evidence that ovary‐ and spermathecae‐related genes as well as genes affecting olfactory behavior, male mating behavior, male‐female attraction and sperm motility may play a crucial role in post‐mating physiology. Based on our findings we also propose potential evolutionary routes toward obligate viviparity. In particular, we propose that the loss of fecundity incurred by viviparity could be offset by benefits arising from enhanced mate discrimination, resource specialization, or modified egg morphology.  相似文献   

5.
The color of lepidopteran eggs often varies by species or egg condition, and parasitoids that attack lepidopteran eggs could therefore potentially use color to obtain information about host identity or quality. The objective of our study was to determine whether females of the egg parasitoid Trichogramma ostriniae Pang & Chen (Hymenoptera: Trichogrammatidae) showed differential responses to egg color when searching for hosts over short distances and when evaluating the suitability of encountered eggs. We examined the wasps’ host‐selection behavior in a Petri dish arena using white, yellow, green, and black clay beads as egg models presented against a green background (to mimic leaf color). In no‐choice tests, bead color had a significant effect on the proportion of tested wasps that accepted a bead for further examination, on the time it took wasps to find and begin examining a bead, and on the time that wasps spent examining the beads. However, bead color had only a marginally significant effect on the proportion of wasps attempting to drill into a bead with their ovipositors, and no significant effect on the amount of time they spent drilling. The wasps also showed significant color preferences when given a choice between two adjacent beads of different colors. The results of the no‐choice and choice trials taken together indicated a color preference ranking of yellow > white > green > black. The wasps’ higher preference for the yellow and white egg models generally corresponds to the white or yellowish‐white egg color of T. ostriniae's target host, the European corn borer moth, Ostrinia nubilalis Hübner (Lepidoptera: Crambidae). The wasps’ strong rejection of black egg models is likely to be an adaptive response that reflects the fact that eggs that are wholly or partially black are often unsuitable for parasitization due to advanced caterpillar development, damage to the egg, or previous parasitization.  相似文献   

6.
Conspecific brood parasites lay eggs in nests of other females of the same species. A variety of methods have been developed and used to detect conspecific brood parasitism (CBP). Traditional methods may be inaccurate in detecting CBP and in revealing its true frequency. On the other hand more accurate molecular methods are expensive and time consuming. Eadie developed a method for revealing CBP based on differences in egg morphology. That method is based on Euclidean distances calculated for pairs of eggs within a clutch using standardized egg measurements (length, width and weight). We tested the applicability of this method in the common pochard Aythya ferina using nests that were identified as parasitized (39 nests) or non‐parasitized (16 nests) based on protein fingerprinting of eggs. We also analyzed whether we can distinguish between parasitic and host eggs in the nest. We found that variation in MED can be explained by parasitism but there was a huge overlap in MED between parasitized and non‐parasitized nests. MED also increased with clutch size. Using discriminant function analysis (DFA) we found that only 76.4% of nests were correctly assigned as parasitized or non‐parasitized and only 68.3% of eggs as parasitic or host eggs. Moreover we found that MED in parasitized nests increased with relatedness of the females that laid eggs in the nest. This finding was supported by positive correlation between MED and estimated relatedness in female–female pairs. Although variation in egg morphology is associated with CBP, it does not provide a reliable clue for distinguishing parasitized nests from non‐parasitized nests in common pochard.  相似文献   

