Effects of male removal on female foraging behavior in the Eurasian treecreeper

In old, spruce-dominated forests of central Finland, Eurasian treecreepers Certhia familiaris divide their territories spatially during the breeding season. Females forage primarily on the upper parts of the tree trunks, while males use the lower parts of the tree trunks. In this study we removed males from eight territories in the early nestling period to see if the mate's absence would change the foraging patterns of the resident female. Widowed females foraged at lower heights, thus behaving more like paired males. These females also spent less time on each tree and on each foraging bout than did paired females. We conclude that male removal facilitated the change in a female's foraging niche and foraging time at the trees. Females may re-optimize their foraging site selection owing to the absence of dominant males and a consequent need to increase their parental care. Behavioral plasticity may be the mechanism of niche partitioning between the sexes in this species. Received: 28 June 1996 / Accepted after revision: 27 March 1997     

Reproductive allocation in female house wrens is not influenced by experimentally altered male attractiveness

The differential allocation hypothesis proposes that females mated to attractive males should invest more resources in their offspring than those mated to less-attractive males, whereas the compensation hypothesis posits that females mated to less-attractive males should invest more resources in their offspring to compensate for lower-quality young. We tested these hypotheses by manipulating attractiveness of male house wrens (Troglodytes aedon) prior to female arrival by adding extra nest sites to territories of some males while leaving control males with only a single nest site. Females laid their eggs sooner in the nests of attractive males, and attractive males were more likely to retain their territory over successive broods and were marginally more likely to obtain a mate for a second brood later in the season than were control males, thereby confirming the effect of our manipulation on male attractiveness. Experimentally enhanced attractiveness also led to increased hematocrit in males. However, there were no consistent differences in the number, size, or quality of eggs laid by females mated to attractive and control males, nor were there any differences in the size, health state, or immune function of nestlings produced from these eggs. There was also no effect of treatment on the number of nestlings surviving to fledging. Collectively, these results are inconsistent with both the differential allocation hypothesis and the compensation hypothesis. Future studies should consider the possibility that the criteria used by females in selecting a mate may vary temporally and be more flexible than generally thought.     

Vocal and visual stimuli enabling copulation behavior in female buntings

Summary The loud and elaborate songs of male songbirds are throught to serve in territorial defense and to stimulate reproductive behavior in the female. We report here that in contrast to several other species, song alone is inadequate to induce sexual (lordosis) behavior in female indigo and lazuli buntings (Passerina cyanea and P. amoena); they require a more elaborate stimulus configuration to develop the full expression of sexual receptivity as indicated by a stereotyped copulation solicitation display. A live male must be present near the female, singing must ensue, and in addition a second and unique vocal utterance must be heard by the female; this is a soft buzzy-sounding vocalization, audible for no more than a few meters, not recognized previously as a critical arousal signal of the male songbird repertoire. Females also utter similar soft sounds in an intimate vocal exchange with the male leading to solicitation of copulation.     

Circadian rhythmicity of locomotor behavior and oviposition in female Teleogryllus commodus

Summary Locomotor activity of the Teleogryllus commodus is under circadian control, with _LL=25.3 h and _DD=23.4 h. In LD 12:12 h, running occurs either exclusively during darkness (57%), mainly in the dark phase (35%), or with substantial activity peaks in both phases (8%). For oviposition, _LL>24 h and _DD<24 h; in LD 12:12 h, 80% of the studied deposited their eggs mainly during the light phase and 20% primarily during darkness. In the discussion, a temporal correlation between stridulation, spermatophore formation, and locomotion, and oviposition is established, which serves to make an encounter and mating between the sexes as likely as possible.Dedicated to Prof. Dr. M. Gersch on the occasion of his 70th birthday     

Aggressive male mating behavior depends on female maturity in Octopus bimaculoides

This laboratory study examined the combined effects of male and female behaviors on the outcome of mating encounters in Octopus bimaculoides. We found that male–male competition for mating opportunities depends on female maturity; the presence of immature females elicited significantly higher levels of aggression between competing males. We conclude that males are able to assess the reproductive status of females. The study also found that immature and mature females resisted male mating attempts to a similar extent but that males that showed more aggression toward male competitors were able to spend more time in contact with females. We suggest that the lack of prominent visual displays in these mating trials indicates the importance of chemical cues in Octopus mating systems, as has been demonstrated for other cephalopods. This study contributes to the growing research on cephalopod mating systems and in particular shows that Octopus mating dynamics may be more behaviorally complex than initially assumed.     

