Nest box use by woodland dormice (Graphiurus murinus): the influence of life cycle and nest box placement

The use of nest boxes by the woodland dormouse, Graphiurus murinus, was investigated over a 13-month period in a riverine forest of the Great Fish River Reserve, South Africa. We predicted that some characteristics of nest box placement would affect nest box use and that the seasonal pattern of nest box use would be linked to the species' life cycle and physiological and socioecological characteristics. Generalized linear models indicated that the time since nest box installation and nest box height above ground positively affected the frequency and intensity of nest box use. Male and female dormice, as well as adults and juveniles, did not differ in the number of nest boxes used and equally occupied individual nest boxes. The percentage of nest boxes used peaked during spring and summer (breeding period) and dropped during winter (hibernation). However, whereas significantly more males were caught during the mating season (spring), the number of females occupying nest boxes was constant during the year. As female dormice successfully bred in the nest boxes, the observed sexual patterns suggest that (artificial) nest sites represent an important resource for females, whereas females seem to constitute the main resource for males, as predicted by the socioecological model.     

The influence of high nestbox density on the common dormouseMuscardinus avellanarius population

The response of common dormouseMuscardinus avellanarius Linnaeus, 1758 population to availability of nest sites was studied by manipulating the nestbox grid and ring-marking dormice. Abundance of adult dormice more than doubled in the 25 × 25 m nestbox grid in comparison to the 50 × 50 m grid, as a result of increased nestbox density from four to 16 boxes/ha. This effect already became apparent in the first year after additional nestboxes were made available and resulted from dormouse immigration, mostly from adjacent areas without nestboxes. In the second and third years, the number of two-year-old and older resident dormice, which had their home ranges in this plot, increased considerably. The average size of dormouse home range decreased by approximately half both in males and females in the 25 × 25 m grid compared to the 50 × 50 m grid. The proportion of breeding adult females did not differ between the two grids in spite of different adult dormouse density. Shortage of secure nest sites was a limiting factor for the common dormouse population abundance in the forest where natural tree hollows were absent, and high nestbox density increased environmental carrying capacity.     

Spacing Behavior of a Non‐Larder‐Hoarding Tamiasciurus: A Study of Mearns's Squirrels in Xeric Coniferous Forests

In ecosystems with seasonal fluctuations in food supply many species use two strategies to store food: larder hoarding and scatter hoarding. However, because species at different geographic locations may experience distinct environmental conditions, differences in hoarding behavior may occur. Tree squirrels in the genus Tamiasciurus display variation in hoarding behavior. Whereas red (Tamiasciurus hudsonicus) and Douglas's (Tamiasciurus douglasii) squirrels in mesic coniferous forests defend territories centered around larder hoards maintaining non‐overlapping home ranges, red squirrels in deciduous forests defend small scatter‐hoarded caches of cones maintaining overlapping home ranges. As in other rodent species, variation in hoarding behavior appears to influence the spacing behavior of red and Douglas's squirrels. In contrast, Mearns's squirrels (Tamiasciurus mearnsi) in xeric coniferous forests neither rely on larder hoards nor appear to display territorial behavior. Unfortunately, little is known about the ecology of this southernmost Tamiasciurus. Using radiotelemetry, we estimated home‐range size, overlap, and maximum distance traveled from nest to examine the spacing behavior of Mearns's squirrels. Similar to scatter‐hoarding rodents, maximum distance traveled from nest was greater for males during mating season, whereas those of females were similar year round. Although no seasonal differences were detected, male home ranges were three times larger during mating season, whereas those of females were smaller and displayed a minor variation between seasons. Home ranges were overlapped year round but contrary to our expectations, overlap was greater during mating season for both sexes, with no detectable relationship between male home‐range size and the number of females overlapped during mating season. Overall, the results appear to support our hypothesis that in the absence of larder hoards, the spacing behavior of Mearns's squirrels should be different from larder‐hoarding congeners and more similar to scatter‐hoarding rodents.     

