Modelled population growth based on reproduction differs from life tables based on age determination in Danish raccoon dogs ( Nyctereutes procyonoides )

The population of raccoon dogs (Nyctereutes procyonoides) in Denmark has increased rapidly from 1995 when the first was recorded until today where 3291 raccoon dogs are trapped, shot by hunters or road killed. The aims of this study are to present the first data on reproduction and life tables of raccoon dogs in Denmark and to compare mortality from modelled life tables with game bag records and sampled raccoon dogs in different age groups. In this study, the uteri of 89 adult females (> 10 months) were examined for placental scars (PSC), and 561 individuals (289 males, 272 females) were aged using pulp cavity width and dental lines in canine teeth. The litter size of raccoon dogs in Denmark is to date the largest litter size recorded in the wild (mean ± SE) 10.8 ± 0.4, range 1–16 pubs and fecundity 8.4 ± 0.6 pubs. The percent-reproducing females are 78–83%, based on dark and all PSC, respectively. A significant difference was found between the proportion of individuals composing the different age groups based on age determination of individuals collected (Ntage) and the modelled number of individuals in age groups based on fecundity and different mortality rate (Ntmodel), X2 = 8, p < 0.05. The discrepancy between the relatively high reproduction and lifetables may be due to older and more experienced animals that avoid culling. A low population density in a newly founded Danish population of raccoon dogs, together with a milder climate where raccoon dogs can forage during the winter, may cause an exceptionally high reproduction in Danish raccoon dogs.     

Reliability of placental scar counts in the Red fox {Vulpes vulpes L.) with special reference to fading of the scars

The use of placental scar counts in determining litter size and pregnancy rate among Red foxes ( Vulpes vulpes L.) is reviewed. There is no experimental material to test the validity of the method, although many authors have discussed it. Recent workers have (1) included uteri from vixens killed later in the season and, (2) become more concerned about effects of resorptions and/or abortions on scar counts, as well as the possibility of scars persisting from earlier pregnancies. In the present study, scar counts overestimated numbers of persisting corpora lutea to a higher degree in old vixens than in yearlings; and scars were shown to fade over winter. Estimates of frequency of breeding among females, based on various combinations of placental scars of different shades, and on date of death, are discussed. The relation between number of scars of different shades, date of death and known minimum litter size in three wild and one captive vixen is compared with the foregoing. Counts of all scars evidently overestimate litter size at birth and breeding frequency, probably because some scars originate at resorption sites and others persist from earlier pregnancies. Only dark scars should be counted early in the season, but those of increasingly lighter shade should be included as winter progresses. A guide to counting placental scars over winter is presented.     

Maternal body size as a morphological constraint on egg size and fecundity in butterflies

It is a widespread notion that in arthropods female reproductive output is strongly affected by female size. In butterflies egg size scales positively with female size across species, suggesting a constraint imposed by maternal size. However, in intraspecific comparisons body size often explains only a minor part of the variation in progeny size. We here include representatives of various butterfly families to test the generality of this phenomenon across butterflies. Phenotypic correlations between egg and maternal body size were inconsistent across species: correlations were non-significant for Pararge aegeria and Lycaena tityrus, significantly positive for Papilio machaon, significantly negative for Araschnia levana, and contradictory for Pieris napi. Thus, there was no general pattern linking egg size to maternal size, e.g., caused by an allometric relationship. Consequently, there was at best limited evidence for maternal size acting as a morphological constraint on egg size within butterfly species. Realized fecundity depended on maternal size in P. napi and A. levana, but not in P. aegeria, suggesting that maternal size may affect egg number more strongly than egg size. Yet, variation in fecundity was primarily explained by variation in longevity as is expected for income breeders. Heritability estimates across species were rather similar for pupal mass (ranging between 0.14 and 0.19), but more variable for egg size (0.17–0.31).     