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

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

Social influence on the correlation between behaviours in young-of-the-year perch

The connection between risk-taking behaviour and exploratory behaviour in young-of-the-year perch (Perca fluviatilis) was studied in aquarium experiments to see whether individual behaviour patterns could be identified in this species and also to investigate how individual behaviour is influenced by their social environment. Risk-taking was defined as the time spent foraging in an open area vs hiding in the vegetation in the presence of a piscivore. Explorative behaviour was measured as latency to enter a passage leading to an unknown area. Groups of four fish were used for the observations, and both behaviours measured were positively correlated with the mean scores of these behaviours in the other group members. Risk-taking and explorative behaviours were correlated only when data was adjusted for the behaviour of the other group members. Individuals that spent more time in the open than their companions also tended to be faster than the others to enter the passage to the unknown area and vice versa. The results indicate that there are consistent individual differences in boldness in perch, but also that behaviour could be modified according to the behaviour of group members.     

High degree of multiple paternity in the viviparous Shiner Perch, <Emphasis Type="Italic">Cymatogaster aggregata</Emphasis>, a fish with long-term female sperm storage

The Shiner Perch (Cymatogaster aggregata Gibbons) exhibits a viviparous reproductive mode and long-term female sperm storage, two biological features that may predispose this fish species for both intense sperm competition and frequent multiple paternity within broods. To test these hypotheses, we used polymorphic microsatellite markers to identify sires and quantify paternal contributions to the progeny arrays of 27 pregnant females from a natural population of C. aggregata. The number of sires per brood ranged from one to eight (mean 4.6), typically with skewed distributions of fertilization success by the fathers but no correlation between sire number and brood size. The extraordinarily high incidences of multiple paternity in this species probably are due in part to high rates of mate encounter, but selection pressures related to the avoidance of maternal–fetal incompatibility may further have promoted the evolution of polyandrous mating behaviors in this female-pregnant species. Our genetic data are consistent with the hypothesis that viviparity, long-term sperm storage, and extreme polyandry are interrelated reproductive phenomena that should promote the evolution of post-copulatory sperm competition and/or cryptic female choice in these fishes.     

Gill lesions in the perch, Anabas testudineus, exposed to monocrotophos

Histopathological effects of sublethal doses of monocrotophos on the gills have been studied by exposing the fish for a period ranging from ten to twenty days. The extent of damage to gills was dependent on the dose and duration of exposure. Histopathological changes in the gills observed were characterized primarily by hemorrhage in the primary and secondary gill lemellae. Degeneration and necrosis of epithelial cells were very prominent. Distortion of the secondary lamellae was very prominent, beginning with disruption of epithelial cells from pillar cells.     

Reproductive success increases with degree of kinship in cooperative coalitions of female red howler monkeys (Alouatta seniculus)

