BIONOMICS AND POPULATION GROWTH STATISTICS OF CYRTOMENUS BERGI
ECOLOGY, BEHAVIOR AND BIONOMICS An Annotated List of … · 2006-01-11 · Setembro, 2000 An. Soc....
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An. Soc. Entomol. Brasil 29(3) 433 Setembro, 2000 ECOLOGY, BEHAVIOR AND BIONOMICS An Annotated List of Insect Herbivores Foraging on the Seedlings of Five Forest Trees in Guyana YVES BASSET 1 AND ELROY CHARLES 2, 3 1 Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Ancon, Panama. Email [email protected] 2 CABI Bioscience: Environment, Ascot, U.K. 3 Present addresses: Department of Natural Sciences, University of Guyana, Guyana and Tropenbos-Guyana Programme, 12E Garnett St, Campbelville, Georgetown, Guyana. Email [email protected] An. Soc. Entomol. Brasil 29(3): 433-452 (2000) Lista Comentada de Insetos Herbívoros Encontrados em Plântulas de Cinco Espécies Florestais na Guiana RESUMO - Uma lista de insetos herbívoros de vida livre colecionados em plantas de cinco espécies de importância econômica, perto de Mabura Hill, Guiana é apresentada. As plantas hospedeiras incluem as seguintes espécies: Chlorocardium rodiei (Scomb) (Lauraceae), Mora gonggrijpii (Kleinh) Sandw. (Caesalpinaceae), Eperua rubiginosa Miq. (Caesalpinaceae), Pentaclethra macroloba (Willd.) Kuntze (Leguminosae) and Catostemma fragrans Benth. Bombacaceae). Cerca de 10.000 plântulas foram observadas com intervalos de um mês durante dois anos. Durante este período, 27.735 insetos pertencentes a 604 espécies foram colecionados. Os insetos mastigadores de folhas foram testados no laboratório, com a finalidade de identificar as espécies não herbívoras e as ocasionais. Os insetos sugadores mais comuns e abundantes são: Psyllidae, Cicadellidae, Derbidae, Membracidae, Achilidae; Galerucinae, Eumolpinae, Alticinae, Cryptocephalinae, Gelechiidae e Entiminae (insetos mastigadores de folhas). As espécies mais comuns (n ≥ 22 indivíduos) são generalistas. Ao início do segundo ano de coleta, metade das árvores mães (“estações”, n=125) foram selecionadas para ser cortadas, imitando o processo madeireiro. Apresenta- se um teste preliminar comparativo da abundância das espécies mais freqüentes durante os dois anos de coleta, assim como entre as estações cortadas e as não cortadas. Os resultados sugerem que perturbações leves podem aumentam a abundância de apenas algumas espécies. Estes padrões não parecem ser similares para espécies congenéricas. Esta investigação visa fornecer informações para o parco estudo da entomofauna herbívora em plântulas numa floresta húmida tropical. PALAVRAS-CHAVE: Insecta, Catostemma, Chlorocardium, Eperua, floresta húmida. ABSTRACT - An annotated list of the free-living insect herbivores collected
Transcript of ECOLOGY, BEHAVIOR AND BIONOMICS An Annotated List of … · 2006-01-11 · Setembro, 2000 An. Soc....
An. Soc. Entomol. Brasil 29(3) 433Setembro, 2000
ECOLOGY, BEHAVIOR AND BIONOMICS
An Annotated List of Insect Herbivores Foraging on theSeedlings of Five Forest Trees in Guyana
YVES BASSET1 AND ELROY CHARLES2, 3
1Smithsonian Tropical Research Institute, Apartado 2072, Balboa,Ancon, Panama. Email [email protected]
2 CABI Bioscience: Environment, Ascot, U.K.3Present addresses: Department of Natural Sciences, University of Guyana,
Guyana and Tropenbos-Guyana Programme, 12E Garnett St,Campbelville, Georgetown, Guyana. Email [email protected]
An. Soc. Entomol. Brasil 29(3): 433-452 (2000)
Lista Comentada de Insetos Herbívoros Encontrados em Plântulasde Cinco Espécies Florestais na Guiana
RESUMO - Uma lista de insetos herbívoros de vida livre colecionados em plantasde cinco espécies de importância econômica, perto de Mabura Hill, Guiana éapresentada. As plantas hospedeiras incluem as seguintes espécies:Chlorocardium rodiei (Scomb) (Lauraceae), Mora gonggrijpii (Kleinh) Sandw.(Caesalpinaceae), Eperua rubiginosa Miq. (Caesalpinaceae), Pentaclethramacroloba (Willd.) Kuntze (Leguminosae) and Catostemma fragrans Benth.Bombacaceae). Cerca de 10.000 plântulas foram observadas com intervalos deum mês durante dois anos. Durante este período, 27.735 insetos pertencentes a604 espécies foram colecionados. Os insetos mastigadores de folhas foramtestados no laboratório, com a finalidade de identificar as espécies não herbívorase as ocasionais. Os insetos sugadores mais comuns e abundantes são: Psyllidae,Cicadellidae, Derbidae, Membracidae, Achilidae; Galerucinae, Eumolpinae,Alticinae, Cryptocephalinae, Gelechiidae e Entiminae (insetos mastigadores defolhas). As espécies mais comuns (n ≥ 22 indivíduos) são generalistas. Aoinício do segundo ano de coleta, metade das árvores mães (“estações”, n=125)foram selecionadas para ser cortadas, imitando o processo madeireiro. Apresenta-se um teste preliminar comparativo da abundância das espécies mais freqüentesdurante os dois anos de coleta, assim como entre as estações cortadas e as nãocortadas. Os resultados sugerem que perturbações leves podem aumentam aabundância de apenas algumas espécies. Estes padrões não parecem ser similarespara espécies congenéricas. Esta investigação visa fornecer informações parao parco estudo da entomofauna herbívora em plântulas numa floresta húmidatropical.
