Reservoir Competence of the Vole,Clethrionomys rufocanus bedfordiae,for Borrelia garinii or Borrelia afzelii

In autumn of 1994 and spring of 1995, we examined Borrelia infection among Microtinae voles, Clethrionomys rufocanus bedfordiae, in Hokkaido, Japan. In BSK culturing of the earlobe tissues of 45 C. rufocanus bedfordiae captured, twelve rodents were positive for Borrelia. Eight isolates were used for the polymerase chain reaction (PCR) and the restriction fragment length polymorphism (RFLP) analysis. According to the results, these isolates were classified into B. garinii or B. afzelii. It is considered that a common vole, C. rufocanus bedfordiae, plays a significant role in the transmission and maintenance of B. garinii and B. afzelii, similar to the role of Apodemus speciosus mice.     

The biology of the voleClethrionomys rufocanus: A review

The biology of the gray-sided voleClethrionomys rufocanus in Hokkaido, concerning taxonomy, morphology, phylogeny, distribution, and natural history, is reviewed. Applied issues in forest management (damage, control and census) are also mentioned. AlthoughClethrionomys rufocanus of Hokkaido was originally identified as a distinct species,Evotomys (=nowClethrionomys) bedfordiae Thomas, 1905, current literature generally refers to the gray-sided vole of Hokkaido asClethrionomys rufocanus or asC. rufocanus bedfordiae (vernacular name, the Bedford’s red-backed vole). The gray-sided vole is the most common small mammal in Hokkaido. It inhabits open areas as well as forests, and mainly feeds on green plants. The gray-sided vole has a high reproductive potential; litter size: 4–7; gestation period: 18–19 days; maturation age: 30–60 days old. Although spring-born individuals usually attain sexual maturity in their summer/fall of birth, their maturation is sometimes suppressed under high densities. The breeding season is generally from April to October, but with some regional variation.Clethrionomys rufocanus has a rather specialized diet (folivorous), particularly during winter when it feeds on bamboo grass. Many predators specialize on the grey-sided vole in Hokkaido; even the red fox, which is a typical generalist predator, selectively feeds on this vole. Damage by voles’ eating bark used to be sever on forest plantations in Hokkaido. Censuses of small rodents have been carried out for management purpose since 1954.     

The effects and limits of territoriality on population regulation in grey red-backed voles,Clethrionomys rufocanus bedfordiae

The effects of breeding territoriality on the stability of grey red-backed vole (Clethrionomys rufocanus bedfordiae) populations were investigated on a control grid and a grid on which the voles were fed, in an outdoor enclosure in Hokkaido, Japan. Vole populations were monitored by live trapping from 1984 to 1986: (1) Population density was 2–7 times greater on the experimental grid to which food was added than on the control grid. Reproductive output was more closely associated with the difference in density between grids than survival or dispersal (immigration and emigration) rates. (2) The number of adult females and pregnancy rate of the experimental population were significantly greater than those of the control one. The difference in the number of adult females between the populations was greater than that in pregnancy rate. (3) The proportion of successful litters and the number of weanlings per litter were not significantly different between the control and experimental population. (4) Adult females held territories on both the control and experimental grid; they were spaced out more than would be expected from random occupation. The territories overlapped more on the experimental grid than on the control grid. (5) Mean territory size of adult females on the experimental grid was about half of that on the control grid. The territory size was correlated negatively with population density. (6) The proportion of trap sites that were used by adult females was significantly greater on the experimental grid than on the control grid. This suggests that adult females on the experimental grid used the area more extensively. This factor, in association with territory size and overlapping of territory, was also important in causing the difference in the number of adult females between the grids. (7) These results call into question the hypothesis that territoriality stabilizes the density in populations of Clethrionomys.     

