Variation in homeward orientation performance in the sleepy lizard (Tiliqua rugosa): effects of sex and reproductive period

Sleepy lizards are monogamous skinks which show high pair fidelity. This study reveals inter- and intrasexual differences in homeward orientation performance in this lizard. Male and female lizards were displaced during three phases of the spring activity period, the pre-pairing, pairing/mating, and post-pairing periods. All groups (with the exception of post-pairing males) were significantly oriented homewards, but males were significantly better oriented towards home than females during the pairing period. Furthermore, males were significantly better homeward oriented during the pre-pairing and pairing periods than in the post-pairing period. Similar results were observed for rate of movement away from the release site. In sleepy lizards, sex-based differences in homing behaviour are unlikely to be attributable to differences in the area of familiarity, or availability of orientation mechanisms. However differences in homing motivation may explain these differences. Males may miss mating if absent from the home range during the pre-pairing and pairing periods, while females may still be able to obtain a mating even when absent. Females however may be more motivated than males to return to the familiar home range during the post-pairing period to ensure efficient feeding during internal embryo development. Received: 16 February 1998 / Accepted after revision: 28 March 1998     

Seasonal variation in the relationship between cellular immune response and badge size in male house sparrows (Passer domesticus)

The immunocompetence handicap hypothesis postulates that secondary sexual traits are honest signals of male quality because steroid hormones (such as corticosteroids and sex steroids), which are supposed to favor the development of secondary sexual traits, may also have immunosuppressive effects. Certain secondary sexual traits are not only used as mate choice signals but also play a role as badges of status. In the house sparrow (Passer domesticus), males have a bib of black feathers which is used both as a signal of social status in male-male interactions and by females when choosing a mate. We investigated the relationships between bib size and cellular immune response in male house sparrows during and outside the reproductive season. Males with large badges were found to have lower levels of immunocompetence, as assessed using a T-cell-mediated immunity assay, during the reproductive season, as predicted by the immunocompetence handicap hypothesis. Conversely, in November, the correlation between badge size and cellular immune response was positive, possibly reflecting the better access to trophic resources of large-badged dominant males in winter flocks. Received: 24 September 1998 / Received in revised form: 2 February 1999 / Accepted: 14 February 1999     

Ontogeny of diet changes in a tropical benthic carnivorous fish, Parupeneus barberinus (Mullidae): relationship between foraging behaviour, habitat use, jaw size, and prey selection

Parupeneus barberinus forages on benthic invertebrates using a wide range of foraging modes, including vigorous digging in the substratum, resulting in considerable disturbance to the benthos. Polychaetes were the most important prey item for all size classes, but fishes less than 120 mm total length consumed more small ostracods and nematodes than did larger fishes. Fishes greater than 120 mm total length consumed mostly bivalves, and fishes over 240 mm total length consumed mostly bivalves and crabs. A morphological examination of the feeding apparatus suggested that the size of important prey items consumed was determined by gape height and jaw width. Prey available to different size classes of fishes was determined by combining information on microhabitat use, foraging behaviours, and prey volumes in the substratum. Small fishes spent more time foraging on the reef flat and slope, compared with larger fishes that foraged mostly on the reef edge and base. In addition smaller fishes foraged mostly in the upper 2 cm of sediment, whereas larger fishes often foraged to depths of 10 cm. Selection ratios showed that different size classes of fishes selectively extracted different prey items from the substratum. Small fishes showed a preference for ostracods whereas large fishes selected for bivalves and crabs. Although polychaetes were the dominant prey item for all size classes, they were consistently selected against.     

Effect of group size and physiological state of a cagemate on reproduction in the parthenogenetic lizard,Cnemidophorous uniparens (Teiidae)

Summary Cnemidophorous uniparens, an all-female, parthenogenetic species of whiptail lizard (Teiidae), were housed in the laboratory under seminatural conditions in groups of four, in pairs, or as isolates. Group structure and an animal's position in the dominance hierarchy affect reproductive effort. Housed in pairs, the dominant animals exhibit a higher rate of reproduction, as measured by number of clutches and length of clutch cycle, than do their subordinate cagemates. Females housed with dihydrotestosterone-treated ovariectomized females lay more clutches than females housed with ovariectomized hormone-untreated females, those housed with intact females, or housed alone.     