PALAVRAS-CHAVE: Insecta, Catostemma, Chlorocardium, Eperua, florestahúmida.
ABSTRACT - An annotated list of the free-living insect herbivores collected
434 Basset & Charles
on the seedlings of five rainforest tree species of economic importance nearMabura Hill, Guyana, is presented. The host plants were Chlorocardium rodiei(Scomb.) (Lauraceae), Mora gonggrijpii (Kleinh.) Sandw. (Caesalpiniaceae),Eperua rubiginosa Miq. (Caesalpiniaceae), Pentaclethra macroloba (Willd.)Kuntze (Leguminosae,) and Catostemma fragrans Benth. (Bombacaceae). Dur-ing the monitoring of approximately 10,000 seedlings at monthly intervals dur-ing two years, 27,735 insect individuals were collected representing 604 spe-cies. Leaf-chewing insects were further tested in captivity, to remove transientand non-feeding species. The most common higher taxa included Psyllidae,Cicadellinae, Derbidae, Membracidae and Achilidae for sap-sucking insects andGalerucinae, Eumolpinae, Alticinae, Cryptocephalinae, Gelechiidae andEntiminae for leaf-chewing insects. Most of the common species collected (n ≥22 individuals) were generalists. At the onset of the second collecting year, halfof the parent trees (“stations”, n = 125) were felled to mimic selective logging.Preliminary tests comparing the abundance of the most common species duringthe successive collecting years and at the non-felled vs. felled stations are alsopresented. These tests suggest that moderate levels of disturbance increased theabundance of a few species only and these patterns were not necessarily similarfor congeneric species. The present survey represents one of the few studies ofinsect herbivores on seedlings in tropical rain forests.
KEY WORDS: Insecta, Catostemma, Chlorocardium, Eperua, logging, rainforest.
Despite much theoretical attention relatedto tree regeneration and maintenance of localtree diversity in rain forests (Janzen 1970),insect communities that feed on seedlings arenot well-known in the tropics. Most studieshave concentrated on the damage and mor-tality sustained by the seedlings (Becker 1983,Clark & Clark 1985, de la Cruz & Dirzo1987), more rarely on a few common insectspecies attacking the seedlings (Folgarait etal. 1995, Gombauld 1996). However, studiesquantifying the whole community of herbiv-ore insect foraging and feeding on seedlingswith adequate sampling effort in the tropicsare practically lacking, with the notable ex-ception of New (1983) studying Acacia seed-lings in Australia.
Typically, insects subsist at low densitieson seedlings (Becker 1993, Basset 1999).Thus, one of the main problems facing ento-mologists may be that surveying adequatenumbers of seedlings for prolonged periods
of time represents a near-impossible task for asingle researcher, particularly if seedlingpatches are rather scattered in the forest. Asolution to this problem is to train and workwith insect parataxonomists (Janzen 1992,Novotny et al. 1997, Basset et al. - in press).This study reports on insects collected onseedlings of five species of rainforest timbertrees near Mabura Hill, central Guyana, withsuch a team of parataxonomists.
As in other large-scale surveys of tropicalinsects (see discussion in, Erwin 1995), theresulting species list that we were able to com-pile with the help of taxonomist colleagues isfrustratingly simple and includes many uni-dentified and undescribed species. However,we are motivated by the paucity of data oncommunities of insect herbivores foraging onseedlings in tropical rain forests and by thereassuring thought that the material has beendeposited in a safe repository and is availablefor further examination.
An. Soc. Entomol. Brasil 29(3) 435Setembro, 2000
Insect sampling was part of a study re-porting on the effects of selective logging oncommunities of insect herbivores at MaburaHill. The present species list is augmented bypreliminary tests of annual variability and ofthe impact of canopy opening for the mostcommon insect species collected. Moredetailed analyses accounting for the effectsof rainfall, leaf production and canopy openingat the specific level or at the level of the insectcommunity will be presented elsewhere.
Material and Methods
Study Site and Study Plants. Insect samplingwas performed in a plot of 0.92 km2 ofunlogged forest (Block 17), in the Camoudicompartment of the logging concession ofDemerara Timbers Limited, 40 km South ofMabura Hill, central Guyana (5°13’N, 58°48'W, altitude = ca. 30 m). The main forest typesin Block 17 include well- and poorly-drainedmixed forests (ter Steege et al. 1996). An-nual rainfall at Mabura Hill is high and vari-able, from 2500 to 3400 mm, while annualair temperature is about 25.9°C.
This study focused on the seedlings andfoliage of felled trees of the following spe-cies, which are either important timber spe-cies in Guyana or relatively common in Block17: Chlorocardium rodiei (Scomb.)(Lauraceae, known locally as ‘Greenheart’);Mora gonggrijpii (Kleinh.) Sandw.(Caesalpiniaceae, ‘Morabukea’); Eperuarubiginosa Miq. (Caesalpiniaceae, ‘WaterWallaba’); Pentaclethra macroloba (Willd.)Kuntze (Leguminosae, ‘Trysil’); Catostemmafragrans Benth. (Bombacaceae, ‘SandBaromalli’). A collecting station was definedas a fixed number of tagged seedlings (40 forChlorocardium and Catostemma, 50 for Moraand Eperua and 15 for Pentaclethra) grow-ing below the parent tree or in its vicinity. Fiftysuch collecting stations were chosen for eachspecies in Block 17 (total of 250 stations and9,750 seedlings). Seedlings which died dur-ing the course of the study were replaced byother non-tagged seedlings growing below theparent tree. Other characteristics of the study
site, stations and plants are detailed elsewhere(Thomas 1999, Basset 1999).
Insect Collecting and Processing. The sam-pling protocols targeted free-living insect her-bivores collected by hand or with small aspi-rators during day-time. This included leaf-chewing (e.g., Chrysomelidae, someCurculionidae, juvenile Lepidoptera, someOrthoptera) and sap-sucking insects(Homoptera and some Heteroptera).Meristem-feeders and stem-boring insectswere not surveyed on a regular basis sincetheir census would have destroyed the seed-lings.
