NATURAL HISTORY AND POPULATION DYNAMICS OF THE LAND SNAIL HELIX TEXTA IN ISRAEL (PULMONATA: HELICIDAE)

Helix texta is endemic to the Mediterranean regions of Israel.It has a seasonal activity pattern which starts in the autumn,with the first rains, and dwindles towards the spring, whenthe snails dig into the ground for a six months long aestivation.A cold spell of 0°C will, however, terminate the activeseason of the adult snail, even in the middle of the rainy season.Survival of the young is very low and most of them (90%) donot survive their first year, because of the winter cold andthe summer drought. Massive predation of adult snails by wild boar was observedin December 1986: within a few days, about 50% of the adultsin the study in the area were eaten. A very rapid growth ofyoung and subadults was observed immediately after this predation.These observations suggest that the extent of recruitment ofnew adults to the population may be partly controlled by existingadults, through a growth-inhibiting pheromone in the mucus.After predation this inhibiting factor disappears, enablinga subsequent rapid growth of the young. In this manner, massive,irregular predation of the adults by a large predator, and changesin juvenile survival, result in sharp fluctuations in the agestructure of the population. The resulting pattern of unstablepopulation dynamics is different from that described for theEuropean species of Helix. (Received 16 January 1989; accepted 17 April 1989)     

POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE MEDITERRANEAN LAND SNAIL ALBINARIA CORRUGATA (PULMONATA: CLAUSILIIDAE)

The amount of gene flow among local populations partly determines the relative importance of genetic drift and natural selection in the differentiation of such populations. Land snails, because of their limited powers for dispersal, may be particularly likely to show such differentiation. In this study, we directly estimate gene flow in Albinaria corrugata, a sedentary, rock-dwelling gastropod from Crete, by mark-recapture studies. In the same area, 23 samples were taken and studied electrophoretically for six polymorphic enzyme loci. The field studies indicate that the population structure corresponds closely to the stepping-stone model: demes are present on limestone boulders that are a few meters apart, and dispersal takes place mainly between adjacent demes. Average deme size (N) is estimated at 29 breeding individuals and the proportion of migrants per generation at 0.195 (Nm = 5.7). We find no reason to assume long-distance dispersal, apart from dispersal along occasional stretches of suitable habitat. Genetic subdivision of the population, as derived from F_ST values, corresponds to the direct estimate only at the lowest spatial level (distance between sample sites < 10 m), where values for Nm of 5.4 and 17.6 were obtained. In contrast, at the larger spatial scales, F_ST values give gene-flow estimates that are incompatible with the expected amount of gene flow at these scales. We explain these discrepancies by arguing that gene flow is in fact extremely limited, making correct estimates of Nm from F_ST impossible at the larger spatial scales. In view of these low levels of gene flow, it is concluded that both genetic drift and natural selection may play important roles in the genetic differentiation of this species, even at the lowest spatial scales.     

中国陆生贝类一新种（柄眼目：坚螺科）

1984年至1986年间作者在云南省南部地区考察,获得陆生贝类标本一批,经鉴定发现一新种,即金平坚螺Camaena jinpingensis、隶于肺螺亚纲、柄眼目、坚螺科、坚螺属。本文对其新种形态特征,栖息环境进行了描述,并对其相近似种也进行了讨论。     

FOOD SELECTION IN THE LAND SNAIL LIMICOLARIA FLAMMEA MULLER (PULMONATA: ACHATINIDAE)

Food consumption by juvenile and adult specimens Limicolariaflammea was measured as dry weight of food eaten. The snailwas fed weekly for 15 months on four types of food (lettuce,potato, apple and carrot) and chalk provided, hence an opportunityto select for food. Juvenile snails showed greater selectivitydepending on whether the food was needed for somatic growthor reproduction. On the other hand, when fed exclusively onone each of the same food items and given no opportunity tochoose, the snails grew better on the same food items they selectedin the choice experiment. The absorption efficiencies (AD) werehigh, the utilisation efficiency (ECI) was 1–13% and showeddistinct peaks corresponding to egg production. Values for ECDwere 1.25–15.5%. Variation in growth rates was due mainlyto utilisation efficiencies of the four foods tested. Largeamounts of chalk were eaten during the period of somatic growthand greatly reduced during the reproductive phase of L. flammea.The significance of these findings to the snails ecology isdiscussed. (Received 9 April 1991; accepted 11 June 1991)     

