Exposure to strange adults does not cause pregnancy disruption or infanticide in the gray-tailed vole

A widely accepted paradigm in mammalian behavioral biology is that exposure to unfamiliar males causes pregnancy disruption in female rodents (commonly known as the Bruce effect). This behavioral phenomenon has been demonstrated in the laboratory with at least 12 species of rodents, primarily within the genus Microtus, and is supposedly an adaptation that provides male perpetrators with reproductive access to females, and functions, for females, as a counterstrategy to infanticide. However, neither the Bruce effect nor its adaptive significance have been tested experimentally in the field. In a controlled field study, we exposed reproducing female gray-tailed voles (Microtus canicaudus) to treatments in which males were removed and replaced by either unfamiliar males or females, and found no significant differences in intervals between parturitions, number of pregnancies, and juvenile recruitment among the treatment and control animals. Thus, we conclude that neither the Bruce effect nor infanticide occurred differentially as a consequence of the treatments in gray-tailed voles. Multimale mating to confuse paternity, and postpartum estrus resulting in simultaneous pregnancy and lactation may deter infanticide and functionally negate any benefits of pregnancy disruption in gray-tailed voles and perhaps other murid rodents with similar mating systems. In light of our results, we recommend field verification for other species of murid rodents that exhibit the Bruce effect in the laboratory before the results are applied to evolutionary theory. Received: 12 May 1998 / Accepted after revision: 6 September 1998     

Pregnancy blockage following multiple-male copulation or exposure at the time of mating in deer mice,Peromyscus maniculatus

Summary Pregnancy blockage resulting from multiple-male copulation or exposure around the time of mating was studied in three experiments on deer mice, Peromyscus maniculatus. First, significantly more females were pregnant after copulating, either with or without disturbance and a delay, with one male than after receiving the same total number of ejaculations from two males. Second, when females received three ejaculations from one male and then were immediately exposed to either a blonde male or a wild-type male for 2 h, the pregnancy rate was lower than when they were exposed to an empty cage. Third, when immediately after mating females were placed for 2 h in the cage of a strange male they were less likely to deliver a litter than when placed in the familiar male's cage. This phenomenon bears at least a superficial resemblence to the Bruce effect but either communality or difference in underlying mechanism remains to be determined. The present multi-male pregnancy block may provide a means for females in unstable social situations to defer reproduction for 4 days or longer when more stable conditions may exist and a single male prevails.     

Length polymorphism at the avpr1a locus is correlated with male reproductive behavior in a natural population of prairie voles (Microtus ochrogaster)

Laboratory studies have shown that vasopressin can influence sociosexual behavior through its action on the vasopressin 1a receptor (V1aR). There is substantial evidence that the length of a microsatellite in the gene (avpr1a) encoding for the V1aR can affect social attachment to females and paternal behavior in male prairie voles under laboratory conditions. However, previous field studies of prairie voles have failed to detect a strong effect of the length of a male’s avpr1a allele on their sociosexual behavior, but these studies are typically much shorter than the average prairie vole breeding lifespan. We examined the relationship between male avpr1a microsatellite allele length and sociosexual behavior in a natural population of prairie voles for 15 weeks, closer to the lifespan of prairie voles in nature. Contrary to predictions, we found that males with the longest avpr1a microsatellite alleles were significantly more likely to sire offspring with more than one female and to sire offspring that survived until trappable age than males with the shortest avpr1a microsatellite allele lengths. This relationship was the strongest for males with the longest tenure on the study site. As in previous field studies, we did not find evidence of a relationship between a male’s avpr1a genotype and any index of social behavior including male residency status or the number of females with which males associate. This is the first study to support the hypothesis that a male’s avpr1a genotype is a factor underlying variation in the genetic mating system of prairie voles under natural conditions.     

