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Population Dynamics and Spatial Dist'ribution of Tlle Terrestrial Snail Ovachlamys Fulgens (Stylommatopbora: Helicarionidae) in a Tropical Environment

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Population Dynamics and Spatial Dist'ribution of Tlle Terrestrial Snail Ovachlamys Fulgens (Stylommatopbora: Helicarionidae) in a Tropical Environment Rev. Biol. Trop., 48(1): 71-87, 2000 www.ucr.ac.cr Www.ots.ac.cr www.ots.duke.edu Population dynamics and spatial dist'ribution of tlle terrestrial snail Ovachlamys fulgens (Stylommatopbora: Helicarionidae) in a tropical environment Zaidett Barrientos Departamento de Malacología, Instituto Nacional de Biodiversidad (INBio), Apdo. 22-3100 Sto. Domingo, Heredia, Costa Rica. Fax (506)2442816, E-mail: [email protected] Received 8-VI-1999. Corrected 9-XI-1999. Accepted 20-XI-1999. Abstract: The introduced snail Ovachlamys fulgens (Stylommatophora: Heliearionidae) oeeurs on cultivated land habitats in Costa Rica, where its macrodistribution seems to be limited by annual mean temperature (20 - 27.6°C) and annual preeipitation (1 530 - 3 034 and 3 420 - 8 000 mm, with no more than six dry months). This species can be found in ¡itter and on vegetation up to 70 cm tal\. Randomquadrat field sampling was done in leaf litter and understory plants every three months for a total of five dates inCentral Costa Rica. At least 150 plots of 2Sx25 cm were analyzed on each date. Abundance of living specimens andeggs was positively correlated with (1) litter abundance anddepth, (2) litter and soil humidity, (3) relative humidity and (4) earlymoming tempera­ ture (6:30 AM), and negatively correlated with temperature later in the moming (10:00 AM). Besides these fac­ tors, living snail abundance was eorrelated with thickness of the herbaeeous vegetation and with the oeeurrence of fueca elephantiphes (in litter and understory). Egg abundanee was also correlated with the sampling date, apparentlybecause of changes in humídity. The correlationpattem of shell abundance was opposite to that of liv­ ing specimens. Population size and number of empty shells throughout the year parallel the rainfall pattem. Reproduetion takes place between May and November (wet season); and up to 92% ofthe specimens can befound aestivating betweenDecember and April (dry season). Clutch size averages three eggs. The maximum density of living specimens was reaehed in December (43.4 1 ind/m2) and the mínimum in March (8.30 ind/m2). Shells decompose in all average of fivemonths. Key words: Landsnail, distribution, microdistribution, Ovachlamys, Helicarionidae, reproduction, demography, shell decomposition, Costa Rica. In Costa Rica,as in many other places, Ín­ by Japan (http://is3.hk.super.net/-csjwv/dyt­ creased human activity has accelerated the in­ f.html), its subtropical weather has the fo11o­ troduction of snail species, enhancing the need wing ranges: average temperature nADc, mi­ to study the niches they occupy as weU as the nimum 13.6-26.1°C, maximum 18.6-3l.1°C, adaptations that allow their establishment in yearly precipitation 2 036.8mm without dry tropicalcountries. Ovachlamysfulgenswas 011- months (data from Naha City, Okinawa lsland) ginally described from Loo Choo (Gude 1900) (http://jin.jcic.or.jp/stat/stats/OlCEN 12.html). an archipelago also known as Líu Chi or Ryuk­ This species has entered other countries in yu Islands (Okinawa being the largestisland of shipments of ornamental plants (F.G. Thomp­ it). The archipelago is currently administrated son, pers. comm. 1995). 1t may have become 72 REVISTA DE BIOLOGÍA TROPICAL established in Costa Rica in the last 15-20 sp. (plantain), Psidium guajava (guava), Psi­ years, with the massive cultivatíon of natíve dium friedrichsthalianum (sour guaya), Yucca and introduced ornamental plants. Currently, elephantipes (yucca), Annona muricata (gua­ O. fulgens is considered an agricultural pest nabana) and Acnistus arborescens (güitite). (Monge-Nájera 1996), but with the exception The only maintenance was mechanical mowing of a few discussions about its taxonomic classi­ (about four times ayear). The site was selected ficationand its life cycle, there appearsto be no because it was the only Costa Rican habitat of informationon the ecology of the species. O. fulgensknown when the study began. Studies done in temperate habitats found Fieldmethods: Considering the body size that the microdistribution of leaf litter pulmo­ of O. fu/gens, site characteristics, ecological nates depends on the amount of lítter (Boag variables and time availability, the lot was di­ and Wishart 1982, LocasciuIlí and Boag 1987) víded into 3 984 quadrats (25x25cm2). Sample and humidíty (Boag 1985). The scanty infor­ size was determined by preliminary sampling mation about tropical snails suggests that well­ and the equatíon presented by Southwood drained soils �e also important (Peake 1968). (1978). Five samplings of at least 150 quadrats This paper about Costa Rican O. fulgens re­ each were done every three months with the ports on: 1- macrodistribution of the species, assistance of a random number generator. 