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Cantareus Aspersus) in Northern New Zealand 60 60 NEW ZEALANDAvailable JOURNAL on-line at: OF http://www.newzealandecology.org/nzje/ ECOLOGY, VOL. 31, NO. 1, 2007 Morphology, distribution and desiccation in the brown garden snail (Cantareus aspersus) in northern New Zealand John K. Perrott1*; Iris I. Levin1,2 and Elizabeth A. Hyde1,3 1 EcoQuest Education Foundation, Ecological Field Centre, East Coast Road, RD 3, Pokeno, New Zealand 2 Department of Biology, Bowdoin College, Brunswick, Maine 04011, USA 3 Department of Natural Resources, University of New Hampshire, Durham, New Hampshire 03824, USA * Author for correspondence (E-mail [email protected]) Published on-line: 21 May 2007 ___________________________________________________________________________________________________________________________________ Abstract: Size, density and distribution of the brown garden snail (Cantareus aspersus) were observed relative to cover at a coastal reserve on the North Island, New Zealand. Cover variables depended on vegetation height and available debris (rubbish, wood, cow dung). Air temperature, ground temperature, and relative humidity were recorded continuously during the field survey for the various cover types. Live snails used debris disproportionately as resting habitat, where relative humidity and temperature were the least variable. Desiccation rates were obtained by dehydrating snails of three size-classes and calculating desiccation rates as a function of water loss over time under controlled conditions in the laboratory. Results suggest that the shell aperture:size ratio is an important ___________________________________________________________________________________________________________________________________factor regarding desiccation stress in C. aspersus, especially in young and small-sized snails. Keywords: Cantareus aspersus; coastal reserve; density; distribution; desiccation; shell size; shell aperture: size ratio Introduction establish from a single individual. In addition, under optimal conditions, C. aspersus can lay up to 2500 The brown garden snail Cantareus aspersus, formerly eggs in a season (Runham 1989). However, under dry Helix aspersa (Barker 1999), is a cosmopolitan pest conditions, juvenile survival and recruitment can be species the biology of which is well understood, but its significantly reduced (Heatwole & Heatwole 1978). invasion of New Zealand indigenous ecosystems is not Water loss in land snails occurs mainly from the mantle well documented. The species occurs widely in New cavity via respiration, through the shell aperture, and Zealand and there is growing concern about its potential through the shell itself depending on age and thickness adverse effects on indigenous ecosystems (Barker (Goodfriend 1986). Pulmonates minimise water loss & Watts 2002). Concerns about negative impacts via three primary means: structure, behaviour and include herbivore pressure, resource competition, physiology (Asami 1993). A snail’s shell and the habitat fouling, and parasite and disease transmission. secretion of a mucus epiphragm over the shell aperture The snail is calciphile and most prevalent in coastal are its primary structural means of protection from environments, especially the scrubland and dune water loss (Machin 1967). Retraction of the entire systems of the upper North Island. The New Zealand body inside the shell or avoiding daytime heat and Department of Conservation is currently considering dryness are behavioural means to reduce water loss strategic options and methodologies for the control (Machin 1967; Cameron 1978). Physiological means of C. aspersus on conservation lands. This study was include aestivation and the retention of pallial fluid conducted at a coastal reserve and provides information within the mantle cavity (Blinn 1964; Riddle 1981; on the effects of desiccation stress on the density and Smith 1981). size distribution of C. aspersus at the Robert Findlay Temporal variation in population density, growth, Wildlife Reserve, Miranda, Firth of Thames. and reproductive activity in land snails is largely Cantareus aspersus was introduced into New controlled by environmental conditions (Albuquerque Zealand during the 1860s and is now the most abundant de Matos 1989, 1990; Elmslie 1989). Shell size and and widespread terrestrial snail species. It is a potential shape can be highly variable, often reflecting age or pest due to its high fecundity and dispersal ability. adaptations to particular habitats. Reproductive output Because it is hermaphroditic, populations are able to is often positively correlated with shell size (Goodfriend New Zealand Journal of Ecology (2007) 31(1): 60-67 ©New Zealand Ecological Society PERROTT ET AL.: DESICCATION STRESS IN CANTAREUS ASPERSUS 61 1986). Because