Seasonal Fluctuations in Rainfall, Food and Abundance of Tropical Insects Author(s): Henk Wolda Source: Journal of Animal Ecology,

Vol. 47, No. 2 (Jun., 1978), pp. 369-381 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/3789 . Accessed: 20/07/2011 11:27
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Journal of Animal Ecology (1978), 47, 369-381

SEASONAL FLUCTUATIONS IN RAINFALL, FOOD AND ABUNDANCE OF TROPICAL INSECTS

BY HENK WOLDA Smithsonian TropicalResearch Institute, P.O. Box 2072 Balboa, Canal Zone

SUMMARY (1) Light trap data, covering over 3 years, are discussed for some groups of insects in the lowland tropical monsoon forest on Barro Colorado Island. The groups concerned are Homoptera, Tettigoniidae and Mantidae. (2) The data are compared with information on rainfall and leaf production. It is shown that, especially for those groups of Homoptera that have nymphs feeding on the foliage, there is a good relation between leaf production and the demography of the species concerned. (3) In all cases the data are discussed in general terms although data per species are available. (4) In some cases anomalies in the rainfall pattern, such as an inch or so of rain in early March, the mid dry season, has a strong positive effect on the populations through increasing leaf production, in other cases it may actually have a negative effect. INTRODUCTION Several studies have provided evidence showing that tropical insects undergo seasonal changes in abundance, at least for those parts of the tropics where wet and dry seasons alternate (Davis 1945; Dobzhansky & Pavan 1950; Owen 1969; Emmel & Leck 1970; Gibbs & Leston 1970; Robinson & Robinson 1970; Fogden 1972; Owen & Chanter 1972; Janzen 1973a, b; Ricklefs 1975; Frith 1975; Janzen & Pond 1975; Bigger 1976; Buskirk & Buskirk 1976; Wolda 1977a) although not all authors agree (Elton 1975). In areas with a pronounced dry season, such as most of Panama, the abundance of most insects in the dry season is relatively low. In areas with a very mild dry season, such as Sarawak, insects decrease in abundance during the drier season and increase during the wetter season (Fogden 1972). Some spiders in New Guinea show the same seasonal pattern (Robinson, Lubin & Robinson 1974). On the other hand, Janzen (1973a, b) finds for an area with a similarly mild dry season in Costa Rica, that a sweepnet sample of insects taken in the dry season contains more insects than such a sample from the wet season, at least in secondary vegetation. It seems likely that the seasonal presence of insect species is synchronized with a seasonal presence of its food, if food availability varies seasonally. For foliage feeders this means that they should usually be present when new leaves are produced. The apparent abundance of foliage in a forest does not necessarily imply an abundance of food for herbivores, as already pointed out by Voute (1957). Plants have evolved defense mechanisms against herbivory while herbivores have evolved means to cope with some of these defenses. As a result of this coevolution, mature leaves often have toxins, are tough and, the best defense of all, have low contents of usable nutrients. Young leaves in many 0021-8790/78/0600-0369$02.00 ? 1978 Blackwell Scientific Publications 369