Social organization and space use in California voles: seasonal,sexual, and age-specific strategies

We intensively monitored space use and movement in Microtus californicus over a 2-year period that included 1 year of high density (maximum 618/ha) and one of low (minimum 5/ha); historically this population has exhibited cycles of 2 or 4 years. Adults of both sexes dispersed at the start of the breeding season, culminating in the establishment of intrasexually exclusive territories. In females, these territories persisted throughout life, except that many young females recruiting during the breeding season established contiguous, overlapping, or adjacent home ranges with their mothers. This female philopatry explains the conclusion of previous workers that females of this species are non-territorial. In the dry (non-breeding) season, females had smaller ranges that often overlapped and were clustered. Adult males moved breeding territories at a modal interval of 6 weeks; this is consistent with their avoidance of inbreeding with philopatric daughters. Ranges overlapped 1–4 adult females at any one time, and a cohort of 7 long-lived males overlapped an average of 16.4 females during their tenure on the grid. The period of maximum overlap with adult females varied among individual males, and did not correlate with the time of maximum body weight. Ranges of males in the dry season overlapped extensively, with persistent associations among some individuals. In the lowdensity year, ranges of some adults failed to overlap intersexually. Juvenile males dispersed gradually between 3 and 13 weeks of age (half before 9 weeks), with some leaving after reaching sexual maturity; a few remained philopatric. Of juvenile females, 47% remained philopatric with the rest disappearing before 9 weeks of age. New understanding of vole social behavior, dispersal, and space use is achieved by focusing on the seasonal dynamics of spatial relationships among individuals with respect to age, sex, and relatedness.     

Intensity of space use reveals conditional sex‐specific effects of prey and conspecific density on home range size

Home range (HR) size variation is often linked to resource abundance, with sex differences expected to relate to sex‐specific fitness consequences. However, studies generally fail to disentangle the effects of the two main drivers of HR size variation, food and conspecific density, and rarely consider how their relative influence change over spatiotemporal scales. We used location data from 77 Eurasian lynx (Lynx lynx) from a 16‐year Scandinavian study to examine HR sizes variation relative to prey and conspecific density at different spatiotemporal scales. By varying the isopleth parameter (intensity of use) defining the HR, we show that sex‐specific effects were conditional on the spatial scale considered. Males had larger HRs than females in all seasons. Females' total HR size declined as prey and conspecific density increased, whereas males' total HR was only affected by conspecific density. However, as the intensity of use within the HR increased (from 90% to 50% isopleth), the relationship between prey density and area showed opposing patterns for females and males; for females, the prey density effect was reduced, while for males, prey became increasingly important. Thus, prey influenced the size of key regions within male HRs, despite total HR size being independent of prey density. Males reduced their HR size during the mating season, likely to remain close to individual females in estrous. Females reduced their HR size postreproduction probably because of movement constrains imposed by dependent young. Our findings highlight the importance of simultaneously considering resources and intraspecific interactions as HR size determinants. We show that sex‐specific demands influence the importance of prey and conspecific density on space use at different spatiotemporal scales. Thus, unless a gradient of space use intensity is examined, factors not related to total HR size might be disregarded despite their importance in determining size of key regions within the HR.     

Home range and mating system of two sympatric field mouse species,Apodemus speciosus andApodemus argenteus

The size and spatial distribution of home ranges in two sympatric field mouse species,Apodemus speciosus andApodemus argenteus, were revealed by the capture-recapture method in a temperate deciduous forest from June 1987 to June 1990. InA. speciosus, the home ranges of males were significantly larger tha those of females, and overlapped with those of other males and females during the breeding season. InA. argenteus, the home range of each male overlapped only that of a single female throughout the year. These results suggested thatA. speciosus was promiscuous or polygynous andA. argenteus was monogamous.     

Energy or information? The role of seed availability for reproductive decisions in edible dormice

The edible dormouse is a specialized seed predator which is highly adapted to the fluctuations of food availability caused by mast seeding of beech and oak trees. Dormice produce young just in time with maximum food availability, and can completely skip reproduction in years with a lack of seeding. Because their decision to reproduce or not in any particular year is made long before the ripe seeds are available, it seems that dormice can anticipate the upcoming mast situation. We tested the hypothesis that the presence of high caloric food in spring affects their reproductive decision. Therefore, we supplementary fed dormice in a field experiment from spring to early summer with sunflower seeds, which also contain a high amount of energy. Supplemental feeding caused significant increases in the proportion of reproducing females and reproductively active males. These results suggest that edible dormice may use the occurrence of an energy rich food resource to predict the autumnal mast situation. Further, our data indicate that the decision to reproduce was not the result of an increased body mass due to the consumption of surplus food, but that sufficient seed abundance acts as an environmental signal to which dormice adjust their reproduction.     