Rural populations of the red fox Vulpes vulpes show little evidence of reproductive senescence

The ageing theory predicts fast and early senescence for fast-living species. We investigated whether the pattern of senescence of a medium-sized, fast-living and heavily-culled mammal, the red fox (Vulpes vulpes), fits this theoretical prediction. We used cross-sectional data from a large-scale culling experiment of red fox conducted over six years in five study sites located in two regions of France to explore the age-related variation in reproductive output. We used both placental scars and embryos counts from 755 vixens’ carcasses aged by the tooth cementum method (age range: 1–10), as proxies for litter size. Mean litter size per vixen was 4.7 ± 1.4. Results from Generalized Additive Mixed Models revealed a significant variation of litter size with age. Litter size peaked at age 4 with 5.0 ± 0.2 placental scars and decreased thereafter by 0.5 cubs per year. Interestingly, we found a different age-specific variation when counting embryos which reached a plateau at age 5–6 (5.5 ± 0.2) and decreased slower than placental scars across older ages, pointing out embryo resorption as a potential physiological mechanism of reproductive senescence in the red fox. Contrary to our expectation, reproductive senescence is weak, occurs late in life and takes place at an age reached by less than 11.7% of the population such that very few females exhibit senescence in these heavily culled populations.     

Reproductive investiment in the scorpion Centruroides vittatus

Among invertebrates, scorpions possess a relatively unique set of reproductive traits. The interrelationships of these traits may have important implications for life history theory, yet there have been few studies of these traits in scorpions. Our data indicate that larger female Centruroides vittatus produce more offspring and have a higher total litter mass than smaller females. There was, however, no significant relationship between offspring size and female or litter size. Mean offspring mass increased with increases in total litter mass and within litter variation in offspring size (coefficients of variation) decreased with increasing total litter mass. These results suggest that large female scorpions with a larger investment in reproduction produced more offspring that were more uniform in size, but not significantly larger, than small females with less investment. The fractional clutch principle and physiological and functional constraints on size and number of offspring are suggested as possible explanations for the relationships we found among offspring size, variation in offspring size and total investment in offspring in C. vittatus.     

Population genetic structure and distribution of introduced American mink (<Emphasis Type="Italic">Mustela vison</Emphasis>) in Spain,based on microsatellite variation

The population genetic structure of an invasive species in Spain, the American mink (Mustela vison), was investigated using microsatellite DNA markers. This semi-aquatic carnivore, originating from North America, was imported into Europe for fur farming since the beginning of the 20th century. Due to massive escapes, farm damages, deliberate releases and/or accidents, feral mink populations were established in the aquatic ecosystems of many European countries, including Spain. We genotyped 155 American mink originating from the Spanish regions Basque Country, Catalonia, Castilla-Leon, Aragon, Valencia and Galicia using 10 polymorphic microsatellite loci to highlight population genetic structure, distribution and dispersal. M. vison populations in Spain appear differentiated and not yet connected by gene flow. Bayesian clustering analyses and spatial analyses of molecular variance detected four inferred clusters, overall coinciding with the sampled geographical localities. Preliminary testing shows moderate to large estimated effective population sizes. Molecular analyses result useful to provide baseline data for further research on the evolution of invasive mink populations, as well as support local management strategies and indirectly benefit the conservation of threatened species in Spain, such as the endangered European mink (Mustela lutreola), and the polecat (Mustela putorius), which share the habitat with the American mink. This paper is dedicated to the memory of Xavier Domingo-Roura.     

Reproductive biology of Rutilus rubilio (Bonaparte, 1837) in Lake Trichonis

The reproductive biology of R. rubilio in lake Trichonis was examined. Absolute fecundity depended on size, age and condition of the female spawners but these relationships were less clear in the case of relative fecundity and egg size. The adaptive significance of the reproductive process of R. rubilio in the oligotrophic conditions of the lake is discussed.     