Effects of Habitat Loss and Fragmentation on the Behavior and Demography of Gray-Tailed Voles

We monitored the short term behavioral and demographic responses of gray-tailed voles (Microtus canicaudus) to the reduction and fragmentation of their habitat. Our objectives were (1) to test whether animals perished or moved into remaining fragments after 70% of their habitat was removed; and (2) to test the null hypothesis that the social structure and demography of animals would not differ between habitats consisting of one large continuous fragment (625 m²), a mosaic of 25 small fragments (each 25 m²) separated by 4 m of bare ground, and control, unmanipulated habitats (1850 m²). We conducted the experiment in 12, 0.2-ha enclosures planted with alfalfa with four replicates for each of two manipulated treatments and a control. A 70% reduction in habitat did not adversely affect adult survival, reproductive rate, juvenile recruitment, or population size. However, an influx of unrelated females into habitat fragments resulted in decreased juvenile recruitment in those fragments. Voles from cleared habitat moved into the remaining habitat and did not measurably affect the resident population. Similarly, the demography of voles did not differ significantly among the large-fragment, small-fragment, and control enclosures. Peak density estimates based on the amount of habitat in each enclosure were 545 animals per hectare in control, 1056 in large-fragment, and 2880 in small-fragment enclosures. Reduced movement of animals among the small fragments was the most obvious effect of habitat fragmentation. Six percent of females and 15% of males moved among small fragments within a week compared to approximately 60% moving comparable distances in large-fragment and control enclosures. Rates of juvenile dispersal and sexual maturation declined throughout the summer on all treatments, were associated with season and density, and were only marginally associated with habitat loss and fragmentation. We conclude that at the time of habitat removal and fragmentation, populations were small enough to accommodate a 70% reduction in habitat and still continue to increase in numbers. The social system of gray-tailed voles was sufficiently flexible to accommodate an influx of animals to withstand densities> 1000 voles per ha. The behavioral and demographic features of gray-tailed voles are similar to those reported for other small mammals, thus confirming the use of voles for ecological model systems in habitat fragmentation studies.     

Female-biased immigration and male peace-keeping in groups of the shell-dwelling cichlid fish Neolamprologus multifasciatus

The small cichlid fish Neolamprologus multifasciatus is endemic to Lake Tanganyika and is unique among fish because it lives in complex social groups with several reproductive males and females, all of which participate in defending their territory against neighbors and intruders. Individuals use empty snail shells for breeding and shelter. Previous parentage analysis using microsatellites suggested occasional exchange of individuals between groups. In field experiments, we found that females showed a higher tendency than males to migrate into territories already occupied by a resident pair. The phenomenon and causes of female-biased immigration were further investigated in aquarium experiments: Nine of 15 females, but only 2 of 15 equally sized males, settled in territories of established pairs. Territorial males exhibited more aggression toward strange males (potential reproductive competitors) than toward strange females (potential additional mates); their females were more aggressive toward strange females (probably competitors for shells) than toward strange males. Apparently, a conflict exists between the sexes regarding the immigration of additional females. This conflict seems to be the selective basis for observed active male interference in aggressive disputes between females in the territory. Interfemale tolerance in a group was greater in the male’s presence than in his absence. Received: 1 February 2000 / Received in revised form: 2 May 2000 / Accepted: 5 May 2000     

Reproductive cycle of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama

The reproductive status of the holothuroid species Isostichopus badionotus (Selenka, 1867) and Holothuria mexicana (Ludwig, 1875) was studied over 16 months in Bocas del Toro (Panama), from November 1999 to February 2001. Sexual reproduction was evaluated by the gonad index method, and by histology of gonad development. In addition, population structure was assessed based on sex ratio, minimum reproductive size, and length and weight distributions of males and females. The sex ratio in both species was 1:1, with a unimodal population distribution composed mainly of mature individuals. The minimum reproductive length and weight were 13-20 cm and 150 g, respectively, for both species, although reproductive individuals 10 cm in length were also found. A consistently higher gonad index was observed in H. mexicana, due to a high proportion of mature females and males and high gonad indices in most monthly samples. Gametogenesis and spawning patterns seemed to occur throughout the year, with periods of enhanced activity. Two periods of maximum reproductive activity were tentatively identified: July-November for I. badionotus and February-July for H. mexicana, but neither species had a single, sharply defined annual spawning event. Further work on these exploited holothuroids should examine the relationships between reproduction and environmental factors and between reproductive status and recruitment.     

