Haematology of the grass snake Natrix natrix natrix L.

1. Haematological parameters and cytomorphological picture of circulating blood of the grass snake Natrix natrix natrix L. were studied. Mean annual values of the haemoglobin concentration, haematocrit level, red cell counts, erythrocytes indices, erythrocyte sedimentation rate, thrombocyte and leucocyte counts, per cent composition of white blood cells and size of morphotic elements were determined for a population sample (N = 154) of the species, considering the sex of snakes. 2. Parameters of red blood cell system (RCC, Hb, Hct) were statistically significantly higher in males, remaining indices showed no statistically significant differences. 3. A strict positive correlation was found between RCC, Hb and Hct in the studied sample. 4. Cytomorphology of blood is presented in microphotographs. 5. In the discussion haematological data for the grass snake and other snakes are compared.     

Seasonal changes of circulating blood parameters in the grass snake Natrix natrix natrix L.

• 1.1. Seasonal changes of circulating blood parameters of Natrix n. natrix were evident and involved both sexes to the same extent.
• 2.2. A significant decrease in red cell count, haematocrit and haemaglobin concentration in the mating period, and an increase in those parameters and mean cell volume in autumn were observed, and haemodilution during winter torpor.
• 3.3. The changes during the breeding season had probably a hormonal background; in winter, they resulted first of all from a decreased erythropoietic activity and, to a lesser extent, from an increased red blood cell breakdown rate. However, the possibility that some erythrocytes were withdrawn from the circulation cannot be excluded.
• 4.4. Winter lymphocytopenia, eosinocytopenia and neutrophilic granulocytosis in females during egg laying were expressions of changes of leucocyte formula.
• 5.5. Seasonal cyclicity was found only with respect to the white cell count in males and the eosinophile fraction in males and females.
• 6.6. Probable reasons for, and mechanisms of the changes in blood composition are discussed.

     

Xanthine:NAD+ oxidoreductase in the liver of grass snake Natrix natrix

The enzyme hydroxylating oxypurines in the liver of grass snake (Natrix natrix, Colubridae) was found to be a stable xanthine:NAD+ oxidoreductase (EC 1.2.1.37). The Michaelis constants for NAD+ and xanthine amounted to 14.4 and 12.3 microM, respectively. The enzyme affinity to hypoxanthine is lower than that to xanthine, but the former substrate is hydroxylated faster than the latter. The enzyme is only slowly and slightly (up to 22%) inhibited by NADH accumulating during xanthine hydroxylation. The above data and the time-course of hypoxanthine----xanthine----uric acid hydroxylation indicated that the kinetic properties of the snake liver enzyme provide in this uricotelic animal fast elimination of superfluous nitrogen derived from protein catabolism.     

Comparison of xanthine:NAD+ oxidoreductase from liver of two uricotelic species: hen Gallus gallus and snake Natrix natrix

1. Kinetic properties of xanthine:NAD+ oxidoreductase from liver of two uricotelic species of vertebrates (hen Gallus gallus and snake Natrix natrix) are compared. 2. Hen enzyme is saturated by hypoxanthine and xanthine at higher concentrations than the snake enzyme. For both species the enzyme-saturating concentration and hydroxylation rate of hypoxanthine are higher than those of xanthine, and the rate of uric acid production in the hypoxanthine----xanthine----uric acid reaction sequence is independent of the initial hypoxanthine concentration. 3. Km's for xanthine are the same, but Km for NAD+ of the hen enzyme is approximately 5-fold lower. The enzyme from both species is inhibited by NADH only slightly and at high non-physiological concentrations.     

The development of the trigeminal jaw adductor musculature in the grass snake Natrix natrix

The ontogenetic development of the jaw adductor musculature in Natrix natrix (L.) is described in detail and related to patterns of ossification in the skull. A comparison with the development of the jaw adductors in the lizard Podarcis sicula reveals some interesting differences of dynamics in the developing head. The problem of the establishment of topological relations of similarity (homology) in developing systems is discussed.
The homologies of the external jaw adductor compartments in lizards and snakes are revised. Early ontogenetic divergence explains the reversal of fibre direction in the anterior portion of the external adductor of snakes, and renders the homologization of that fibre bundle with part of the external adductor in lizards impossible.     

Motor reactivity of isolated heart of the grass-snake (Natrix natrix L.)

Stimulation of the vagus nerve with a volley of electric impulses changed the action of grass-snake heart producing a negative chronotropic and inotropic effect. The effect of vagal stimulation was not different from the effect of acetylcholine administration and it was absent in the presence of atropine and hexamethonium. It was not possible to demonstrate sympathetic nervous fibres in the stimulated segment of the vagus nerve and trials of finding a separate nerve increasing the heart rate were unsuccessful. Parasympathicotonic agents caused bradycardia and a fall in the amplitude of cardiac contractions, and in sufficiently high doses they arrested the heart in diastole. The action of muscarine-like agents was stronger than that of nicotine, and the anticholinergic action of tubocurarine was weaker than that of atropine. Catecholamines exerted a positive inotropic and chronotropic effect which was completely blocked by propranolol in some tests only.     