7.
Recent evidence suggests that blue‐green coloration of bird eggshells may be related to female and/or egg phenotypic quality, and that such colour may affect parental effort and therefore the nutritional environment of developing nestlings. Here we suggest that these relationships and the signal function of eggshell coloration would affect the outcome of coevolution between avian brood parasites and their hosts in at least three different non‐exclusive evolutionary pathways. First, by laying blue‐green coloured eggs, cuckoo females may exploit possible sensory biases of their hosts, constraining the evolution of parasitic egg recognition, and thus avoid rejection. Second, because of the relatively high costs of laying blue eggs, cuckoo females may be limited in their ability to mimic costly blue‐green eggs of their hosts because cuckoo females lay many more eggs than their hosts. Furthermore, costs associated with foreign egg recognition errors would be relatively higher for hosts laying blue eggs. Third, cuckoos may use coloration of host eggs for selecting individuals or specific hosts of appropriate phenotypic quality (i.e. parental abilities). We here explored some predictions emerging from the above scenarios and found partial support for two of them by studying egg coloration of European cuckoos (Cuculus canorus) and that of their 25 main hosts, as well as parasitism and rejection rate of hosts. Cuckoo hosts parasitized with more blue, green, and ultraviolet cuckoo eggs, or those laying more blue‐green eggs, were more prone to accept experimental parasitism with artificial cuckoo eggs. In addition, coloration of cuckoo eggs is more variable when parasitizing hosts laying bluer‐greener eggs, even after controlling for the effect of host egg coloration (i.e. degree of egg matching). Globally, our results are consistent with the proposed hypothesis that host egg traits that are related to phenotypic quality of hosts, such as egg coloration, may have important implications for the coevolutionary interaction between hosts and brood parasites. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 154–168.  相似文献   

8.
According to foraging theory, female parasitoids should alter their host choice in response to cues that indicate a limitation of resources. We tested whether females of the polyembryonic parasitoid Ageniaspis fuscicollis (Hymenoptera: Encyrtidae), which attack egg batches of small ermine moths (Lepidoptera: Yponomeutidae), would alter their host acceptance pattern in response to different pre‐patch experience. We kept females of the parasitoid prior to a patch visit under different conditions, which should indicate different levels of competition for hosts. With increased competition as pre‐patch experience, females laid more eggs per host egg and self‐superparasitized more often, and the resultant egg distributions showed a trend from more regular distributions to increasingly Poisson and aggregated distributions. Consequently, females with a pre‐patch experience that would indicate low competition for hosts had the most even egg distributions. We conclude that pre‐patch experience of competitors may lead to a significant change of mutual interference patterns in egg‐laying A. fuscicollis wasps.  相似文献   

9.
Avian communal breeding systems generate alternative behavioural strategies for females, resulting in differences in reproductive success. Identifying eggs of different females in such systems is problematic, however, due to egg destruction before incubation, difficulty of capturing adults, and/or inaccuracy of egg identification based on egg morphometry. Here, we describe a technique that uses electrophoresis of yolk proteins to determine egg ownership, which we applied to communally breeding guira cuckoos (Guira guira). Validation of the method included identical yolk protein banding patterns in all eggs of the same female, but different patterns in eggs of different females in budgerigars (Melopsittacus undulatus), and identical patterns in yolk follicles of the same females in guira cuckoos. We applied the protocol to 195 guira cuckoo eggs from 34 joint nests in 2 years. All multiple guira cuckoo eggs laid on the same day in single nests had distinct banding patterns of yolk proteins, practically eliminating the possibility of more than one female being represented by the same pattern. Some identical banding patterns were repeated in different days within a nesting bout, indicating that some females laid several eggs in shared nests. Identical patterns occasionally occurred in renestings of groups, indicating that some females lay eggs in consecutive nestings. Yolk protein electrophoresis is a useful tool to identify egg maternity in other circumstances, such as polygynous mating systems with joint nests and intraspecific parasitism. Additionally, it is an alternative method for species where electrophoresis of egg white proteins does not show sufficient polymorphism.  相似文献   