Reproductive biology of a small isopod symbiont living on a large isopod host: from the maternal marsupium to the protective grip of guarding males

Mating systems of many symbiotic crustaceans are characterised by a high degree of mate guarding. A peculiar case of mate guarding has been reported for small symbiotic janirid isopods where males mate with immature females. Field samples of individual hosts and laboratory experiments were conducted to reveal the mating behaviour of the symbiont in a natural environment, that is, on their hosts. Along the coast of the Magellan Strait, Chile, the janirid isopod Iais pubescens was frequently found on the shore-living isopod Exosphaeroma gigas. Symbiont prevalence (percent hosts occupied) was high at eight of the nine sampling sites. Mean symbiont intensity was very low at one site (<<1 individual host^-1), intermediate at two sites (1-10 individuals host^-1) and high at the other sites (10-40 individuals host^-1). The mean sex ratio (males:females) was male biased at most sampling sites (n=7). Females of I. pubescens reached substantially larger sizes (1.5-3.0 mm body length, BL) than males (1.1-1.9 mm BL). The majority of males were carrying small juveniles (66.15%), and males with juveniles were significantly larger than males without juveniles - this suggests that males prefer virgin juveniles to adult females and that they compete for small juveniles. In laboratory observations, males were seen to manipulate the marsupium of adult females that were about to release small juveniles. Males obtained virgin juveniles in this manner. Juveniles were carried for ~7 days, and they moulted shortly before being fertilised and released by males. The high proportion of juveniles carried by males in the field (68.2%) supports previous observations that males initially are not able to distinguish male and female juveniles. It is suggested that the mating system of symbiotic janirid isopods with long-term sperm storage and continuous receptivity in females and male mating with virgin females has evolved in response to highly unpredictable encounter probabilities between the sexes. Mate guarding and manipulation of small virgin juveniles may be favoured on the highly mobile hosts of symbiotic janirid isopods. Furthermore, adult females may gain by leaving their emerging offspring in the protective grip of guarding males, thereby reinforcing the maintenance of this peculiar mating system.     

Field-testing the protandry-based mating system for the lizard, Anolis carolinesis: does the model organism have the right model?

Using the lizard Anolis carolinensis as its subject, an early and enduring model of psychobiology was presented in which neuroendocrines and social behavior interact to coordinate reproduction between the sexes. The adaptive context for the model was protandry, here defined as a "male-first" emergence pattern from winter dormancy to the onset of breeding. In the protandry scenario, males emerge, become reproductive, and contest for territories. Then, pre-reproductive females emerge to settle on male territories, whereby ovarian recrudescence is facilitated by the behavior of courting males. A collateral inference of the protandry sequence is that females are choosing their future mates from among local males and their territories (i.e., intersexual selection). The model and its assumptions, though never validated by field data, have served as the paradigm for many laboratory experiments on A. carolinensis. Recently, however, field studies described an intrasexually selected mating system for A. carolinensis, without evidence of direct female choice. Differences between the selective inferences from the two perspectives led us to test in the field a number of protandry-based characteristics previously ascribed to A. carolinensis. We monitored free-ranging adults across the winter-to-breeding period using data on sex ratios, social behavior, gut contents, sex steroids, and gonadal condition. We found no evidence of protandry, and little support for any of the associated assumptions. Instead, the sexes broadly overlapped in their temporal transition from winter retreats to an active status in the habitat, with average male reproductive development about 1-2 weeks in advance of females. We replace the traditional protandry-based A. carolinensis paradigm with a realistic onset sequence into the breeding season and a new model for the species' mating system. Our study underscores the need for field validation when laboratory-generated data are fitted to adaptive paradigms.     

