The distribution and status of the adder (Vipera berus L) in Scotland determined from questionnaire surveys

Abstract. During 1992 two questionnaire surveys, aimed at farmers and sections of the general public, were conducted to determine the distribution and status of the adder ( Vipera berus L.) in Scotland. In total, 94% of the Scottish mainland and inner isles were covered by the two surveys. The adder appears to be widespread throughout Scotland, although it is absent from much of the Central Valley, the Outer Hebrides and Northern Isles, and from much of the mountainous region between Inverness and Glasgow.
The distribution of the adder in Scotland was compared with the distributions of different habitat types defined in the ITE Land Class survey. Adders were strongly associated with areas of heterogeneous land use and negatively associated with intensively arable areas and rugged mountainous areas. Strong evidence was found to suggest that perceived adder abundance had declined during the last 10 years. Although the evidence suggesting that the distribution of adders in Scotland had changed was less strong, the results of the Farm Survey did indicate that it had contracted. The perceived change in status of the adder in Scotland during the last 10 years was compared with the observed changes in land use over the same period.     

Phylogeography and Pleistocene refugia of the adder (Vipera berus) as inferred from mitochondrial DNA sequence data

In order to contribute to the debate about southern glacial refugia used by temperate species and more northern refugia used by boreal or cold-temperate species, we examined the phylogeography of a widespread snake species (Vipera berus) inhabiting Europe up to the Arctic Circle. The analysis of the mitochondrial DNA (mtDNA) sequence variation in 1043 bp of the cytochrome b gene and in 918 bp of the noncoding control region was performed with phylogenetic approaches. Our results suggest that both the duplicated control region and cytochrome b evolve at a similar rate in this species. Phylogenetic analysis showed that V. berus is divided into three major mitochondrial lineages, probably resulting from an Italian, a Balkan and a Northern (from France to Russia) refugial area in Eastern Europe, near the Carpathian Mountains. In addition, the Northern clade presents an important substructure, suggesting two sequential colonization events in Europe. First, the continent was colonized from the three main refugial areas mentioned above during the Lower-Mid Pleistocene. Second, recolonization of most of Europe most likely originated from several refugia located outside of the Mediterranean peninsulas (Carpathian region, east of the Carpathians, France and possibly Hungary) during the Mid-Late Pleistocene, while populations within the Italian and Balkan Peninsulas fluctuated only slightly in distribution range, with larger lowland populations during glacial times and with refugial mountain populations during interglacials, as in the present time. The phylogeographical structure revealed in our study suggests complex recolonization dynamics of the European continent by V. berus, characterized by latitudinal as well as altitudinal range shifts, driven by both climatic changes and competition with related species.     

Reproductive success, mortality and sexual size dimorphism in the adder, Vipera berus

From 1981 to 1986 an isolated adder population was studied in the extreme south of Sweden. During this period 48 adult males and 44 adult females were marked. Male adders did not grow as large as the females. Large males had a significantly higher annual mating success and were engaged in more combats than smaller males. The mean length of recaptured males was significantly lower than that of those not recaptured, indicating a higher motality of larger males. Females brood size was positively correlated with body size. In females there was no difference in mean length of recaptured vs not recaptured individuals. The adder is one of the few snake species with male combat where males are smaller than females. I suggest that this is due to stronger selective advantages for large body size in females than in males.     

Opposing fitness consequences of colour pattern in male and female snakes

Local populations of the adder, Vipera berus, are polymorphic for dorsal colour pattern, containing both melanistic (black) and zig-zag patterned individuals. Colour patterns in snakes influence crypsis and thermoregulatory capacity and therefore may be subjected to natural selection. To find an explanation for the maintenance of this polymorphism I examined temporal and spatial variation in morph frequency, and tested for differential selection among morphs using data from a six year capture-mark-recapture study. The data derive from six groups of islands in the Baltic Sea off the Swedish east coast, two mainland localities near the coast, and one inland locality. Morph frequency did not change over time within a population but varied among populations: melanistic individuals were not found at the inland locality, but comprised from 17 to 62% of the coastal and island populations. Adders frequently moved between islands within a group, but the tendency to disperse was independent of morph. These results suggest that the polymorphism is stable and maintained by a deterministic process. Scar frequency was twice as high among melanistic as among zig-zag snakes, and melanistic individuals were easier to capture, indicating that predation may be higher on the melanistic morph. Colour morphs did not differ in body size, but analysis of recapture data shows evidence for differential survival among morphs. Zig-zag males survived better than melanistic males, but the relative survival rates of morphs were reversed in females. This difference was consistent through time and may be due to sexual differences in behaviour, with melanism increasing predation intensity when associated with male but not with female behaviour. Opposing fitness consequences of colour pattern in the two sexes may help maintain colour polymorphism within populations of Vipera berus.     