Possible relationships between morphology,territory quality,and skin color of American Kestrels1

ABSTRACT Carotenoid‐based coloration of skin and plumage has been found to be correlated with individual quality in many species of birds during the breeding season. However, less is known about the possible role of these signals during the nonbreeding season, particularly among nonpasserines that defend winter territories. American Kestrels (Falco sparverius) are sexually dimorphic raptors that defend winter territories and possess carotenoid‐based morphological features known to be correlated with individual quality. Much is known about winter territory use and habitat segregation by male and female kestrels, but possible relationships among morphological features, individual quality, and habitat quality have not been examined. Our objective was to examine possible relationships between morphology, territory quality, and skin color of American Kestrels. Male kestrels had brighter skin than females, and the skin color of male kestrels was positively correlated with size (wing chord and tail length) and territorial quality (hunting territories with less canopy and more grass cover). No such relationships were found for female kestrels. Skin color appears to be an honest indicator of quality for male American Kestrels and may serve both intersexual (territory acquisition) and intersexual (mate choice) functions during the breeding and nonbreeding seasons.     

A time series and geographical analysis of population dynamics of the red-backed vole in Hokkaido,Japan

Summary A time series and geographical analysis of the long term (25-years and 16-years) census data of the red-backed vole, Clethrionomys rufocanus bedfordiae, populations in Hokkaido, Japan was carried out. Eighty seven populations from all over Hokkaido were grouped into the following 4 types according to their flucturation patterns: type A: low density-constant type; type B: medium density-gradual increase type; type C: medium density-wavy change type; type D: high density-large amplitude type. The border between type B and type C was not clear. Although the distribution of the fluctuation types did not show any geographical cline, the high density type (type D) occurred in limited areas where the winter was severe and spring came later and almost all the populations in the Oshima peninsula were the low density type (type A). The most common type of population in Hokkaido, type C, greatly fluctuated from year to year with the maximum density level/the minimum one being over 10. The periodicity of the peak years was not statistically significant, although peaks often occurred at 3 or 4 year intervals. The population grew from spring to autumn in almost all years so that the pattern of the seasonal change was qualitatively constant. The population density levels in spring were not significantly different between peak years and others, while those in autumn were greatly different between them, so that the population growth rates from spring to autumn were variable from year to year. This differential pattern was also found between high and low density areas.     

Identification of motivational state in adult male Atlantic walruses inferred from changes in movement and diving behavior

Motivational changes in animals are likely to be detectable retrospectively through observed changes in behavior. Breeding represents one of the strongest motivational states in mammals, and its timing is often tied to a seasonally optimal suite of environmental and physical conditions. While seasonal changes in behavior may be directly observable in some species, for others that breed cryptically or in difficult to access areas, detecting behavioral changes may only be feasible using data collected remotely. Herein, we explore whether behavioral changes can be used to infer motivational state for a wild, free‐ and wide‐ranging high arctic marine mammal, adult male Atlantic walruses (Odobenus rosmarus rosmarus). Using satellite‐relayed location and dive data from 23 adult male walruses instrumented in the Svalbard Archipelago, we identify seasonal movement to discrete geographic regions deep into winter pack ice. Adult male walrus diving behavior underwent marked seasonal movements between geographical areas that coincided with changes in light regime. At offshore wintering sites adult males (n = 4) shifted from a summer pattern of deep, long benthic dives to much shallower diving. Some males performed similar shallow, winter dive behavior at coastal locations (n = 12) suggesting that breeding might also occur around the coast of Svalbard. However, interpretation of behavioral changes of these coastal individuals was challenging. The presumed breeding sites at the winter off‐shore locations were situated in areas where polynyas are known to occur, making them a predictable resource even if they are located deep inside the winter pack‐ice. We demonstrate that remotely collected behavioral data can be used to identify seasonally explicit changes in the behavior of cryptic species.     