A 13-year population study of the edible dormouse <Emphasis Type="Italic">Glis glis</Emphasis> in Britain

Edible dormice Glis glis Linnaeus, 1766 were monitored from 1996–2008 inclusive. This has been the first population study of this non-native species since its introduction to Britain in 1902. Dormice were present in nest boxes from May or June until October or November. Numbers recorded varied greatly from year to year, and breeding failure occurred in 6 of the 13 years, apparently in response to environmental cues such as food availability. Litter sizes declined with growth of the young but some of the largest litters resulted from females aggregating their offspring. Males used more nest boxes than females, some of whom showed clear site fidelity from year to year. Survival rates were high and some individual lifespans exceeded 9 years. Longevity, compensating for periodic reproductive failure, and an extremely short active season, make the life history of edible dormice exceptional among small rodents, perhaps unique. Population density, rate of population increase and evidence for local emigration in non-breeding years are discussed, issues of particular significance in populations of recently-established introduced species such as this.     

Mate Choice and Mating Pattern in a Stream Goby of the Genus Rhinogobius

The mate choice and mating pattern of a benthic goby Rhinogobius sp. CB (cross band type) were investigated in the Kamo River, Shikoku, Japan. During the breeding season, gravid females assumed a nuptial color and either males or females initiated a courtship display. Males preferentially courted a female of similar size to lead her to his nest, whereas females courted more frequently when they encountered a large male. Eggs in any one nest were always at the same developmental stage. Sampling data of nesting males and females indicated that, in more than half the nests, males gathered more than one female before spawning. In some nests with eggs, two or three females had spent ovaries, indicating that the eggs were laid by multiple females within a short span of time. However, a comparison between the total number of eggs which females would spawn in one nest and the number of eggs actually deposited suggested that eggs were contributed by one female in most nests. This low level of polygyny in spite of multiple female availability is attributed to a limited available spawning area of the nest.     

Male mate choice in the chameleon grasshopper (Kosciuscola tristis)

In many species, males can increase their fitness by mating with the highest quality females. Female quality can be indicated by cues, such as body size, age and mating status. In the alpine grasshopper Kosciuscola tristis, males can be found riding on subadult females early in the season, and as the season progresses, males engage in fights over ovipositing females. These observations suggest that males may be competing for females that are either unmated (early season) or sperm‐depleted (late season). We thus hypothesised that male K. tristis may be choosy in relation to female mating status, and specifically, we predicted that males prefer females that are unmated. We conducted behavioural experiments in which males were given the choice of two females, one mated and one unmated. Contrary to our prediction, males did not mate preferentially with unmated females. However, copulation duration with unmated females was, on average, 24 times the length of copulation with mated females. While female K. tristis can reject mates, we did not observe any evidence of overt female choice during our trials. Females may gain additional benefits from mating multiply and may therefore not readily reject males. While our experiment cannot definitively disentangle female from male control over copulation duration, we suggest that males choose to invest more time in copula with unmated females, perhaps for paternity assurance, and that male mate assessment occurs during copulation rather than beforehand.     

Vagabond males and sedentary females: spatial ecology and mating system of the speckled rattlesnake (Crotalus mitchellii)

We used radiotelemetric data and behavioural observations to characterize seasonal (mating versus post‐mating seasons) and sexual variation in movement patterns, as well as to examine some of the ecological factors contributing to the evolution of the mating system in a venomous predator from the Mojave Desert of North America, the speckled rattlesnake, Crotalus mitchellii. Mating occurs in spring from late April to early June, shortly after emergence from hibernation, when snakes are predictably aggregated around the dens. Males and females travelled further per unit time in the mating season compared to the post‐mating season. Males also travelled longer distances per unit time than females in the mating and post‐mating seasons, and males with larger home ranges during the mating season had more potential mating partners. The results obtained suggest that males actively locate females during the mating season, and that the drastic increase in distance travelled by males during the mating season may be caused by strong male–male competition for access to females, probably because of the limited availability of sexually receptive females. Furthermore, males fight for access to females, and males of larger size are more likely to acquire females. Therefore, sexual selection apparently acts on two different male phenotypic traits: investment in mate‐searching activities and male body size. The present study demonstrates that combining quantitative spatial analyses and behavioural observations in an explicit temporal context can significantly advance our understanding of the ecology and evolution of organismal mating systems. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 681–695.     

