Life Cycle and Field Abundance of the Snail Succinea Costaricana (Stylommatophora: Succineidae), a Tropical Agricultural Pest
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Rev. Biol. Trop.,43(1-3): 181-188,1995 Life cycle and field abundance of the snail Succinea costaricana (Stylommatophora: Succineidae), a tropical agricultural pest Carlos Villalobos M. \ Julián Monge-Nájera1.2, Zaidett Barrientos3and José Franc04 I Revista de Biología Tropical, Escuela de Biología, Universidad de Costa Rica, San José, Costa Rica. Centro de Investigación General, UNED, San José, Costa Rica. Instituto Nacional de Biodiversidad (INBIO), Apdo. 22-3100 Sto. Domingo, Heredia, Costa Rica. Coalición Costarricense de Iniciativas de Desarrollo,CINDE, Apdo. 7170-1000, San José, Costa Rica. (Revised 4-VIII-1994. Accepted 16-1-1995) Abstraet: The neotropical terrestrial snail Succinea costaricana has become a quarantenary pest in ornamental plants (Dracaena marginata, Dracaenaceae). Specimens were collected in GuápiJes, Limón, Costa Rica, where they reached a density of 282900 individualslha. In the field, reproduction is continuous (as is rainfall) and eggs, young and copu lating pairs are found mainly under moist litter. The population concentrated on plants rather than on the soil and greatly decreased after several control methods were applied. The life cycle was studied in the laboratory. The follow ing are mean values: 7 eggs/cluster; lA mm egg diameter; 0.25 and 0.84 mm embryo and newborn shell length, respectively; 11 days (embryo development at 24 oC); 16 mm/min locomotion speed (young). The animals can repro duce at 12 weeks (lifespan: 44 weeks). The pigmentation pattern is defined after seven weeks. Key words: Land snail, reproduction, life cycle, growth, quarantenary pest, population, spatial distribution. Most knowledge about the cosmopolitan ter La Palma and Guácimo, near Guápiles, Limón restrÍal snail genus Succinea Drapamaud 1801 Province, Costa Rica (von Martens 1892- (Succineidae) is based on European or North 1901). American species. The few studies on tropical This paper provides the first observations species are mostly local taxonomic papers about the life cycle and field abundance of the (Lanzieri 1966, Hertlein and Allison 1968) or species, which were studied taking advantage morphological studies (Patterson 1989). of an abnormal population increase of this Sorne species are agricultural pests, causing species in a farm during 1992 (Monge-Nájera damage to cucumber, tomato, lettuce, chrysan 1995). themus, carnations, roses and tulips. Sometimes they are carriers of diseases or intermediate hosts of parasites (Godan 1983). MATERIAL AND METHODS Taxa are particularly difficult to identify, in part because shell morphology is very similar in several species (von Martens 1892-1901, Study site: The study site was an ornamen Godan 1983). tal plant farm (Fig. 1) located at La Roxana de The neotropical species Succinea costari Guápiles, Limón Province, Costa Rica (altitude cana von Martens 1898 has been recorded from 106 m a.s.l.). The region has a tropical rainy Costa Rica through Mexico. It was described climate and the following yearly means: from specimens collected in the XIX century at 24.5°C, 4440 mm rainfall and 87 % relative 182 REVISTA DE BIOLOGIA TROPICAL Fig. l. The study !lite: organization of plant rows in the fann that currently covers what was a tropical rain forest in Limón, Costa Rica. humidity (Costa Rican Meteorological Insti dom numbers. The four observers participated tute). Soil pH is 4.6 and the:gtean calcium con in all types of counts to avoid individual bias. centration is 122 ppm (A&L Southern An estimate of density was done the night of Agricultural Laboratories, Florida, U.S.A.). March 13, 1992 (20:30-24:30 hr) when the The main crops of this farm are Dracaena mar population reached pest proportions, by count ginata and Dracaena deremensis (Dracaena ing snails and shells in plants and in a 20 cm ceae). radius hemicircle of soil at the base of each In the three years previous to this study, leaf plant. This was done with flashlights checking litter was normally abandoned on the plants only the accesible side of each plant (they are and soil to reduce costs and as an organic fertil planted in tight double rows, Fig. 1). A random izer. S. costaricana and another snail species, procedure was also used to decide if the west or became abundant under the litter. Pesticide east side of each row was examined. applications, basically aimed at the control of To assess changes in abundance throughout thrips (Thysanoptera), caused sorne mortality the year, visits were made to the field every among snails. The chemicals used include two months from March 1992 through May metamidofos OP, malation OP, metoMil C, 1993. Fixed time collections