10.
We collected gravid king ratsnakes (Elaphe carinata) from three geographically separated populations in Chenzhou (CZ), Lishui (LS) and Dinghai (DH) of China to study the geographical variation in female reproductive traits and trade‐offs between the size and number of eggs. Not all reproductive traits varied among the three populations. Of the traits examined, five (egg‐laying date, post‐oviposition body mass, clutch size, egg mass and egg width) differed among the three populations. The egg‐laying date, ranging from late June to early August, varied among populations in a geographically continuous trend, with females at the most northern latitude (DH) laying eggs latest, and females at the most southern latitude (CZ) laying eggs earliest. Such a trend was less evident or even absent in the other traits that differed among the three populations. CZ and DH females, although separated by a distance of approximately 1100 km as the crow flies, were similar to each other in most traits examined. LS females were distinguished from CZ and DH females by the fact that they laid a greater number of eggs, but these were smaller. The egg size–number trade‐off was evident in each of the three populations and, at a given level of relative fecundity, egg mass was significantly greater in the DH population than in the LS population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 701–709.  相似文献   

11.
Social groups of the joint‐laying Pukeko Porphyrio porphyrio melanotus typically contain one or two breeding females. Male Pukeko mated to two females father more offspring and therefore benefit from this mating arrangement; however, primary females should not prefer this system, because fewer eggs hatch per female in the larger joint clutches. Here, we investigated male response to simulated egg destruction, a common female competitive tactic observed in other joint‐laying species. In response to egg removal, males reduced the consistency of their incubation and in some cases nests were abandoned. Such decreases in paternal effort could eliminate any putative advantage gained by a female that destroys the eggs of a co‐nester. Our study demonstrates facultative adjustments in paternal care in a joint‐laying species and suggests that primary females may be limited in their ability to monopolize reproduction.  相似文献   

12.
One of the hypotheses to explain egg colour variation in birds lays in the context of sexual selection, where egg colour may signal the female’s physical condition and antioxidant capacity. We tested one of the assumptions following from this hypothesis, that eggshell pigment deposition should be limited for females. The study was conducted in a captive house sparrow (Passer domesticus) population over several years under constant environmental conditions. This multi-brooded species lays eggs which vary in ground colour (biliverdin pigment) and in the intensity and distribution of brownish-red spots (protoporphyrin pigment). Spot darkness, spread and ground colour diminished along the laying sequence, suggesting that the deposition of both pigments was limiting for females over the short term. Also, the proportion of eggs with biliverdin diminished in consecutive clutches laid by the same female over the breeding season, suggesting a long-term cost of biliverdin deposition. On the other hand, spots were darker at the end of the breeding season, indicating that protoporphyrin deposition was probably not limited over the long term. This result could indicate a lower capacity for calcium deposition over the long term, which was compensated for by darker spots. Female age also significantly affected the proportion of bluish eggs and spot patterns. Egg pigmentation decreased with age, indicating that senescing passerine females lay less pigmented eggs. Clutch size was positively related to the proportion of bluish eggs and to spot patterns (darker and more evenly spotted). These results are in accordance with assumptions for the sexual selection hypothesis.  相似文献   

13.
Abstract. 1. Harem polygyny can have fitness benefits and costs on females. In bark beetles of the genus Ips the latter may include within‐harem competition between larvae. However, earlier competition between females for male care and mating opportunities may also influence oviposition behaviour. There has been relatively little investigation into the relationship between harem size and initial egg output. The present study investigated this relationship in the bark beetle Ips grandicollis. 2. The measure of egg output used was the number of eggs in the gallery with the most eggs in each harem. Mean (±SE) harem size of 242 observed harems was 3.25 ± 0.10. A curvilinear relationship was found between egg output and harem size, with females in smaller harems (one to four females) laying more eggs with increased harem size. However, females in larger harems (five to seven females) laid fewer eggs as harem size increased. The optimal harem size (in terms of number of eggs laid) was close to four females. 3. We found no evidence from a behavioural assay that females could preferentially choose unmated males over mated males with harems of two females. Additionally, the distribution of harem sizes suggests that females distribute themselves among males randomly. 4. The results suggest that harem size has effects on female reproduction that extend beyond larval competition and influence patterns of oviposition. The mechanism that determines why egg laying is greatest at intermediate levels is unknown. There is no evidence that smaller harems belong to lower quality males, but females may adjust egg‐laying behaviour in large harems as a result of reduced male attendance or anticipated larval competition.  相似文献   