Determinants of reproductive success in voles: space use in relation to food and litter size manipulation

Spacing behaviour of female mammals is suggested to depend on the distribution and abundance of food. In addition, food limitation has been found to constrain the reproductive success of females. However, whether females maximize their reproductive success by adjusting space use in relation to current food availability and reproductive effort (e.g. litter size) has not been experimentally studied. We examined these questions by manipulating simultaneously food resources (control vs. food supplementation) and litter sizes (control vs. plus two pups) of territorial female bank voles (Clethrionomys glareolus) in large outdoor enclosures. Females with supplementary food had smaller home ranges (foraging area) and home range overlaps than control females, whereas litter size manipulation had no effect on space use. In contrast, the size of territory (exclusive area) was not affected by food supplementation or litter size manipulation. As we have previously shown elsewhere, extra food increases the reproductive success of bank vole females in terms of size and proportion of weaned offspring. According to the present data, greater overlap of female home ranges had a negative effect on reproductive success of females, particularly on survival of offspring. We conclude that higher food availability increases the reproductive success of bank vole females, and this effect may be mediated through lower vulnerability of offspring to direct killing and/or detrimental effects from other females in the population. Moreover, it seems that when density of conspecifics is controlled for, home range sizes of females, but not territoriality, is related to food resources in Clethrionomys voles.     

Age specific fecundity,litter size,and sex ratio in the chacoan peccary (Catagonus wagneri)

Ten years of data on the reproductive biology of the Chacoan peccary (Catagonus wagneri) were analyzed to determine average litter size, sex ratio, timing of births, and individual and age-specific fecundity. Data were obtained from a captive herd of chacoan peccaries located in the western Paraguayan Chaco. Births peak in the austral spring months of September, October, and November, with fewer litters born during the dry season months of June, July, and August. The average litter size was 2.4 with a sex ratio of 56:44 (M:F). There was no significant difference in litter size among individual females or among females of different ages. Finally, there was some individual variation in the age at which sows produced their first litter, but no discernible variation among sows in the average time between litters. Zoo Biol 16:301–307, 1997. © 1997 Wiley-Liss, Inc.     

The interplay between natural and sexual selection in the evolution of sexual size dimorphism in Sceloporus lizards (Squamata: Phrynosomatidae)

Sexual size dimorphism (SSD) evolves because body size is usually related to reproductive success through different pathways in females and males. Female body size is strongly correlated with fecundity, while in males, body size is correlated with mating success. In many lizard species, males are larger than females, whereas in others, females are the larger sex, suggesting that selection on fecundity has been stronger than sexual selection on males. As placental development or egg retention requires more space within the abdominal cavity, it has been suggested that females of viviparous lizards have larger abdomens or body size than their oviparous relatives. Thus, it would be expected that females of viviparous species attain larger sizes than their oviparous relatives, generating more biased patterns of SSD. We test these predictions using lizards of the genus Sceloporus. After controlling for phylogenetic effects, our results confirm a strong relationship between female body size and fecundity, suggesting that selection for higher fecundity has had a main role in the evolution of female body size. However, oviparous and viviparous females exhibit similar sizes and allometric relationships. Even though there is a strong effect of body size on female fecundity, once phylogenetic effects are considered, we find that the slope of male on female body size is significantly larger than one, providing evidence of greater evolutionary divergence of male body size. These results suggest that the relative impact of sexual selection acting on males has been stronger than fecundity selection acting on females within Sceloporus lizards.     

The influence of American mink odour on the spatial distribution and behaviour of water voles

The perception and assessment of predation risk often cause changes in the activities of animals and induce behavioural responses that may in turn affect their movements and distribution. To simulate high predation risk in a midfield pond riparian habitat, we used fresh faeces from ranch American mink Neovison vison and recorded behavioural responses of water voles Arvicola amphibius. In areas where mink odour was deployed, the numbers of captured vole individuals and their trappability were significantly lower than in control areas. Several voles migrated from the zones with deployed mink faeces to the areas without faeces, thus proving that increased predation risk affects the distribution of individuals in a population. The response to mink odour was much more pronounced in females than in males; in areas with deployed mink faeces, not a single female was trapped. We conclude that although American mink is a non‐native, invasive predator, water voles respond to mink odour by reducing their activity and/or by avoiding places with higher predation risk.     