Rainfall,food and tropical insects cases lack such effective defenses. Their toxin levels are lower, they are less tough and have a higher nutrient content than mature leaves (Feeny 1970). This is also true for species of trees on Barro Colorado Island, the area to be discussed in the present paper (C. Milton, personal communication). Fogden (1972) finds that the seasonal cycle of caterpillars and orthopteroid insects is well correlated with that of leaf production, He also finds that parasitic Hymenoptera increase during the caterpillar season. Janzen & Schoener (1968) find in the dry season that differences between areas in the abundance of insects correlate with differences in the production of leaves and shoots. During the season when most leaves are mature, the insects may depend entirely on the few new leaves available (Feeny 1970; Rockwood 1974) and one might expect slight changes in leaf production, especially during the off-season, to have a major effect on the demography of insect populations. The data to be discussed in the present paper support this hypothesis. 370 PROCEDURES The data on insect abundance to be discussed were collected by means of two lighttraps inside the forest on Barro Colorado Island (BCI), Panama Canal Zone. One trap was in the canopy, the other some 2 m above ground level. They were set up by N. Smythe as part of the Smithsonian Environmental Science Program (ESP) and have been in operation since June 1973, each producing one sample of insects per day. As I have discussed before (Wolda 1977b), there are good reasons to believe that, at least for many species, the catches in the lighttraps do provide reliable information on real changes in population size for that part of that population that flies, i.e. the adults. The insects to be discussed here are mostly Homoptera, all of them sapsuckers, but variable in where they get the sap from, especially in the nymphal stages. Other groups to be mentioned are katydids (Tettigoniidae, mostly leaf eaters) and mantids (predators). In all these cases the data are available per species, but in the present paper I will concentrate on higher taxa. As a result of variable weather conditions, the numbers caught vary from day to day and because of the depressing effect of the full moon (cf. Wolda 1977a) there is a variation with a period of about 4 weeks which reflects changes in trappability of the insects rather than real changes in the populations. For these reasons the data are presented per period of 4 weeks, of which there are thirteen per year. Rainfall data are from tables produced by the Meteorological and Hydrographical Branch of the Panama Canal Company, which operates a rain gauge on BCI, and from unpublished data of Smythe and Windsor. Leaf flush data are available since August 1974. The sampling programme was set up by R. Foster, as part of the ESP, and the actual sampling was done by Bonifacio de Leon. Because of trees dying and others being added to the programme, the actual number of trees included in the census varied somewhat. At the moment there are 372 individual trees being sampled every week, excluding the palms, belonging to 182 species. This is not a random sample since common species are underrepresented while rare species are overrepresented. Nevertheless, it is felt that the data do give a fairly good impression of leaf flush in the forest in general. Leaf flush was scored in each tree as 'complete', i.e. the entire tree was producing new leaves, as 'partial' with new leaves on only some branches, or as 'absent'. The presence of many lianas and epiphytes makes scoring often very difficult and the 'partial' category especially should be treated with caution. In the present paper I will only be using the number of trees scored as in complete flush, expressed as a percentage of all trees sampled.

HENK WOLDA

371

An impression the agingof leavescan be obtainedby comparingthe productionof of new leaveswithlitterfall.The data on litterfallwerealso obtainedthroughthe ESP. RESULTS Beforediscussing results,a wordof cautionis in order.The conclusionsto be given any betweenvariables observed theforestfor a periodof almost4 in arebasedon correlations yearsor, as is the case for leaf flush,for not even 3 years.It will be shownthat thereare betweentheseyearsandan attemptwillbe madeto explainsuchdifferences differences in to the insectsby referring corresponding differences othervariables.Sucha procedure in canbe rathertrickywithso fewdatapoints.However,as willbe shown,thecorrelations to be discussed makesense.Theconclusionsto be givenshouldbe considered do hypotheses to be testedwith the data to be collectedin the next few years.
Rainfall, leaf production

in The relevantdata are presented Fig. 1. Almost all of the annualrainfallfalls from to Decemberwith only occasionalshowersduringthe dry season. Good mid-dry May seasonshowersoccurredin week 10, in earlyMarch,both in 1974and 1975.No rain to good showersoccurred duringthedry speakof fell duringthe 1976dryseasonand several season of 1977.The 1976rainy season startedearly,but is the drieston record.It also endedearly(in week47, late November)and the followingdry seasonlasteduntil after week 19 (mid May). Leafproduction seemsto be ratherstronglyrelatedwithrainfall.The 'normal'picture for a forestsuchas thaton BCIseemsto be a majorleafflushat thebeginning therainy of
30 E

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1973 1974 1975

,
1976 1977

FIG. 1. Rainfall, leaf fall and leaf production in the forest on Barro Colorado Island.

