Population differentiation in female sex pheromone and male preferences in a solitary bee

Population differentiation in female mating signals and associated male preferences can drive reproductive isolation among segregated populations. We tested this assumption by investigating intraspecific variation in female sex pheromone and associated male odour preferences among distant populations in the solitary bee Colletes cunicularius (L.) by using quantitative gas chromatography and by performing field bioassays with synthetic blends of key sex pheromone compounds. We found significant differences in sex pheromone blends among the bee populations, and the divergence in odour blends correlated positively with geographic distance, suggesting that genetic divergence among distant populations can affect sex pheromone chemistry. Our behavioural experiments, however, demonstrate that synthetic copies of allopatric female sex pheromones were cross-attractive to patrolling males from distant populations, making reproductive isolation by non-recognition of mating signals among populations unlikely. Our data also show that patrolling male bees from different populations preferred odour types from allopatric populations at the two sites of bioassays. These male preferences are not expected to select for changes in the female sex pheromone, but may influence the evolution of floral odour in sexually deceptive orchids of the genus Ophrys that are pollinated by C. cunicularius males.     

Satellite tracking reveals a dichotomy in migration strategies among juvenile loggerhead turtles in the Northwest Atlantic

Few data are available on the movements and behavior of immature Atlantic loggerhead sea turtles (Caretta caretta) from their seasonal neritic foraging grounds within the western north Atlantic. These waters provide developmental habitat for loggerheads originating from several western Atlantic nesting stocks. We examined the long-term movements of 23 immature loggerheads (16 wild-caught and seven headstart turtles) characterizing their seasonal distribution, habitat use, site fidelity, and the oceanographic conditions encountered during their migrations. We identified two movement strategies: (1) a seasonal shelf-constrained north–south migratory pattern; and (2) a year-round oceanic dispersal strategy where turtles travel in the Gulf Stream to the North Atlantic and their northern dispersal is limited by the 10–15°C isotherm. When sea surface temperatures dropped below 20°C, neritic turtles began a migration south of Cape Hatteras, North Carolina (USA) where they established fidelity to the waters between North Carolina’s Outer Banks and the western edge of the Gulf Stream along outer continental shelf. Two turtles traveled as far south as Florida. Several turtles returned to their seasonal foraging grounds during subsequent summers. Northern movements were associated with both increased sea surface temperature (>21°C) and increased primary productivity. Our results indicate strong seasonal and interannual philopatry to the waters of Virginia (summer foraging habitat) and North Carolina (winter habitat). We suggest that the waters of Virginia and North Carolina provide important seasonal habitat and serve as a seasonal migratory pathway for immature loggerhead sea turtles. North Carolina’s Cape Hatteras acts as a seasonal “migratory bottleneck” for this species; special management consideration should be given to this region. Six turtles spent time farther from the continental shelf. Three entered the Gulf Stream near Cape Hatteras, traveling in the current to the northwest Atlantic. Two of these turtles remained within an oceanic habitat from 1 to 3 years and were associated with mesoscale features and frontal systems. The ability of large benthic subadults to resume an oceanic lifestyle for extended periods indicates plasticity in habitat use and migratory strategies. Therefore, traditional life history models for loggerhead sea turtles should be reevaluated.     

Polyandrous mating in treetops: how male competition and female choice interact to determine an unusual carnivore mating system

The diversity of mammalian mating systems is primarily shaped by sex-specific reproductive strategies. In the present study, we explored determinants and consequences of a unique mating system exhibited by fossas (Cryptoprocta ferox), the largest Malagasy carnivore, where females mate polyandrously on traditional mating trees, and males exhibit intrasexual size dimorphism. Males face both contest and scramble competition, and inter-sexual size dimorphism can be pronounced, but its magnitude depends on the male morph. Using a continuous behavioral observation of six estrous females over 4 years, we investigated correlates of male contest competition and female choice based on 316 copulations. Furthermore, we assessed correlates of male scramble competition based on testes size and movement data obtained from GPS tracking. We found that females dominated males regardless of their smaller size and that females actively solicited copulations. Heavy males had highest mating success during the female’s peak mating activity, but were discriminated against afterwards. Female choice and male–male competition thus converged to generate a mating advantage for heavier males. Our results suggest that females actively seek polyandrous matings, presumably for indirect genetic benefits. Since body mass is the major determinant of male mating success and is at the same time dependent on the degree of sociality and associated hunting mode of the respective male morph, a male’s feeding ecology is likely to influence its reproductive tactic. A combination of benefits from female polyandry and the consequences of different subsistence strategies may thus ultimately explain this unusual mating system.     

