Long-term population dynamics in a Mediterranean aquatic snake

A review of several long-term studies has recently suggested that snakes might be declining in large parts of the world. Additional data from other long-term studies are therefore urgently needed in order to assess the generalities of such suggested declines. Based on a 20-year study, we analyzed demographic data on adult dice snakes (Natrix tessellata) studied in central Italy between 1985 and 2004. Both male and female dice snakes were relatively long-lived, with no significant differences in longevity between the sexes. Individual males and females were observed over a maximum of 10 and 14 years, respectively. However, the among-year recapture rates between the year the snakes were initially captured and the subsequent year (i.e., year 1 to year 2) was significantly lower (45%) than the among-year recapture rates during subsequent years (74%; i.e., year 2 to year 3), suggesting that a large proportion of the snakes at first capture were in fact not resident within our study area, and hence many snakes were migrating in and out of our 2-km stream study site, with no inter-sexual difference in dispersal rates. Sex ratio was virtually equal if we consider the study period as a whole. Significant annual fluctuations were, however, observed through the study. In 1985–1990, 1993–1995, 1998 and 1999 the sex ratio was male-biased, whereas in 2000–2004 female-biased. In terms of both survival and recapture probabilities, model selection showed that Akaike’s information criterion favored the model incorporating body size, with the model incorporating year having an intermediate likelihood, and the model with sex included being the most disfavored. Total population number estimates suggest an average 86 adult individuals along the 2 km of stream with only minor annual variations. However, a significant decrease in the number of males occurred during the last 6 years of our study. Thus, further monitoring of this population is warranted in light of the decline of snake populations reported recently.     

Chick parasitism by blowflies affects feeding rates in a Mediterranean population of blue tits

Offspring fitness depends on interactions between parental care and environmental constraints. It has been suggested that in altricial birds parents are able to compensate for the detrimental effects of ectoparasites by improving food provisioning. We tested this prediction in a population of blue tits highly parasitized by blowfly larvae. The frequency of parental feeding visits was significantly higher in parasitized broods than in broods experimentally deparasitized. Despite a strong increase in parental care, chicks of parasitized broods were lighter, smaller, and more anaemic than chicks in deparasitized broods. Parents invest more in feeding parasitized young but cannot fully compensate for the negative effects of parasites, hence young are in poor condition at fledging.     

Long-term stability of allozyme frequencies in a wood lemming, Myopus schisticolor, population with a biased sex ratio and density fluctuations

Wood lemming (Myopus schisticolor) populations are characterized by female biased sex ratios and cyclic variations in population size. Both of these characteristics are assumed to reduce genetic variation and thus affect the evolutionary adaptation of the species. We addressed these questions by studying the genetic structure of a wood lemming population from eastern Finland by isozyme markers during a 21-year period, which corresponds to 40-50 generations. Contingency tests showed that genotypic proportions conformed to Hardy-Weinberg equilibrium in each of the four sampling years. Among the temporal replicates, allele frequencies differed most by 0.14 and were not significant. Genetic variation was also stable and fairly high with a mean observed heterozygosity of H = 0.057. Variability in the Heinavesi population was higher than previously reported in wood lemming. The difference was mainly caused by variation at a phosphoglucomutase locus that was monomorphic in earlier studies. Significant linkage disequilibrium was observed in three of the comparisons but the disequilibrium did not appear consistently in all years. This pattern was also evidenced by the variance components, which indicated that selection favoured for specific allele pairs only in few subsamples.     

Estimating genetic drift and effective population size from temporal shifts in dominant gene marker frequencies

Measurement of temporal change in allele frequencies represents an indirect method for estimating the genetically effective size of populations. When allele frequencies are estimated for gene markers that display dominant gene expression, such as, e.g. random amplified polymorphic DNA (RAPD) and amplified fragment length polymorphism (AFLP) markers, the estimates can be seriously biased. We quantify bias for previous allele frequency estimators and present a new expression that is generally less biased and provides a more precise assessment of temporal allele frequency change. We further develop an estimator for effective population size that is appropriate when dealing with dominant gene markers. Comparison with estimates based on codominantly expressed genes, such as allozymes or microsatellites, indicates that about twice as many loci or sampled individuals are required when using dominant markers to achieve the same precision.     