Most of the sampling protocol was per-formed by trained assistants. From October1996 to September 1997, 11 monthly insectsurveys were organized (Year 1). During Oc-tober 1997, half of the parent trees at the sta-tions were felled (n = 125). This felling mim-icked a situation of selective logging, whereonly particular areas in the forest are cut anddamaged. The size of the gaps created werebetween 175 m2 and 600 m2 [as measured withRunkle’s (1981) method], and most were be-tween 300-400 m2, an area mostly equivalentto “medium-sized” gaps in Charles (1998).From January to November 1998, 11 otherinsect surveys were performed (Year 2). Dur-ing both years of sampling, the following pro-tocol was used. During each survey, all thetagged seedlings of the 250 collecting stationswere inspected once, during day-time. As faras possible, insects flying away were recordedto the insect family. On average, one assistantspent at least 30 minutes at each collectingstation, carefully inspecting each tagged seed-ling. Overall, sampling effort during the twostudy years amounted to 1,114 person-daysof field work.
Juveniles of leaf-chewing insects (all cat-erpillars) were collected and reared withyoung foliage from seedlings grown for thispurpose. Juveniles of sap-sucking insects werenot collected but recorded to the nearest in-sect family. Leaf-chewing insects were keptin plastic vials with young leaves of the host-plant species there were collected from. The
436 Basset & Charles
vials were kept for 3-4 days in Block 17 andrecords of leaf damage and frass were subse-quently checked for. Insect species responsi-ble for obvious damage were later assignedto the “feeding” category, others, includingdead insects, to the “non-feeding” category.Only the former were assigned tomorphospecies and are included in the presentlist. Insect specimens were mounted, assignedto morphospecies on the basis of morphologi-cal characters and examination of genitalia,issued with a unique specimen access numberand recorded in a database. Whenever possi-ble, taxonomist colleagues examined the ma-terial further (see acknowledgments), whichwas deposited in the collections of the Centrefor Biodiversity, University of Guyana,Georgetown.
Whenever possible, feeding in situ of leaf-chewing insects was also recorded for thestudy hosts. For sap-sucking insects, hostrecords could only be ascertained in the fieldfor xylem-feeding species (e.g., exudation ofdroplets for some Cicadellinae). From thisinformation, as well as from distributionalrecords, both leaf-chewing and sap-suckinginsects were classified as “specialist” and“generalist” categories in calculating Lloyd’sindex of patchiness (see Basset 1999, for fur-ther details). A species was considered to bea “specialist” if 80% or more of its individu-als were collected on a single host species. Inthe present context, a specialist should beconsidered to be a species that showed a clearpreference for one of the five host speciesstudied, but without implication ofmonophagy.
Insect species were considered “common”if a total of 22 individuals per species werecollected during the two study years (i.e., atleast one individual collected per survey onaverage). For these common species, we testedwhether their abundance was significantlydifferent between Year 1 and Year 2 of sam-pling with a Mann-Whitney test. Data werepooled per survey to ensure that sample sizewas large enough for this test. Significancelevels were corrected with the Bonferonimethod to account for the number of simulta-
neous tests. For generalist species, recordsfrom all study hosts were considered; for spe-cialists, only records from the major host wereconsidered. Further, it was tested whether,during Year 2, the abundance of common spe-cies was significantly different between non-felled stations vs. felled stations with aWilcoxon signed ranks test. As previously, thedata were pooled per survey, Bonferoni cor-rections were applied and records from allhosts were considered unless the speciestested was a specialist. The Mann-Whitneytest explored whether the annual variabilityof species was high. A significant differencemay result from differences in rainfall, leafproduction, felling of stations or other factorsbetween study years. The Wilcoxon test ex-plored whether a significant difference couldbe more specifically related to a species’ re-sponse to the felling of stations.
Results
A synoptic list of the insect herbivorescollected on each study host is presented inAppendix I. Species identified at least to thegeneric level, along with common unidenti-fied species are listed first by highertaxonomical order (order, family, subfamily),then by alphabetical order. In total, 27,735insect individuals were collected during bothstudy years, including juveniles, damaged andlost specimens. This included 3,148 leaf-chewing insects representing at least 179 spe-cies and 24,587 sap-sucking insects represent-ing at least 425 species. The most commonhigher taxa included Psyllidae, Cicadellinae(particularly Cicadellinae, Coelidiinae andIdiocerinae), Derbidae, Membracidae (par-ticularly Smiliinae) and Achilidae for sap-sucking insects and Chrysomelidae (particu-larly Galerucinae, Eumolpinae, Alticinae andCryptocephalinae), Curculionidae (particu-larly Entiminae) and Gelechiidae for leaf-chewing insects. Aphididae were sometimescollected from Mora gonggrijpii in theMabura Hill area, but not from the taggedseedlings in Block 17. Orthoptera were col-lected rarely on seedlings and included
An. Soc. Entomol. Brasil 29(3) 437Setembro, 2000
nymphs and non-feeding specimens ofAcrididae and Pyrgomorphidae. On an iso-lated occasion, larvae of sawflies (Hymenop-tera) were collected from non-tagged seed-lings of Catostemma fragrans but could notbe reared to adult stage. No Thysanoptera andPhasmoptera were collected from the taggedseedlings.