Evaluation of evolutionary mechanisms proposed to promote cooperative behavior depends on the relative influence of the behavior on the reproductive success of individuals, the reproductive success of the group in which they interact behaviorally, and the degree of gene correlation among cooperators. The genetic relationship within cooperative coalitions of female red howler monkeys was examined for three populations with different densities and growth rates. Patterns of gene correlation change within coalitions is documented using data from the mitochondrial and nuclear genomes, and long-term census monitoring. Differences in fecundity and infant survivorship within and between groups of unrelated (rˉ=0) and related (rˉ≥ 0.25) females are compared. Females that emigrate from their natal groups form coalitions with other migrant females. These coalitions attempt to establish a territory and, once successful in producing offspring, exclude other females from feeding resources. Females in these coalitions had different mtDNA haplotypes and a genetically estimated mean r of 0, supporting demographic data on emigration patterns indicating that these females rarely have the opportunity to form coalitions with kin. Patterns of recruitment and rate of matriline development within social groups supported behavioral data indicating that females actively attempt to promote their own matriline as breeders over that of other females, and that some matrilines are more successful at this than others. Mean r among females was significantly higher in coalitions established as social groups for several generations (rˉ=0.44). In these groups, females all shared the same mtDNA haplotype, and mtDNA haplotype divergence was significantly higher between than within groups. Females in coalitions with kin had significantly higher reproductive success than females in unrelated coalitions in all populations. This difference was not a function of coalition size, number of males, socionomic sex ratio, or primiparity, although anecdotal evidence suggests that allomothering may compensate for inept new mothers in related coalitions more often than in unrelated ones. Differences in territory quality could not be ruled out as a potential causal factor in the saturated populations, but were unlikely in the low-density, growing population. There were substantial differences among long-established coalitions in overall reproductive output in all three populations, and this was significantly correlated with the number of breeding females. Increase in coalition size was a function of both group age and the behavioral tolerance among females. Regardless of the underlying reasons for the patterns observed, reproductive success clearly increases with degree of gene correlation among females within cooperative coalitions, and coalitions that recruit more daughters produce more offspring. The nature of the cooperative relationship among group females directly influences both of these outcomes. This is associated with substantial genetic differentiation among social groups within populations, creating conditions in which genetic tendencies towards cooperative behavior can become tightly associated with group reproductive success. Received: 15 September 1999 / Revised: 27 April 2000 / Accepted: 27 May 2000     

Effect of sperm depletion and starvation on female mating behavior in the water strider,Aquarius remigis

The resolution of intersexual conflict over mating should be dependent on the current state of each individual. In this study, I used a factorial design to examine the influence of two physiological factors, sperm depletion and food deprivation, on resistance to mating by females of the water strider, Aquarius remigis. Females employ several different mate-resisting tactics during an encounter with a male. Five measures of female resistance to mating were identified: jumping, rolling, dunking, time spent dunking, and struggle duration. Jump, roll, and dunk rates were highly correlated with each other and combined into one metric of resistance to mating (PC1) using principal components analysis. Time per dunk (T/D) and struggle duration were also analyzed. Discrete male behaviors during the struggle could not be identified. Two measures of female resistance, PC1 and T/D, were significantly lower in sperm-depleted females than in sperm-replenished females. Struggle duration did not differ between the two treatments. Starvation had no effect on any of the measures of resistance. Sperm depletion significantly enhanced the probability of mating (54% vs. 24% for replenished females), while starvation had no effect on the probability of mating. I pooled all the females and compared females that mated with those that did not mate. Nonmating females resisted significantly more than mating females in all three measures of resistance. Path analysis indicated that PC1 was the only measure of resistance that was significantly negatively related to the probability of mating. Almost half (46%) of sperm-depleted females showed no resistance to males, while only 3% of sperm-replenished females were nonresistant. When nonresisters were removed from the analysis, sperm depletion had no effect on any of the measures of female resistance to mating and no effect on the probability of mating. In A. remigis, female resistance appears to be a yes/no phenomenon with respect to sperm depletion and not affected directly by starvation. Received: 2 September 1994/Accepted after revision: 9 September 1995     

The effect of mating on female sexual behavior of Teleogryllus commodus Walker

Summary Virgin Teleogryllus commodus females show rhythmic locomotor activity in light-dark (LD) and constant light (LL) conditions. A taped 3-h calling song stimulates them into further walking, and they lay few eggs. Successful mating suppresses or reduces circadian-controlled walking, but in LL-conditions the free-run still persists. Mated females no longer respond to the 3-h calling song. The number of eggs laid within 24–72 h after insemination increases, and the three effects of mating occur independently of one another. The possible physiological basis of these changes and their ecological and behavioral significance under field conditions are discussed.     