14.
Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third (preferred hosts) or fourth (non-preferred hosts) instar larvae of Spodoptera littoralis (Boisd.) were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female' s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female' s life span, oviposition rate and host size from which the female was derived. These results suggest that: (i) M. rufiventris wasp is a weak synovigenic species; (ii) the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; (iii) the egg load of the newly emerged female is significantly less than the realized fecundity; and (iv) because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.  相似文献   

15.
Although female insects generally gain reproductive benefits from mating frequently, females do not mate unlimited numbers of times. This study asks whether the limit on female mating rate is imposed by trade‐offs between reproduction and survival. Female Gryllus vocalis were given the opportunity to mate 5, 10, or 15 times with novel males, and the effects on daily fecundity (egg production), fertility (proportion of eggs that were fertilized), and female post‐experimental longevity were measured. Females that mated 10 times laid more eggs and had a higher proportion of fertile eggs than females that mated 5 times. However, females that mated 15 times did not lay significantly more eggs or have a higher proportion of fertile eggs than females that mated 10 times. Although number of matings did not affect the date that females laid their last egg, mating more times was associated with a prolonged period of laying fertile eggs. Number of matings did not affect female post‐experimental longevity. Thus, there was no trade‐off between female reproductive effort and survival, even when females mated very large numbers of times. When females were allowed to mate ad libitum, the average number of times that females mated was greater than the number of times that confers maximal fitness. The lack of cost to mating explains why females might be willing to mate beyond the point of diminishing reproductive returns.  相似文献   

16.
ABSTRACT Recent work suggests that avian egg color could be a sexually selected signal to males that provides information about female condition, female genetic quality, or maternal investment in eggs. Theory predicts that egg color should influence male investment if it is an honest signal of the marginal fitness returns on paternal investment; a male should invest more in a colorful clutch if that investment increases offspring success more than an equivalent investment in a less colorful clutch. Some experimental support for this hypothesis has been found for species that lay blue eggs containing the pigment biliverdin, a potentially costly antioxidant. However, the brown eggshell pigment protoporphyrin, a pro‐oxidant associated with poor female condition, has received less attention as a potential predictor of female quality or investment. We performed a cross‐fostering experiment with House Wrens (Troglodytes aedon) in southwest Michigan in 2007 to test whether brown egg color was related to female condition or maternal investment, and whether male provisioning of nestlings was related to egg color. We swapped entire clutches between nests and measured egg characteristics and parental provisioning rates. We found that brighter eggs (i.e., those with less brown pigment) were heavier, and that nestlings that hatched from brighter eggs were fed at higher rates by their foster mothers, but not by their foster fathers. This pattern is consistent with the hypothesis that egg color is a potential signal of egg quality and female investment, but we found no evidence of a male response to this potential signal. This lack of a response could be the result of methodological limitations, a nonadaptive biological constraint, or adaptive indifference because chicks from brighter eggs do not actually yield increasing marginal returns on paternal investment. Clearly, additional study is needed to differentiate among these possibilities.  相似文献   

17.
Summary Samples of ovigerous female Ceratoserolis trilobitoides carrying newly spawned (stage A1) eggs in the marsupium were obtained from Borge Bay; Signy Island (South Orkney Islands), the continental shelf near Elephant Island (South Shetland Islands), and two sites in the inner Weddell Sea. The dry mass of a newly spawned egg was greater at the inner Weddell Sea sites even when the larger size of the females from the Weddell Sea and the positive relationship between egg size and female size were taken into account. Larger eggs contained more yolk, and there was a slight but significant tendency for larger eggs to have a lower percentage nitrogen content. Eggs from Ceratoserolis meridionalis were smaller in size than sympatric Ceratoserolis trilobitoides, but of similar composition. The reason for a greater investment per egg by females at higher latitudes is not clear, but it may be related to a slightly longer development period, itself associated with the lower water temperatures in the inner Weddell Sea.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation  相似文献   