The assessment of female reproductive state during courtship and scramble competition in the fiddler crab, Uca paradussumieri

Male fiddler crabs, Uca paradussumieri, mate underground during a 4- to 7-day period each full and new moon. As soon as the tide recedes, males enter the burrows of females that will ovulate the following day ('pre-ovigerous' females). Males copulate with and guard these females until they ovulate. When interrupted by an intruding male, the first male to reach the female is usually able to defend her and successfully mate with her. In fiddler crabs, females mate multiply and there is last male sperm precedence. Before each semi-lunar mating period, male U. paradussumieri were more likely to court females with whom they would later mate than other nearby females with whom they did not mate. This suggests that males collect information on female reproductive state prior to the females becoming ovigerous. In this species, aggression was common between males that courted the same female. When previously courted females were approached by other males, the initial courter attempted to forcefully disrupt the courtship. This behavior may allow males the exclusive use of information on female reproductive condition. It also suggests a type of scramble competition between males over females. Furthermore, it indicates that males are able to locate receptive females prior to their becoming ovigerous. The shorter guarding period observed in this species, as compared with other fiddler crabs, is caused by females rejecting longer guarding periods. Male ability to assess female reproductive status may therefore be advantageous because it increases male mating success within a scramble type of competitive polygyny.     

Male dominance,female choice and male copulatory behavior in two species of voles (Microtus ochrogaster and Microtus montanus)

Summary Female preferences for dominant males in prairie and montane voles were analyzed in two different test situations. In the first, prairie vole females preferentially spent time in proximity to dominant versus subordinate males which were housed behind a wire mesh screen. In a two-male tether test prairie voles females both preferentially associated and mated with dominant males. Montane voles, on the other hand, showed no preference in either test situation. The baseline copulatory behavior of naive montane vole males which became dominant differed significantly from those which became subordinate; no such differences were evident in the baseline copulatory behavior of naive praire voles. One hour following dominance testing, there was no difference evident between dominant and subordinate males of either species but there was a general facilitation of male copulatory behavior in both species, the effect being of greater magnitude in montane voles.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Reproduction in a simultaneous hermaphroditic shrimp, Lysmata wurdemanni: any two will do?

The caridean shrimp Lysmata wurdemanni (Gibbes) displays protandric simultaneous hermaphroditism with out-crossing, but not all males become simultaneous hermaphrodites (euhermaphrodites). In this laboratory study, we attempted to determine why some shrimp remain males. In our experiment, we grew L. wurdemanni from postlarvae to adults in several group sizes and observed their reproductive function. We found that all shrimp reared in isolation become euhermaphrodites. When cultured in a group, the proportion of shrimp remaining male decreased with increasing group size. Except for those that mated within a day, inter-molt euhermaphrodite-phase shrimp (with or without embryos) and inter-molt male-phase shrimp fertilized eggs successfully. On the other hand, euhermaphrodite shrimp can only mate as females and have their eggs fertilized during a narrow post-molt window (less than 12 h) in each molt cycle (10 days). The fertilization rate of male-euhermaphrodite pairs was similar to that of euhermaphrodite-euhermaphrodite pairs. There are at least two non-exclusive explanations for the persistence of male shrimp in a group. In certain group compositions, an individual may gain more reproductive fitness as a large male with multiple mating partners than as a small female with low clutch size. Alternatively, the presence of male-phase individuals, with variable molt-cycle duration (5-8 days), may be necessary to ensure mating. This study is the first direct experimental demonstration of social control of sex change in the decapod crustaceans.     

Reproduction in a simultaneous hermaphroditic shrimp, Lysmata wurdemanni: any two will do?

The caridean shrimp Lysmata wurdemanni (Gibbes) displays protandric simultaneous hermaphroditism with out-crossing, but not all males become simultaneous hermaphrodites (euhermaphrodites). In this laboratory study, we attempted to determine why some shrimp remain males. In our experiment, we grew L. wurdemanni from post-larvae to adults in several group sizes and observed their reproductive function. We found that all shrimp reared in isolation become euhermaphrodites. When cultured in a group, the proportion of shrimp remaining male decreased with increasing group size. Except for those that mated within a day, inter-molt euhermaphrodite-phase shrimp (with or without embryos) and inter-molt male-phase shrimp fertilized eggs successfully. On the other hand, euhermaphrodite shrimp can only mate as females and have their eggs fertilized during a narrow post-molt window (less than 12 h.) in each molt cycle (10 days). The fertilization rate of male-euhermaphrodite pairs was similar to that of euhermaphrodite-euhermaphrodite pairs. There are at least two non-exclusive explanations for the persistence of male shrimp in a group. In certain group compositions, an individual may gain more reproductive fitness as a large male with multiple mate partners than as a small female with low clutch size. Alternatively, the presence of male-phase individuals, with variable molt-cycle duration (5-8 days), may be necessary to ensure mating. This study is the first direct experimental demonstration of social control of sex change in the decapod crustaceans.     

