Specific dynamic action and postprandial thermophily in juvenile northern water snakes,Nerodia sipedon

1. We measured oxygen consumption of seven juvenile Northern water snakes, Nerodia sipedon, at 3, 19, 41, and 89 h after feeding. 2. Oxygen consumption 19 h after eating was significantly higher than at 3, 41, and 89 h after feeding and the values at 3 h post-feeding were significantly higher than those at 41 and 89 h. 3. We monitored substrate temperature preference in nine juvenile N. sipedon at 2000, 0000, 0800, 1200, and 1600 h for 3 days. 4. After feeding and at night snakes selected warmer substrate temperatures.     

Genetic analysis of the mating system and opportunity for sexual selection in northern water snakes (Nerodia sipedon)

We used data collected over 3 years at two study sites to quantifythe rates and consequences of multiple paternity and to determinethe opportunity for selection on male and female northern watersnakes (Nerodia sipedon). We sampled litters from 45 femalesthat gave birth to 811 offspring. Using eight microsatelliteDNA loci (probability of exclusion of nonparental males >0.99), we assigned paternity to 93% of neonates from one studypopulation and 69% of neonates from the other population. Observationsof participation in mating aggregations predicted individual reproductive success poorly for two reasons. First, males regularlycourted nonreproductive females. Second, more than half ofall sexually mature males obtained no reproductive successeach year, despite the fact that many of them participatedin mating aggregations. The number of sires per litter ranged from one to five, with 58% of all litters sired by more thanone male. Multiple paternity increased with female size, apparentlyboth because bigger females mated with more males and becausethe larger litters of big females provide paternity opportunitiesto more males. Multiple paternity was also more prevalent inyears with shorter mating seasons. We detected no advantage to multiple paternity in reducing either the number of unfertilizedovules or stillborn young. Despite the majority of males siringno young each year, some males fathered young with as manyas three different females in one year. Male reproductive successincreased by more than 10 offspring for each additional mate,whereas female success increased by fewer than 2 offspring foreach additional mate. The opportunity for sexual selectionwas more than five times higher in males than females.     

A test of the thermal coadaptation hypothesis with black rat snakes (Elaphe obsoleta) and northern water snakes (Nerodia sipedon)

• 1.The thermal coadaptation hypothesis predicts that (1) ectotherms experiencing a narrow range of body temperatures in the wild will evolve to perform well over a narrow range of body temperatures and that (2) the optimal temperature for performance will be equal to the preferred body temperature of the species.
• 2.We tested the predictions of the thermal coadaptation hypothesis with black rat snakes (Elaphe obsoleta) and northern water snakes (Nerodia sipedon) because black rat snakes experience lower and more variable body temperatures than northern water snakes at our study site.
• 3.We measured swimming speed, tongue-flicking speed, and striking speed in black rat snakes, and swimming speed and tongue-flicking speed in northern water snakes.
• 4.Adult water snakes generally had narrower performance breadths and higher optimum performance temperatures than adult black rat snakes.
• 5.Performance breadths were the same for swimming, tongue flicking, and striking within adult black rat snakes, but performance optima for these behaviours differed significantly. Performance breadths differed and performance optima were the same for swimming and tongue flicking within adult northern water snakes.
• 6.The relative swimming performance of neonates of the two species was similar in breadth to that of adults, but the thermal optimum for neonate black rat snakes was higher than that of adults.
• 7.Overall, our results provided support for the thermal coadaptation hypothesis.

     

Microgeographic population genetic structure in the northern water snake, Nerodia sipedon sipedon detected using microsatellite DNA loci

We describe the isolation and genetic characterization of eight microsatellite DNA loci from the northern water snake, Nerodia sipedon sipedon and use these loci to analyse levels of genetic differentiation between local (< 2 km apart) populations of these snakes in Ontario. These loci are variable, with expected heterozygosities ranging from 0.28 to 0.91, and can correctly exclude nonsires in parentage analyses with a high probability (0.998). Population analyses reveal significant deviation from expected heterozygosity levels for one population, probably a result of a null allele(s) at a single locus and small but significant levels of genetic differentiation among all three populations. This demonstrates that microgeographic genetic structure exists in this species, possibly due to limited dispersal.     

Histological aspects of natural eustrongyloid infections of the northern water snake, Nerodia sipedon

Encapsulated as well as unencapsulated fourth stage larvae of Eustrongylides sp. were found in four northern water snakes, Nerodia sipedon, from Virginia. Unencapsulated nematodes occurred within muscle, subcutaneous tissue and in the coelom. Encapsulated nematodes occurred in three forms: recently encapsulated, degenerating, or sclerotic; they were found on the mesentery, in muscle and subcutaneous tissue. Encapsulation was associated with muscular hypotrophy, fibroplasia and eosinophilia.     