Most species were rare and many repre-sented only by singletons. However, 61 spe-cies were considered to be common, includ-ing 51 species of sap-sucking and 10 speciesof leaf-chewing insects. About 85% of thesecommon species were generalist (52generalists and nine specialists). Mostgeneralist species were represented by xylem-feeding Cicadellinae, by presumably phloem-feeding Cixiidae, Achilidae and Derbidae, andby leaf-chewing Eumolpinae and Entiminae.Specialist species included one species ofPseudococcidae, two of Psyllidae, three ofMembracidae, one of Galerucinae, one ofCryptocephalinae and one of Gelechiidae.Lepidoptera larvae were relatively rare, rep-resenting only 6% of the total insect individu-als collected. It is difficult to comment on theactual damage that seedlings sustained fromsap-sucking insects. In addition to intake ofsap, many sap-sucking species may be vectorof various plant diseases (Nielson 1968), butthis has been quantified rarely in studies as-sessing seedling mortality in rain forests(Clark & Clark 1985, Folgarait et al. 1995).Leaf damage due to leaf-chewing insects onseedlings was rather low, estimated to be lessthan 5%. Leaf damage other than by leaf-mi-ners on Chlorocardium rodiei was rare, andit is probable that this is due to a better chemi-cal protection from chewing- rather than sap-sucking insects (Basset 1999). Seedlings ofMora gonggrijpii were attacked byCryptocephalus esuriens Suffrian, notably,but rarely sustained high damage. The samewas true of the unidentified species ofGelechiidae (TORT001) and Galerucinae(CHRY007) attacking Pentaclethramacroloba and Catostemma fragrans, respec-tively. Seedlings of Eperua rubiginosa wereattacked by various species, notably an uni-
dentified Eumolpinae (CHRY008), but noneof them were unusually common during thestudy years at the study site. However, seed-lings of E. rubiginosa were attacked by a spe-cies of bud-galling Cecidomyiidae, which wascommon but not included in the present cen-sus.
The foliage of seedlings was also shelterto several herbivore species feeding on seedsof the study hosts. For example, Scolytidaeattacking the seeds of C. rodiei (Hammondet al. 1994) were common on conspecificseedlings, but not included in the present cen-sus. Many of the weevil species ofConotrachelus and Zygopinae, whose adultsoccasionally perform maturation feeding onseedlings as reported here, may be boringconspecific seeds at the larval stage.
Whereas the number of leaf-chewing in-sects did not increase notably between bothstudy years (1,611 and 1,537 individuals col-lected, respectively), that of sap-sucking in-sects increased from 7,412 to 17,175 individu-als between Year 1 and 2, respectively. Thisdifference was due mainly to an increase ofthe specialist psyllid Isogonoceraia sp. andits nymphs, feeding on Eperua rubiginosa.The abundance of this species was also sig-nificantly higher at felled vs. non-felled sta-tions during the second year of collecting(Appendix I). The abundance of most of thecommon species did not change significantlybetween collecting years (62% of species withtest not significant, see Appendix I) or be-tween the non-felled vs. felled stations dur-ing Year 2 (70% of species with test not sig-nificant). The abundance of some specialistas well as generalist species was affected bythe collecting years or the felling treatment.A trend was noted for specialist species to bemore sensitive to collecting years than weregeneralists, but this was not significant (G-test, G = 3.65, P = 0.056).
When the number of cases for which atleast one of the two tests performed was sig-nificant (n = 29 species), the most commonsituation occurred when the abundance of thespecies increased both during the second col-lecting year and at felled stations (n = 11 spe
438 Basset & Charles
cies or 38% of cases). The second com-mon situation included a significant decreaseduring the second collecting year but no sig-nificant change at the felled stations (n= 8 spe-cies or 28% of cases). Only one species, thecixiid Pintalia sp. (CIXI002) showed a sig-nificant decrease both during the second col-lecting year and at felled stations. Annual vari-ability, when significant, induced mixed re-sponses from insects: the abundance of 14species increased, whereas the abundance ofnine others decreased. However, when the ef-fect of the felling treatment was significant,the abundance of the species often increased(16 species with higher abundance againsttwo species with lower abundance; G-test, G= 4.34, P < 0.05).
Finally, the trends (or their absence) in thechange of abundance of insect species eitherbetween collecting years or between non-felled and felled stations were not necessar-ily similar for congeneric species. This wasobvious when comparing the results of the twotests for different species in the following gen-era: Plectoderes, Pintalia, Amblyscarta,Ladoffa, Dasmeusa, Soosilius and nr Oragua,for which this comparison was possible (Ap-pendix I).
Discussion
As far as we are aware, the present listrepresents one of the very first attempts tocharacterise the entire community of insectherbivores foraging on seedlings in a tropicalrain forest with a suitable sample size. As simi-lar compilations become available from othersites in the Amazon basin or elsewhere in theNeotropical region, knowledge of the ecol-ogy of many herbivore species foraging in theforest understorey of tropical forests maygreatly improve. Further, such compilationsmay also prompt taxonomic revisions anddescriptions of particular insect groups, suchas some tribes in the Derbidae, Cixiidae orEumolpinae.
The most abundant leaf-chewing speciesthat were collected on seedlings feed readilyon the host from which they were collected
and were sometimes observed feeding in situ.Although this could be only ascertained in situfor a few xylem-feeding species, it is prob-able that most of the very common species ofsap-sucking insects also feed on the seedlingsof the study hosts. Most of these insect spe-cies feeding on seedling appeared to begeneralist species. The general impression ofthe study system is that many insect speciesmay feed on the seedlings, but probably oc-casionally only, and few reach densities thatmay be detrimental to their hosts, with thepossible exception of the spread of plant dis-eases, particularly by sap-sucking insects(Nielson 1968). The implications of high lev-els of generalist insects in the present systemwith reference to models of tree regenerationin tropical rain forests are discussed elsewhere(Basset 1999).
Relatively few larvae of Lepidoptera werecollected from the seedlings. This may be re-lated to the infrequency of their leaf flush and,as such, seedlings being a poor food resourcefor most insect herbivores (Basset 1999).Moving from one seedling to another in searchof young foliage may be risky for lepidop-teran larvae, but less so for alate adults suchas chrysomelids and cixiids, for example. Thisinterpretation is reinforced further by the sig-nificantly higher abundance of Lepidopteralarvae in parent trees, which offer more abun-dant food resources, in comparison withconspecific seedlings (Basset et al. 1999).