Mating frequency, colony size, polyethism and sex ratio in fungus-growing ants (Attini)

The mating frequency of queens was estimated for eight attine ant species, Myrmicocrypta ednaella, Apterostigma mayri, Cyphomyrmex costatus, C. rimosus (four lower attines), Trachymyrmex isthmicus, Serico-myrmex amabalis, Acromyrmex octospinosus and Atta colombica (four higher attines), and correlated to colony size, worker polyethism, and sex ratio. Mating frequency was calculated from within-colony relatedness estimated by CAP-PCR DNA fingerprinting. Most queens of lower attines and T. isthmicus mated with only one male, while those of the three higher attines mated with multiple males. Mating frequency was positively correlated with colony size. Polyethism among workers was dependent on worker age in lower attines but on body size in higher attines, suggesting some correlation between mating frequency (i.e., within-colony gene diversity) and caste complexity. The sex ratio was biased toward females in species where the mating frequency equaled one, but toward males in species where the mating frequency was greater than two. Changing in nest site from ground surface to deep underground may have facilitated the evolution of large colony size in Attini, and this may have resulted in the evolution of polyandry (a queen mates with multiple males). With the evolution of polyandry in higher attines, Atta and Acromyrmex in particular have generated high genetic diversity within their colonies and complex social structures. Received: 26 October 1999 / Revised: 25 May 2000 / Accepted: 24 June 2000     

The relationship between epilithic periphyton (biofilm) bound metals and metals bound to sediments in freshwater systems

Surficial sediments and epilithic periphyton (biofilm) were sampled from six sites on the River Churnet and five sites on the River Manifold in Staffordshire and analysed for cadmium (Cd), copper (Cu), lead (Pb) and zinc (Zn). The sites demonstrated a wide range of sediment trace metal concentrations determined by inductively coupled plasma optical emission spectrometry (ICP-OES) and atomic absorbtion spectroscopy (AAS). Biofilm was removed from the substrate using physical abrasion and 0.005 M ethylenediaminoethanetetra-acetic acid (EDTA) extractant. The European Standards, Measurements and Testing Programmes (BCR) operationally defined geochemical speciation scheme was used to determine the exchangeable, acid soluble fraction of the sediments. Significant positive correlations were determined between the EDTA extractable biofilm and the exchangeable sediment fraction for Cd, Cu and Zn but not for Pb. Natural epilithic periphyton may be a potential metal biomonitor particularly of Cu, Cd and Zn in aquatic systems and provide supporting information in relation to potential sediment toxicity.     

The relationship between social dominance,subcutaneous fat,and annual survival in wintering white-throated sparrows (Zonotrichia albicollis)

Summary We used multivariate analysis to identify factors correlated with level of subcutaneous fat (a form of stored energy) in a migratory, wintering population of white-throated sparrows. Dominant birds, residents from previous years, and birds residing in certain regions of the study area tended to have high mean fat levels during January and February. On the basis of differences in levels of fat, dominant prior residents could probably survive 50% longer without food than subordinate newcomers. An additional analysis revealed that dominant sparrows returned more frequently to the study area than subordinates, a result that might indicate higher survival. Offprint requests to: W.H. Piper (at the present address)     

Estimating paternity with spatial behaviour and DNA fingerprinting in the striped plateau lizard, Sceloporus virgatus (Phrynosomatidae)