Gastrointestinal responses to feeding in a frequently feeding colubrid snake (Natrix maura)

Ectotherm vertebrates show physiological mechanisms that reduce metabolic costs during prolonged fasting. Once feeding, these animals adopt a wide variety of metabolic responses such as changes in gastrointestinal organ masses. Up-regulatory responses after feeding have been widely explored in infrequently feeding snakes like pythons, whereas few studies have been devoted to frequently feeding snakes. In this study, we have considered the gastrointestinal responses after feeding in a frequent feeder, the viperine snake Natrix maura, in the Ebro Delta rice fields. In this habitat, viperine snakes are exposed to long periods of food deprivation due to the lack of available prey as a consequence of the man-induced rice cycle. We weighed prey items and full gut masses, and measured length of combined esophagus and stomach, and intestine of viperine snakes belonging to a wide range of sizes. Snakes concentrate foraging activity when rice fields were flooded. In this period, gut masses increased. Likewise, intestines increased in length during the feeding period, which suggests that viperine snakes probably experience a postfeeding hypertrophy of their small intestines that contributes to their larger length. Once the intestine length was corrected for the snake size, it was shown that adults present longer intestines than immature snakes, reflecting an increase in the posterior part of the body linked to the gonads development. This study contributes to explore the physiological responses to feeding in frequently feeding snakes modelled by abrupt shifts of food availability.     

Gm and Km frequencies in a Portugese population

Summary Serum samples from 79 healthy subjects and 31 patients with type I amyloid neuropathy were typed for nine Gm markers. No significant associations were found between any of the Gm phenotypes and the disease. The frequencies of the various phenotypes showed satisfactory agreement with the Hardy-Weinberg distribution. One individual had an unusual Gm phenotype.Publication no. 526 from the Department of Basic and Clinical Immunology and Microbiology, Medical University of South Carolina. Research supported in part by USPHS Grants CA-25746 and AI-07063, and by South Carolina State Appropriations for Research     

Climate-change-related shifts in annual phenology of a temperate snake during the last 20 years

Global warming is thought to be a far-reaching threat to biodiversity, and is supposed to influence several aspects of the ecology of animals. Global warming should influence especially the ectotherm vertebrates, which depend directly from the external thermal conditions for their activities and performances. Here, we analyze the changes in phenology which have occurred in the last 20 years in a marked population of vipers, Vipera aspis, and we try to relate these changes with the intervening climatic changes. We analyzed three metrics of viper's annual phenology: (i) annual onset of above-ground activity (hereby AOA); (ii) annual onset of feeding period (AOF); (iii) annual onset of the hibernation (AOH). The annual variations of these three phenological metrics were correlated to five variables of climatic data: (1) mean annual air temperature, (2) mean February air temperature, (3) mean July air temperature, (4) yearly number of rainy days, and (5) yearly number of days with rainstorm. We observed a statistically significant reduction of AOA values from >28 days between 1987 and 1997, to approximately 20 days from 1998 to 2011, with a similar statistical trend also found for AOF values. The number of days of delay in entering hibernation increased significantly since 1998. Three sets of relationships between climatic variables and metrics of viper phenology were statistically significant, i.e. the correlation (i) between annual mean temperature and AOA (negative), (ii) between annual mean temperature and AOF (negative), and (iii) between annual mean temperature and AOH (positive). The percent of field days (between 20th February and 20th March) with no viper observed also decreased significantly over the years. Our study showed that three different traits of the annual phenology of a Mediterranean snake are shifting in the 20+ years of monitoring, and that there is correlational evidence that these shifts are linked to intervening climate change.     

Long-term shifts in the cyclicity of outbreaks of a forest-defoliating insect

Optimisation of reproductive investment is crucial for Darwinian fitness, and detailed long-term studies are especially suited to unravel reproductive allocation strategies. Allocation strategies depend on the timing of resource acquisition, the timing of resource allocation, and trade-offs between different life-history traits. A distinction can be made between capital breeders that fuel reproduction with stored resources and income breeders that use recently acquired resources. In capital breeders, but not in income breeders, energy allocation may be decoupled from energy acquisition. Here, we tested the influence of extrinsic (weather conditions) and intrinsic (female characteristics) factors during energy storage, vitellogenesis and early gestation on reproductive investment, including litter mass, litter size, offspring mass and the litter size and offspring mass trade-off. We used data from a long-term study of the viviparous lizard, Lacerta (Zootoca) vivipara. In terms of extrinsic factors, rainfall during vitellogenesis was positively correlated with litter size and mass, but temperature did not affect reproductive investment. With respect to intrinsic factors, litter size and mass were positively correlated with current body size and postpartum body condition of the previous year, but negatively with parturition date of the previous year. Offspring mass was negatively correlated with litter size, and the strength of this trade-off decreased with the degree of individual variation in resource acquisition, which confirms theoretical predictions. The combined effects of past intrinsic factors and current weather conditions suggest that common lizards combine both recently acquired and stored resources to fuel reproduction. The effect of past energy store points out a trade-off between current and future reproduction.     