Sink populations in carnivore management: cougar demography and immigration in a hunted population.

Carnivores are widely hunted for both sport and population control, especially where they conflict with human interests. It is widely believed that sport hunting is effective in reducing carnivore populations and related human-carnivore conflicts, while maintaining viable populations. However, the way in which carnivore populations respond to harvest can vary greatly depending on their social structure, reproductive strategies, and dispersal patterns. For example, hunted cougar (Puma concolor) populations have shown a great degree of resiliency. Although hunting cougars on a broad geographic scale (> 2000 km2) has reduced densities, hunting of smaller areas (i.e., game management units, < 1000 km2), could conceivably fail because of increased immigration from adjacent source areas. We monitored a heavily hunted population from 2001 to 2006 to test for the effects of hunting at a small scale (< 1000 km2) and to gauge whether population control was achieved (lambda < or = 1.0) or if hunting losses were negated by increased immigration allowing the population to remain stable or increase (lambda > or = 1.0). The observed growth rate of 1.00 was significantly higher than our predicted survival/fecundity growth rates (using a Leslie matrix) of 0.89 (deterministic) and 0.84 (stochastic), with the difference representing an 11-16% annual immigration rate. We observed no decline in density of the total population or the adult population, but a significant decrease in the average age of independent males. We found that the male component of the population was increasing (observed male population growth rate, lambda(OM) = 1.09), masking a decrease in the female component (lambda(OF) = 0.91). Our data support the compensatory immigration sink hypothesis; cougar removal in small game management areas (< 1000 km2) increased immigration and recruitment of younger animals from adjacent areas, resulting in little or no reduction in local cougar densities and a shift in population structure toward younger animals. Hunting in high-quality habitats may create an attractive sink, leading to misinterpretation of population trends and masking population declines in the sink and surrounding source areas.     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Post-mating odor in females of the solitary bee, Andrena nigroaenea (Apoidea, Andrenidae), inhibits male mating behavior

We investigated odor changes and their behavioral significance in the solitary, ground-nesting bee Andrena nigroaenea. We used gas chromatography with electroantennographic detection and performed behavioral tests with males in the field using natural odor samples and synthetic compounds. We found that only cuticle extracts of young females elicited copulation attempts in the males. We demonstrated that among the 17 compounds which triggered electroantennographic responses, all-trans-farnesyl hexanoate and all-trans-farnesol were significantly more abundant in unattractive cuticle extracts of A. nigroaenea females. Dufour’s gland extracts of these females also contained greater amounts of both compounds. In bioassays using synthetic farnesyl hexanoate and farnesol we found that these compounds inhibit copulation behavior in the males. Farnesyl hexanoate is probably synthesized in Dufour’s gland and used by females for lining brood cells. We interpret the semiochemical function of farnesyl hexanoate and its precursor farnesol to have evolved secondarily. As an outcome of sexual selection, it facilitates the discrimination by males of receptive females from nesting and thus already mated individuals. The dual function of these compounds represents an elegant parsimony in the chemical communication system of this insect. Received: 19 January 2000 / Revised: 29 May 2000 / Accepted: 24 June 2000     