Reproductive cycle and maternal effects on offspring size and number in the neogastropod Buccinum undatum (L.)

This paper examines relationships between maternal phenotype and offspring phenotype in the commercially exploited marine neogastropod Buccinum undatum (L.). The study also aimed to estimate the effects of female size on subsequent recruitment, and to determine the reproductive cycle of B. undatum in Swedish waters. The evaluation of gonad development in field-sampled animals from August 1998 to February 2000 revealed a definite annual reproductive cycle with a single major egg-laying period in the autumn. In addition, adult whelks of varying sizes from two locations and in two consecutive years were kept in aquaria to study the influence of female size on fecundity. Twenty-seven females deposited eggs in the laboratory. Offspring quality, measured as egg and hatchling size, was independent of female size, although larger females produced more eggs and hatchlings than smaller females, and thus probably account for a disproportionately large share of potential recruitment; the relative fecundity (number of eggs or hatchlings per gram of body weight) was constant with female size. I found no evidence for a longer time interval between successive egg-layings in larger females, i.e. mature females of all sizes probably spawn equally often. The present results therefore indicate that spawning-stock biomass is probably a better measure of spawning-stock size than the number of mature individuals in the population. Effects of an individual female's characteristics on stock productivity have largely been overlooked by fishery managers but could help to explain the weakness often found in stock-recruitment relationships.     

Foraging decisions of American robins: somatic and reproductive tradeoffs

Summary Breeding units (occupants of a nest including at least one reproductive female) within two free-living populations of the prairie vole, Microtus ochrogaster, were monitored by live-trapping at nest during two 28-h periods each week from October 1980 to March 1984. Data are presented for 281 breeding units from all seasons, at high and low population densities and during breeding and nonbreeding periods. Fifty percent of the breeding units were monogamous (single resident reproductive male and female), 27% consisted of a single reproductive female with no resident adult male and 23% included more than one resident adult male and/or female (complex units). Monogamous units were present in the same proportions during breeding and nonbreeding periods. The number of monogamous units was significantly greater at low population densities than at high densities. During winter there were relatively more complex units and fewer single female units than during the rest of the year. Monogamous pairs remained together for an average of 42 days. Seventy-eight percent of these pairs were disbanded by the death of one or both members. There were few overlaps of the home ranges of adjacent breeding units. Significantly more nests were visited by nonresident males than by females, and the intervals between visits by males were significantly shorter than those for visits by females. Males visited single female units significantly more often than units with one or more resident males. Survival of juveniles was generally very low; 38% and 34% of young males and females, respectively, that were trapped survived until 30 days of age. Of young females remaining at the natal nest at low population densities, only 17.6% were reproductively activated; 77.1% of such females became reproductively activated at high densities. All young females that dispersed from the natal nest became reproductive.     

Size-assortative pairing across three developmental stages in the Zeus bug, Phoreticovelia disparata

The mechanisms underlying size-assortative pairing have received considerable attention. Typically, pairing is assumed to occur at, or just prior to, the adult phase of the life cycle. However, in many invertebrates, males commence associations with juvenile females who are more than a single moult away from sexual maturity. These species are ideal to explore the importance of reproductive and survival benefits as mechanisms driving size-assortative pairing. In the Zeus bug, Phoreticovelia disparata, adult males are found riding on juvenile (fourth and fifth instar) and adult females—a behaviour that is costly for females but has survival benefits for males. Using a combination of field collections and laboratory manipulations, we show that pairing is size-assortative both within and between female age classes and that riding males are smaller than non-riding males. In a series of mating trials, we revealed that males attempt to ride any female but that their riding success is dependent on female age. We also provide the first direct evidence of female resistance to male riding attempts in P. disparata. We propose that size-assortative pairing arises through adaptations that have evolved to minimise the potential costs of sexual conflict. We suggest that the selective pressure on males to maximise survival benefits is sufficiently high that it outweighs the reproductive benefits of discriminating against fourth instar females. Finally, given that female resistance is under direct selection in juvenile females, it is likely to be the main form of selective pressure for adult females.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