Long-term progression of inbreeding depression in a Mediterranean ornithophilous shrub

The ornithophilous species Anagyris foetida L. is a Mediterranean shrub with highly fragmented populations and a mixed mating system. In a previous study, we analyzed the first 3 years of the life cycle of two progenies (selfed and outcrossed) grown from seed obtained by hand pollination and planted in an experimental garden in 2005. In that study, we found that inbreeding depression (ID) was manifested both reproductively and vegetatively throughout the life cycle, with male reproductive function being the most affected trait. In the present study, our main aim was to check the progression of the two progenies 12 years after transplantation. For this we analyzed their survival and their vegetative and reproductive traits. According to our results, levels of ID were similar to those obtained in the previous study, with some factors decreased and thus varying with the age of the studied individuals. Vegetative parameters were found to have a greater influence than reproductive ones (δ = 0.56 vs. δ = 0.36) on overall ID. As indicated by the global ID (δ = 0.72) the populations have a mating system that is intermediate between outcrossing and a mixed system. Furthermore, the lower male reproductive capacity of the selfed individuals has been maintained over time. Our study also demonstrates the importance of studying the ID value throughout the life cycle of plants.     

Respiratory consequences of feeding in the snake Python molorus

Snakes can ingest large meals and exhibit marked increases in metabolic rate during digestion. Because postprandial oxygen consumption in some snakes may surpass that attained during exercise, studies of digestion offers an alternative avenue to understand the cardio-respiratory responses to elevated metabolic rate in reptiles. The effects of feeding on metabolic rate, arterial oxygen levels, and arterial acid-base status in the snake Python molorus are described. Four snakes (180-250 g) were cannulated in the dorsal aorta and blood samples were obtained during 72 h following ingestion of a meal (rat pups) exceeding 20% of body weight. Oxygen consumption increased from a fasting value of 1.71 +/- 0.08 to 5.54 +/- 0.42 ml kg-1 min-1 at 48 h following feeding, and the respiratory gas exchange ratio increased from 0.67 +/- 0.02 to a maximum of 0.92 +/- 0.03 at 32 h. Plasma lactate was always less than 0.5 mM, so the postprandial increase in metabolic rate was met by aerobic respiration. In fasting animals, arterial PO2 was 66 +/- 4 mmHg and haemoglobin-O2 saturation was 92 +/- 3%; similar values were recorded during digestion, but haematocrit decreased from 15.8 +/- 1.0 to 9.8 +/- 0.8 due to repeated blood sampling. Plasma [HCO3-] increased from a fasting level of 19.3 +/- 0.8 to 25.8 +/- 1.0 mmol l-1 at 24 h after feeding. However, because arterial PCO2 increased from 21.1 +/- 0.5 to 27.9 +/- 1.4 mmHg, there was no significant change in arterial pH from the fasting value of 7.52 +/- 0.01. Acid-base status returned to pre-feeding levels at 72 h following feeding. The increased arterial PCO2 is most likely explained by a reduction in ventilation relative to metabolism, but we predict that lung PO2 does not decrease below 115 mmHg. Although ingestion of large meals is associated with large metabolic changes in pythons, the attendant changes in blood gases are relatively small. In particular, the small changes in plasma [HCO3-] and stable pH show that pythons respond very differently to digestion than alligators where very large alkaline tides have been observed. It is unclear why pythons and alligators differ in the magnitude of their responses, but given these interspecific differences it seems worthwhile to describe arterial blood gases during digestion in other species of ectothermic vertebrates.     

Pathogen interactions, population cycles, and phase shifts

Interspecific pathogen interactions can profoundly affect pathogen population dynamics and the efficacy of control strategies. However, many pathogens exhibit cyclic abundance patterns (e.g., seasonality), and temporal asynchrony between interacting pathogens could reduce the impact of those interactions. Here we use an extension of our previously published model to investigate the effects of cycles on pathogen interaction. We demonstrate that host immune memory can maintain the impact of an interaction, even when the effector pathogen abundance is low or the pathogen is absent. Paradoxically, immune memory can result in pathogens interacting more strongly when temporally out of phase. We find that interactions between species can result in changes to the temporal pattern of the affected species. We further demonstrate that this may be observed in a natural host-pathogen system. Given the continuing debate regarding the relevance of pathogen interactions in natural systems and increasing concern about treatment strategies for coinfections, both the discovery of a shift in cycle in empirical data and the mechanism by which we identified it are important. Finally, because the model structure used here is analogous to models of a simple predator-prey system, we also consider the consequences of these findings in the context of that system.     