Since insect seasonality was low duringcollecting years (Basset - in press), it was pos-sible to pool the insect data to ensure that sam-ple size was high enough for the analyses.However, insect densities on seedlings weregenuinely low, presumably because seedlingsrepresent a marginal food resource for mostof the species collected (Basset - in press).Thus, it is possible that lack of significantchange in the abundance of many insect spe-cies either between years or between non-felled and felled stations may result purelyfrom the low abundance of the insects. None-theless, the abundance of some insect specieswas significantly different between the twocollecting years. The last surveys of Year 1
An. Soc. Entomol. Brasil 29(3) 439Setembro, 2000A
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3620
162
1211
47
n.s.
n.s.
Gen
eral
ist
Gen
. sp.
?D
ER
B02
130
1911
86
23
11n.
s.n.
s.G
ener
alis
tO
ther
Der
bida
e66
spp
.27
513
813
749
8273
2447
--
-K
inna
rida
eSo
uthi
a sp
. nr o
ppos
itaC
IXI0
0369
4722
312
67
23n.
s.n.
s.G
ener
alis
tF
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rida
eN
r Lo
cron
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LG
002
11
0-
1-
--
--
-
440 Basset & CharlesN
r Sc
aral
is s
p.FU
LG
001
11
0-
-1
--
--
-A
chili
xiid
aeG
en. s
p. ?
AC
CX
001
44
0-
1-
3-
--
-A
chili
dae
Plec
tode
res
colla
ris
F.A
CH
I002
132
5478
2716
1427
48n.
s.n.
s.G
ener
alis
tPl
ecto
dere
s sp
.A
CH
I023
352
334
815
17
In.
s.G
ener
alis
tPl
ecto
dere
s sp
.A
CH
I025
20
2-
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--
--
-Pl
ecto
dere
s sp
.A
CH
I026
10
1-
1-
--
--
-Se
via
bica
rina
ta F
.A
CH
I001
117
6156
613
2248
28n.
s.D
Gen
eral
ist
Sevi
a co
nsim
ile F
enna
hA
CH
I006
166
102
42
17
--
Kno
wn
from
Trin
idad
(L.
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rien,
pers
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mm
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r C
aton
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p.A
CH
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11
01
--
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--
-G
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p. ?
AC
HI0
0911
47
--
31
7-
--
Oth
er A
chili
dae
19 s
pp.
5431
237
1113
518
--
-D
icty
opha
rida
eTa
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mul
iebr
is W
alke
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ICT
006
20
2-
1-
-1
--
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ropa
ferr
ifera
(W
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ICT
001
148
63
13
61
--
-To
ropa
pic
ta W
alke
rD
ICT
002
31
2-
1-
-2
--
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sp.
DIC
T00
41
10
--
1-
--
--
Oth
er D
icty
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ridae
3 sp
p.11
29
--
33
5-
--
Cix
iidae
Ben
nare
lla s
p.A
CC
X00
218
126
15-
21
--
--
Pin
talia
sp.
CIX
I006
669
554
115
128
7385
130
253
Dn.
s.G
ener
alis
tP
inta
lia s
p.C
IXI0
0241
535
560
101
1282
2519
5D
DG
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tP
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p.C
IXI0
0911
091
1923
1235
1921
Dn.
s.G
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alis
tN
r B
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ra s
p.A
CH
I018
21
12
--
--
--
-N
r B
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ra s
p.A
CH
I020
10
1-
--
1-
--
-G
en. s
p. ?
CIX
I008
2216
6-
36
85
n.s.
n.s.
Gen
eral
ist
Oth
er C
ixiid
ae11
spp
.88
5434
1121
1810
28-
--
Tro
pidu
chid
aeG
en.
sp.?
TR
OP0
012
11
-1
--
1-
--
Gen
. sp
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RO
P004
10
1-
-1
--
--
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lati
dae
An. Soc. Entomol. Brasil 29(3) 441Setembro, 2000N
r A
norm
enis
sp.
FLAT
012
60
61
-2
12
--
-N
r Loc
rona
sp.
FLAT
004
31
2-
2-
-1
--
-N
r P
oeki
llopt
era
sp.
FLAT
016
10
1-
--
-1
--
-O
ther
Fla
tidae
5 sp
p.14
86
25
10
6-
--
Issi
dae
Gen
. sp.
?IS
SI00
211
56
11
13
5-
--
Oth
er I
ssid
ae9
spp.
209
112
24
48
--
-A
cana
loni
idae
3 sp
p.5
32
3-
--
1-
--
Nog
odin
idae
Nog
odin
a re
ticul
ata
F.N
OG
O00
114
68
32
32
4-
--
Gen
. sp
.?N
OG
O00
21
01
--
-1
--
--
Cic
adid
aeG
en. s
p. ?
CIC
D00
21
01
--
--
1-
--
Cer
copi
dae
Gen
. sp.
?C
ER
C00
111
74
22
21
4-
--
Oth
er C
erco
pida
e5
spp.
2311
124
35
65
--
-M
embr
acid
aeE
ndoi
asti
nae
Scyt
odep
sa s
p.M
EM
B01
244
1034
18
-12
23n.
s.n.
s.G
ener
alis
tTr
opid
aspi
s sp
.M
EM
B04
93
03
-1
2-
--
--
Trop
idas
pis
sp.
ME
MB
106
10
1-
-1
--
--
-O
ther
End
oias
tinae
2 sp
p.8
08
11
71
--
--
Steg
aspi
nae
Steg
aspi
s fr
ondi
tia L
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EM
B04
33
12
-2
--
1-
--
Nr
Lyco
dere
s sp
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EM
B05
14
04
--
11
2-
--
Dar
nina
eC
ypho
tes
sp.
ME
MB
095
10
1-
-1
--
--
-D
arni
s la
tera
lis F
.M
EM
B00
781
2952
815
319
36n.
s.n.
s.G
ener
alis
tN
r St
icto
pelta
sp.
ME
MB
022
11
0-
1-
--
--
-H
eter
onot
inae
Het
eron
otus
nr
trid
ens
Bur
mei
ster
ME
MB
024
10
1-
--
-1
--
-M
embr
acin
ae
442 Basset & CharlesB
olbo
nota
sp.