Seasonal and spatial pattern of shelter use by badgers<Emphasis Type="Italic">Meles meles</Emphasis> in Białowieża Primeval Forest (Poland)

In 1997–2001, we investigated the use of day-time shelters by radio-collared badgersMeles meles (Linnaeus, 1758) in the Białowieża Primeval Forest, eastern Poland. Each social group of badgers utilised, on average, 9 different shelters per territory (range: 4–20). The main setts, occupied for breeding and winter sleep, were also most frequently used for day-time rest throughout the year (73% of days). Badgers living in the pristine oldgrowth stands utilised larger number of shelters and spent more days in hollow trees (mainly limeTilia cordata), compared to badgers inhabiting younger secondary tree stands. Number of shelters used by individuals varied between seasons and depended on sex and age of animals. In summer, badgers used more shelters than in spring and autumn. In winter, they stayed in their main setts only. Adult males occupied more shelters and spent fewer days in the main sett than other badgers. In spring, females rearing young used only the main setts. The average underground space used by badgers within the main sett was 128 m². It was largest in summer and smallest in winter, and also varied between males and females. We proposed that, in a low-density population, badgers used several setts and other daily shelters to reduce energy expenditure when exploring their large territories and foraging. Furthermore, setts may play a role of marking sites. Analysis of the biogeographical pattern of sett use by European badgers showed that the number of setts used by social groups increased with increasing territory size, whereas the density of setts (n setts/km²) was negatively correlated with territory size. We proposed that different factors could shape the utilisation of setts by badgers in low- and high-density populations.     

Changes in scale circulus spacings of an endangered Atlantic salmon Salmo salar population: evidence of a shift in marine migration?

Post‐smolt scale circulus spacing patterns for two Atlantic salmon Salmo salar populations from the Southern Upland (SU) of Nova Scotia, Canada, were compared with spacings from two endangered populations from the inner Bay of Fundy (iBoF) Nova Scotia and New Brunswick, to determine if growth patterns differed among these populations, and if growth patterns had changed as the abundance of these populations declined. An analysis of numbers of marine circuli from scales of post‐smolts and one‐sea‐winter adults of known age indicated that circuli were deposited at a rate of about one circulus per week in summer and slowed to one every 2 weeks in winter. During the summer and the autumn, mean circulus spacing in the iBoF populations, known to have occupied the outer Bay of Fundy during these seasons, was lower than in the SU populations, which are known to migrate to the North Atlantic. Similar circulus spacing patterns within SU populations is suggestive of a common marine distribution for these populations. In contrast, a cluster analysis revealed that within the geographically intermediate Big Salmon River (iBoF), some individuals exhibited wider spacing patterns that resemble the distant migrating SU populations, while others exhibited narrower spacing similar to other iBoF S. salar. Within the Big Salmon River, the relative abundance of the wider and the narrower spacing patterns varied in the earlier years, but all fish sampled since 1999, exhibited wider spacings similar to distant migrating SU S. salar. The apparent disappearance of the narrower pattern, characteristic of localized migration and indicative of historical iBoF populations, suggests that local migration may not presently be a successful strategy for these populations.     

Over-wintering strategies of the Lesser Sheathbill Chionis minor in an impoverished and insular environment

Summary The Lesser Sheathbill Chionis minor is an opportunistic predator and scavenger that breeds on sub-Antarctic islands during the summer season, when many other seabirds are present, especially penguins on which it is kleptoparasitic. It is a poor flyer and over-winters on its breeding grounds, where it faces a diminished food supply and low temperatures. Its foraging behaviour was suspected to be plastic enough to respond to such a variable environment, and we show how this species enlarged its niche to survive in winter. Population movements, dietary changes and monthly weight gains were quantified during a 13-month period of observation. Mortality rates, calculated over a 19-year period of banding, appeared to be age dependent: they decrease, due to competition, from juveniles to subadults, adult non-breeders, and breeders. One of the survival strategies described here is, to our knowledge, unique among birds. After the Crested Penguin colonies have been deserted, some adult sheathbills (mainly females) remained territorial, extending their territory size and diet while their partners moved to previously undefended zones in King Penguin colonies (permanent and large in the Crozet archipelago), where they competed for a winter territory and subsequently associated with another mate. After this winter mating, mobile sheathbills returned the following summer to their breeding territory and previous mate.     