The association between spatial proximity and paternity was studied in a population of the striped plateau lizard, Sceloporus virgatus. The relationship between estimated mating success and male phenotypic traits was examined for a sample of 55 males. DNA samples were obtained from 13 female-offspring families. The males with the closest spatial proximity to each female were tested as possible sires within each family. Fingerprinting with two multilocus hypervariable minisatellite probes revealed a strong correspondence between male-female spatial proximity and actual paternity. Paternity could be assigned for 72 of the 100 hatchlings. Most hatchlings with identifiable sires were attributed to a male with the highest category of spatial proximity to the mother. However, there was a low to moderate level of multiple paternity within clutches, and for some clutches probable sires could not be identified even though the most likely behavioural candidates were tested. Thus, nonterritorial males or other males lacking strong social and spatial relationships with females may achieve some degree of reproductive success. Analysis of mating success revealed that male success increased with body size, up to a point beyond which larger size conferred no advantage. Received: 7 January 1997 / Accepted after revision: 16 June 1997     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

Intraspecific parasitism and split sex ratios in a monogynous and monandrous ant (Leptothorax nylanderi)

Sex ratios were bimodally distributed in a population of the monogynous and monandrous ant Leptothorax nylanderi during each of 3 study years. The population-wide investment ratios suggested worker control of sex allocation. Nest-level variation in the proportional investment in virgin queens was not affected by the presence or absence of a queen and only slightly by collecting year, but was correlated with nest size, total sexual investment and, unexpectedly, with differences in nestmate relatedness: small, low-investment nests and nests with several worker lineages produced male-biased sex ratios. Colonies containing several worker lineages arise from usurpation of mature colonies by unrelated founding queens and the fusion of unrelated colonies under strong nest site limitation. In contrast to facultatively polygynous and polyandrous species of social insects, where workers can maximize their inclusive fitness by adjusting sex ratios according to the degree of relatedness asymmetry, workers in mixed colonies of L. nylanderi do not benefit from manipulating sex allocation, as here relatedness asymmetries appear to be the same as in homogeneous colonies. Received: 7 December 1999 / Received in revised form: 29 February 2000 / Accepted: 13 March 2000     

The functional significance of infanticide in a population of California ground squirrels (Spermophilus beecheyi)

Naturally-occurring infanticide was observed in a population of California ground squirrels (Spermophilus beecheyi). In four seasons, 40 infanticides were observed. All victims were post-emergent pups. Of 37 killings in which the killer was sexed 36 were by females. All infanticidal females were mothers at the time they killed, but in no case was a mother seen to kill or harm her own young. The victim was cannibalized in 22 cases and taken immediately into the killer’s burrow in 16 others. In no case did killers gain access to the victimized mother’s burrow or territory and female pups were not killed preferentially over males. In light of evolutionary explanations, infanticide in this population may best fit the resource exploitation hypothesis, in which killers commit infanticide to gain a nutritional benefit. Resource competition is a possible auxiliary explanation, since any time a female kills unrelated young she is eliminating possible competitors to her own offspring. This behavior could confer a fitness advantage on killers or it could be an adaptively neutral, alternative feeding strategy. More data are necessary to distinguish between these hypotheses. Received: 26 January 1995/Accepted after revision: 9 September 1995     

Social cohesion and foraging decrease with group size in fallow deer (Dama dama)

We studied the impact of group size on foraging behaviour and level of movement synchronisation among female herdmates of a fallow deer population in Central Italy. Both proportion of foraging events and movement synchronisation decreased with increasing group size. The proportion of foraging events was higher for animals on the edge of the group than for deer in the centre of the group; hence, there appears to be a trade-off between protection against predators and foraging interference, both of which decrease from the centre to the periphery of the group. This is the first time this type of behaviour has been recorded for wild ungulates. As expected, we also found that the movement of peripheral animals was less synchronised than that of central animals. Consequently, peripheral animals may lose contact with their herdmates and split off the group. We conclude that social inequalities may lead to conflicting requirements among group members and instability of large groups. Movement synchronisation (as a function of group size) appears to interact with habitat openness to produce variations of group size (which appear to be adaptive for individuals) as an emergent property of these aggregations.     