ME
MB
032
33
0-
--
3-
--
-B
olbo
nota
sp.
ME
MB
089
10
1-
-1
--
--
-C
ampy
lenc
hia
sp.
ME
MB
093
10
1-
--
1-
--
-E
ncho
phyl
lum
sp.
ME
MB
088
90
92
-1
33
--
-Le
iosc
yta
sp.
ME
MB
030
61
5-
2-
13
--
-Le
iosc
yta
sp.
ME
MB
071
210
2120
--
-1
--
-M
embr
acis
nr
folia
ta L
.M
EM
B04
87
16
3-
-2
2-
--
Mem
brac
is s
p.M
EM
B02
647
542
1-
237
7I
ISp
ecia
list
on P
mM
embr
acis
sp.
ME
MB
041
11
0-
--
-1
--
-N
assu
na s
p.M
EM
B05
02
02
1-
-1
--
--
Not
ocer
a sp
.M
EM
B00
933
429
65
69
7I
n.s.
Gen
eral
ist
Pot
nia
glad
iato
r W
alke
rM
EM
B01
59
36
3-
3-
3-
--
Spho
ngop
horu
s nr
que
lini F
ron.
ME
MB
054
10
1-
--
1-
--
-Su
ndar
ion
sp.
ME
MB
091
10
1-
--
1-
--
-N
r A
cono
phor
a sp
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EM
B03
56
51
-5
1-
--
--
Nr
Cym
bom
orph
a sp
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EM
B04
71
10
--
--
1-
--
Oth
er M
embr
acin
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spp.
211
202
22
141
--
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iliin
aeA
mas
tris
sp.
ME
MB
029
11
0-
--
1-
--
-A
mas
tris
sp.
ME
MB
062
10
1-
--
1-
--
-A
phet
ea s
p.M
EM
B00
516
313
--
-16
--
--
Aph
etea
sp.
ME
MB
039
32
1-
1-
2-
--
-H
arm
onid
es s
p.M
EM
B00
211
110
3-
11
6-
--
Har
mon
ides
sp.
ME
MB
010
10
1-
1-
--
--
-H
arm
onid
es s
p.M
EM
B02
313
211
11
42
5-
--
Har
mon
ides
sp.
ME
MB
031
11
01
--
--
--
-M
embr
acid
aeSm
iliin
aeH
arm
onid
es s
p.M
EM
B08
12
02
1-
--
1-
--
Har
mon
ides
sp.
ME
MB
087
40
4-
12
-1
--
-Sm
ilior
hach
is s
p.M
EM
B00
321
424
190
1212
617
77
II
Spec
ialis
t on
Pm
Nr
Hor
iola
sp.
ME
MB
004
330
1731
31
3318
257
21I
n.s.
Spec
ialis
t on
Pm
Oth
er S
mili
nae
22 s
pp.
6413
5110
289
1710
--
-
An. Soc. Entomol. Brasil 29(3) 443Setembro, 2000O
ther
Mem
brac
idae
18 s
pp.
5721
3618
109
1010
--
-A
etal
ioni
dae
Gen
. sp.
?A
ETA
001
80
8-
8-
--
--
-C
icad
ellid
aeL
edri
nae
Xed
reot
a tu
berc
ulat
a (O
sbor
n)C
ICA
023
185
132
33
19
--
-A
galli
nae
Aga
llia
sp.
CIC
A06
118
117
-4
4-
10-
--
Aga
llia
sp.
CIC
A17
01
10
1-
--
--
--
Idio
ceri
nae
Chi
asm
odol
on s
p.C
ICA
020
4231
113
1114
68
n.s.
n.s.
Gen
eral
ist
Chi
asm
odol
on s
p.C
ICA
065
209
111
5-
113
--
-G
en. s
p. ?
CIC
A03
639
1227
1215
42
6n.
s.n.
s.G
ener
alis
tO
ther
Idi
ocer
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12 s
pp.
247
173
86
16
--
-C
icad
ellid
aeG
ypon
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Bar
bata
na e
xter
a Fr
eyta
gC
ICA
029
54
12
2-
1-
--
-C
urta
ra s
p.C
ICA
027
10
1-
--
1-
--
-G
ypon
a bu
lbos
a D
eLon
g&
Fre
ytag
CIC
A00
816
214
19
22
2-
--
Gyp
ona
flavo
limba
taM
etca
lfC
ICA
018
209
1114
-2
-4
--
-G
ypon
a fu
nda
DeL
ong
CIC
A11
22
02
-1
1-
--
--
Gyp
ona
glau
ca F
.C
ICA
030
32
1-
11
-1
--
-G
ypon
a of
fa D
eLon
g &
Fre
ytag
CIC
A16
91
01
--
1-
--
--
Pol
ana
sp.
CIC
A00
71
01
1-
--
--
--
Oth
er G
ypon
inae
5 sp
p.7
25
14
01
1-
--
Coe
lidiin
aeB
alub
a pa
ralle
la N
iels
onC
ICA
016
379
281
46
719
n.s.
IG
ener
alis
tD
ocal
idia
o’r
eilly
Nie
lson
CIC
A06
03
12
--
-1
2-
--
Doc
alid
ia s
p.C
ICA
024
4212
307
34
325
n.s.
IG
ener
alis
tD
ocal
idia
sp.
CIC
A12
97
34
-1
11
4-
--
Doc
alid
ia s
p.C
ICA
130
55
01
--
13
--
-
444 Basset & CharlesD
ocal
idia
sp.
CIC
A13
12
20
--
11
--
--
Par
acar
inol
idia
sp.
CIC
A07
810
46
6-
2-
2-
--
Cic
adel
lidae
Coe
lidiin
aeSt
alol
olid
ia s
p.C
ICA
052
11
0-
-1
--
--
-N
r D
ocal
idia
sp.
CIC
A04
013
58
4-
11
7-
--
Oth
er C
oelid
iinae
11 s
pp.