Adaptive variation in head size in Vipera berus L. populations

To prove that predators are morphologically adapted to the size of their prey one has to demonstrate that the morphological variation in the trophic apparatus is related to the prey size distribution and that the variation in the trait has some effect on individual fitness. I have studied geographic variation in relative head length (RHL) of adders, Vipera berus , on the Swedish mainland and on groups of islands in the Baltic Sea, and the relationship between RHL and physical condition, a character related to fitness. I also examined the relationship between RHL and sex and colour morph. Relative head length of adders was smallest on the mainland and increased on the islands with increasing body size of the main prey, Microtus agrestis , suggesting stabilizing selection for head size within each population. There was no difference in RHL between sexes or colour morphs. However, physical condition was positively correlated with RHL, indicating directional selection for larger heads. The observed pattern is interpreted as an evolutionary response to the geographic variation in body size of the main prey species and the smaller number of alternative prey species available on islands.     

Determinants of reproductive success in female adders,Vipera berus

Summary Female lifetime reproductive success in a small population of individually-marked adders in southern Sweden was studied over a period of seven years. Reproductive characteristics varied little from year to year and were consistent through time in individual females. Most females mature at four years of age and reproduce every two years. The total number of offspring produced by a female depends on her adult body size (and thus, litter size) and longevity (and thus, number of litters per lifetime). Adult body size in females is influenced mainly by subadult growth rates. Offspring size depends on maternal body size and a tradeoff between offspring size and offspring number. Maternal age does not affect litter sizes and offspring sizes except through ontogenetic changes in maternal body size.Survival of females after parturition is low because of the high energy costs of reproduction, compounded by low feeding rates of gravid females because of their sedentary behaviour at this time. About one-half of females produce only a single litter during their lifetimes, although some females live to produce four or five litters. On a proximate basis, rates of energy accumulation for growth (in subadults) and reproduction (in adults) may be the most important determinants of fitness in female adders.     

Reproductive success and helper effects in the cooperatively breeding grey-crowned babbler

Cooperative breeding, where some individuals help to raise offspring that are not their own, is a relatively rare social system in birds. We studied the breeding biology of a declining cooperative breeder, the grey-crowned babbler Pomatostomus temporalis , with the aim of isolating the social factors that affect its reproductive success. Most breeding pairs were assisted by philopatric offspring, although pairs could breed successfully without helpers. Females laid up to four clutches (usually three eggs per clutch) per season. Male (but not female) helpers increased the number of young fledged from individual nests and the likelihood of re-nesting, resulting in higher seasonal fledgling production. Helper effects on brood size and fledgling production were greater in the second year of the study, which was also characterized by higher nest failure. This suggests that helpers enhance reproduction more in poor conditions. Our study demonstrates the interacting effects of social and ecological factors on reproductive success, and that retention of offspring is not always beneficial for the breeders in cooperative species.     