Fitness costs of neighborhood disruption in translocations of a solitary mammal

Translocation is used to reestablish wild populations of animals, but translocation projects often do not meet their objectives because postrelease mortality of animals is high. One reason for translocation failure is that the behavioral or ecological requirements of released animals are unmet. Maintaining founder-group social relationships during release can affect reestablishment of social species. Solitary territorial species with stable neighbors (restricted dispersal and lifetime occupation of a home range) of the same species may also benefit from the maintenance of these social relationships during translocation. We translocated Stephens' kangaroo rats (Dipodomys stephensi), a solitary species listed as endangered under the U.S. Endangered Species Act, with and without neighboring kangaroo rats. We compared the settlement (establishment of a stable home range) decisions and fitness of kangaroo rats between the 2 treatments. Kangaroo rats translocated with neighbors traveled shorter distances before establishing territories, had higher survival rates, and had significantly higher reproductive success than kangaroo rats translocated without neighbors. Number of offspring was 24-fold higher for kangaroo rats translocated with neighbors than those translocated without neighbors. Differences in behavior following release may partially explain differences in survival between the 2 groups. Immediately following release, animals translocated with neighbors fought less and spent significantly more time foraging and digging burrows than animals translocated without neighbors. Our results indicate that even for solitary species, maintaining relationships among members of a translocated group of animals can influence translocation success. This study is the first empirical demonstration of the fitness consequences of disrupting social relationships among territorial neighbors.     

Host choice and offspring sex allocation in a solitary parasitic wasp

Summary Females of the parasitic wasp Antrocephalus pandens can detect differences in the quality of their hosts (pupae of Corcyra cephalonica, a stored-product moth) and allocate offspring of either sex accordingly. Larger and younger hosts are accepted more often in both dead and live hosts; more female offspring emerge from the perceived better hosts, while more males emerge from the smaller, older ones. These patterns are consistent with a sex allocation strategy by the mother, since females from a given size host tend to be larger than males and larger females produce more eggs. However, when wasps lay their eggs in groups of hosts of different size and age rather than encountering them one at a time, no difference in number or sex ratio of offspring is detected between groups. This result and evidence from the change in offspring sex ratio with female age and with numbers of females foraging on a group of hosts are interpreted and discussed in the context of sex allocation (Charnov 1979) and local mate competition (LMC, Hamilton 1967) theories.     

Local genetic structure and relatedness in a solitary mammal, Neotoma fuscipes

We used DNA microsatellites to investigate temporal and spatial patterns of local genetic differentiation and relatedness in a solitary mammal, the dusky-footed woodrat (Neotoma fuscipes). Patterns of genetic variation were measured relative to spatial clusters, or neighborhoods, of woodrats. We detected significant genetic differentiation among woodrat neighborhoods in two populations spanning multiple habitat types and densities. Estimates of θ _ST among neighborhoods ranged 0.034–0.075 and were comparable to levels reported in social mammals. Genetic differentiation at such a local scale is noteworthy because it occurred in the absence of any physical barriers to gene flow, suggesting that the patterns observed are linked to the nonrandom patterns of mating and dispersal that characterize woodrat social structure. Genetic differentiation and relatedness among neighborhoods were even higher when only resident females were analyzed. These results are consistent with a pattern of female philopatry and male-biased dispersal in woodrats. Geographic distance and relatedness were inversely correlated in adult females at intermediate densities, but not at low densities. Nonetheless, matrilineal genetic structure was apparent even at low woodrat densities based on estimates of θ _ST among neighborhoods of resident females that were significantly greater than zero and consistently greater than estimates including all individuals. In summary, this study demonstrates a matrilineal genetic structure in dusky-footed woodrats. In addition, our results support the idea that intermediate densities may be better at facilitating the formation of spatial kin clusters than either extreme. An erratum to this article can be found at     

Strategic male signalling effort in a desert-dwelling fish

Males often use elaborate courtship displays to attract females for mating. Much attention, in this regard, has been focused on trying to understand the causes and consequences of signal variation among males. Far less, by contrast, is known about within-individual variation in signal expression and, in particular, the extent to which males may be able to strategically adjust their signalling output to try to maximise their reproductive returns. Here, we experimentally investigated male courtship effort in a fish, the Australian desert goby, Chlamydogobius eremius. When offered a simultaneous choice between a large and a small female, male gobies spent significantly more time associating with, and courting, the former, probably because larger females are also more fecund. Male signalling patterns were also investigated under a sequential choice scenario, with females presented one at a time. When first offered a female, male courtship was not affected by female size. However, males adjusted their courtship effort towards a second female depending on the size of the female encountered previously. In particular, males that were first offered a large female significantly reduced their courtship effort when presented with a subsequent, smaller, female. Our findings suggest that males may be able to respond adaptively to differences in female quality, and strategically adjust their signalling effort accordingly.     