Allele frequencies in a domestic cat population in a country area of northern Moravia,Czechoslovakia

The frequencies of alleles responsible for colour of cat fur were studied by method of house-to-house survey in a population of domestic cats in northern Moravia, Czechoslovakia.Of the total of 249 adult cats examined, 114 animals were males and 135 females, the sex ratio being 0.458. The following allelic frequencies were recorded: 0=0.016, a=0.646, t^b=0.288 and S=0.237. No other alleles were recorded in the phenotype. The results are compared with other populations in Czechoslovakia and other European countries.     

Rain,prey and predators: climatically driven shifts in frog abundance modify reproductive allometry in a tropical snake

To predict the impacts of climate change on animal populations, we need long-term data sets on the effects of annual climatic variation on the demographic traits (growth, survival, reproductive output) that determine population viability. One frequent complication is that fecundity also depends upon maternal body size, a trait that often spans a wide range within a single population. During an eight-year field study, we measured annual variation in weather conditions, frog abundance and snake reproduction on a floodplain in the Australian wet-dry tropics. Frog numbers varied considerably from year to year, and were highest in years with hotter wetter conditions during the monsoonal season (“wet season”). Mean maternal body sizes, egg sizes and post-partum maternal body conditions of frog-eating snakes (keelback, Tropidonophis mairii, Colubridae) showed no significant annual variation over this period, but mean clutch sizes were higher in years with higher prey abundance. Larger females were more sensitive to frog abundance in this respect than were smaller conspecifics, so that the rate at which fecundity increased with body size varied among years, and was highest when prey availability was greatest. Thus, the link between female body size and reproductive output varied among years, with climatic factors modifying the relative reproductive rates of larger (older) versus smaller (younger) animals within the keelback population.     

Types and frequencies of Q-variant chromosomes in a Japanese population

Summary The types of Q-variant bands were determined by a combination of numerical designations setting five levels for both the size of bands and the intensity of fluorescence. This scoring system was used in a study of the frequencies of Q variants in 400 Japanese individuals: variant bands were observed in seven specific autosome pairs of Nos. 3,4,13,14,15,21, and 22. The number of variants per individual ranged from 0 to 8, and the mean was 3.83±1.86. The incidence of Q variants according to the types of variant bands was determined in specific chromosomes.A low frequency of No. 3 chromosome variants and a high frequency of a long Y in males seems to be characteristic for Japanese populations.Variation in the length of the long arm of Y (Yq) was analyzed in a total of 157 men. The relative length of Yq, which was determined by a ratio of Yq/21q, ranged from 0.98 to 2.27, with an average of 1.56±0.25. The length of pale band Yq11 was relatively constant between individuals, with an average of 0.64±0.08. Therefore, it was clear that the variation in the Yq length was the result mainly of a variation in the length of the brilliant band Yq12. However, a slight tendency for the length of band Yq11 to increase in proportionally to the total length of the Yq was revealed. In this study special consideration was paid to the reliable analysis of Q-band heteromorphism, and the factors or obstacles preventing such analysis have been discussed briefly.     

Maximum-likelihood estimation of molecular haplotype frequencies in a diploid population

Molecular techniques allow the survey of a large number of linked polymorphic loci in random samples from diploid populations. However, the gametic phase of haplotypes is usually unknown when diploid individuals are heterozygous at more than one locus. To overcome this difficulty, we implement an expectation-maximization (EM) algorithm leading to maximum-likelihood estimates of molecular haplotype frequencies under the assumption of Hardy-Weinberg proportions. The performance of the algorithm is evaluated for simulated data representing both DNA sequences and highly polymorphic loci with different levels of recombination. As expected, the EM algorithm is found to perform best for large samples, regardless of recombination rates among loci. To ensure finding the global maximum likelihood estimate, the EM algorithm should be started from several initial conditions. The present approach appears to be useful for the analysis of nuclear DNA sequences or highly variable loci. Although the algorithm, in principle, can accommodate an arbitrary number of loci, there are practical limitations because the computing time grows exponentially with the number of polymorphic loci. Although the algorithm, in principle, can accommodate an arbitrary number of loci, there are practical limitations because the computing time grows exponentially with the number of polymorphic loci.