Reproductive success of Macrotermes bellicosus (Isoptera, Macrotermitinae) in two neighbouring habitats

The fungus-cultivating, mound-building termite Macrotermes bellicosus can reach high densities in African savannah habitats, whereas in forests it is comparatively rare and is only found in relatively open areas. Earlier studies in the Comoé National Park (Côte d'Ivoire, West Africa) suggested that this might be because ambient temperatures are lower in the forest than the savannah. Therefore, forests seem to be suboptimal habitats. During 3 consecutive years we measured relevant fitness parameters in both habitats to test this hypothesis. Colonies in the savannah had higher reproductive outputs. They reproduced annually, producing high numbers of offspring (alates), whereas forest colonies reproduced on average only once every 3?years and even then only low numbers of alates were produced. Annual growth of mound height, which can be regarded as an indicator of colony growth, varied considerably between individual colonies and years. Nevertheless, in two years the growth rate of mounds in the shrub savannah was higher than that of mounds in the gallery forest. Unexpectedly, survival probability as calculated by Kaplan Meier analysis was much lower for colonies in the shrub savannah than for colonies in the gallery forest. Using these data in a computer simulation we calculated lifetime reproductive success (LRS) for an average colony in both habitats. As LRS was much higher for colonies in the shrub savannah than in the gallery forest, these results confirmed our suggestion that the forest is a suboptimal habitat for M. bellicosus.     

Reproductive investment under fluctuating predation risk: Microtine rodents and small mustelids

Summary We studied the reproductive investment of microtine rodents (bank vole (Clethrionomys glareolus),Microtus epiroticus andMicrotus agrestis) in western Finland under predation risk from small mustelids. During 1984–1992, the yearly mean litter size of overwintered bank voles was smaller at high least weasel and stoat densities than at low densities (close to 3 versus 4–5). In addition, the annual mean litter size of young bank voles was negatively correlated to the least weasel density. In youngM. agrestis voles, the yearly late summer litter size was negatively associated with the autumn density of small mustelids. In the crash phase of the vole cycle (1989 and 1992), we removed small mustelids (mainly least weasels) from four unfenced areas in late April to late May and studied the reproduction of voles in four removal and comparable control areas (each 2–4 km²). Reduction of small mustelids significantly increased the proportion of pregnant bank vole females, but not that of pregnantMicrotus vole females. We conclude that predation risk apparently reduced reproductive investment of free-living bank vole females; these voles appear to trade their current parental investment against future survival and reproductive prospects. Accordingly, the presence of small mustelids (or their scent) may slow down the reproductive rate of voles. As antipredatory behaviours occurred on a large scale, our results add evidence to the hypothesis that crashes in multiannual vole cycles are driven by small mustelid predators.     

Reproductive life history variation among colour morphs of the pygmy grasshopper Tetrix subulata

Individuals of pygmy grasshoppers ( Tetrix subulata [L.] Orthoptera: Tetrigidae) exhibit genetically coded discontinuous variation in colour pattern. To determine whether reproductive performance is likely to be affected by colour pattern, this study investigated variation in body size and reproductive life-history characteristics among individuals belonging to five different colour morphs. The proportion of reproductive females (i.e. females with eggs) declined significantly as the season progressed (from 100% in mid-May to 40% in mid-June), but no such seasonal trend was apparent for body size, clutch size or egg size. Colour morphs differed significantly in body size, and these size differences accounted for most of the variation in clutch size and egg size. Colour morphs also differed in the regression of egg size on clutch size, suggesting that trade-offs between number and size of offspring might vary among morphs. Finally, I found a negative relationship across colour morphs between the proportion of females with eggs and average clutch size. This suggests that individuals belonging to certain colour morphs produce a relatively large number of clutches per unit time, at the expense of fewer offspring in each clutch, compared to other morphs. Collectively, my results indicate that different colour morphs of T. subulata may have different reproductive strategies. These differences may reflect variation in thermoregulatory capacity or differences in probability of survival induced by visual predators.     

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.     

Factors Affecting Reproduction and Mortality Among Baboons in the Okavango Delta,Botswana

We present results of a 10-year study of free-ranging gray-footed chacma baboons (Papio ursinus griseipes) in the Okavango Delta of Botswana. The majority of deaths among adult females and juveniles were due to predation, while infants were more likely to die of infanticide. There were strong seasonal effects on birth and mortality, with the majority of conceptions occurring during the period of highest rainfall. Mortality due to predation and infanticide was highest during the 3-mo period when flooding was at its peak, when the group was more scattered and constrained to move along predictable routes. The reproductive parameters most likely to be associated with superior competitive ability—interbirth interval and infant growth rates—conferred a slight fitness advantage on high-ranking females. However, it was counterbalanced by the effects of infanticide and predation. Infanticide affected high- and low-ranking females more than middle-ranking females, while predation affected females of all ranks relatively equally. As a result, there were few rank-related differences in estimated female lifetime reproductive success.     