Rainfall,food and tropical insects season in May-June, with very little production during the later part of the rainy season and during the dry season. This is the general impression obtained in previous years and very similar to what Frankie, Baker & Opler (1974) report for a climatologically similar, although slightly drier, site in Costa Rica. Old leaves fall mainly during the dry season, especially in January and February. As can be seen in Fig. 1, apart from the major peak in leaf flush in May-June, there are minor peaks in October and in January. The actual picture deviates in some important aspects from this generalized 'normal' year. The rain at week 10 in 1974 was followed by many trees and shrubs producing a flush of new leaves. Unfortunately I have only casual notes about this as the sampling programme had not started yet. The mid dry season rain of 1975 and the first rains of the rainy seasons of 1975 and 1976 were followed by an increase in leaf production. This increase starts less than a week after the rain and reaches a peak some 2 to 3 weeks after the rain. It is interesting to see that the rains of January and February 1977 did not have an appreciable effect on leaf production. It seems that a completely dry period of some length is required for a good shower to have any effect. It is conceivable that the subjective appreciation of what constitutes a 'complete' and what a 'partial' leaf flush changes with time. One might think that the large difference between 1975 and 1976 could be explained by a change in scoring and would thus be an artifact. I do not think this is so, at least it cannot be the entire explanation. First of all, the same observer collected all the data and he is an experienced woodsman. Secondly, even if one takes complete and partial flushes together, 1976 does not even reach half the level of 1975. In fact, the partial and complete flushes scored in 1976 combined reach a considerably lower level than the complete scores alone in 1975. Thirdly, the leaves produced in 1976 fell in the littertraps in early 1977 and litterfall then was quite a bit less than during the dry season of 1976. Undoubtedly, the difference between the two years is, at least partly, real. The minor peaks in leaf flush in October 1974 and January 1975 are much higher than the corresponding peaks in the other years, for some reason. Production in late Augustearly September 1974 was extremely low. Leaf production then went up sharply, correlated with, and may be as a result of, extensive rains after a drier period. The next, narrow, peak in leaf flush in week 47 (late November) also coincides with major rains after some weeks with little rain. It thus seems that relatively minor variations in rainfall can cause quite important variations in leaf production and, therefore, in food for herbivores. In so far as the availability of food for a particular herbivore is correlated with the presence and/or quantity of young leaves in general, one would predict that the numbers would increase some 2-3 weeks after the mid dry season rains in 1974, i.e. there should be many more in period 4 than in the previous three periods. One would also expect such insects to be more abundant during the late wet season of 1974 and the dry season of 1975, than they are during the similar periods in other years. The expected effect of the rains in mid dry season in 1975 is difficult to formulate. If above a certain minimum level of leaf flush the numbers are no longer governed by the actual amount of new leaves present, which does not seem unlikely, the rains in March 1975 may not have any effect at all. These expectations hold especially for those species which depend both as nymphs and adults on new leaves for food. For many fulgoroids this is not true. For these, such as the Derbidae, where the nymphs feed on fungi in rotting wood, the effect of the rain may be quite different. These species seem to be near-absent in the dry season. Those species that use rain as a clue to emerge from diapause or to start reproducing, might find the situation 372

HENK WOLDA

373

in March quite unsuitable for reproduction as there is little good food for the nymphs. For such species the rains in the dry season might have a negative effect on their abundance. Homoptera For several major groups within the Homoptera, the number of individuals caught per period of 4 weeks is plotted in Figs 2 and 3. Some relevant statistics are given in Tables 1, 2 and 3. These are the mean number of individuals per week for the first part of the dry season (Table 1), for the end of the dry season (Table 2) and for the early rainy season (Table 3). The statistics are, in a sense, questionable. In order to get sufficient degrees of freedom to make the standard errors meaningful, I had to take mean numbers per week rather than per 4 week period. This means that the variances not only include random error, but also systematic errors such as effects of the lunar cycle. Any real changes in numbers during the period in question also show as an increased variance, making the demonstration of significant differences between means more difficult, however, it also means that one may confidently accept differences that turn out to be significant statistically in spite of this. 1. TABLE Meannumberof individuals weekfor the first 12 weeksof eachyear, per withstandard error,for somegroupsof Homoptera caughtin thelighttrapson Barro ColoradoIsland
1974 1 33+ 1 96 Cicadoidea 2.00+0-39 Membracoidea 0.50+0 15 Cercopoidea 7.50+ 196 Psylloidea 1175+ 2.06 Cicadellidae 2-08+058 Delphacidae (Fulgoroidea Delph.) 23-67+5.64 1142+ 2-65 Achilidae 4.83+ 148 Cixiidae 2-50+085 Derbidae 0 Tropiduchidae 3.33+ 120 Flatidae 0-42+0-15 Dictyopharidae 1975 1 42 + 0-53 483+1 04 183 +052 64-42+26-29 4542+ 8.84 3.67+1.51 9308+ 10-84 2608 + 377 1650+2-25 2-83+ 032 1.42+ 048 32-33+5 55 6.58+ 159 1976 1977 1 25 + 0.43 3.50+ 1 79 1 50+0-67 1.83+0-34 0-92+0.23 2.42+ 093 1 58?0.58 3.17+0.88 13-67+181 ** 4-33? 128 0.58+0.23 3.17+ 161 5025 + 593 ** 20-17+835 6.17+ 2.72 12.- +209 15-25+1.83 *** 1.50+0.72 5.25+ 187 * 0-50+ 023 0.83+ 0.30 0.50+ 029 8.58+365 13-58+240 1-58+0-92 2.17+0.47