Prior encounters with the opposite sex affect male and female mating behavior in a wolf spider (Araneae,Lycosidae)

Mate availability can vary widely in nature depending upon population density and sex ratio and can affect the ability of individuals to be selective in mate choice. We tested the effects of prior encounters with the opposite sex (i.e., exposure to the opposite sex either with or without mating) on subsequent mating behavior in two experiments that manipulated mate availability for both males and females in the wolf spider, Hogna helluo. The probability of mating in the experimental trial depended upon whether the prior encounter involved mating or not, and males and females responded in opposite directions. Exposure without mating resulted in a higher subsequent frequency of mating for females and a lower subsequent frequency of mating for males, while prior mating experience resulted in a lower frequency of female remating and a higher frequency of male remating. Prior exposure without mating did not affect female aggression. However, mated females engaged in precopulatory cannibalism more frequently than virgins. Mated males escaped postcopulatory cannibalism more frequently than virgins. Our results show that males respond to exposure without mating in the expected manner. However, prior mating (1 week earlier) had unexpected effects on males, which may be due to mated males being of higher quality. There were little or no effects of the size of the prior exposure individual or mate on subsequent mating behaviors. Further research is needed to determine why different species use different degrees of prior information in mate choice.     

Modeling effects of toxin exposure in fish on long-term population size, with an application to selenium toxicity in bluegill (Lepomis macrochirus)

A primary goal in ecotoxicology is the prediction of population-level effects of contaminant exposure based on individual-level response. Assessment of toxicity at the population level has predominately focused on the population growth rate (PGR), but the PGR may not be a relevant toxicological endpoint for populations at equilibrium. Equilibrium population size may be a more meaningful endpoint than the PGR because a population with smaller equilibrium size is more susceptible to the negative effects of environmental variability. We address the individual-to-population extrapolation problem with modeling utilizing classical mathematical theory. We developed and analyzed a general model applicable to many freshwater fish species, that includes density-dependent juvenile survival and additional juvenile mortality due to toxicity exposure, and we quantified effect on equilibrium population size as a means of assessing toxicity. Individual-level effects are typically greater than population-level effects until the individual effect is large, due to compensatory density-dependent relationships. These effects are sensitive to the recruitment potential of a population, in particular the low-density first-year survival rate S_b. Assuming high S_b could result in underestimating effects of population-level toxicity. The equilibrium size depends directly on S_b, the reproductive potential, the toxin concentration at which mean mortality is 50% (LC₅₀), and the rate at which individual mortality increases with increasing toxin concentration. More experimental data are needed to decrease the uncertainty in estimating these parameters. We then used existing data for selenium toxicity in bluegill sunfish to parameterize a simulation version of the model as an example to assess the effects of environmental stochasticity on toxicity response. Effects of environmental variability resulted in simulated extinctions at much lower toxin concentrations than predicted deterministically.     

Resource allocation varies with parental sex and brood size in the asynchronously hatching green-rumped parrotlet (<Emphasis Type="Italic">Forpus passerinus</Emphasis>)

When eggs hatch asynchronously, offspring arising from last-hatched eggs often exhibit a competitive disadvantage compared with their older, larger nestmates. Strong sibling competition might result in a pattern of resource allocation favoring larger nestlings, but active food allocation towards smaller offspring may compensate for the negative effects of asynchronous hatching. We examined patterns of resource allocation by green-rumped parrotlet parents to small and large broods under control and food-supplemented conditions. There was no difference between parents and among brood sizes in visit rate or number of feeds delivered, although females spent marginally more time in the nest than males. Both male and female parents preferentially fed offspring that had a higher begging effort than the remainder of the brood. Mean begging levels did not differ between small and large broods, but smaller offspring begged more than their older nestmates in large broods. Male parents fed small offspring less often in both brood sizes. Female parents fed offspring evenly in small broods, while in large broods they fed smaller offspring more frequently, with the exception of the very last hatched individual. These data suggest male parrotlets exhibit a feeding preference for larger offspring—possibly arising from the outcome of sibling competition—but that females practice active food allocation, particularly in larger brood sizes. These differential patterns of resource allocation between the sexes are consistent with other studies of parrots and may reflect some level of female compensation for the limitations imposed on smaller offspring by hatching asynchrony.     