TAXONOMIC REVIEW OF THE TREE SNAIL AMPHIDROMUS ALBERS, 1850 (PULMONATA: CAMAENIDAE) IN THAILAND AND ADJACENT AREAS: SUBGENUS AMPHIDROMUS

Extensive series of Amphidromus Albers, 1850 were collectedfrom low-altitude areas (30–500 m above sea level)in Thailand, Laos, Malaysia, Singapore and Indonesia between1993 and 2004. Data on shell colour, genital and spermatophoremorphology, and radular structure are presented. Five speciesof Amphidromus (Amphidromus) are reviewed, of which four occurin Thailand. Intraspecific variation between populations ofAmphidromus (Amphidromus) atricallosus (Gould, 1843) is considered;three subspecies are accepted and a new subspecies described.Subspecies of A. (A.) inversus (Müller, 1774) and A. (A.)schomburgki (Pfeiffer, 1861) are reconsidered, and a new subspeciesof the latter is described. A dichotomous key to the speciesand subspecies of Amphidromus (Amphidromus) in Thailand is provided. (Received 7 May 2004; accepted 31 March 2005)     

中国陆生贝类一新种(肺螺亚纲:柄眼目:拟阿勇蛞蝓科)

作者于1981～1983年,参加中国科学院横断山综合考察队,赴云南地区进行考察,又于1993～1994年,赴滇南一带调查,获得陆生贝类标本一批,经整理鉴定,得拟阿勇蛤蝓科一新种：即云龙隐带螺Cryptozonayunlongensissp.nov.。本文对新种的形态特征作了详细描述,并对生殖系统进行了解剖,对颚片和齿舌作了电子显微镜观察,与其近似种古怪隐带螺Cryptozonaerratica（Heude,1882）进行了比较。     

THE LIFE CYCLE, POPULATION DYNAMICS, GROWTH AND SECONDARY PRODUCTION OF THE SNAIL BRADYBAENA FRUTICUM (MULLER, 1774) (GASTROPODA PULMONATA) IN NORTHERN GREECE

The life cycle, population dynamics, growth and secondary productionof the pulmonate land snail Bradybaena fruticum were studiedin northern Greece. The demographic analysis of the populationsof B. fruticum revealed that a) two cohorts exist in the fieldthroughout the year b) the reproductive period started in thebeginning of summer and c) growth was increased during springand autumn. According to von Bertallanffy' method B. fruticum needs 5 yearsto attain its maximum size in the field of 25.40 mm. Mortalityrate increases and life expectancy decreases with increasingage. Net reproductive rate (Ro) was equal to 3. 15 and per capitarate of increase (r^c) was equal to 1. Annual secondary production, calculated by Hynes' size frequencymethod, revealed a mean annual density of 5.9 individuals/m²,a mean standing crop (B) of 8.09 g/m²/year and an annual production(P) of 1.92 ± 0.11g/m²/year. Annual turnover ratio (P/B)was equal to 2.37. (Received 25 May 1989; accepted 17 March 1989)     

贵州洞穴陆生贝类一新种(肺螺亚纲,柄眼目,烟管螺科)

记述了贵州省洞穴烟管螺科真管螺属1新种,即荔波真管螺Euphaedusa libonensis sp.nov．,标本采自贵州省荔波县董背洞内。对新种进行了详细描述,并与近似种做了比较。模式标本保存在中国科学院动物研究所无脊椎动物标本馆。     

ASPECTS OF THE LIFE CYCLE, POPULATION DYNAMICS,GROWTH AND SECONDARY PRODUCTION OF THE EDIBLE SNAIL HELIX LUCORUM LINNAEUS, 1758 (GASTROPODA, PULMONATA) IN GREECE

This paper examines the biology and ecology of Helix lucorumL. which lives in mainland Greece, as well as its growth andsecondary production. A demographic study revealed that (a)3 cohorts exist at any time during the year (when adults ofall generations belong to the same cohort) (b) egg-laying andhatching occur during the months of July and August respectively,(c) the most rapid growth takes place during spring. Study ofH. lucorum genitalia in relation to age showed that the snailsaxe sexually mature 3 years after hatching, when the largestdiameter of their shell (D) is equal to or greater than 35 mm Von Bertallanffy's method suggests that Helix lucorum may liveup to 14 years or more in order to reach its possible maximumsize (48.80 mm) The study of relative growth of D in relation to P_s (peristomesurface) of Helix lucorum shows that D grows faster than P_swhen D12.50 mm; juveniles change their growth rate when theirD arrives at 22.05 mm, and growth becomes slower when adultsarrive at 36.27 mm Annual secondary production calculated by the size frequencymethod gave a mean annual density of 3.39 individuals per m²,a mean annual crop (biomass) of 4.04 g-m^–2 and an annualproduction (P) of 5.02 g · m^–2. The annual turnoverratio (P//b) is equal to 1.24 (Received 23 June 1987;     