Corolla herbivory, pollination success and fruit predation in complex flowers: an experimental study with Linaria lilacina (Scrophulariaceae)

BACKGROUND AND AIMS: Herbivory on floral structures has been postulated to influence the evolution of floral traits in some plant species, and may also be an important factor influencing the occurrence and outcome of subsequent biotic interactions related to floral display. In particular, corolla herbivory may affect structures differentially involved in flower selection by pollinators and fruit predators (specifically, those ovopositing in ovaries prior to fruit development); hence floral herbivores may influence the relationships between these mutualistic and antagonistic agents. METHODS: The effects of corolla herbivory in Linaria lilacina (Scrophulariaceae), a plant species with complex flowers, were considered in relation to plant interactions with pollinators and fruit predators. Tests were made as to whether experimentally created differences in flower structure (resembling those occurring naturally) may translate into differences in reproductive output in terms of fruit or seed production. KEY RESULTS: Flowers with modified corollas, particularly those with lower lips removed, were less likely to be selected by pollinators than control flowers, and were less likely to be successfully visited and pollinated. As a consequence, fruit production was also less likely in these modified flowers. However, none of the experimental treatments affected the likelihood of visitation by fruit predators. CONCLUSIONS: Since floral herbivory may affect pollinator visitation rates and reduce seed production, differences among plants in the proportion of flowers affected by herbivory and in the intensity of the damage inflicted on affected flowers may result in different opportunities for reproduction for plants in different seasons.     

Male mating success, reproductive success and multiple paternity in a natural population of sand lizards: behavioural and molecular genetics data

Sand lizard Lacerta agilis females characteristically mate with several males which, in staged mating experiments, results in multiple paternity of the offspring. In order to investigate multiple paternity in a natural population and interpret male reproductive behaviours in terms of sired young, we sampled the blood of females, potential fathers and hatchlings, and determined paternity using multilocus DNA fingerprinting as well as the variation at a single locus detected by the probe (TC) _n . The paternity analyses were preceded by a laboratory experiment in which we established that the parental alleles identified by the single locus probe were inherited in a Mendelian way. Our molecular data demonstrated that 12 out of 13 males (92%) that sired offspring were correctly identified from the 56 sexually mature males in the population. Also smaller males were accepted as sexual partners by the females, but sired fewer young in competition with larger males and were less able to maintain prolonged post-copulatory mate guarding. This may result in that some sexually successful males are only observed inside a female's home range, but never in pair-association with the female.     

Vertebral number in adders, Vipera berus: direct and indirect effects on growth

Given the importance of body size, and thereby growth rate, for many reproductive parameters in snakes, morphological traits conferring an advantage in terms of growth may be important targets of selection. Studies have demonstrated effects of vertebral number of growth rate in garter snakes. In this study effects of total number of body vertebrae and of number of abnormal body vertebrae (obtained by counting number of ventral scutes and number of abnormal scutes) on growth rate in free-ranging male and female adders, Vipera berus (L.), are examined by calculating directional performance gradients (estimating linear effects) and stabilizing performance gradients (estimating curvilinear effects). After controlling for body size (SVL) female adders demonstrated a significant positive directional gradient for vertebral number, and a significant interaction between body size and vertebral number, showing that females with more vertebrae have higher size-specific growth rates, and that this effect is strongest among small, fast growing individuals. Females also showed a weak stabilizing effect of abnormal vertebrae. Males, on the other hand, showed a positive directional gradient for number of abnormal vertebrae, whereas no effect of vertebral number was observed. Indirect effects of the same variables were evaluated by use of path analysis. Generally, indirect effects were weak and did not substantially increase the amount of explained variance in growth rate. Field data showed that the correlation between vertebral number and growth rate in females was stronger in years with higher overall growth rate. To evaluate whether vertebral number and food availability show an interactive effect I used captive born juvenile adders in an experiment with two different food levels. The experiment confirmed the field data. No relationship between vertebral number and growth was observed in the low food level group, whereas in the high food level group a significant positive correlation was demonstrated. Finally, the heritability of vertebral number was examined using a mother-offspring regression and a full-sib analysis. The estimated heritabilities were 0.30 and 0.39, respectively. From these results it is concluded that both vertebral number and abnormal vertebral number may significantly affect growth in adders, but that this effect may differ between sexes and among years.     