The sex in short supply for matings varies over small spatial scales in a katydid (Kawanaphilanartee, Orthoptera: Tettigoniidae)

In katydids such as Kawanaphilanartee, a female bias in the operational sex ratio (OSR) results in female competition for mates and male choice of mates. Previous work showed that the excess of sexually active females occurs when food availability is low, in part because less food increases the propensity of females to mate as they forage for the large edible spermatophores produced by males. In this study with K.nartee, a pollen-feeding species, we estimate natural variation in numbers of sexually active males and females by assessing male calling activity and the propensity of females to respond to experimental calling males. We found an excess of sexually active males at a site with many flowers and an excess of sexually active females at a site with few flowers about 900 m away. Between-site differences in gut masses of calling males were consistent with the hypothesis that pollen availability controls OSR. Finally, at a third site where flowers were at first scarce, we found that the initial excess in sexually active females changed to an excess of sexually active males after a clump of grass-trees flowered. The mean gut mass of all sampled males from this site increased after flowering. The large variation in OSR that we document for K. nartee highlights the importance of identifying the appropriate spatial and temporal scales over which OSRs are measured in studies of factors controlling sexual selection. Received: 13 May 1997 / Accepted after revision: 27 October 1997     

Fluctuating sexual dimorphism and differential hibernation by sex in a primate, the gray mouse lemur (Microcebus murinus)

The aim of this study was to investigate reproductive strategies and their consequences in gray mouse lemurs (Microcebus murinus), small solitary nocturnal primates endemic to Madagascar. Previous reports of sexual dimorphism in favor of males and females, respectively, a high potential for sperm competition and pheromonal suppression of mating activity among captive males, led us to investigate mechanisms of intrasexual competition in a wild population. Based on 3 years of mark-recapture data, we demonstrate that sexual dimorphism in this species fluctuated annually as a result of independent changes in male and female body mass. Male body mass increased significantly prior to the short annual mating season. Because their testes increased by 100% in the same period and because their canines are not larger than those of females, we suggest that large male size may be advantageous in searching for estrous females and in enabling them to sustain periods of short-term torpor. In contrast to reports from captive colonies, we found no evidence for two morphologically distinct classes of males. Finally, we also show that most adult males are active throughout the cool dry season that precedes the mating season, whereas most adult females hibernate for several months. This is in contrast to other solitary hibernating mammals, where males typically emerge 1–2 weeks before females. Thus, this first extended field study of M.␣murinus clarified previous conflicting reports on sexual dimorphism and male reproductive strategies in this primitive primate by showing that their apparent deviation from predictions of sexual selection theory is brought about by specific environmental conditions which result in sex-specific life history tactics not previously described for mammals. A general conclusion is that sexual selection can operate more strongly on males without resulting in sexual dimorphism because of independent selection on the same traits in females. Received: 6 July 1997 / Accepted after revision: 28 March 1998     

Male care of infants in a siamang (Symphalangus syndactylus) population including socially monogamous and polyandrous groups

While male parental care is uncommon in mammals, siamang (Symphalangus syndactylus) males provide care for infants in the form of infant carrying. I collected behavioral data from a cohort of five wild siamang infants from early infancy until age 15–24 months to identify factors affecting male care and to assess the consequences of male care for males, females, and infants in a population including socially monogamous groups and polyandrous groups. There was substantial variation in male caring behavior. All males in polyandrous groups provided care for infants, but males in socially monogamous groups provided substantially more care than males in polyandrous groups, even when the combined effort of all males in a group was considered. These results suggest that polyandry in siamangs is unlikely to be promoted by the need for “helpers.” Infants receiving more care from males did not receive more care overall because females compensated for increases in male care by reducing their own caring effort. There was no significant relationship between indicators of male–female social bond strength and male time spent carrying infants, and the onset of male care was not associated with a change in copulation rates. Females providing more care for infants had significantly longer interbirth intervals. Male care may reduce the energetic costs of reproduction for females, permitting higher female reproductive rates.