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1 975 ; 1973; 1975), Uetz, 1 979; Wolda Rev.Bio1.Trop., 30.(1): 27-34, 1982. Analysis of the leaf litterarthropod fauna of a lowland tropical evergreen forest site (LaSelv a, Costa Rica) Susan Lieberman and Charles F. Dock Allan Hancock Foundation, University of Southern California. University Park, Los Angeles, California 9007. (Received for publication August 13, 19.81) Abstract: Leaf litter arthropod population changes were monitored by means of extensive can trapping over a thirteen month period at a lowland evergreen forest site in Costa Rica. Samples were taken within an undisturbed forest plot and an adjacent fallow cacao plantation. Comparisons of abundance were made between nocturna! and diurna! samples as well as between months. The dominant arthropod groups in the samples in order of abundance were the Formicidae, Apterygota, Orthoptera and Arachnida. Total numbers of individuals in all major groups were consistently greater in the initial stages of the rainy season and remained relatively constant throughout the remainder of the year. It is suggested that numbers in crease in response to the onset be the rainy season. Greater numbers of individuals were collected in the undisturbed forest than the fallow cacao plantation. Siptificant diel differepces in abundance were fo und for many of the major groups inc1uded in the study. Maximum arthropod abundimce corresponded to the time of greatest leaf litter depth within the study plots. ..:. Historically, studies of tropical arthropods trap samples. Additionally , differences in ar­ have been qualitative an d descriptive in nature thropod abundance between undisturbed forest (Allen, 1926 ; Davis, 1945 ; Strickland, 1945). and disturbed forest planted in cacao are More recently, insect abundance has been analyzed. The potential relationship between studied by many workers, concentrating on the arthropod numbers and rainfall patterns is also understory and canopy (J anzen, 1973 ; Mont­ exarnined. gomery and I..ubin, 1979; Buskirk and Buskirk, 1976). Though the litter floor has begun to MATERIAL AN D METHODS receive more attention (Anderson, 1975 ; Elton, 1973; 1975), most studies have concentrated The study area was Finca La Selva, Heredia on individual taxa, in particular beetles, mites, Province, Costa Rica . This area is a lowland and spiders (Meijer, 1974; Baars, 1979; evergreen wet forest (Holdridge , 1967) at an I Mitchell, 1963; Stanton, 1979 ; Uetz and altitude of 60 m, with a mean annual rainfall of Unzicker, 1976; Uetz, 1979; Hanlon and 3969 mm. Two dry months of J anuary-Febru­ Anderson, 1972). Bigger (1976) documented ary usually average less than 200 mm rainfall, the existence of oscillations in tropical insect while the wet months of March-December populations in general, and considere d preda­ average more than 200 mm. The study was tor-prey relations to be a causative factor. carried out at two sites: one in a relatively Wolda (1977 ; 1979) has shown seasonal fluctu­ undisturbed fo rest remnant, the other in an ations in tropical insect populations that may adjacent plantation that was once planted with relate to rainfall and food abundance. cacao trees for agricultural purposes but has In this report we evaluate seasonal and diel been fallow for at least 20 years. variation in arthropod abundance of tropical The arthropods were trapped using a series forest leaf litter, based on multiple monthly pit of one gallon pit traps. The pit traps were laid 27 28 REVISTA DE BIOLOGIATROPICAL out at 10 m intervals in two transect lines, one Lepidoptera, Coleoptera, other identifiable lar­ in undisturbed forest, the other in the cacao vae , and unidentifiable larvae. The data base for plantation. f;ach trap had a top that could be the separation was too small, and no sienificant removed during trapping operations, and con­ differences between them were found. Conse­ tained pichc acid solution to fIx all specimens q�ently all larvaeare grouped together through­ at time of capture. Traps were operated at both out the paper. The miscellaneous category plots once a month for a period of 13 months, includes Ephemeroptera, Neuroptera, and from February 1974 to February 1975. Sam­ Lepidoptera. The unidentified category pling spanned a continuous four-day period in includes all arthropod individuals that could each of the 13 months. The traps were cleared not be accurately identified. every 6-10 hours day and night (mean satTlpling period: 9.02 ± 3.7 hours), the picric acid 'was RESULTS replaced, and the arthropods removed and preserved. A total of 21 ,944 individual arthropods we re In the laboratory, can trap samples were collected, of which 34.8% were ants, 22.7% evaluated. Each sample was recorded with Orthoptera, 18.0% Apterygota, 5.0% spiders, respect to numbers ofvarious arthropod groups and the remaining 19.5% was comprised of all and their respective sizes. Most arthropods were other groups combined. classifIed to order (Table 1). Ants we re sepa­ Figure 1 shows the average numbers of rated from other Hymenoptera ; arachnids were individuals collected per months for the major groups. Total monthly rainfall and rainfall during the sampling periods are also presented. Numbers for all major groups peaked in spring and subsequently declined in summer. With the exception of the Apterygota, numbers re­ mained relatively constant over the remainder � ! of the year. Apterygota exhibited a further .... decline in numbers in late fall and winter. 