Multivariate patterns of geographic variation between the island and mainland populations of the Eastern grass snake (Natrix natrix natrix)

The substantial racial variation between the mainland and island populations of the Eastern grass snake ( Natrix natrix natrix ) is analysed by a range of multivariate methods, including principal component and canonical analysis. These techniques reveal a complex pattern of geographic variation which include sharp transition zones, gradual clines, a wide range of divergence of island populations and greater divergence per distance in the south than in the north. These patterns relate to the phylogenesis of this "incipient" species, and its post-Pleistocene range expansion as presented here and elsewhere. These racial patterns do not generally relate to physiographic features, conventional subspecies or CURRENT physical or biotic factors.     

Individual success in mating balls of the grass snake, Natrix natrix: size is important

Natricine colubrid snakes, including the grass snake, Natrix natrix , are frequently involved in complex social behaviour during the reproductive season. During these social behaviours, several males may simultaneously court a single female, resulting in a 'ball'of mating snakes in which each male 'combats'with rival males by 'tail wrestling'(see Madsen & Shine, 1993). I performed some experiments in outdoor enclosures for testing the male-male competition and the determinants of mating success in male grass snakes involved in such 'ball'aggregations. I demonstrated that competition between males occurred both when a single female was available to multiple males and when two females were simultaneously available to males. The larger males achieved more copulations than the smaller ones, thus demonstrating that body size is a crucial determinant of the individual mating success. It was not clear which aspect of male body size is the most important in determining success in these mating 'balls', but it was evident that the age of the 'fighting'male was not correlated with mating success. Larger females attracted more males than smaller ones, both in the field and in the enclosure. Furthermore, when the size difference between available females in the cage was high, only the largest female was courted and coupled.     

Bluffing and Waiting: Handling Effects and Post-Release Immobility in a Death-Feigning Snake (Natrix natrix)

Tonic immobility is a common response of animals to capture by a predator; in some cases, the behaviour is elaborated into death-feigning. Death-feigning is usually interpreted as a last-resort anti-predator tactic that depends on the predator ceasing or slowing its attack when the prey is apparently dead, thus buying time for the prey to escape if the predator's attention is directed elsewhere, even momentarily. I tested the effects of different handling regimes on the expression of death-feigning in the grass snake ( Natrix natrix ). In one test, degree of handling had a significant effect on frequency of death-feigning in small snakes, with snakes that received more handling feigning death more often. A second test showed that different initial handling regimes also affected the probability of death-feigning in large snakes, with snakes initially held by the tail feigning death least frequently and those initially held by the head most frequently. Thus, increased apparent threat to a vulnerable part of the body is more likely to result in death-feigning. It remains to be seen whether death-feigning can reduce the threat posed by real predators, but immobility is frequently carried over into the post-release period, during which the animal presumably weighs its chances and awaits an opportunity to escape. Regardless of cause, death-feigning snakes exhibited significantly more frequent and longer post-release immobility than did non-death-feigning snakes, which typically fled immediately upon release. Overall, small snakes feigned death less frequently than adults and were more likely to flee upon release, suggesting that immobility is a riskier anti-predator defence strategy for them.     

The contribution of small collections to species distribution modelling: A case study from Fuireneae (Cyperaceae)

The recent and rapid digitization of biodiversity data from natural history collection (NHC) archives has enriched collections based data repositories; this data continues to inform studies of species' geographic distributions. Here we investigate the relative impact of plant data from small natural history collections (collections with < 100,000 specimens) on species distributional models in an effort to document the potential of data from small NHCs to contribute to and inform biodiversity research. We modelled suitable habitat of five test case species from Fuireneae (Cyperaceae) in the United States using specimen records available via the Global Biodiversity Information Facility and that of data ready to mobilize from two regional small herbaria. Data were partitioned into three datasets based on their source: 1) collections-based records from large NHCs accessed GBIF, 2) collections-based records from small NHCs accessed from GBIF, and 3) collections-based records from two small regional herbaria not yet mobilized to GBIF. We extracted and evaluated the ecological niche represented for each of the three datasets by applying dataset occurrences to 14 environmental factors, and we modelled habitat suitability using Maxent to compare the represented distribution of the environmental values among the datasets. Our analyses indicate that the data from small NHCs contributed unique information in both geographic and environmental space. When data from small collections were combined with data from large collections, species models of the ecological niche resulted in more refined predictions of habitat suitability, indicating that small collections can contribute unique occurrence data which enhance species distribution models by bridging geographic collection gaps and shifting modelled predictions of suitable habitat. Inclusion of specimen records from small collections in ongoing digitization efforts is essential for generating informed models of a species' niche and distribution.     

SDMvspecies: a software for creating virtual species for species distribution modelling

The following is a report on software developed to create virtual species for the study of species distribution modelling (SDM). SDMvspecies provides several methods to create virtual species. The package is designed to be simple and intuitive, even for users who are not familiar with the R language. SDMvspecies is available online free of charge from < http://cran.r‐project.org/web/packages/sdmvspecies/ >.