Lack of postprandial thermophily in diamondback water snakes,Nerodia rhombifera

We compared the preferred body temperature (T_b) of fasted and satiated diamondback water snakes (Nerodia rhombifera) in linear thigmothermal gradients. We found no significant postprandial thermophily. Thermoregulatory precision varied among 6 days of testing and was most precise during the first day (24–48 hr after feeding).     

Dynamic changes in body form during swimming in the water snake Nerodia sipedon

Video records of swimming water snakes show that during moderate to rapid swimming, the rear half to two-thirds of the trunk is compressed laterally, approaching the body form of some sea snakes. Body form of swimming snakes differed significantly from their shape when resting on a flat surface or when anesthetized and suspended in water. The extent of lateral flattening is positively correlated with swimming speed, a relationship generally supported by tests of trunk models in a flow tank. In Nerodia, the ability to temporarily flatten the trunk depends on kinetic costovertebral joints, a large compressible body cavity, and the absence of ventral skeletal support - features found in most snakes. Histological studies and manipulations of partially dissected preserved specimens showed that the resting angle of the ribs is maintained by localized elastic hypertrophy of the costovertebral capsular ligament. Trunk form during swimming in Nerodia is proposed to arise from anteromedial movement of the distal rib powered by deep muscles acting in concert with those proposed to generate undulation of the vertebral column.     

Variation in seasonal ultrastructure of sexual granules in the renal sexual segment of the Northern Water Snake, Nerodia sipedon sipedon

The renal sexual segment of the kidney (RSS) can be found in many male squamate reptiles, encompassing the distal region of the nephron and, in some cases, collecting ducts. This sexually dimorphic structure exhibits varying degrees of hypertrophy and regression throughout the year. Although researchers have been aware of and have investigated this unique structure for over a century, its ultimate function remains under discussion. In many studies hypertrophy and regression of the RSS have been correlated with testicular activity and androgen secretion. As in most of the snakes studied to date, the male Northern Water Snake (Nerodia s. sipedon) does not exhibit a dramatic cycle of hypertrophy and regression, as reported in lizards. Following the initial hypertrophy at maturity, the male Northern Water Snake maintains a level of RSS hypertrophy throughout the year. Variations in the appearance and makeup of the sexual granules provide an identifiable and quantifiable seasonal pattern that can be correlated with the concentration of plasma androgens. In the Northern Water Snake, plasma androgens are elevated upon emergence and the RSS epithelial cells are filled with solid granules. As androgen levels decline during spring, sexual granule content appears to be breaking down (utilized?), becoming diffuse in appearance. By mid‐ to late summer androgen synthesis is at a maximum, increasing circulating androgens and stimulating the development and return of the solid granules. This study utilized electron microscopy and steroid radioimmunoassay to examine seasonal cycles of sex granules, in terms of development, maintenance, and regression, correlated with plasma androgen concentration. In addition, this investigation provides evidence of a possible secondary source of androgen secretion. J. Morphol. 261:70–80, 2004. © 2004 Wiley‐Liss, Inc.     

Sexual size dimorphism and male combat in snakes

Summary This paper reviews published literature on snakes to test the hypothesis that large male size, relative to female size, evolves because of the advantage it confers in male combat. Analysis of the data reveals a high correlation between the occurrence of male combat, and sexual dimorphism in which the male is the larger sex. This correlation holds (i) within the total sample of snake species (n=224), (ii) within the family Colubridae (n=134), and (iii) in a comparison between the eight families of snakes for which data are available. These results strongly support the hypothesis that large male size is an adaptation to intrasexual competition. The analysis also shows that females are larger than males in about 66% of snake species, that male combat is known in only about 15% of species, and that both sexual size dimorphism and the incidence of male combat tend to be distributed along taxonomic lines.     

Intraspecific variation in ejaculate traits of the northern watersnake (Nerodia sipedon)

Sperm competition occurs when the sperm of more than one male compete to fertilize the eggs of a female. In reptiles, sperm competition is particularly prevalent and is an important agent of sexual selection in males. Spermatogenesis in reptiles can be energetically expensive, suggesting that there may be costs to producing high-quality ejaculates. The northern watersnake Nerodia sipedon has a mating system characterized by aggregations in which a single female mates with multiple males, resulting in high levels of multiple paternities. Under these circumstances sperm competition is likely important, and selection should favour sperm and ejaculate traits that enhance a male's reproductive success. In this study, we examined intraspecific variation in ejaculate quality (sperm length, motility, sperm density, spermactocrit) in male watersnakes and determined whether ejaculate traits varied with body size and condition, using both size-corrected mass and haematocrit as indices of condition. We found large variation among males in all these traits, except for sperm length. Although there was significant variation in sperm length among males, the majority of variation in sperm length occurred within rather than among individuals. Males with high haematocrit had sperm that were less variable with respect to length, and large males produced ejaculates that were less concentrated with respect to sperm than small males. The lack of condition dependence of most ejaculate traits is consistent with previous studies that indicate that male reproductive effort in this species is generally not energy limited, perhaps because of opportunistic foraging during the mating season.     