Proximal Spacing Between Individuals in a Group of Woolly Monkeys (Lagothrix lagotricha) in Tinigua National Park, Colombia

I studied proximal spacing within a group of woolly monkeys (Lagothrix lagotricha) during 7 months at Parque Nacional Tinigua, Colombia. I collected a total of 1188 instantaneous samples on focal individuals, recording the number and age/sex class of individuals that were in contact with, <2 m from, <5 m from the focal animal. The results indicate that proximate spacing reflects social affinities and is related to mother–infant relationship and social grooming. Subadult females and adult males are the sex/age classes with the lowest number of individuals in proximity. There are low proximity between adult females and between adult males and high frequencies of nearness between mother and offspring. Associations between males and females were usually low, but in some cases males showed preferences for a given female. There was a relatively gradual increase in spacing between mothers and their offspring as they became older. Old juvenile males were associated chiefly with other males—mostly subadults—whereas juvenile females maintained some proximity only to their mothers. There are also differences in spacing behavior according to different activity types.     

Winter territory prospecting is associated with life‐history stage but not activity in a passerine

Finding a high quality territory is essential for many animals to reproduce successfully. Despite its importance for fitness, we know little about the process of territory prospecting in wild birds, and whether individual traits and behaviours, such as personality, co‐vary with territory prospecting. Here, we use long‐term data from a wild, insular house sparrow Passer domesticus population to test three hypotheses about territory fidelity and prospecting: 1) house sparrows show high territory fidelity between years and also during winter. 2) Individuals will prospect for a breeding territory during their first winter whereas older, more experienced individuals will keep a territory from previous years and will, therefore, show no or reduced winter territory prospecting. 3) More active behavioural types will prospect more than less active behavioural types. We use data from four winters from automatically, daily recorded nest‐box visits of 188 birds of known age. The number of nest‐boxes that each individual visited within each winter was used as a proxy of winter territory prospecting. We show that house sparrows visit multiple nest‐boxes during their first winter, whereas older individuals keep territories year‐round and, potentially because of this, indeed show reduced winter territory prospecting. Activity was not associated with the number of nest‐boxes visited. Further research is needed to investigate whether time of territory and mate acquisition differs among individuals and the possible effect on lifetime fitness.     

Competitive abilities of oystercatchers ( Haematopus ostralegus ) occupying territories of different quality

In territorial species, habitat heterogeneity results in some individuals occupying poor quality sites while others occupy high quality sites. Floaters (mature nonbreeders) may accept a low quality territory, because it is the best they can get and defend (‘inferior phenotype hypothesis’), or because it is a strategic alternative for a high quality territory in the long run (‘queue hypothesis’). Oystercatcher Haematopus ostralegus territories differ consistently in the amount of offspring produced each year and this is related to the distance between the nesting and feeding territories. The inferior phenotype hypothesis was previously rejected on the basis of the absence of morphometric differences (assumed to indicate competitive abilities) among breeders. We investigated social dominance, in the field and in captivity, in relation to the quality of the breeding territory. In the field, birds with high-quality territories won more often compared to those occupying low-quality territories. However, this difference was not apparent in a small dataset of captive birds. These results are discussed in the framework of the long-term fitness prospects of settling in a high or low quality territory.     

The role of vole populations in prevalence of the parasite (Echinococcus multilocularis) in foxes