surface area to volume ratios decrease In aestivation, C. aspersus can survive water loss equal with increasing size, one would predict that conditions to 50% of its body weight (Barnes 1986). Understanding of greater desiccation stress would favour larger the occurrence and frequency of aestivation in relation snails (Nevo et al. 1983; Goodfriend 1986). This is to desiccation stress may allow wildlife managers to not found at the faunal level as both large and small design more effective control measures. snails occur in dry and wet conditions (Goodfriend In general, moisture conditions appear to be the 1986) even though within species, larger snails have best-documented environmental correlate of shell size greater survivorship compared with smaller snails in across a wide range of terrestrial land snail species low-humidity environments (Heatwole & Heatwole (Goodfriend 1986). Higher levels of moisture tend 1978; Knights 1979). to correlate with increased snail densities, growth Variation in shell aperture area relative to shell rates, and larger adult sizes (Brown 1913; Sacchi size has usually been interpreted in the context either 1965; Gould 1984). High snail densities probably of regulation of water loss or of reduction in predation correlate with both increased snail fecundity and pressure (Goodfriend 1986). Machin (1967) compared more efficient snail matings. These will be due in part three sympatric land snail species and found that, to more frequent or extended periods of activity and when inactive, the mean water loss through the shell higher food intake rates. However, there are many aperture increased with mean aperture size, regardless possible confounding variables such as calcium levels, of the presence of an epiphragm. Water loss through population density, and food availability. C. aspersus the shell aperture is proportional to aperture area represents an abundant, non-toxic, easily captured (Goodfriend 1986). C. aspersus is typically active prey, and therefore they are consumed by a wide in wet conditions and at night, when moisture loss range of predators. Avian and mammal predators have is supposedly less than during daytime. Activity and little difficulty extracting the snail from its shell. The population dynamics of C. aspersus are often dictated function of the shell, multipurpose or not, appears to by climate, as well as by availability of cover sites (tall be primarily related to protection from insect predators vegetation and debris). and water conservation. In this paper we investigate The functional significance of cover availability C. aspersus densities, distributions, and morphology is key to understanding population dynamics of in relation to cover type and desiccation stress. We test C. aspersus. The density and distribution of suitable the following predictions: (1) snail densities and size cover sites may regulate population structure in several distributions vary according to cover type, (2) smaller ways. High-density cover sites can contribute to high C. aspersus have elevated desiccation rates, and (3) snail densities by providing greater spatial and temporal C. aspersus with reduced shell aperture:size ratios have opportunities for egg-laying, less clumping and greater reduced desiccation rates. potential to avoid avian predators, greater opportunity to escape adverse weather conditions, increased activity due to greater likelihood of finding cover, and increased fecundity due to increased activity. Aestivation, Methods grouping and homing behaviours in C. aspersus may compensate to some degree in low-density cover sites. Study site Considering that exotic weedy species often compete The Robert Findlay Wildlife Reserve, about 25 ha, is best with native species in disturbed environments, situated at Miranda (37º13’S and 175º23’E; Fig. 1), the unpredictability of cover in coastal environments on the south-western coast of the Firth of Thames in may benefit C. aspersus via reduced competition for northern New Zealand. The reserve is an internationally space. Therefore, while it should not be surprising important habitat for shorebirds, providing a stopover to find calciphiles like C. aspersus predominately in and feeding site for up to 25 000 birds at a time, most coastal environments, their success in these habitats of which are migratory wading species. The reserve may be primarily due to their broad environmental is open to the public and largely comprised of pasture tolerances, high fecundity, and adaptability to stochastic grass. Up to 90 head of domestic cattle are grazed environments. periodically throughout the year on 15 of the 25 ha, In low-density cover sites, snails can manage but were not present during this study. Cattle grazing water loss by grouping together and by forming
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