Attraction of the sexes inFormica lugubris Zett (Hymenoptera: Formicidae)

Summary Sexuals ofFormica lugubris fly to mating places, where females attract males by using a sex pheromone. Females collected on the nest surface before departing on a mating flight are much less attractive than those collected on the mating place after the mating flight, suggesting that the mating flight triggers the release of the sex pheromone. Olfactory cues are essential for males to locate females while they patrol. Males probably use visual cues to locate females once they have alighted nearby them. Males are also attracted by aggregations of other males on the ground, probably because one or several females are likely to be close to male aggregations.     

Effects of size and sex on the courting success and foraging behaviour of Embiotoca jacksoni

To test life-history theory that body size and sex should influence how animals allocate time to foraging versus reproductive activities, we measured the effects of size and sex on courting success and foraging behaviour of black surfperch Embiotoca jacksoni off Santa Catalina Island, southern California. Observations of focal fish were made while snorkelling, during which the length of each fish (estimated to the nearest cm), total duration of courting encounters and foraging rates were recorded. We made observations during and outside the mating season. Courtship occurred only between pairs and its duration increased with the size of both the male and female. Although males would court females that were smaller or larger than themselves, pairs that were closely matched in size had long courting sessions, whereas those that differed considerably in size courted only briefly. Small fish foraged more than larger fish, both during and outside the mating season. Males and females foraged at similar rates outside of the mating season, but during the mating season males reduced their foraging rates to less than half that seen outside of the mating season, whereas females continued to forage at the same rate. This decrease in foraging rate of males during the mating season was seen in all sizes of males but was proportionally greatest in the largest males. These observations indicate that males trade off time spent on foraging for time spent courting during the mating season, whereas females do not.     

Body size and alternative mating tactics in the beewolf Philanthus zebratus (Hymenoptera; Sphecidae)

A study of the mating behaviour of males of the beewolf Philanthus zebratus revealed that in one population males display variability in mating tactics and that this variability is related to male body size. There was a tendency for large males to patrol the airspace above the nesting area while smaller males were territorial adjacent to it. The mean sizes of the two groups of males were significantly different, although the size ranges of the two groups overlapped. Only 2.5% of the males were observed to undertake both mating tactics, at different times. Observations are presented on daily and seasonal activity patterns and on the relative location of nests, territories, and patrolling males. A second population, with lower nest density, was observed for several days, revealing only territorial males. It is suggested that the presence of patrolling males is related to the higher nest density of the one population. The fact that patrolling males tend to be relatively large is possibly related to flight energetics or simply to the ability of large males to seize females, which are usually larger than males, in mid-air.     

Size Assortative Mating and Reproductive Success of the Funnel-Web Spider, Agelena limbata (Araneae; Agelenidae)

Field observations and laboratory experiments were conducted to assess the relation between male size and reproductive success in the funnel-web spider, Agelena limbata Thorell (Agelenidae), in 2 years. In this species, the body size of males is similar to that of females. In the field, size assortative mating occurred in both years. In 1 year, partial correlation coefficient analysis indicates that male cephalothorax width is a beter predictor of the copulated female cephalothorax width than of the date of pairing. In laboratory experiments, females tended to reject courting males that were smaller in relative body size, and males that were larger in relative body size had greater copulation success. Consequently female rejection of smaller courting males has some contribution to size assortative mating. Since larger females deposited more numerous eggs in the field, larger males are expected to have a higher reproductive success.     

Size-mediated dominance and aggressive behavior of male Japanese fluvial sculpin Cottus pollux (Pisces: Cottidae) reduce nest-site abundance and mating success of conspecific rivals

Despite growing evidence for plasticity in the mating patterns of nest-holding animals in relation to the changes in nest abundance, the effects of aggressive interaction by dominant males on nest availability for conspecific rivals remains unclear. To quantify the effects of male–male competition on nest-site choice and mating success of the male Japanese fluvial sculpin Cottus pollux, we conducted experiments on 5 males from different 5 size classes under both sufficient and shortage nest-abundance conditions. Nest-choice experiments showed that both male size class and nest-abundance condition had significant effects on the nesting rates of males. Following the nest-choice experiments, 10 gravid females were added in the experimental tanks. Mating experiments revealed that male size, nesting rate before addition of females, and the number of courtship attempts on females were valid variables of male mating success, regardless of nest-abundance conditions. After achieving initial mating success, the largest nesting male exhibited more frequent aggressive interaction with other conspecific males than he did before obtaining eggs in his nest. Our results suggest that size-mediated dominance and aggressive behavior of males may disrupt nest acquisition of other conspecific males, and may consequently result in extreme variation in mating success among males even under sufficient nest-abundance conditions.     