5 D. More individuals were trapped du ring the � i day among the Apterygota, Diptera and the Miscellaneous group, than at night. More indi­ i � viduals were trapped at night than during the day among the ants, spiders, Isopoda, other arachnids, and Chilopoda and Diplopoda. Fur­ thermore , more individuals were found in the forest plot than in the cacao for the Or­ thoptera, spiders, and Homoptera. Table 1 lists the results of multiple analyses of variance for each arthropod group's abundance. The three factors in the analyses ofvariance were months of the study, forest versus cacao site, and day versus night. The purpose of such anovas is to test if there are significant effects due to any of these factors, and to test if there are any Fig. 1 Seasonal abundances of predominant arthropods and rainfall patterns for the same interactions between the factors.The table lists periodo the probability that there is a significant effect; probability values greater than .05 are not listed divided into spiders, ]l1ites, and other arachnids. and are not considered significant. The three­ Chilopoda and Diplopodawere placed together, way multiple anova used allowed for an adjust­ as were snails, worms, and leeches (all relatively ment for unequal sample sizes. The condition uncommon). Though snails, worms ana leeches ofhomoscedasticity was met. are not arthropods, they were the only other Significant effects exist for month of trap­ invertebrates found and were thus included. ping for all arthropod groups except larvae, The larvae were initially separated into Diptera, miscellaneous, and unidentifIed (all very rare). L1EBERMAN el DOCK: Arthropodfau na, La Selva, Costa Rica 29 Except fo r the Hemiptera (p = .006), Isoptera For ants, the average abundance in May is (p = .03), and the Dennaptera (p = .007), these significantIy different from a11 other months; effects are significant at the .0001 level or March and April fonn a cluster, as do March. better. September, November, and the fo11owingJanu­ Interactions between sampling�month and ary. Sorne taxa have only two groups of months forest versus cacao occur in the following that cluster (e.g. 'spiders and mites) , while groups: Orthoptera, mites, Isopoda, Coleoptera, others evince a greater complexity, with up to Apterygota, Homoptera, Hemiptera, and five such clusters (Diptera, Apterygota). Isoptera. Interactions also occur between Figure 3 relates directly to Table 1. The month and time of day for ants, Orthoptera, abundances in the forest and cacao plots, Diptera, Isopoda, Coleoptera, Apterygota, separated into day and night collections, are Arachnida, Homoptera, Hemiptera, and tabulated. They are tabulated only for those Dermaptera. For many groups, the difference arthropods with significant effects dile to either between day-night abundances depends on the the forest-cacao factor, the day-night factor or month (and possibly the associated rainfall). both. For other groups, the diffe rences between Table 2 represents the Pearson produ ct­ abundances in the fo rest versus cacao plots also moment correlations between the numbers of depend on the month of the year. Three-way individuals of the arthropod groups. Those interactions between month, forest-cacao, and correlations significant at the .001 and .0001 day-night occur only for the Apterygota and levels are indicated. Such stríngent significance the Homoptera. levels were deemed necessary due to the large Figure 2 represents the results of Duncan's sample sizes. multiple-range a posteriori test. This test was DISCUSSION run o subsequent to the three-way multiple anovas on aH abundance variables for which Many studies have shown that pit traps (also there was a significant effect due to month. In caBed can traps or pitfall traps) are the most Figure 2, lines connect those months for which efficient way of coHecting litter arthropods and the means (average abundance of the group) are evaluating thcir abundances over time (Baars, not significantly different at the .05 level. 1979; Meijer, 1974; Mitchell, 1963; Uetz and Unzicker, 1976). Specifically,. Meijer (1974) .ITHIOII'OD MONTHS OooUl'S 1-_-. ,1,-, ,'1-.,,'1 -,_', I'�-, rI .�-"" I -"..' 'I-� " I-..--" I'M- , r1 ...-1"' 1....,- ,. ¡-.,, ', 1---1 and Denboer (1971) have founa pittraps to be .L-...L. , vtiry successful as measures of relative abundance and particularly useful in comparing populations in differe nt sites, even though trap efficiency varíes from species to species and habitat to habitat (Mitchell, 1963). Baars (1979) cautions that many traps are needed in micro-spatially heterogeneous habi­ tats. The large sample size in this study reflects this consideration. The numbers recorded in this study must be viewed comparatively; Le.
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