Folliculogenesis, onset of puberty and fecundity of mink (Mustela vision Schreb) selectively bred for docility or aggressiveness

It has been suggested that selective breeding of animals for docile behavior is correlated with early onset of puberty and improved fertility. We wished to test the hypothesis that mink bred for docility would show earlier onset of puberty and greater fecundity than mink bred for aggressiveness. We used farm-raised, 7-mo-old mink females that had been selectively bred for 7 to 10 generations on the basis of behavior towards humans. Onset of puberty was estimated once (between 15 and 20 December) by vaginal smears and was said to start wtih preponderance of cornified epithelial cells in the cytological specimen. Fecundity was measured by litter size and rate of folliculogenesis, with and without hCG stimulation, by histomorphometric examination of ovaries and uteri. A total of 43/100 (43%) docile females achieved proestrus and estrus as compared to 16/136 (12%) of the aggressive ones. Overall pregnancy rate, survival to 5 d after whelping and litter size did not differ between the docile and aggressive females. Docile females showed significantly higher numbers (P < 0.05 and < 0.001) of growing, maturing and atretic follicles than the aggressive ones, however the latter showed a highly significant (P < 0.001) folliculogenic response to hCG. The response of the ovary of aggressive females to hCG is particularly dramatic because in most aggressive females the ovaries contained none or only few follicles of any kind. The inhibition of folliculogenesis in aggressive mink is similar to that reported in Silverblue and Sapphire mink homozygous for the Stuart factor. The paradoxical response to hCG may be an indication that selective breeding for aggression may be correlated with the disturbance at the early stages of folliculogenesis which creates a deficiency of follicles that are sensitive to LH-type stimulation.     

Hormonal manipulation of offspring number: maternal effort and reproductive costs

We used exogenous gonadotropin hormones to physiologically enlarge litter size in the bank vole (Clethrionomys glareolus). This method allowed the study design to include possible production costs of reproduction and a trade-off between offspring number and body size at birth. Furthermore, progeny rearing and survival and postpartum survival of the females took place in outdoor enclosures to capture salient naturalistic effects that might be present during the fall and early winter. The aim of the study was to assess the effects of the manipulation on the growth and survival of the offspring and on the reproductive effort, survival, and future fecundity of the mothers. Mean offspring body size was smaller in enlarged litters compared to control litters at weaning, but the differences disappeared by the winter. Differences in litter sizes disappeared before weaning age due to higher mortality in enlarged litters. In addition to the effects of the litter size, offspring performance was probably also influenced by the ability of the mother to support the litter. Experimental females had higher reproductive effort at birth, and they also tended to have higher mortality during nursing. Combined effects of high reproductive effort at birth and high investment in nursing the litter entailed costs for the experimental females in terms of decreased probability of producing a second litter and a decreased body mass gain. Thus, enlarged litter size had both survival and fecundity costs for the mothers. Our results suggest that the evolution of litter size and reproductive effort is determined by reproductive costs for the mothers as well as by a trade-off between offspring number and quality.     

Potential Impact of an Exotic Mammal on Rocky Intertidal Communities of Northwestern Spain

Being the interface of sea and land, the coast can be invaded by introduced species coming from either of these two worlds. Recent reviews of coastal invasions emphasize the human-mediated transport of non-indigenous marine plants and invertebrates, forgetting the potential role of invaders of terrestrial origin. By studying the diet of the introduced American mink (Mustela vison) on a rocky shore of southwestern Europe, we draw attention to the potential impact on intertidal communities of exotic species coming from inland. We analysed 199 mink faeces collected in August 1997 and August 1999 in Baiona, a coastal and urban area of northern Spain recently invaded by minks. The diet of the species was based almost exclusively on crabs (45.4% of individual prey) and fish (53.3%). Most crabs were marbled crabs (Pachygrapsus marmoratus) and most fish were adult blennies (Coryphoblennius galerita and Lipophrys pholis). Given its energy requirements (about 1250 kJ/day), a single mink will consume during the month of August approximately 945 blennies and 496 crabs. Although we lack accurate data on mink abundance, a cautious estimation (4 mink/km before dispersal), supported by field observations, suggests that predation in August may reach 3780 blennies and 1984 crabs per km of shoreline. This predation pressure could affect the numbers of blennies and (less probably) crabs, indirectly benefiting the populations of their prey, that is, sessile invertebrates and snails. More field research is needed, but our results suggest that an exotic non-marine top predator such as the American mink could affect intertidal communities in Eurasia.     