Seasonal differences in the feeding ecology and behavior of male edible dormice (Glis glis)

Mammalian hibernators undergo dramatic seasonal changes of food intake and the use of their gastrointestinal tract. During several months of hibernation fat-storing hibernators do not use their intestinal tract for nutritional intake. However, during the rest of the year they have to increase their energy intake in order to compensate high reproductive investment and store sufficient body fat to survive the following hibernation period. Edible dormice (Glis glis) are obligate fat-storing hibernators which hibernate in Germany from September until June. Males incur high energetic costs during mating and as soon as reproduction is terminated they have to accumulate high quantities of fat to survive hibernation. In order to understand how fat-storing hibernators like edible dormice cope with these energetically demanding situations, we measured body mass changes of captured male edible dormice in the field and studied their feeding ecology. Furthermore, we measured seasonal changes in food ingestion and assimilation rates by feeding experiments carried out in captivity.Results of this study revealed that during the mating season males significantly lowered their body mass, while food ingestion and assimilation rates remained constant. The body mass reduction showed that they used their body fat reserves to pay at least part of the energetic costs of reproduction. During the pre-hibernation fattening period males increased their body mass but held their assimilation rates on a constant level. Nevertheless, they increased the amount of ingested food and subsequently the amount of energy intake. Furthermore, they changed their dietary spectrum in the field by turning to lipid-rich seeds. These behavioral adaptations enable them to restore their energy losses during reproduction and to accumulate sufficient body fat to survive hibernation.     

Social and foraging components of the home range in Antechinus stuartii (Dasyuridae: Marsupialia)

The home range of Antechinus stuartii has two components. A small area (0.94 ha for males, 0.38 ha for females) is used for foraging, while a much greater social range (mean for males in a given year may exeed 5 ha; the mean for females never exceeded 3 ha) encompasses communal nests, which are used by both males and females. The foraging range of females is stable throughout the year. The foraging range may not coincide at all, or only partially coincide with her social range. By contrast, male foraging ranges drift through the year, and generally lie within their social range. Intersexual differences reflect the different selection pressures on the sexes. Females require access to resources to sustain a costly lactation, while males require a knowledge of many possible sites for mating and early resolution of agonistic interactions with other males with which they cohabit during mating.     

Determinants of male mating success in a natural population of a stream goby of the genus Rhinogobius

The mating success of individually identified males of a stream goby, Rhinogobius sp. CB (cross‐band type), was recorded over one breeding season. One to 3 year‐old males were active in mating, but 3 year‐old males, which accounted for 36% of the population, amounted to c . 70% of the total males guarding eggs in the nest. Three quarters of the breeding males had only one brooding cycle, but the others had two or three. All the latter males changed their nest sites between cycles within a riffle. In 1 and 2 year‐old males, the number of brooding cycles contributed more to the mating success than egg mass size in one brooding cycle. For mating success of 3 year‐old males, the egg mass size in one brooding cycle, which can be enhanced by spawning with a large female or multiple females, was as important as the number of brooding cycles. These male reproductive tactics could be attributed to the age‐related ability of nest construction and mate acquisition.     

Nest Building and Mating in Relation to Male Size in Corkwing Wrasse, Symphodus Melops

This study examines nest building and mating in relation to size in male corkwing wrasse, Symphodus melops, in the absence of male–male competition. We kept males of varying size together with mature females in tanks containing adequate materials and sites for nest construction. Our results indicate that the size of territorial males does not influence the ability to construct a nest. However, larger males initiated nest building earlier than smaller males and nest size correlated positively with male size. Furthermore, male size did not affect whether or not a nesting male succeeded in mating, indicating that male size was not essential for female acceptance (i.e. whether or not a female is willing to mate) of a male in the current situation.