GROWTH AND REPRODUCTION OF THE MINUTE LAND SNAIL PUNCTUM PYGMAEUM (DRAPARNAUD)

Reproductive characters and growth of Punctum pygmaeum wereinvestigated under laboratory conditions. Eggs were laid singlyat intervals of 2–8 days. Egg size averaged 0.41 by 0.50mm. Due to a variable period of egg retention, newly depositedeggs contained differently developed embryos resulting in anincubation period varying from 1 to 34 days. Each snail on averagelaid 6.0 eggs (range 1–16 eggs) during its life. Egg productionrate and hatching success in individuals kept in isolation werethe same as in individuals kept in pairs. Hatchlings measured0.49 mm in shell breadth. Snails grew throughout their life,but the growth rate declined significantly at a shell breadthof 1.3–1.4 mm, the point at which the onset of sexualmaturity occurred. Comparison of reproductive characters ofsnails living in leaf litter revealed that very small-sizedspecies produce few, but in relation to their shell size largeeggs, which they oviposit singly. ^*Present address: Zoological Institute, University of Basel,Rhein-sprung 9. CH-4051 Basel, Switzertand (Received 23 May 1988; accepted 11 July 1988)     

PHYLOGENETIC RELATIONSHIPS OF THE CAMAENIDAE (PULMONATA: STYLOMMATOPHORA: HELICOIDEA)

The land snail family Camaenidae is defined by the lack of characters,such as dart sac and glands, which are present in related familiesAlthough the Camaenidae is regarded as monophyletic, no synapomorphiesunite the family It is possible that the Camaenidae is para-or polyphyletic This was investigated using a cladistic analysisof the super-family Helicoidea which was based on anatomicalcharacters from previous studies by Tilher (1989) Camaenidaewas split into two geographical groups (American and Australasian).Successive weighting generated a single most parsimonious cladogram(length 198, c.i. 0.61, r.i. 0.64), which suggested that theCamaenidae is a polyphyletic taxon. Evidence from this analysisdemonstrated that the American component of the Camaenidae isclosely related to the Hehcidae and Helminthoglyptidae, andthe Australasian component is sister group to the Bradybaenidae ^*Current address Department oJ Zoology. James Cook Universityof North Queensland. Townsville. Queensland 4811, Australia (Received 30 July 1994; accepted 20 June 1995)     

THE INFLUENCE OF INGESTIVE CONDITIONING ON FOOD CHOICES IN THE LAND SNAIL HELIX ASPERSAMULLER (GASTROPODA: PULMONATA: STYLOMMATOPHORA)

An experiment of food choice in one-day-old naïve landsnail Helix aspersa Müller was carried out, involving threephases. First, a training period on a monophagous diet of leafdiscs of Taraxacum officinale or Urtica dioica was given for1, 5, 10, 15 or 30 days. The area of leaf discs consumed byeach snail during this stage was assessed. In the intermediatephase, the animals were not fed for 24hours. Finally, food choicewas tested over 24 hours, when animals could eat leaf discsof each species of plant usedfor training. A coefficient offood preference for Urtica dioica was calculated (UPC = areaof Vrtica leaf disc consumed/area of Urtica and Taraxacum discsof leaves consumed for 24 hours). The length of the training phase did not reduce the percentageof individuals preferring the plant previously consumed. Onthe contrary, after 15 days of monophagous diet, animals formerlyfed on Urtica showed a coefficient of food preference for Urticasignificantly higher than Taraxacum fed animals, and after 30days of training this phenomenon was more pronounced. Thus,during the choice test, snails ate more of the previously eatenplant leaf. Our experiment highlighted the feeding behaviourplasticity of H. aspersa. The significant correlation betweenthe area of Urtica eaten during the training phase and the coefficientof food preference for Urtica is discussed. (Received 10 April 1994; accepted 10 January 1995)     

SEED BANK,SURVIVORSHIP, AND SIZE DISTRIBUTION OF A NEBRASKA POPULATION OF IMPATIENS CAPENSIS (BALSAMINACEAE)