The economics of harem maintenance in the sac-winged bat, Saccopteryx bilineata (Emballonuridae)

Saccopteryx bilineata has a polygynous mating system in which males defend females in a harem territory. Harem defense and courtship include energetically costly flight maneuvers and hovering displays. We tested if (1) harem males have a greater field metabolic rate than non-harem males or females and if (2) the field metabolic rate of harem males is correlated with the number of females in a harem territory. We measured the energy budget in 32 S. bilineata with the doubly labeled water method and compared these estimates with behavioral observations in the daytime roost. Among adult bats, field metabolic rate varied with body mass by an exponent of approximately two. We found no significant difference in field metabolic rate or mass-specific field metabolic rate between harem and non-harem males. The mass-specific field metabolic rate of harem-males increased with harem size. The latter finding supports the hypothesis that the energy costs of courtship display and territorial defense influence the energy budget of harem males. Overall, field metabolic rates of S. bilineata were lower than those of similarly sized bats of the temperate zone and only 2.3 times above the basal metabolic rate recorded for this species. We suggest that male S. bilineata did not take advantage of their metabolic capacity because a prudent allocation of energy to activities of harem maintenance is an adaptive strategy for males in this mating system.     

A field test of the effects of familiarity and relatedness on social associations and reproduction in prairie voles

Inbreeding depression is a well-documented phenomenon. In animals, one means of avoiding the costs of inbreeding is through the recognition and avoidance of kin as mates. Prairie voles (Microtus ochrogaster) are short-lived, socially monogamous rodents that demonstrate inbreeding depression in the laboratory. Field data indicate that pair formation in nature is opportunistic but pairing among close relatives seems uncommon. We examined the role of relatedness and familiarity on prairie vole social associations and reproduction by placing adult voles into 0.1-ha enclosures with familiar siblings, unfamiliar siblings, and unrelated, unfamiliar conspecifics. Live-trapping data indicated that indices of social pair bonding were random with respect to relatedness and familiarity. Among females whose litters were sired by a single male, litters were significantly more likely to be sired by unfamiliar than familiar males, but the number of litters sired by males that were unrelated to their partner was not different from the number of litters sired by males that were related to their partner. Additionally, females that produced offspring with familiar siblings were significantly more likely to have litters with multiple paternity than females not producing offspring with familiar siblings. However, multiple paternity was not influenced by relatedness of sires. Finally, older individuals were more likely to produce offspring with each other than with younger individuals. Our findings suggest that prior association is a more important mechanism of inbreeding avoidance than phenotype matching for prairie voles mating under ecologically relevant conditions.     

A histological study of reproduction in the serpulids<Emphasis Type="Italic"> Pomatoceros triqueter</Emphasis> and<Emphasis Type="Italic"> Pomatoceros lamarckii</Emphasis> (Annelida: Polychaeta)

The reproductive cycles of the serpulid polychaetes Pomatoceros lamarckii (Quatrefages, 1865, as Vermilia) and P. triqueter (Linnaeus, 1767, as Serpula triquetra) were studied by histological examination during 1999 and 2000 at Bantry Bay, south-west Ireland. Gametogenesis, sex ratios and hermaphroditism were also investigated. The male/female ratio in P. lamarckii was approximately equal, but in P. triqueter it deviated significantly in favour of females. There was no significant variation in the sex ratios throughout the year in either species. Results confirm that the species are protandric hermaphrodites. In P. lamarckii, but not in P. triqueter, sexual dimorphism was exhibited in body size, the females being larger than the males of the same species. In both species, the proportion of females increased with increasing body size. Simultaneous hermaphroditism was recorded in P. lamarckii, but not in P. triqueter. There was no clearly defined annual gametogenic cycle in either species. Both species appear to have an extended reproductive season, with numerous small-scale peaks in reproductive maturity that can vary annually. Spawning was broadly synchronous between sexes. Visual observation alone (i.e. without histology) was insufficient to accurately assess reproductive condition in individuals not in, or close to, a ripe state.