Reduction in locomotor ability as a cost of reproduction in gravid snakes

Summary Recent studies suggest that lower survival among gravid squamate reptiles may be partially the result of decreased locomotor ability during gestation. In this study, we compared the speed and endurance of female garter snakes (Thamnophis marcianus), before, during, and after pregnancy. Gravid snakes had significantly lower locomotor performance than did non-gravid females, and performance varied among stages of gestation, reaching a minimum 0–6 weeks prior to parturition. Both number of offspring and relative clutch mass were inversely correlated with locomotor performance; as females increased these traits, locomotor ability decreased. If reduced locomotor performance results in greater risk of predation and/or lowered foraging ability, then natural selection (operating via differential mortality or feeding rates of gravid females) may result in important constraints on both clutch size and relative clutch mass in squamates.     

The strange occurrence of male cannibalism on adult females in snakes

Cannibalism is widespread in the animal kingdom. The most common, well-known cases are found in some arthropod systems where the larger females occasionally consume the smaller males before, during, or after mating (i.e., sexual cannibalism). The killing and consumption of females by males are, on the other hand, seemingly much rarer and theoretically more difficult to explain, at least among adults, because the consumption of females would result in the loss of potential mates for males. Here, we use observations submitted to a community science reporting system to describe three instances of adult males feeding on adult females in the Montpellier snake, Malpolon monspessulanus, in the south of France. We discuss our observations in the theoretical framework of cannibalism to provide potential explanations for these observations and briefly review empirical findings of cannibalism in vertebrates, which confirms that this sex-reversed form of sexual cannibalism is rare in nature.     

Sexual selection and adult male orangutan long calls

Long calls given by adult male orangutans (Pongo pygmaeus) have been hypothesized to mediate interindividual spacing among male orangutans and to attract female orangutans, to high-ranking mates over long distances. To test these two hypotheses I conducted observations of calling behaviour and a series of experimental playbacks of recorded long calls. The findings were consistent with the male-spacing hypothesis but did not support the mate-attraction hypothesis. Experimental playbacks demonstrate that long calls regulate spacing between males through an approach-avoidance system based on dominance relationships: the highest-ranking adult male approaches calls, while lower-ranking, males avoid calls. The results of playbacks also reveal that sexually active females do not move toward long calls, and therefore do not indicate that long calls function in attracting mates. This latter finding suggests that female mate choice may not have been an important selective factor during the evolution of long calls.     

Standard and digestive metabolism in the banded water snake, Nerodia fasciata fasciata

Estimating energy costs by respirometry is fundamental to many studies of the ecology, behavior and evolution of reptiles. However, traditional respirometry procedures seldom incorporate objective techniques for removal of outliers from estimates of metabolic parameters. We demonstrate how computer-automated respirometry equipment, which records many respiratory measurements over short intervals, can be coupled with mathematical procedures to produce robust estimates of pre- and post-prandial metabolism in banded water snakes (Nerodia fasciata fasciata). Standard metabolic rate of N. f. fasciata was estimated to be 1.21 ml O2/h (mass = 30.21 +/- 0.74 g) at 25 degrees C. After ingestion of a fish equaling 20% of their body mass, snakes exhibited a fivefold increase in metabolic rate with peak O2 consumption rate (VO2) reaching 6.5 ml O2/h. Total cost of digestion was 5.44 kJ, equivalent to approximately 21% of the energy in the meal. Repeated measurements of metabolism in the same individuals revealed that our methods yielded similar results, even when individuals exhibited different patterns of VO2 variation between respiratory trials. Our results underscore the importance of obtaining many VO2 measurements, coupled with objective removal of outlier values from estimates of metabolic rate, especially when metabolic values are to be interpreted in a comparative context.     

Resource allocation to growth,reproduction and survival in Gladiolus : The cost of male function

Theoretical models of life-history evolution assume trade-offs between present and future reproduction and/or survival. Models of the evolution of sex assume trade-offs between male function and female function. Generally, experiments designed to evaluate the cost of reproduction on other functions tend to ignore male function. The present work on Gladiolus takes into account simultaneously the different primary functions of the plant and separates sexual reproduction into one male component (pollen production) and one female component (seed production). The study of environmental (within-clone), between-clone and genetic correlations using strains of Gladiolus and principal component analysis show that trade-offs exist between male function, female function and survival, including both characters of plant vigour, perennation (corm production) and vegetative propagation (cormel production). Phenotypic correlations, using different species and species-hybrids, have been obtained which confirm these results. In particular, these results underline the importance of the impact of pollen production on the other functions.