2- microdistribution of individuals, shells and Sampling was done in the following dates: eggs, 3- the yearly changes in abundance, 4- 1= December 18; 1992 (beginning of dry reliability of the sampling procedure and 5- season), 159 quadrats. shell decay. 2= March 6-14; 1993 (dry season), 153 quadrats. 3= June 8-18; 1993 (wet season), 150 qua­ MATERIALS AND METHODS drats. 4= September 7-14; 1993 (wet season), Macrodistribution: The geographic distri­ 150 quadrats. bution was plotted from the database of the De­ 1= December 7-19; 1993 (beginning of partment of Malacology, Instituto Nacional de dry season), 150 quadrats. Biodiversidad (INBio) in Heredia, Costa Rica. Ca­ In each quadrat, quantitative and qualitati­ lJecting sires inelude forested and cultivatedareas. ve variables that potentiaUy affected the abun­ dance (or presence) of snails, shells and eggs Microdistribution: were measured (Tables 2-6). Al molluscs seen without special optical equipment were col­ Study area: Field work was done in an ur­ lected for 5 min in each quadrat (see Villalobos ban lot (12x18m2) in Pavas, San José, Costa et al. 1995, Emberton et al. 1996). The number Rica (9°56' 45"N, 84°07' 15"W, 1075 masl)o of live individuals, shells and eggs of O. ful­ The biotic area classification is "subtropical, gens were recorded in each quadrat, as was the tropical, moist with 5-6 dry months" (Herrera shell diameter of dead and live individuals and Gómez 1993). The soil is slightly alkaline (cIassified as "active" or "aestivating"). Live (pH = 6.6) and caIcium-rich (Ca+=17.1 o. fu/genswere returnedto the originalsite af­ cmoI(+ )/1) (Centro de Investigaciones Agronó­ ter measuring. Plant cover was measured with micas, Universidad de Costa Rica). The site, a fixed point pattern (lcm between points) and originally cleared for a coffee plantation, beca­ soil moisture by weíght differen�e after oven me an urban lot wÍth secondary growth and drying for 48hr at 90°C (moist weight dívided grasses (Pennisetum purpureum). In 1982 an by dry weight). Litter was divided in three le­ orchard was planted, mainly: Pasea america­ veIs: 21 closest to the soil, ¡eaves were black na (avocado), Citros spp. (lemon, tangerine, and retained more humidity and could only be sour orange), Mangiferaindica (mango), Musa classifiedas monocotyledons and dicotyledons BARRIENTOS: Population dynamics oC a tropical landsnail 73 b�cause of decomposition, Z2 brown dead lea­ 1993, on 39 randomly selected 25x25cm2 qua­ ves identifiable to family or species level; and drats. After the standard visual count, all the Z3 freshly fallen leaves, green-yellow-brown, vegetation and soil to a depth of 1 cm were co­ in many cases the layer was artificially produ­ llected and examined in the laboratory under ced when trees were routinely pruned. To lOx magnification. The correction factor was analyze the effect of plants, only the most calculated as the total number of specimens di­ abundant species of each quadrat was conside­ vided by the mean number of visually counted red. When abundance was assessed the follo­ specimens. wing rating was used O::absent, l=absent-scar­ Shell decomposition: The decomposition ce, 2=scarce, 3=scarce-regular, 4=regular, time was measured for 23 shells, 5 mm or mo­ 5=regular-abundant, 6=abundant, 7=very re in diameter, placed simultaneously in a abundant. Abundance of fungal hyphae was li­ mesh cage left in the field; this prevented ex­ mited to those visible to tbe naked eye. posure to macrofauna yet allowed the action of Microhabitat temperature was measured on water and other climatic factors. Shells were each sampling day at around 06:30 and 10:00 examined monthly and classified as either in­ hours on areas with trees and without them at tact, faded or broken. four levels: 1) in air 1 m aboye ground, 2) in her­ Vo ucher O. fulgens were deposited in the baceous vegetation 3 cm aboye ground, 3) insi­ Museum of Natural History, University of Flo­ de litter and 4) at a depth of 0.5 cm in the soil. rida and Instituto Nacional de Biodiversidad Spearman's correlations were applied to (INBio) catalogue numbers 1474249 (17 speci­ estimate the correlation of the 34 quantitative mens) and 1474252 (19 specimens). Summary variables with the number of shells, living statistics follow this format: mean (sample size, snails and eggs of O. fulgensper quadrat. Chi­ ±standarddeviation, minimum-maximum). square was used for the analysis of the qualita­ tive variables, but as the amount of cells with less than five data was high the data were mo­ RESULTS dified as follows: the dependent variables we­ re classified as O = absent and 1 = present, all Macrodistribution: The distribution of O. grasses were merged into a single category, the fulgensin Costa Rica (Fig. 1) is correlated with same was done for Myrtaceae species, and ca­ human presence and specially with agriculture tegories with low frequencies were discarded. (Table 1). Only in Pejibaye de Cartago and in Demograpby: The snails were classified Río Claro de Puntarenas (Table 1) was it found as: Neonates (shell diameter �2.5mm), Juveni­ in a secondary forest, othersamples were collec­ les (shell diameter 2.6 - 4.9 mm) and Adults ted in gardens or plantations.
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