* * * ** *** ** *** ** *** ***

* * ** ** **

** *

between successive differences by yearsareindicated *(P<0-05), ** (P<0-01) or *** Signifcant (P<0 001).

Before discussing the statistics I will, briefly, mention some salient points of Figs 2 and 3. The cicadas are highly seasonal. There are three common dry season species, all of which are rare in the traps. Of the nine rainy season species some are found for the first time in April, others in May. Some disappear in August, others stay on until October. After that no cicadas are caught, or heard, until the beginning of the dry season. The nymphs are subterranean and could, conceivably, be affected by oddities in the rainfall pattern either through changes in soil moisture or through changes in activity of the plant roots on which they feed. The thirteen species of spittlebugs (Cercopoidea) that were caught, four of them in large or very large numbers, are all rainy season species with only some individuals extending into the dry season. They usually do not appear in the trap until early June, well after the rains have started in early May. Several of these species, especially the common ones, have subterranean nymphs. The Psyllids have two peaks in abundance, one late in the rainy season and one early in

374
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Rainfall,food and tropical insects

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FIG.2. Seasonal distribution, in number of individuals per 4 weeks, of four groups of Homoptera in a set of two light traps on BCI.

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FIG. 3. Seasonal distribution, in number of individuals per 4 weeks of three groups of Homoptera

in a set of two light traps in BCI.

HENK WOLDA

375

thedryseason.Of thenineteenspeciesscored,fourpeakin thedryseason.Thenymphsof psyllidsaremostlyout on the foliageor in gallson thefoliageandone mightexpectthem of to be rathersensitiveto changesin the availability new leaves. Thereare severalspeciesof membracids involved,all of whichshouldbe classifiedas rainyseasonspecies,althoughsome also occurduringpartof the dry season.They,and theirnymphs,feedon leavesandon new shootsand one mightexpectthemto respondto changesin the seasonalpatternof productionof new leavesand shoots. The delphacids wereexcludedfromthe total fulgoroidsbecausemost of the speciesfly of of The individuals somedelphacid speciesin veryerratically. occurrence macropterous to the trapis almostcomparable that of alatesof ants caughtduringtheirmatingflight. On a particular days, manymay be caught,whileduringthe day, or on a few successive nextfew weeksone maynot get anyat all. However,theyareall rainyseasonspecies,with a majorpeakat the beginningof the rainyseason(Fig. 3). Therestof thefulgoroidsand thecicadellids relatively in abundance are low duringthe aroundJulyandAugust.Nymphsof cicadellids are dryseasonandreachtheirmaximum usuallyout on the foliage,but possiblymainlyon shrubsand herbs,plantsthatwerenot in included thecensusfor leafproduction. Fulgoroidnymphshavevariousfeedinghabits, whichI will mentionbelow. It is expectedthat, in 1974,the numberof individuals mightbe higherin weeks 13-16 (period 4) than in weeks 1-12 (periods 1-3), because of the effects of the rains that occurredduringearly Marchof that year. This is confirmedby the analysisfor several cicadellids,fulgoroidsand even the groups of Homoptera(Table2). For membracids, the of aftertherain, delphacids, meannumber individuals weekincreased per significantly in spite of the dry weatherduring that period, presumablybecause of the increased of the but availability (undocumented) food. In thecercopids changeis a decrease, I doubt that this small difference any real meaning,in spite of the statistics.In the cicadas has therewas an overseven-fold but The is increase, thiswas not significant. increase entirely dueto a newspeciesof Taphura whichappeared earlierin 1974thanin otheryears.On the psyllids the rain and correspondingleaf flush had no effect, presumablybecause it occurredtowardsthe end of theirseasonalpeak.
TABLE2.