2510
154
23
79
--
-X
esto
ceph
alin
aeX
esto
ceph
alus
des
erto
rum
(Ber
g)C
ICA
144
260
267
--
-19
n.s.
n.s.
Gen
eral
ist
Cic
adel
linae
Acr
ocam
psa
palli
pes
F.C
ICA
005
327
2511
41
97
n.s.
n.s.
Gen
eral
ist
Am
blys
cart
a in
venu
sta
You
ngC
ICA
006
147
2512
229
2728
2538
II
Gen
eral
ist
Am
blys
cart
a sp
.C
ICA
012
351
3413
15
412
II
Gen
eral
ist
Am
blys
cart
a sp
.C
ICA
011
31
2-
--
3-
--
-C
ardi
osca
rta
quad
rifa
scia
ta L
.CIC
A02
525
1114
511
1-
8n.
s.n.
s.G
ener
alis
tD
asm
eusa
pau
pera
ta Y
oung
CIC
A00
116
838
130
2520
3120
72I
IG
ener
alis
tD
asm
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sp.
CIC
A07
674
4628
38
336
24n.
s.n.
s.G
ener
alis
tLa
doffa
agu
ilari
Loz
ada
CIC
A05
813
112
11
73
1-
--
Lado
ffa c
omiti
s Y
oung
CIC
A05
424
420
62
31
12n.
s.n.
s.G
ener
alis
t; F:
Cf
Lado
ffa ig
nota
Wal
ker
CIC
A00
348
1236
93
176
13n.
s.I
Gen
eral
ist;
F: P
mLa
doffa
sp.
CIC
A07
710
55
24
12
1-
--
Mac
ugon
alia
moe
sta
(F.)
CIC
A06
262
620
3910
92
2D
n.s.
Gen
eral
ist
Poe
cilo
scar
ta c
ardi
nalis
F.
CIC
A03
317
611
35
21
6-
--
Soos
iulu
s fa
bric
ii M
etca
lfC
ICA
004
1172
603
569
404
122
217
165
264
n.s.
n.s.
Gen
eral
ist;
F: C
r, M
g, E
r, C
fSo
osiu
lus
inte
rpol
is Y
oung
CIC
A08
114
865
8337
2930
2824
n.s.
n.s.
Gen
eral
ist;
F: E
rN
r O
ragu
a sp
.C
ICA
010
254
203
5141
1682
3283
Dn.
s.G
ener
alis
tN
r O
ragu
a sp
.C
ICA
080
5938
219
422
519
n.s.
n.s.
Gen
eral
ist
Oth
er C
icad
ellin
ae10
spp
.16
88
14
44
3-
--
Cic
adel
lidae
Typ
hloc
ybin
aeJo
rum
a co
ccin
ea M
cAte
eC
ICA
002
290
291
24
-22
n.s.
n.s.
Gen
eral
ist
Joru
ma
nr a
pica
ta C
aldw
ellC
ICA
022
122
101
-2
-9
--
-
An. Soc. Entomol. Brasil 29(3) 445Setembro, 2000G
en. s
p. ?
CIC
A00
915
69
21
2-
10-
--
Oth
er T
yphl
ocyb
inae
15 s
pp.
493
464
63
630
--
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elto
ceph
alin
aeM
atto
gros
sus
colo
noid
es(L
inna
vuor
i)C
ICA
165
10
11
--
--
--
-O
sbor
nellu
s sp
.C
ICA
015
73
42
3-
2-
--
-P
lano
ceph
alus
fla
vico
sta
(Sta
l.)C
ICA
143
70
73
--
-4
--
-Ta
peri
nha
sp.
CIC
A04
38
44
11
12
3-
--
Oth
er D
elto
ceph
alin
ae8
spp.
104
62
02
33
--
-
Oth
er C
icad
ellid
ae11
spp
.29
821
6-
34
16-
--
Psy
llida
eE
upha
leru
s sp
.PS
YL
001
118
311
5-
118
--
-I
ISp
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list
on M
g; F
: M
g (n
ymph
s)
Isog
onoc
erai
a sp
.PS
YL
003
2932
698
2234
6211
526
8212
61I
ISp
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list
on E
r; F:
Er
(nym
phs)
Not
ophy
llura
sp.
PSY
L00
51
10
1-
--
--
--
Gen
. sp
.?PS
YL
007
20
2-
--
-2
--
-A
leyr
odid
aeat
leas
t 8 s
pp.
4121
206
210
122
--
-P
seud
ococ
cida
eG
en. s
p. ?
CO
CC
002
149
147
214
9-
--
-D
n.s.
Spec
ialis
t on
Cr
Oth
er P
seud
ococ
cida
e3
spp.
100
108
-1
1-
--
-H
ET
ER
OP
TE
RA
Cor
eida
ePa
chyl
is n
r la
ticor
nis
F.C
OR
E00
215
96
11
13-
--
Nr
Aca
ntho
ceph
ala
sp.
CO
RE
003
144
102
-8
13
--
-N
r M
eroc
oris
sp.
CO
RE
009
20
2-
2-
--
--
-N
r P
eran
thus
sp.
CO
RE
001
303
2712
11
79
II
Gen
eral
ist
Oth
er C
orei
dae
6 sp
p.18
414
112
12
2-
--
Lyg
aeid
aeG
en.
sp.?
LYG
A00
11
10
1-
--
--
--
Gen
. sp
.?LY
GA
002
10
11
--
--
--
-
446 Basset & CharlesC
ydni
dae
Gen
. sp
.?C
YD
N00
16
60
-1
12
2-
--
Pen
tato
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oxia
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11
31
--
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dess
a sp
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NT
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4311
325
412
517
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Gen
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611
29
11
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8-
--
Ede
ssa
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T00
74
13
1-
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--
Mor
mid
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n (L
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--
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r E
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NT
009
11
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ther
Pen
tato
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142
127
22
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--
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001
253
122
131
615
630
655
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Gen
eral
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ella
rida
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oris
sp.