18.
Sacha Haywood 《Ibis》2016,158(1):195-198
Sensory mechanisms controlling avian clutch size have diversified into distinct types, according to the nature of the input that is used to disrupt the growth of ovarian follicles and hence halt egg‐laying. In an article on brood parasitism, Lyon (2003) claimed that female American Coots Fulica americana can reduce their clutch size on the basis of visual cues in response to eggs laid in their nests by other females; in this species, therefore, egg counting would be used to control clutch size. After a close examination of the physiological determination of clutch size in American Coots, I show that seven of 17 parasitized clutches were smaller than the range controlled through the mechanism using an input to disrupt follicular growth (7–10 eggs per clutch). My reanalysis suggests that American Coots are incapable of adjusting clutch size via counting and re‐asserts that a species that can count eggs has yet to be found among birds that rely upon their own body heat for incubation.  相似文献   

19.
1. Whether the potential fecundity of herbivorous insects is realised or not may depend on female behaviour, which in turn may be influenced by host plant acceptability. Female Bupalus piniarius were observed to discriminate against needles growing out the year following defoliation (current‐year needles) of its host plant Pinus sylvestris. 2. It was hypothesised that the discriminatory behaviour was due to current‐year needles being less secure as a substrate. Field and laboratory experiments were designed to test this hypothesis and to estimate the discrepancy between potential and realised fecundity when females were offered defoliated branches. 3. In a laboratory oviposition experiment, B. piniarius females were exposed to branches bearing either current‐year needles only or both mature and current‐year needles. Daily oviposition rate, egg batch size, longevity, and mature eggs and fat retained at death were recorded for each female. In field experiments, the rate at which eggs dropped from expanding needles and the capacity of neonate larvae hatching from the dropped eggs to colonise a tree were assessed. 4. Significantly fewer eggs were laid when females were exposed to defoliated branches. 5. Twenty‐six and 16% of the eggs laid on current‐year needles dropped from the needles in 1998 and 1999 respectively, whereas no eggs dropped from mature needles in 1998 and only one egg (< 1%) dropped in 1999. 6. A very small proportion of larvae hatching on the forest floor (simulated egg drop) was able to recolonise host trees. 7. These results emphasise the importance of oviposition behaviour on realised fecundity when analysing insect population dynamics. In the case of B. piniarius, egg placement, although a minor detail during the normal course of events, became of key importance when defoliation deprived females of their preferred egg attachment site.  相似文献   

20.
Pepper weevils, Anthonomus eugenii Cano (Coleoptera: Curculionidae), feed and oviposit in flower buds and small fruits of plants in the genus Capsicum, as well as several species of Solanum (Solanaceae). Females chew a small hole into the fruit, deposit a single egg within the cavity, and seal the hole with a clear anal secretion that hardens into an ‘oviposition plug’. Female oviposition behavior was studied in a series of small‐arena bioassays to determine whether previous oviposition in Jalapeño pepper fruit deterred subsequent oviposition and to determine what specific cues from an infested fruit influence female behavior. In choice and no‐choice tests, females preferred clean fruit to fruit that had received four eggs 24 h previously (i.e., infested fruit), whether the fruit was infested with conspecific eggs or their own eggs. Further bioassays demonstrated that the presence of female frass, or oviposition plugs alone, in the absence of eggs or any fruit damage, was sufficient to deter oviposition. In addition, females given the choice between an infested fruit with the oviposition plug removed or an unaltered infested fruit preferred the fruit with no plugs, even when eggs, frass, and feeding damage were still present. To determine whether females would avoid infested peppers under more natural conditions, we quantified oviposition on infested and uninfested sentinel pepper fruit within individually caged plants and on clean and infested plants caged together. Females consistently laid more eggs on clean fruit than on infested fruits and moved within and among pepper plants to search for more acceptable oviposition sites. We conclude that oviposition plugs, along with contaminated female, but not male, frass contain a deterrent that, in the absence of any other cue, is enough to alert a female that a patch is occupied.  相似文献   

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