Simultaneous denitrifying phosphorus accumulation in a sequencing batch reactor

In order to achieve simultaneous nitrogen and phosphorus removal in the biological treatment process, denitrifying phosphorus accumulation (DNPA) and its affecting factors were studied in a sequencing batch reactor (SBR) with synthetic wastewater. The results showed that when acetate was used as the sole carbon resource in the influent, the sludge acclimatized under anaerobic/aerobic operation had good phosphorus removal ability. Denitrifying phosphorus accumulation was observed soon when fed with nitrate instead of aeration following the anaerobic stage, which is a vital premise to DNPA. If DNPA sludge is fed with nitrate prior to the anaerobic stage, the DNPA would weaken or even disappear. At the high concentration of nitrate fed in the anoxic stage, the longer anoxic time needed, the better the DNPA was. Induced DNPA did not disappear even though an aerobic stage followed the anoxic stage, but the shorter the aerobic stage lasted, the higher the proportions of phosphorus removal via DNPA to total removal.     

Modelling the motion of an internal solitary wave over a bottom ridge in a stratified fluid

A series of laboratory experiments has been carried out to investigate the passage of an internal solitary wave of depression over a bottom ridge, in a two-layer fluid system for which the upper and lower layer is linearly-stratified and homogeneous respectively. Density, velocity and vorticity fields induced by the wave propagation over the ridge have been measured simultaneously at three locations, namely upstream, downstream and over the ridge crest, for a wide range of model parameters. Results are presented to show that wave breaking may occur for a sufficiently large wave amplitude and a strong ridge blockage factor, with accompanying mixing and overturning. Density field data are presented (i) to illustrate the overturning and mixing processes that accompany the wave breaking and (ii) to quantify the degree of mixing in terms of the wave and ridge parameters. For weak encounters, good agreement is obtained between the laboratory experimental results (velocity and vorticity fields induced by the wave propagation) and the predictions of a recently-developed fully nonlinear theory. Discrepancies between theory and experiment are discussed for cases in which breaking and mixing occur.     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

Kinship structure and mating system in a solitary subterranean rodent, the silvery mole-rat

The African mole-rats (Bathyergidae, Rodentia) are subterranean rodents that have evolved a wide range of social organisations varying from solitary to eusocial. Due to some unusual features of their breeding and social systems, much attention has been focused on social species, with solitary species being virtually ignored. Here, we present the first genetic study on the mating system of any solitary mole-rats, the silvery mole-rat (Heliophobius argenteocinereus) - a species which is assumed to be monogamous. Microsatellite markers were used to analyse the mating system and the kinship structure in populations in southern Malawi. Isolation by distance between individuals was apparent in two studied populations, but not in the third, probably, as a result of barriers limiting dispersal in the latter population. Polygyny was found to be a mating system in this population, where a strongly female-biased adult sex ratio was present. In this case, large distances between the burrow systems of mating partners exclude the possibility of belowground searching for mates, suggesting that the males might seek females aboveground. Interestingly, among analysed litters from various localities, one multiple-sired litter was found. Therefore, the results suggest that the mating system in potentially monogamous solitary subterranean rodents may be much more variable than expected and can differ among populations.     