The growth and survivorship of a population of Impatiens capensis were followed in eastern Nebraska. Soil cores were taken within the population to estimate the size of the seed bank. The number of seeds in the soil remained relatively constant from April through October, with a mean of 200 seeds/m². The seed bank size corresponded to observed seedling densities in the following years. Twenty permanent plots were established within the population and censused for three years, 1983–1985. Seedling recruitment to the population was similar in 1983 and 1984 but 30% higher in 1985. The seedling densities of plots were uncorrelated from year to year. Survivorship in 1983 followed a Deevey Type I pattern. Almost one-third of the population survived until chasmogamous reproduction. Plant size (stem height) was measured monthly on all individuals during 1983. The frequency distribution of stem height was positively skewed as long as growth was exponential. When the growth rate slowed, the distribution became more normal.     

THE EFFECTS OF POPULATION DENSITY ON GROWTH RATE IN LIMICOLARIA FLAMMEA MULLER (PULMONATA: ACHATINIDAE)

Density effects on growth and sexual maturity in laboratory-keptLimicolaria flammea MÜller are described; high densityresults in low growth, higher mortality and a delay in sexualmaturity. (Received 9 April 1991; accepted 11 June 1991)     

CONSUMPTION OF AMPHIPODS BY THE NEW ZEALAND LAND SNAIL WAINUIA URNULA (PULMONATA: RHYTIDIDAE)

An unusual carnivorous diet was documented for the rhytididsnail Wainuia urnula urnula (Pfeiffer) from the southern North Island,New Zealand. Eighty-two percent of 315 samples of faeces or gutcontents contained remains of landhoppers (Crustacea: Amphipoda).Earthworms (Oligochaeta) were the second most common food typeidentified, but occurred in only 4 percent of samples. In the laboratory,W. urnula urnula captured landhoppers by rapidly everting theTVU-section odontophore beneath the prey and immediately drawingit into the mouth in a single action. Diet samples from the allopatriccongeners W. edwardi (Suter) (n 5 51) and W. clarki Powell (n5 7) contained no remains of landhoppers, and these snails didnot eat landhoppers in the laboratory. The diet of these specieswas dominated by earthworms. The subspecies W. u. nasuta ateboth earthworms and amphipods with similar frequency (n 5 40).Individual teeth on the radula of W. urnula urnula had a simpleblade-like cusp like other Wainuia species, but there were fewerteeth per row and fewer rows on the whole radula than in otherspecies. W. u. nasuta had a similar number of rows of teethto W. clarki and W. edwardi, but the number of teeth per rowwas intermediate between these species and W. u. urnula. Recordsof pulmonate species preying on arthropods are rare, yet inthe case of W. urnula this specialization has been accompaniedby minimal morphological change. (Received 22 December 1998; accepted 11 March 1999)     

STARVATION, DESICCATION AND USE OF ALLOSPERM IN THE HERMAPHRODITE FRESHWATER SNAIL BIOMPHALARIA GLABRATA (GASTROPODA: PULMONATA)

Pairs of albino and pigmented snails were used to test the hypothesisthat the hermaphrodite freshwater gastropod Biomphalaria glabratawas able to conserve and use allosperm, despite periods of desiccationand starvation. In laboratory experiments, lots of twenty snailswere subject to 0, 5, 10, 15 AND 20 days starvation; similarlots of 20 snails were subjected to 0, 2, 4, 6, 8, 10 weeksof desiccation at R.H. 100%. After all periods of starvationand desiccation, albino parents were still producing significantnumbers of pigmented offspring, suggesting preferential cross-fertilizationusing stored allosperm. (Received 30 September 1993; accepted 23 December 1993)     

A NEW GENUS AND SPECIES OF PLANORBIDAE (GASTROPODA: HETEROBRANCHIA: PULMONATA) FROM NORTHERN AUSTRALIA

A new genus and species of Planorbidae, Kessneria papillosa n.gen.and sp. from the Northern Territory, Australia, are described.The new genus is morphologically most similar to Leichhardtia,another genus restricted to Northern Australia, from which itdiffers in a number of important shell and anatomical characters.The shell of the new genus differs from all other planorbidsin having a raised, papillate protoconch, and is also uniqueamongst Australian high-spired genera in being umbilicate and inhaving a sinuate outer lip. (Received 12 May 2000; accepted 25 September 2000)