Testosterone-induced depression of male parental behavior in the barn swallow: female compensation and effects on seasonal fitness

In birds, many aspects of male socio-sexual and parental behavior are influenced by androgens, most notably testosterone (T). We report the effects of subcutaneous T-implants in male barn swallows (Hirundo rustica) on male and female parental behavior and on seasonal reproductive success. Males were assigned to one of three experimental groups: (i) implanted with a T-filled Silastic tube; (ii) implanted with an empty Silastic tube; and (iii) not implanted. T-implanted males provided a smaller proportion of feedings (number of feedings by the male/total number of feedings by both parents) and fed nestlings less frequently (number of feedings/h) than males of the other two groups. Females paired to T-implanted males fed nestlings significantly more often than females paired with unimplanted males. Females almost fully compensated for their mates' shortfall, and this resulted in similar combined feeding efforts among treatments. Reproductive success in their first broods or during the entire breeding season was unaffected by T- treatment. These results confirm earlier reports of the suppressive effects of T on male parental behavior. However, they are inconsistent with current ESS models that predict partial compensation as the optimal response by one individual to reduction of parental effort by its mate in monogamous, biparental systems.     

Reproductive traits in dicyemids

Several characters involved in the life cycle in dicyemids were examined to understand reproductive strategy and adaptations to cephalopod hosts. In most dicyemids distinctly small numbers of sperms are produced in a hermaphroditic gonad (infusorigen). The number of eggs and sperms are roughly equal (means of the number of sperm:egg=1:1.58). An inverse proportional relationship was found between the number of infusorigens and the gametes, suggesting a trade-off between them. Fecundity was positively correlated with the body size of adult stages (nematogens and rhombogens). Fecundity of a single dicyemid is not very high compared with that of the other endoparasite taxa, but a total reproductive capacity per community is high, because a great number of individuals multiply asexually in the renal sac. The size of mature infusoriform embryos (dispersal embryos) that develop from fertilized eggs was not correlated with their adult sizes, but the size of embryos was correlated with the maximum mantle length of the host octopus species. Although at present the process of infecting new hosts is still unknown, the size of the infusoriform embryo is likely determined by host-specific factors in this process. The size of vermiform embryos that are asexually formed from agametes was positively correlated with size of the adults; however, the number of vermiform embryos present in the axial cell of adults was not correlated with size of the adults. A correlation was not found between maximum mantle length of the host cephalopod species and length of the adult dicyemids. In dicyemid species the size of adults appears to be constrained by the renal habitat, including renal-pancreatic complex and branchial hearts of each host cephalopod species. Size thus may be determined by the volume of the renal sac, the diameter of the renal tubules, or the depth of folding in the surface of glandular renal appendages of cephalopods.     

Reproductive behavior of three tube-building peracarid crustaceans: the amphipods Jassa falcata and Ampithoe valida and the tanaid Tanais cavolinii

Laboratory observations indicate that Jassa falcata, Ampithoe valida, and Tanais cavolinii have quite similar patterns of behavior: males attend females until ovulation and copulation occur, then the male and female separate. Females tend to remain inside their own tubes, and males tend to move between the tubes of receptive females. This pattern of behavior (called “cruising males”) may be common in crustaceans that exhibit some fidelity to a specific site, especially if the females produce several broods in succession, do not store sperm, and reproduce asynchronously relative to each other. Male J. falcata have a terminal molt which is marked by the presence of a specific sexually dimorphic characteristic, the “thumb” on the second gnathopod. Alternating ovulatory and anovulatory molts occur in T. cavolinii.     