Mean numberof individuals week, with standarderror,for the per fourthperiod,weeks13-16,of eachyear,for somegroupsof Homoptera caught in the light trapson BarroColoradoIsland
1974 Cicadoidea
Membracoidea Cercopoidea Psylloidea Cicadellidae Delphacidae Fulgoroidea (-Delph.) Achilidae Cixiidae Derbidae

1975 2.50+ 0.96

8.75+ 1.70 0 4.50+ 1-89* 21 75+ 3-77* 0.50+ 0.50 8275 + 21-05 34-75+ 9.66 27-25+11-26 1.75+ 0.85

1976 0.50+0.20
6.75 +2.56 0.25 +0.25 4.50+1.19 14-75+4-59 3-50+ 104 50-25 + 686 9.50+1.04 16.0 +2-31 0-50+0.50*

8.33+ 4.54
12.75+ 1.97*** ** 0 7.75+ 2-93 44.0 + 8-22** 5-25+ 1.11* 84-25 + 2135* 54.0 +11.29** 20-50+ 4.57** 3.50+ 1.66

Tropiduchidae
Flatidae Dictyopharidae

0.50+ 0.29
25-50+ 9.99* 3.0 + 2.04

0.50+ 0.50
25-25+ 5.81 4.75+ 1.80

1.0 +0-41
20.0 +4-56 2.25+0 85

Significant differences with the previous 12 weeks (Table 1) are indicated after each number with *(P<0-05), **(P<001) or ***(P<0001).

376
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1976 1977 1974 1975 1973 FIG.4. Seasonal distribution, in number of individuals per 4 weeks, of three families of Fulgoroidea in a set of two light traps on BCI.

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FIG.5. Seasonal distribution, in number of individuals per 4 weeks, of three families of Fulgoroidea in a set of two light traps on BCI.

HENK WOLDA

377

For the families within the fulgoroids the results vary. The increase is significant only in the Achilidae, Cixiidae and Flatidae (Figs 4 and 5). Most, if not all, species of derbids have nymphs that feed on fungi in rotting wood and one would not expect them to respond to a flush of new leaves or a temporary moistening of logs. Many achilids and cixiids may have similar feeding habits although some nymphs may be out on the foliage. In actual fact, only some species showed a clearcut relation with the rains in early March 1974. In the Cixiidae it is only two common species that show this response, viz. Pintalia erecta Metc. and Muirolonia metallica Fowl. Among the Achilidae the response is only clear in members of the genus Catonia and a few others. In most species of the families that do have their nymphs feeding on young leaves, viz. Tropiduchidae, Flatidae and Dictyopharidae, the response is there. It is highly significant in the Flatidae, and it is a six-fold, albeit nonsignificant, increase in the Dictyopharidae. The Tropiduchidae are not common enough in the trap to expect any significant results. TABLE Meannumberof individuals week,with standard 3. error,for periods5 per caughtin through9 (weeks17 to 36), for eachyear,for some groupsof Homoptera the light trapson BarroColoradoIsland
1974 1975 1976 26-25+ 6.08 + 19-70 3.52 23-85+ 6.65 7.50+ 3.33 141-25 24-92 + 45-35+ 20-87 25310 + 36-08 81 20+ 11 95 77-50+ 11-86 29-05+ 5-40 1.0 + 0.41 42-40+ 6-57 14-05+ 3.05 or **(P< 001).