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T00
24
22
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--
--
--
Gen
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001
11
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--
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Gen
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AB
002
11
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sp.?
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32
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61
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Stei
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RA
005
20
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alop
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CH
RY
088
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list
on M
g; F
: M
g
An. Soc. Entomol. Brasil 29(3) 447Setembro, 2000E
umol
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911
65
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811
355
582
1053
345
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Gen
eral
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F: M
g, E
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sp.?
CH
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127
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HR
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032
302
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42
5D
n.s.
Gen
eral
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Gen
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HR
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524
519
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Mg
Oth
er E
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746
715
3630
1135
--
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atin
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r La
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HR
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31
10
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--
--
--
Gal
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Gen
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737
221
216
02
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236
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Spec
ialis
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--
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044
43
12
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53
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1610
67
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50
52
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CH
RY
014
4337
61
152
223
n.s.
n.s.
Gen
eral
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ther
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spp
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3735
96
1810
29-
--
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11
10
--
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CU
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004
33
0-
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--
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ntim
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Com
psus
sp.
CU
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012
149
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32
36
--
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sp.?
CU
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003
494
45-
61
2319
II
Gen
eral
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Oth
er E
ntim
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7 sp
p.53
944
1520
313
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--
Hyl
obin
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us s
p.C
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11
01
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er H
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13
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1-
--
--
448 Basset & CharlesP
rion
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Pri
onom
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sp.
CU
RC
017
30
33
--
--
--
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sp.
CU
RC
016
32
1-
11
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--
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CU
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058
20
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1-
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sp.
CU
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002
11
0-
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CU
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036
10
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-1
--
--
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rach
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sp.
CU
RC
040
10
11
--
--
--
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onot
rach
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sp.
CU
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057
10
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1-
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s sp
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UR
C01
81
10
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015
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p.C
UR
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92
11
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--
--
Oth
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rypt
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5 sp
p.5
14
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11
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--
Zyg
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CU
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005
279
182
45
610
n.s.
n.s.
Gen
eral
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Oth
er Z
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15 s
pp.
317
241
215
13
--
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Gen
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32
11
--
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--
Gen
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01
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41
10
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p.2
02
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--
--
--
DIP
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--
--
--
--
--
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ot c
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--
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--
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mon
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cial
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af-m
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sp.?
--
--
--
--
--
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ot c
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on s
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leaf
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An. Soc. Entomol. Brasil 29(3) 449Setembro, 2000G
en.
sp.?
--
--
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4024
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tric
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12
--
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--
--
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001
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LIM
A00
21
10
--
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--
Pyr
alid
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--
--
--
--
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GE
OM
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ther
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14
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HE
SP00
22
11
--
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--
--
Lyc
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mer
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CA
001
42
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--
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on
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ua a
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ican
a (J
oly
1996
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31
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nd C
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--
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11
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--
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ast 4
spp
.5
32
-2
-2
1-
--
450 Basset & Charles
and a few of Year 2 were performed during anEl Niño event. Many changes in insectabundance may result from the rainfall fac-tor, and particularly from its interaction withthe leaf production of the host -plants.
However, a few species apparently ben-efited from an increase in canopy opennessafter the felling of the stations and were sig-nificantly more abundant at the felled stations,as shown by the preliminary tests reportedhere. Even if the gaps created were relativelymodest in size (< 400 m2 for most), this mayhave been important for some heliophilousspecies, such as some Cicadellidae,Membracidae, Pentatomidae or Entiminae, inthe otherwise dark and shady understorey ofBlock 17. Alternatively, higher canopy open-ness may have increased the leaf productionof seedlings and the abundance of insects thatmay depend more directly on the presence ofyoung foliage, such as some Psyllidae andChrysomelidae. This, in addition to the influ-ence of rainfall, will be examined elsewhere.
Interestingly, the present compilation andpreliminary tests suggest that congeneric in-sect species may not necessarily respond in asimilar way to changes induced by rainfall andcanopy openness. This further suggests thatthe ecology and requirements of these spe-cies may be different and that these factorsneed to be examined at the specific level. Evengeneralist species in this study system displaypreferences for particular host-plants or mi-cro-habitats (Basset - in press). It may be ex-pected that insect species will be finely dis-tributed on different resources/micro-habitatsin the forest understorey, although their dis-tribution may be even finer in the canopy. Inthese conditions, insect species are likely todisplay a variety of responses to natural andman-induced forest disturbance (Charles1998), which may parallel their phenotypicand genetic diversity.
Acknowledgments
Milton Allicock, Beverley Daniels, ShawnFiffee, Eustace Francis, Brentnal Hoyte,Henry James, Wade Klencian, Linsford La
Goudou, Troy Lashley, Marlon Mclennan,Dayl Thompson, Terrence Washington, Mar-tin Williams and Orin Williams collected,mounted and helped with the assignment ofmost of the insect material. Wilfred Jarvis,Guyana Forestry Commission, felled the par-ent trees in Block 17 with impressive skills.David Hammond, Raquel Thomas, Val Brownand the Tropenbos-Guyana staff helped withvarious conceptual and logistical aspects ofthe project. Other much needed administra-tive and logistical help was provided by theTropenbos - Guyana Programme, the Inter-national Institute of Entomology, the Over-seas Development Administration and Dem-erara Timbers Limited. Taxonomic help wasprovided by Hector Barrios, John Brown,Rodney Caviochioli, Caroline Chaboo, RexCocroft, Michael Cox, Paul Freytag, DavidHollis, Mervin Nielson, Jacqueline Miller,Lois O’Brien, Dave Rider, Eva Sprecher andMick Webb. Comments by Hector Barrios,Neil Springate and Vojtech Novotny improvedthe manuscript. Work to develop an efficientmethod of data archival was possible throughU.S. National Science Foundation grant DEB-97-07928. The project was financially sup-ported by the Darwin Initiative for the Sur-vival of the Species and British Airways (con-servation travel scheme).
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