The effect of reproductive condition on the foraging behavior of female hoary bats,Lasiurus cinereus

Summary Female mammals experience larg changes in time and energy budgets associated with reproduction and these may influence the foraging strategies of individuals. I studied the changes in foraging behavior associated with reproduction in female hoary bats, Lasiurus cinereus. As lactation progressed, individuals departed to forage earlier in the evening and spent more time foraging per night and less time roosting with their young. Foraging time increased by at least 73% between early lactation and fledging and then declined as the young became independent. Females with two young foraged for longer than did those with one and females with pre- and postfledging young foraged in different habitats. The changes in foraging time suggest that foraging activity of female L. cinereus is constrained and individuals act as time minimizers, adjusting their foraging behavior to meet current energy demand. Predation risk is unlikely to constrain the behavior of these bats. However, maximizing energy intake throughout lactation may not be the optimal strategy because storing excess energy increases flight cost and may reduce foraging efficiency. The need to keep newborn young warm may also influence foraging time. Such constraints, causing changes in foraging activity, may alter the availability of habitats and prey and must be considered when modelling foraging strategies. In addition, changes in flight time may significantly alter the energy budgets of bats in different stages of reproduction.     

Male calling behavior,female discrimination and acoustic interference in the Neotropical treefrog Hyla microcephala under realistic acoustic conditions

Summary Anuran choruses are acoustically complex assemblages of calling males. Little is known about the behavior of males or females in such natural sound environments. I studied calling behavior of males of Hyla microcephala in nature by using an interactive computer-based system that allowed me to simulate call interruptions by a number of males. I also monitored the calling behavior of groups of four to six males. When a male is interrupted by the call of another frog, he increases the spacing between the notes of his call. Responses of this kind are strongest to the loudest neighbor, and some males may ignore interruptions by all but a single close male. Interruptions using synthetic calls with silent gaps indicated that males respond vocally to reductions in sound intensity as brief as 20 ms. This ability helps to explain how males can rapidly alternate notes during pairwise interactions. Amounts of acoustic overlap between pairs of males in the choruses were usually below 10% of an individual's total calling time during bouts. The time a male spent calling that was free of acoustic interference by any other male ranged from 34–92% of his total calling time. When group size was decreased, this unobstructed calling time increased. Previous research showed that females of H. microcephala discriminate against calls that overlap so that the call pulse-train structure is degraded. Here I show that a 6 dB difference in intensity between the overlapped calls is sufficient to reduce the degradative effect of call interference. Females were also given a choice between interfering calls broadcast from two adjacent and two widely separated speakers. An angular separation between speakers of 120° was insufficient to elicit a preference for the separated sources. Together, data on behavior of males and females indicated that males actively reduce acoustic interference with those loud individuals most likely to degrade seriously the temporal structure of their calls.     

Evidence for a female sex pheromone mediating male trailing behavior in the red-sided garter snake, Thamnophis sirtalis parietalis

Summary. Male garter snakes locate females during the breeding season utilizing conspecific trailing behavior. It has been hypothesized that the female-derived chemical cue responsible for mediating male reproductive trailing behavior is the sexual attractiveness pheromone, a previously characterized contact pheromone responsible for releasing male courtship behavior. To examine this hypothesis, we tested the response of male red-sided garter snakes, Thamnophis sirtalis parietalis, to pheromone trails produced by males, females and ‘she-males’. She-males are a small subset of males in the natural population that are morphologically and behaviorally similar to other males except that they produce and express the sexual attractiveness pheromone during the breeding season. When tested on a Y-maze, males were found to detect and follow the skin lipid trails of females and she-males, but displayed no behavioral responses to male trails. In addition, males were unable to discriminate between she-male and female trails when given a choice. These results are consistent with the hypothesis that the sexual attractiveness pheromone is the chemical cue primarily utilized by males to mediate reproductive trailing behavior.     

Mating in the red-sided garter snake,Thamnophis sirtalis parietalis: differential effects on male and female sexual behavior

Summary Female red-sided garter snakes, Thamnophis sirtalis parietalis, become unattractive to most males after mating in the field and in the laboratory. Male red-sided garter snakes vary in their latencies to court attractive females following copulation, with courtship resuming in minutes to hours. Unsuccessful males in mating balls disperse from mating pairs, but are not residually inhibited from courting attractive females. These patterns of behavior indicate that males have evolved mechanisms to maximize opportunities for copulation with several females, while females mate only once per season.