31-65+ 9.63 Cicadoidea 53.0 + 1097 13-85+ 2.43 13-95+ 1.92 Membracoidea + 14-20 4.46 28-70+ 7.84 Cercopoidea 3.40+ 0.74 3.45+ 0.93 Psylloidea + 140-95 27.0 201 79+ 17-33 Cicadellidea 85-42+ 22-50 99-30+ 57-80 Delphacidae 15690 + 24-61** 322-10+4203 Fulgoroidea (-Delphac.) 60-05+ 9.93* 116-50+17-86 Achilidae 39-75+ 8.57* 10560 + 23-16 Cixiidae 7.0 + 1.73* 14-45+ 2.45 Derbidae 0.65+ 0.23** 2.25+ 0.46* Tropiduchidae 48-80+ 5.85 36-75+ 5.81 Flatidae 14-45+ 3.46 7.55+ 1.57 Dictyopharidae between consecutive differences yearsareindicated *(P< 005) by Significant

It is also expected, because of the relatively high level of leaf production, that the number of individuals during the early part of the dry season should be much higher in 1975 than in the other years. As shown in Table 1, the numbers are significantly higher in 1975 than in 1974 during the first three periods. It is not found in cicadas, in the erratic delphacids and in the derbids, whose nymphs do not feed on young leaves. In the achilids and cixiids only some species, the same as the ones mentioned above, are responsible for the relationship shown. In the cercopids the increase is significant, but I doubt that it has anything to do with the leaf flush. In the three families with nymphs on the foliage (Fig. 5), the increase is spectacular. In fact, the mean number of individuals per week during the first 12 weeks of 1975 was about the same as during the maximum in the preceeding rainy season (Table 3). It seems that the increased production of new leaves was sufficient to prevent a decrease in abundance during the dry season, whatever the weather. In early 1976 the numbers are lower again in many groups, as one would expect with the lower level of leaf production then. However, there are a few notes of caution here, showing that one should not expect one environmental factor to determine changes in numbers all by itself, without any help from other factors. For one thing, the decrease did not occur in the Cixiidae. Partly this is due to other species being involved, but this is certainly not the

Rainfall, food and tropical insects entire story. The numbers present during some part of the year are probably a function of the circumstances during that period and of the numbers present at the beginning of that period. During the peak of the rainy season in 1975 (Table 3) the number of cixiids was much higher than in 1974 and a spill-over of individuals into the dry season may have caused the higher numbers during early 1976. This may also be the explanation for the relatively high levels during that period of flatids and dictyopharids. The rain during early March 1975 in no instance had a significant effect on the numbers. The only two significant cases are decreases (Table 2), in psyllids, indicating the decrease at the end of their season, and in cicadellids. It is conceivable that species which, in a given year, can start building up their population at a relatively early date, reach higher levels of abundance later in the year than they do in other years. One might expect, on this basis, that in 1974 numbers of those groups which responded positively to the early March rains, to be higher than in 1973. For early 1973 there are no insect data, but weather data suggest no reason why insects should h ve had an early start then, as they had in 1974. Here only the period of week 23 to 36 can be compared, as data for period 17 to 36 (Table 3) are not available for 1973. The only significant differences found are in the Flatidae (from 17-5+ 2-6 in 1973 to 30-5 + 4-5 in 1974; P<0-05) and in the Derbidae (from 291 +6-5 to 12.2+3-3; P<005). In the Dictopharidae there was also an increase, from 6-4 to 8-2 per week) but this was not statistically significant. In 1975 conditions, again, were favourable early in the year and numbers increased again, significantly so in several groups (Table 3). In the Dictyopharidae the increase was not significant, but the level reached in 1975 was significantly higher than that in 1973. It thus seems that, at least sometimes, an early start of the season may have a favourable effect on the numbers reached later on. The Derbidae merit special discussion. They decreased from 1973 to 1974 and increased after that. As explained above, they are not expected to respond to the production of new leaves. However, they may respond to rain by emerging from their hiding places in order to reproduce. If dry weather reproduction is unsuccessful, mid dry season rains could have a negative effect on the numbers. The data support this hypothesis. From 1973 to 1974 numbers decreased significantly (see above), presumably because of the early March rains in 1974. In 1975 those rains again occurred, but were less intense, and may not have had such a strong effect, and numbers increased somewhat. In 1976 dry season rains did not occur and the number of derbids reached a high level, about the same as that in 1973. 378 Other groups of insects Data are available, or are becoming available, for a number of other groups of insects. Of these I will only briefly discuss here the Tettigoniidae (katydids), as representatives of leaf eating herbivores, and mantids as a group of predators. The data are plotted in Fig. 6. Among the species of katydids caught only two are predatory and represented only with a total of four individuals. All the others are, probably, mostly folivorous. As Fig. 6 shows, the effect of the mid dry season rains, if any, was negative. An explanation will have to wait until one knows more about the biology of the species involved. One might expect a positive response to increased leaf production during certain times of the year, but this was clearly not the case. Mantids are predators and their relationship with leaf production, if any, is expecte-' to be only indirect through increased abundance of their prey. I do not know enough about their diet to be able to measure directly the abundance of their prey items. However, data on general flying insect abundance are available from a set of three traps at a location