Gender differences in activity patterns of American mink <Emphasis Type="Italic">Neovison vison</Emphasis> in Germany

Circannual and circadian activity patterns of American mink Neovison vison were studied in a fishpond area in northern Germany. Fourteen mink were radiotracked, nine of them (five males, four females) for at least 6 months, some of them even up to 21 months. Both sexes have reduced activity rates in winter months and showed a considerable increase of activity during mating season (March). In summer months—during the pup raising period—female activity continued to be high. Male mink, on the other hand, were less active with the exception of August and September. During this time, dispersal of juveniles takes place with resident males increasingly having to defend their territories. Circadian activity rhythms differed markedly between sexes. All investigated females exhibited a perennial diurnal pattern. Female activity during twilight and night was significantly lower than during daylight. Three investigated male mink showed a typical nocturnal course throughout the year. Daylight activity of these animals was very low, increased during dusk and dawn activity rates and stayed at a higher level during the night. Two males behaved arrhythmic; they did not prefer any time of the day for activity. The observed temporal strategies were discussed in context of intra-sexual territoriality, sexual dimorphism in body size and different environmental factors.     

Reproductive Ecology and Cub Survival of Florida Black Bears

Abstract: We investigated reproductive ecology and cub survival of Florida black bears (Ursus americanus floridanus) in Ocala National Forest and the adjacent residential area of Lynne, Florida, USA, 1999-2003. We documented production of 81 cubs from 39 litters. Average litter size was 2.08 ± 0.11 (SE) cubs. The mean age of first reproduction was 3.25 ± 0.27 years. Excluding females that reproduced in consecutive years due to litter loss, interlitter interval was 2.11 ± 0.11 years. The mean annual fecundity rate was 0.57 ± 0.06. We used expandable radiocollars to monitor the fate of 41 bear cubs. The probability of cubs surviving to 9 months of age was 0.46 ± 0.09 and did not differ between cohorts or study locations. The most important causes of cub mortality included infanticide and mortality caused directly or indirectly by collisions with vehicles. Our results indicate that reproductive rates of female black bears in the Ocala study area are comparable to those reported for other black bear populations from eastern United States, but cub survival rates are lower than those reported for most black bear populations. Management of Florida black bears should emphasize strategies to reduce the mortality of cubs.     

Female fecundity traits in wild populations of African annual fish: the role of the aridity gradient

The evolution of life history is shaped by life expectancy. Life‐history traits coevolve, and optimal states for particular traits are constrained by trade‐offs with other life‐history traits. Life histories contrast among species, but may also diverge intraspecifically, at the level of populations. We studied the evolution of female reproductive allocation strategy, using natural populations of two sympatric species of African annual fishes, Nothobranchius furzeri and Nothobranchius orthonotus. These species inhabit pools in the Mozambican savanna that are formed in the rainy season and persist for only 2–10 months. Using 207 female N. furzeri from 11 populations and 243 female N. orthonotus from 14 populations, we tested the effects of genetic background (intraspecific lineage) and life expectancy (position on the aridity gradient determining maximum duration of their temporary habitat) on female fecundity traits. First, we found that variation in female body mass was small within populations, but varied considerably among populations. Second, we found that fecundity was largely defined by female body mass and that females spawned most of their eggs in the morning. Third, we found that the trade‐off between egg size and egg number varied among lineages of N. furzeri and this outcome has been confirmed by data from two separate years. Overall, we demonstrate that local conditions were important determinants for Nothobranchius growth and fecundity and that eggs size in arid region was less limited by female fecundity than in humid region.     