Effects of population fluctuation of the gray-sided vole(Clethrionomys rufocanus) on the prevalence (infection rates) of the parasiteEchinococcus multilocularis in red fox(Vulpes vulpes) populations was investigated from 1985 to 1992 in eastern Hokkaido (Abashiri, Nemuro, and Kushiro area), Japan. This parasite needs two hosts to complete its life cycle; the gray-sided vole as its intermediate host and the red fox as its final host. We found that: (1) Infection rates in foxes depended on the current-year abundance of voles in all three study areas, particularly in Abashiri. (2) In addition to this direct density-dependence, delayed density-dependence between the infection rate and the prior-year abundance of voles was detected in Nemuro and in Kushiro. (3) The regional differences in density-dependence pattern were related to regional differences in the winter food habits of red foxes: in Abashiri the proportion of voles in the fox’s diet greatly decreases in winter, while the proportion remains high in winter in Nemuro and in Kushiro, probably because of shallower snowpack. These results suggest that infection rates in foxes in Abashiri were less influenced by the prior-year prevalence, since the infection cycle might be interrupted in winter, when voles became less important in fox’s diet. In contrast, the state of the prevalence may carry over from year to year in Nemuro and in Kushiro, because red foxes continue to eat a considerable amount of voles throughout year. The regionally contrasted results for the relationship between infection rate in foxes and vole abundance were parallel to the regional difference in fluctuation pattern of vole populations, which are highly variable in Abashiri area, but less variable in Kushiro-Nemuro area. Drastic change in vole populations appears to affect the host-parasite system.     

Planting of fungus onto hibernating workers of the fungus-growing ant Mycetosoritis clorindae (Attini, Formicidae)

We describe a peculiar fungus-coating behavior of the attine ant Mycetosoritis clorindae, where workers plant fungal mycelium on hibernating nestmates. Hibernating nestmates become ultimately enveloped in a live mycelial coat, remain motionless in this coated state, and essentially become integrated into the garden matrix. The shallow nest architecture of M. clorindae (depth of main garden is 15–30 cm) in southern Brazil forces the ants to overwinter at relatively low temperatures in the topmost soil layer. Fungal coating may help the ants to survive the prolonged periods of immobility during winter. Fungus-planting on attine adults is so far unknown from other attine species, but the behavior parallels the planting of mycelium on larvae and pupae occurring in many attine species. Planting of mycelium on adult nestmates may have been overlooked so far in attine ants because this behavior may occur only in dormant nests, which are least frequently collected. The possible adaptive functions of fungus coatings of hibernating adults and developing brood are likely similar, including for example physical protection, prevention of desiccation, shielding against parasites and predators (e.g., army ants), or defense against diseases.     

A comparison of intergroup behavior inCebus albifrons andC. apella

Comparative studies of free-ranging groups ofCebus albifrons andCebus apella in eastern Colombia showed strong differences in the manner in which groups of each species interacted.Cebus albifrons troops were large and multi-male; they showed strongly aggressive tendencies towards neighboring groups of the same species and they defended a territory with little overlap of neighbors, both by means of long-distance spacing calls and by fighting. In contrast,Cebus apella troops were small, sometimes containing only one adult male; they exhibited pacific interactions and curiosity towards neighboring groups of the same species and did not defend a territory, allowing extensive overlapping with their neighbors. Such social differences in closely related species which are found in identical habitats call into question purely ecological interpretations for such differences.     

Territorial behavior of Western Sandpipers on their nonbreeding grounds: effect of sex and foraging interference

ABSTRACT Nonbreeding shorebirds often alternate social structure between anonymous flocks and territorial behavior in response to different environmental factors. To evaluate specific drivers for one species, we studied the spacing behavior of wintering Western Sandpipers (Calidris mauri) at Bahía Santa María, northwestern Mexico, using behavioral observations. The density, population structure, and territorial behavior of Western Sandpipers differed among three habitat types. Cattail marshes supported 110 birds per ha, 49% males, and no territorial birds. Mangrove flats supported 288 birds per ha, 58% males, and 5% territorial birds. Brackish flats supported 365 birds per ha, 76% males, and 7% territorial birds. Territories consisted of rectangular strips (5–9 m long, N= 77). Territory length was not related to either bird density or number of territorial birds by plot, but was positively related to nearest bird distance. Aggression rate was inversely related to territory length, suggesting that territory length is set by the costs of defense. Foraging rate was independent of territory length, and prey densities in territories did not differ from those in areas used by nonterritorial birds. Males were more likely to be territorial and had a higher aggression rate than females, suggesting that males, which forage more on surface prey, were more affected by foraging interference. Our results suggest that the territorial behavior of Western Sandpipers in our study was an opportunistic strategy to reduce foraging interference. The variation in spacing behavior we documented provides evidence that interference competition affects the social structure of Western Sandpipers during the nonbreeding season.     