HENK WOLDA

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calledthe 'weir',at some200m fromthe trapsdiscussed thispaper(Wolda1977a).The in weirtrapsproducethe total dryweightof insectssmallerthan 5 mm and the numbers of insectslargerthan5 mm.Thelattercategorymayincludethemorecommonpreyitemsof themantids.Theseasonof themantidsapparently fromperiod5 to 10,i.e. fromweek17 is to 40. For thisperiodthe dataon generalinsectabundance, insectsflyingat nightand i.e. in withthetotalnumber mantidscaught of captured thetraps,is givenin Table4, together of with the duringthat period.Thesedata suggestthat the abundance mantidscorrelates abundanceof their prey. However, the suggestedcorrelation,which may very well a couldbe spurious. Thereareonly threedatapointsand the represent realphenomenon, measureusedfor preyabundance at best, not very good. is,
400-

o 200-

-I
-

200-

1973 1974 1975 1976 1977 FIG.6. Seasonal distribution, in number of individuals per 4 weeks, of two groups of Orthoptera in a set of two light traps on BCI. TABLE4. Relationbetween insectabundance, measured lighttraps,and as general by

the abundance Mantoidea,also measured light traps,duringthe periodfrom of by week 17 to week40, whenmantidshad theirpeakin abundance
Year 1974
1975 1976

Dry weightof insects smaller than5 mm(g) 16-98

16-04 39-01

Numberof insects largerthan5 mm 3647

2368 5244

Numberof mantids 338

182 422

DISCUSSION It has been shown that some groupsof insectsmay be ratherstrictlycontrolledin their of seasonalityand theirabundance the availability food in a tropicalforest,and that by even smallaberrations the rainfallpatternmay havea strongeffecton the availability in of food, its seasonalityand quantity.It has becomeobvious that the generalquestion aboutseasonalchangesin theabundance insectsin a tropicalforesthas to be rephrased of and mademore precise.Questionsabout distribution the forest,kinds of insectsand in differencesbetweenyears have to be clarified.Data from only one year can be very as for misleading, is readilydemonstrated someinsectgroupsas in Fig. 5. Takingtheyear