were done during protiofos OP, metiocarb C and endosulfan, but mornings on plant rows (5 min) and the soil at no detailed records were kept by the farmer their bases (5 min). This was done on the regarding time, frequency and other application accessible side of plants and on the undisturbed parameters. soil below (workers crush shells while walking between rows). The animals were counted in Field methods: All counts mentioned in the laboratory. this section were done on individually num To assess the reliability of sampling, a ran bered plant rows selected with a table of ran- dom selection of plants and a 20 cm radius VILLALOBOS et al.: Succinea costaricana: an agricultura! pest 183 hemicircle in their bases were sampled by the TABLE I normal procedure and then taken in sealed bags to the laboratory, where plants were dissected Evaluation of the diurnal sampling procedure (individuals collected per 5 min interval by each collector): descriptive and soil examined at ten magnifications to statistics of counted and real number of Succinea extract overlooked snails. costaricanain a Costa Rican farm (n=6 replications of each count) Life cycle in the laboratory: Specimens for x SO Min Max the laboratory were collected in March, 1992. Plant In the University of Costa Rica (San José City) Alive they were kept in plastic containers (diameter Counted 0.3 0.5 O I 7.0 cm; height 5.8 cm) with holes on the lid. A Total 5.2 3.1 I 9 soil layer 1 cm deep, fromthe original location, Dead Counted 0.8 1.3 O 3 covered the container' s bottom. The animals Total 3.7 2.8 I 9 were fed 2 X 4 cm pieces of lettuce twice a week. Eighteen individuals isolated since Soil their eclosion were selected to determine age Alive Counted 0.2 0.4 O at first reproduction. From these, twelve were Total 0.2 0.4 O used to study egg production. Shell length of Dead embryos with a maximum age of 24 hr was Counted 4.2 2.1 2 7 measured with a millimetric ruler and a Total 7.0 5.5 O 14 estereomicroscope. In neonates (aged 12 hr or X Mean, SO' standard deviation, Min minimum, Max maxi less) shell length was obtained at ten magnifi mum cations with a vernier caliper (0.05 mm preci sion). For establishing growth rate the shell plants and alive animals on soil because the lenght of 26 individuals was measured week sampling test count was zero, but the number ly. Illustrations are based on camera lucida alive on plants and dead on soil must be multi drawings and represent specimens which con plied by 3.7 and 2.3, respectively. tracted to sorne extent during fixation. Voucher specimens are deposited in INBio Density: The number of living S. costari (Heredia, Costa Rica, catalogue numbers cana in the farm can be estimated from the INBIOCRIOO1464008, INBIOCRIOO1 46401 O night count, which was made by plant and not and INBIOCRIOOI464012). Additional vouchers will be deposited in the Universities TABLE 2 of Costa Rica (San José) and Florida Evaluation of the nocturnal sampling procedure (Gainesville). (individuals per plant or soil at plant base): descriptive statistics of counted and real number of Succinea costaricana in a Costa Rican farm (n=6 replica RESULTS tions of each count) x SO Min Max AH descriptive statistics are presented as Plants mean ± standard deviation (minimum-maxi Alive mum). Counted 0. 6 1.3 O 6 Total 2.2 1.7 O 8 Dead Sampling procedure: Diurnal counts did Counted O O O O not differ significantly from the real number of Total l.l 1.0 O O snails on the soil (Table 1, Mann-Whitney U, Soil p>O.05) but underestimated populations on the Alive Counted O O O O plants by a factor of 17 in living animals and of Total 0.5 0. 9 O 3 4.6 in shells (Mann-Whitney.u tests, p<O.05). Oead In contrast, nocturnal counts underestimated Counted 2.4 2.3 O 8 the population in aH cases (Table 2, Mann Total 5.5 3.0 I 12 Whitney U tests, p<O.05). The amount. of x Mean, SO standard deviation, Min minimum, Max maxi underestimation is unknown for shells' on mum 184 REVISTA DEBIOLOGIA TROPICAL TABLE3 for day and night, the proportions in the plant are 81.1 % (night) and 77.4 % (day), which is NumberofSuccinea costaricana snailsper planl half accordinglo a noclurnal visualcount on 314 planls not statistically different (Chi-square p>O.05. night N= 314, day N=297). x SD Min Max Abundance: In March of 1992 there were Soil 0.242 0.558 o 5 large numbers of dead animalson plants and of Stems 0.799 1.314 O 8 Leaves 0.236 0.905 O 10 � ,-------------------------------, x Mean, SD standard deviation, Min minimum, Max maxi mum. � �------------------------------� MAR MAV JUL SEP NO\f JAN MAR ),lAy 1� 1993 100,-------------------------------, ..... R MAV JUL SEP NQV JAN .......... R V 11)92 1993 100 �-----------------------------, 90 '0 MAR MAY JUl SEP NQV JAN MAR UAY 1992 ,., 500 "AA UAY JUl SEP NOV JAN MARUAV '992 1993 400 Fig.