Intrasexual selection in Mirza coquereli : evidence for scramble competition polygyny in a solitary primate

The primates of Madagascar (Lemuriformes) deviate from fundamental predictions of sexual selection theory in that polygynous species lack sexual dimorphism, have even adult sex ratios and often live in female-dominated societies. It has been hypothesized that intrasexual selection in these species is either reduced or primarily focused on traits related to scramble competition. The goal of this study was to examine these hypotheses by studying the mating system of a solitary nocturnal species, Mirzacoquereli. During a 4-year field study in western Madagascar, I captured and followed 88 individually marked animals. I found that adult males were significantly larger than females, providing the first evidence for sexual size dimorphism in lemurs. In addition, the adult sex ratio was biased in favour of females in 3 out of 4 years. There was no significant sex difference in canine size, however. Males showed pronounced seasonal variation in testis size with a 5-fold increase before and during the short annual mating season. During the mating season, males had more injuries than females and more than quadrupled their home ranges, overlapping with those of more than ten females, but also with about the same number of rivals. Only about one social interaction per 10 h of observation was recorded, but none of them were matings. Together, these results indicate that these solitary lemurs are clearly subject to intrasexual selection and that male-male competition is primarily, but not exclusively, of the scramble type. In addition, they suggest that the above-mentioned idiosyncracies may be limited to group-living lemurs, that social systems of solitary primates are more diverse than previously thought, and that the temporal distribution of receptive females is responsible for this particular male mating strategy. Received: 11 January 1997 / Accepted after revision: 18 April 1997     

Linking male qualities to multiple display traits: an example in a fish with exclusive male care

Recent theoretical models predict that the relative allocation to advertisement and parental care depends on whether paternal care is necessary for offspring survival: In species with exclusive male care, male investment in attraction is expected to reliably indicate paternal care effort and male phenotypic quality. Previous research, yielding contrasting results, has considered how one trait involved in mate attraction interacts with parental care or a specific aspect of male quality. In the blenny Salaria pavo, we perform a comprehensive analysis of the interplay between overall male attractiveness and male quality, the latter in terms of fertility, condition, and parental care. In this fish, males are larger than females, exhibit two sexually dimorphic traits (head crest and anal glands), and solely care for eggs. We generated a male attractiveness index through principal component analyses of morphological traits and quantified parental effort as the total time spent in egg care. In addition, we analyzed the relationships between specific components of attractiveness and male qualities. In agreement with theory predictions, we found that male overall attractiveness is a reliable indicator of fertility, in terms of sperm number, but is unrelated to body condition and parental care effort, with males able to perform high levels of care regardless of their level of advertisement. However, the relative expression of head crest area appears positively related to sperm number but is traded-off with parental care effort. These findings underline the need, in addressing real patterns, to consider interactions between multiple aspects of male display and quality.     

Sex in the dark: determinants and consequences of mixed male mating tactics in<Emphasis Type="Italic"> Microcebus murinus</Emphasis>, a small solitary nocturnal primate

In mammals with solitary females, the potential for males to monopolize matings is relatively low, and scramble competition polygyny is presumed to be the predominant mating system. However, combinations of male traits and mating tactics within this type of polygyny have been described. The main aim of our study was to identify the relative importance of, and interactions among, potential determinants of contrasting male reproductive tactics, and to determine their consequences for male reproductive success in a small solitary nocturnal Malagasy primate, the gray mouse lemur (Microcebus murinus). We studied their mating behavior over three consecutive annual mating seasons. In addition, we determined the genetic relationships among more than 300 study animals to quantify the reproductive success of individual males. We found that, with a given relatively low overall monopolization potential, successful male mouse lemurs roamed extensively in search of mates, had superior finding ability and mated as early as possible. However, contest competition was important too, as temporary monopolization was also possible. Males exhibited different mating tactics, and heavier males had a higher reproductive success, although most litters had mixed paternities. Switching between tactics depended on short-term local variation in monopolization potential determined by a pronounced dynamic in fertilization probability, number of alternative mating opportunities, and the operational sex ratio. This study also revealed that the dynamics of these determinants, as well as the mutual interactions between them, necessitate a detailed knowledge of the mating behavior of a species to infer the impact of determinants of alternative mating tactics.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by S. AlbertsThis revised version was published online in August 2004 with corrections to Figure 2.