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from June 1973 to June 1974, one finds a pronounced decrease in numbers throughout the dry season. Taking the year from June 1974 to June 1975, one concludes that these insects are virtually non-seasonal in their abundance. Comparing the dry season with the rainy season by taking one sample in each, can be very misleading. A sample in early April can have more insects than those from October if the April sample is collected after a mid dry season rain and the October sample is late in the rainy season with little or no new leaves produced. The changes during a season are as important as differences between seasons. Janzen (1973b) suggests that sapsucking insects may be more sensitive to changes in availability of young leaves and shoots than folivores. The present data show that here too one has to be more precise. For some sapsuckers he is right, for others he is not. Whether all folivores are as the tettigoniids discussed in the present paper I do not know until data on such other groups become available. ACKNOWLEDGMENTS Thanks are due to all those who contributed in collecting data for the Environmental Science Program of the Smithsonian Institution, or who helped setting it up, such as Nick Smythe, Robin Foster, Don Windsor and, especially, since he did most of the actual collecting of data on leaf production and litter fall, and since he operated the light traps, to Bonifacio de Leon. I gratefully acknowledge the help in sorting the insects given by Miguel Estribi and Saturnino Martinez. REFERENCES
Bigger, M. (1976). Oscillations of tropical insect populations. Nature, London, 259, 207-9. Buskirk, R. E. & Buskirk, W. H. (1976). Changes in arthropod abundance in a highland Costa Rican forest. American Midland Naturalist, 95, 288-98. Davis, D. E. (1945). The annual cycle of plants, mosquitos, birds, and mammals in two Brazilian forests. Ecological Monographs, 15, 243-95. Dobzhansky, T. & Pavan, C. (1950). Local and seasonal variations in relative frequencies of species of Drosophila in Brazil. Journal of Animal Ecology, 19, 1-14. Elton, Ch.S. (1975). Conservation and the low population density of invertebrates inside neotropical rainforest. Biological Conservation,7, 3-15. Emmel, Th. C. & Leck, Ch.F. (1970). Seasonal changes in the organization of tropical rain forest butterfly populations in Panama. Journal of Research on the Lepidoptera,8, 133-52 Feeny, P. (1970). Seasonal changes in oak leaf tannins and nutrients as a cause of spring feeding by winter moth caterpillars. Ecology, 51, 565-81 Fogden, M.P.L. (1972). The seasonality and population dynamics of equatorial forest birds in Sarawak. Ibis, 114, 307-43. Frankie, G. W., Baker, H. O. & Opler, P. A. (1974). Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. Journal of Ecology, 62, 881-913. Frith, D. W. (1975). A preliminary study of insect abundance on West Island, Aldabra Atoll, Indian Ocean. Transactionsof the Royal Entomological Society of London, 127, 209-26. Gibbs, D. G. & Leston, D. (1970). Insect phenology in a forest cocoa-farm locality in West Africa. Journal of Applied Ecology, 7, 519-48. Janzen, D. H. (1973a). Sweep samples of tropical foliage insects: Description of study sites, with data on species abundances and size distributions. Ecology, 54, 659-86. Janzen, D. H. (1973b). Sweep samples of tropical foliage insects: Effects of seasons, vegetation types, elevation, time of day and insularity. Ecology, 54, 687-708. Janzen, D. H. & Pond, C. M. (1975). A comparison, by sweep sampling, of the arthropod fauna of secondary vegetation in Michigan, England and Costa Rica. Transactions of the Royal Entomological Society of London, 127, 33-50. Janzen, D. H. & Schoener, Th. W. (1968). Differences in insect abundance and diversity between wetter and drier sites during a tropical dry season. Ecology, 49, 96-110. Owen, D. F. (1969). Species diversity and seasonal abundance in tropical Sphingidae (Lepidoptera). Proceedings of the Royal Entomological Society of London, (A) 44, 10-12.

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Owen, D. F. & Chanter, D. 0. (1972). Species diversity and seasonal abundance in Charaxes butterflies (Nymphalidae). Journal of Entomology, (A) 46, 135-43. Ricklefs, R. E. (1975). Seasonal occurrence of night-flying insects on Barro Colorado Island, Panama Canal Zone. Journal of the New York Entomological Society, 83, 19-32. Robinson, M. H. & Robinson, B. (1970). Prey caught by a sample population of the spider Argiope argentata (Araneae: Araneidae) in Panama: a year's census data. Zoological Journal Linnean Society, 49, 345-57. Robinson, M. H., Lubin, Y. D. & Robinson B. (1974). Phenology, natural history and species diversity of web-building spiders on three transects at Wau, New Guinea. Pacific Insects, 16, 117-63. Rockwood, L. L. (1974). Seasonal changes in the susceptibility of Crescentia alata leaves to the flea beetle Oedionychussp. Ecology, 55, 142-8. Voute, A. D. (1957). Regulierung der Bevolkerungsdichte von schadlichen Insekten auf geringer H6he durch die Nahrpflanze (Myelophilus piniperda L., Retinia buoliana Schff. Diprion sertifer Geoffr.) Zeitschrift fur angewandteEntomologie, 41, 172-8. Wolda, H. (1977a). Fluctuations in abundance of some Homoptera in a neotropical forest. Proceedings of the third Symposiumof Tropical Ecology, (Lubumbashi). (in press). Wolda, H. (1977b). Fluctuations in abundance of tropical insects. American Naturalist (in press).

(Received 22 March 1977)