Territory occupancy rate of goshawk and gyrfalcon: no evidence of delayed numerical response to grouse numbers

Two recent studies on territory occupancy rates of goshawk Accipiter gentilis and gyrfalcon Falco rusticolus report a 2–3-year-delayed numerical response to grouse numbers, which is a requirement for a hypothesis of predator-generated grouse cycles. The time lags were assumed to reflect the average age of sexual maturity in the raptor species. In southern Norway, however, subadult (two-year-old) goshawk hens occupied only 18–25% of territories where occupancy was not recorded in the preceding year, and there was no significant relationship between the proportion of subadults among recruits and grouse indices two years earlier. We argue that territory occupancy rates are not appropriate indices of total raptor population levels, but rather reflect the proportion of territorial pairs that attempt to nest. Because this depends on the body condition of the hens, fluctuations in other important winter resident prey species (most important for the goshawk) and winter weather (most important for the gyrfalcon) should also be addressed. During 1988–2006, the annual proportion of goshawk territories with recorded nesting attempts in southern Norway was most closely related to the preceding autumn’s population indices of black grouse Tetrao tetrix and mountain hare Lepus timidus, whereas the annual proportion of gyrfalcon territories with observations of falcons or with confirmed breeding attempts in central Norway were best explained by population indices of willow grouse Lagopus lagopus and ptarmigan L. mutus from the previous autumn, and by December temperatures. Hence, our studies do not support the predation hypothesis for grouse cycles.     

Lifetime reproductive success in reproductively suppressed female voles

A population of the grey red-backed vole, Clethrionomys rufocanus bedfordiae, was investigated on a 1 ha control grid and a 1 ha grid on which the voles were fed within a 2.1 ha outdoor enclosure in Hokkaido, Japan by live trapping from 1984 to 1986, for testing the Reproductive Suppression Model of Wasser and Barash (1983)-females can optimize their lifetime reproductive success by suppressing reproduction when future conditions for the survival of offspring are likely to be sufficiently better than present ones as to exceed the costs of the suppression itself. Age at the first pregnancy more varied in a higher density population on the experimental grid and females could be classified into the early and the late reproductive type in two generations (A: females born from February to June 1985; B: females born from September to November 1985). Lifetime reproductive success (the number of pregnancies, the number of successful litters, and the number of offspring) was not different between the early and the late reproducing females. The late reproducing females lived for longer periods than the early reproducing females, so that the loss by delayed start of reproduction was compensated for by a longer life span. Life span was not different between offspring of the early and the late reproducing females. These facts supported the Reproductive Suppression Model.     

On the ecology and behavior ofCebus albifrons in eastern Colombia: II. Behavior

A 503-contact-hr study of a 35 member group ofCebus albifrons was conducted in eastern Colombia in 1977 and 1978. The reactions of the group often centered on the largest adult male, which seemed dominant over all other members of the group. This “alpha” male reacted to perceived danger by putting himself between the group and the danger where he threatened and observed until the other members left the scene. Many members of the group sought the alpha's physical support while using threatening behaviors. Much tolerance towards juveniles and infants was shown by adult and subadult males which included carrying young animals out of danger. Territorial behavior was observed towards neighboring groups. Many behaviors were similar to other species ofCebus.     

High site fidelity in Northern Wheatears Oenanthe oenanthe wintering in Africa revealed through colour marking

Wintering Northern Wheatears Oenanthe oenanthe in the Sahel region of Northern Nigeria held small (approximately 70?m diameter) distinct territories during the study period, and territory size did not differ between adult and first winter birds. Evidence suggests that Wheatears may maintain small territories for a significant duration of the winter, similar to many other migrants.