Modelling space use and dispersal of mammals in real landscapes: a tool for conservation

Aim To explore the usefulness of Spatially Explicit Population Models (SEPMs), incorporating dispersal, as tools for animal conservation, as illustrated by the contrasting cases of four British mammals. Methods For each of the four species (American mink, Mustela vison, pine marten, Martes martes, dormouse, Muscardinus avellanarius and water vole, Arvicola terrestris) a spatial dynamics model was developed based on an integrated geographical information system (GIS) population model that linked space use to the incidence of the species. Each model had, first, a GIS, which stored environmental, habitat and animal population information, and secondly, an individual‐based population dynamics module, which simulated home range formation, individual life histories and dispersal within the GIS‐held landscape. Results The four models illustrated different interactions between species life‐history variables and the landscape, particularly with respect to dispersal. As water voles and dormice occupy home ranges that are small relative to blocks of their habitat, they were most effectively modelled in terms of the dynamics of local populations within habitat blocks but linked by dispersal. In contrast, because the home ranges of American mink and pine marten are large relative to blocks of habitat, they were best modelled as individuals moving through a landscape of more or less useful patches of habitat. For the water vole, the most significant predictors of population size were the carrying capacity of each habitat and the annual number of litters. For the dormouse, the likelihood of catastrophe and the upper limit to dispersal movement were the key variables determining persistence. Adult mortality and home‐range size were the only significant partial correlates of total population size for the American mink. Adult mortality was also a significant correlate of total population size in the pine marten, as were litter size and juvenile mortality. In neither the marten nor the mink was dispersal distance a significant factor in determining their persistence in the landscape. Main conclusions At a landscape scale it is difficult to measure animal distributions directly and yet conservation planning often necessitates knowledge of where, and in what numbers, animals are found, and how their distributions will be affected by interventions. SEPMs offer a useful tool for predicting this, and for refining conservation plans before irreversible decisions are taken in practice.     

The cost of maturing early in a solitary carnivore

Central to the theory of life history evolution is the existence of trade-offs between different traits, such as the trade-off between early maturity and an extended period of body growth. Based on analysis of the reproductive tracts of harvested Eurasian lynx (Lynx lynx) females in Norway, we find that females that mature early are generally heavier than those that postpone maturation. A higher proportion of 1.5-year-old females showed signs of ovulation in areas with high prey density, where they were also heavier. Further, we show that female Eurasian lynx that mature early have the same number of placental scars (an index of breeding investment and litter size) as older females, suggesting that they have a relatively high investment in their first litter. This induces a cost in terms of body weight development, as those females that had matured at the age of 1.5 years were substantially lighter by the age of 2.5 years than those that postponed breeding. This effect tended to be more pronounced in areas with low prey density. We discuss to what extent this might affect their future fitness prospects, and suggest that such costs of maturing early in terms of body weight development might be high in terrestrial large carnivores due to a prolonged period of postnatal care.     

Fecundity and oviposition ofEucelatoria bryani, a gregarious parasitoid ofHelicoverpa zea andHeliothis virescens

We examined longevity, fecundity, and oviposition strategies ofEucelatoria bryani Sabrosky (Diptera: Tachinidae), a gregarious endoparasitoid ofHelicoverpa zea (Boddie) andHeliothis virescens (F.) (Lepidoptera: Noctuidae). Longevity of adult femaleE. bryani was not related to body size. In contrast to longevity, largerE. bryani females had greater potential fecundity than smaller females, as determined by the number of embryonated eggs present in the common oviduct. However, female parasitoid size did not affect primary clutch size (number of eggs deposited in a host). Because embryos in eggs located in the ovisac were larger than those located elsewhere in the common oviduct, maximum primary clutch size may be physiologically limited by the number of fully mature eggs a female has available at one time.E. bryani females adjusted primary clutch size in response to host size, for bothH. zea andH. virescens. This adjustment appears to be adaptive because females did not overexploit hosts by depositing more larvae than a host could support. Adult emergence was not related to host size. Although host weight positively influencedE. bryani progeny weight, increases in progeny size with host size were counterbalanced by increases in primary clutch size with host size.