Variation in number of ventral scales in snakes: effects on body size, growth rate and survival in the adder, Vipera berus

The relationship between number of ventral scales, correlating with number of body vertebrae, and body size of adders, Vipera berus , was investigated using captive-born young and wild-caught adults (snout-vent length > 400 mm) from six populations in eastern Sweden. Females had significantly more ventral scales, and were larger, than males in all populations. Among adult individuals, snout-vent length was highly positively correlated with the number of ventral scales when differences due to sex and locality were controlled for. The same pattern was true for newborn individuals when differences due to litter and sex were controlled for. The influence of number of ventral scales on survival and growth rate as possible causes of this positive correlation was examined. Mean number of ventral scales was lower in new-born snakes than in adults, indicating selection against individuals with a low scale count. Since this selection appears to take place in the early juvenile phase, it is inadequate to explain the relationship between adult body size and number of ventral scales. However, individuals with many ventral scales had significantly higher growth rates than individuals with few ventrals. This suggests that individuals with many ventral scales enjoy a higher growth rate and therefore are able to reach a larger size than their conspecifics of the same age with few ventral scales. This would explain the positive relationship between body size and ventral scale number observed in new-born and adult snakes.     

Reproductive success of cat fleas,Ctenocephalides felis,on calves as unusual hosts

Abstract. Cat fleas, Ctenocephalides felis, were released onto calves as unusual hosts, and sampled at intervals for histological examination. Egg output from fleas on age-matched male and female calves was monitored. Using indicators of reproductive maturation and regression together with egg output data, the reproductive success and fertility of cat fleas on male and female calves were estimated. Comparisons were made with fleas taken from cats. The mean egg output of fleas on the bull calf was highly significantly different from that on the age-matched female calf: 28.14 ± 2.96 (SE) eggs/h compared with 16.21 ± 1.96 (SE) eggs/h. A higher proportion of sampled fleas (83.0%) was reproductively mature on the feline hosts compared with the calves (45.4-62.5%). Blue bodies resulting from oocyte resorption were present in the ovarioles of 10.4-19.0% of fleas sampled from the calves. No blue bodies were present in fleas removed from cats. Eggs laid by fleas on calves were viable and larvae were reared to adulthood. The mean percentage hatching success on the age-matched male and female calves was 46.7% and 51.7%. This represents a reduction in viability of 28-33% compared with eggs laid by fleas on cats. Factors which may account for reduced reproductive maturation of fleas on calves, including protein content of the host's blood, are discussed.     

Group size,composition, and reproductive success in wild common marmosets (Callithrix jacchus)

Data from published sources about size and composition of wild common marmoset groups (Callithrix jacchus) were analyzed to see if the number of juveniles in a group is closely related to the number of other group members. Mean group size was 8.7 members including 4.4 adults (1.8 females, 2.5 males), 2.9 subadults, and 1.4 juveniles. The number of juveniles was significantly positively correlated to the number of adult males. Groups with one or two adult males had significantly fewer juveniles (mean: 1:1 juveniles) than groups containing more than two adult males (mean: 2.0 juveniles). Apart from a different number of subadults, results showed obvious similarities between common marmosets and tamarins of the genus Saguinus in size and composition of subgroups of adults as well as the key role of adult males in mediating the reproductive success of a breeding female. Common marmoset females seem to gain direct fitness benefits in increased reproductive success from the presence of a larger number of adult males. Whether or not other group members get fitness benefits depends on the reproductive strategy of adult males (monogamy vs. polyandry), their kinship, and on the genetic relationship of nonbreeders to the offspring of the breeding female. © 1995 Wiley-Liss, Inc.