weevil diversity and seasonally in tropical panama as deduced from light-trap catches

Weevil Diversity andSeasonally in Tropical Panama

as Deduced from Light-Trap Catches(Coleoptera: Curculionoidea)

HENK WOLDA,CHARLES W. O'BRIEN,

andHENRY P. STOCKWELL

I

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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y • N U M B E R 5 9 0

Weevil Diversity andSeasonality in Tropical Panama


Henk Wolda, Charles W. O'Brien,and Henry P. Stockwell

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1998

A B S T R A C T

Wolda, Henk, Charles W. O'Brien, and Henry P. Stockwell. Weevil Diversity and Seasonalityin Tropical Panama as Deduced from Light-Trap Catches (Coleoptera: Curculionoidea).Smithsonian Contributions to Zoology, number 590, 79 pages, 9 tables, 27 figures,1998.—Weevils were collected with light traps at seven localities in the Republic of Panama,varying in altitude from sea level to 2200 m, in climate from sharply seasonal to virtuallynonseasonal, and in habitat from natural tropical forest to areas strongly disturbed by humans.Although only an estimated 25-40 percent of the species of weevils present in an area wereattracted to light, a total of 2086 species was nonetheless obtained in the traps. On BarroColorado Island (BCI), the canopy trap caught more individuals but fewer species than the trapnear the ground. Species richness (alpha-diversity) varied greatly between sites, BCI being therichest and the high-altitude site of Guadalupe Arriba being the poorest. Using the logseries asan arbitrary but useful basis for comparison, there were too many rare species and too fewspecies of intermediate abundances at all sites. Between-site (beta-) diversity was also large,with one-third to two-thirds of the species at each site being only observed at that site, whereasspecies occurring at four or more sites were very rare.

Descriptors of seasonal patterns are proposed that were borrowed from circular statistics, suchas Mean Vector and Mean Week. The former indicates the degree of seasonality, i.e., theconcentration of the individuals in a year, which ranges in value from zero (uniform distribution)to unity (all individuals occurring at the same time), whereas the latter indicates the circularmean of the seasonal distribution. These were used in conjunction with other seasonalitymeasures, such as Peak Week, which is the mode of the seasonal distribution. For all six siteswith at least one year of data, these measures were calculated for each year for all species withat least 10 individuals in that year. A very rich variation in seasonal patterns was observedamong species, ranging from species with very short seasons to species occurring year-round,sometimes without any clear seasonal peaks. At the climatically seasonal sites and at one lessseasonal site, most species exhibited their maximum abundance at the beginning of the rainyseason. However, at all sites some species were active or even had their mean or maximumabundance at any time of the year. Most species demonstrated very similar seasonal patterns insuccessive years, apart from shifts of a few weeks related to the actual beginning of the rainyseason, but there were some clear exceptions. Similarly, for most species that occurred at morethan one site in reasonable numbers the seasonal patterns were rather similar in those differentsites in spite of differences in habitat, seasonality, or altitude. However, there were a number ofspecies with spectacular differences in seasonal patterns at different sites. In some cases suchdifferences could partly be attributed to differences between the sites, in others they could not.Species with an intermediate degree of seasonality showed a higher variability between sitesthan between years within sites.

The results were compared with those obtained for weevils from temperate areas. To the bestof our knowledge, this is the first study on between-year and between-site comparisons inseasonality with a large number of tropical insect species.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and isrecorded in the Institution's annual report, Annals of the Smithsonian Institution. SERIES COVERDESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication DataWolda, HenkWeevil diversity and seasonality in tropical Panama as deduced from light-trap catches (Coleoptera: Curculionoidea)

/ Henk Wolda, Charles W. O'Brien, and Henry P. Stockwell.p. cm.—(Smithsonian contributions to zoology ; no. 590)Includes bibliographical references (p . ) .1. Beetles—Seasonal distribution—Panama. I. O'Brien, Charles William. II. Stockwell , Henry P.

III. Title. IV. Series.QL1.S54 no. 590 [QL587.P2] 590 s-dc21 [595.76'8'097287] 97-43032

CIP

© The paper used in this publication meets the minimum requirements of the AmericanNational Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.

Contents

Page

Introduction 1Acknowledgments 2

Materials and Methods 2Study Sites 2Collecting Methods 3Data Analysis 5

Diversity 5Seasonality 6

Results 7General Remarks 7

Effect of Trap Design and of the Dying Tachygalia Tree 7Diversity 8

Per Year 8Per Week and Per Day 11Families and Subfamilies 13Between-Site Diversity 13Canopy vs. Ground Level 15

Discussion of Diversity 16Species Richness in Panama Compared with Nontropical Areas 16Frequency Distributions of Species Abundances 18Between-Year Comparisons 19Between-Site Comparisons 19Vertical Stratification 19

Seasonality 21Weevils as a Group 21Between-Species Comparisons 21Between-Year Within-Site Comparisons 23Within-Species Between-Site Comparisons 29

Discussion of Seasonality 35Appendix: List of Species of Weevils Collected by Light Traps in Seven Localities

in Panama 42Literature Cited 76

in

Weevil Diversity andSeasonality in Tropical Panama


Henk Wolda, Charles W. O'Brien,and Henry P. Stockwell

Introduction

It is a well-established fact that, as in temperate species,seasonality is a common phenomenon among tropical insects.Since the classic studies by Dobzhansky and Pavan (1950) andBigger (1976) a fair number of papers have been publishedshowing that tropical insect species range from aseasonal tosharply seasonal even in relatively aseasonal climates, avariation much larger than that found in the temperate zone. Fora review see Wolda (1988). For some species there isinformation on between-year similarities in seasonal patterns(Patil and Thontadarya, 1983; Wolda, 1982, 1983c, 1989)suggesting that between-year differences in seasonal abun-dance patterns do exist, but that they are small, comparable tosimilar differences occurring in the temperate zone where aseason may start a few weeks earlier or later as a consequenceof a variation in the weather in spring. Similarly, informationon between-site differences in seasonality of individual tropicalinsect species is rare. Agarwala and Bhattacharya (1993) foundthat the seasonal abundance patterns of the aphid Toxopteraaurantii Boyes de Fonscolombe in India was very different intwo climatically different sites. Aouad (1988) observed thehydrophilid beetle Berosus qffinis Brulle in Morocco to be

Henk Wolda, Henry P. Stockwell, Smithsonian Tropical ResearchInstitute, P.O. Box 2072, Balboa, Republic of Panama; Charles W.O'Brien, Florida A. & M. University, Entomology-Biocontrol,Tallahassee, FL 32307-4100, U.S.A.Review Chairman: Ira Rubinoff, Smithsonian Tropical ResearchInstitute (STRI), Apartado 2072, Balboa, Republic of Panama.Reviewers: R.S. Anderson, Canadian Museum of Nature, Ottowa,Canada: H.R. Burke, Texas A. & M. University, College Station,Texas, 77843; A. T. Howden, Carleton College, Northfleld, Minnesota,55057; D.M. Windsor (STRI).

univoltine in temporary ponds and bivoltine in permanentponds. Arbeille (1987) found an effect of fire on voltinism insome cockroaches in the Ivory Coast. Reddy and Krishnamur-thy (1976-1977) observed between-site differences in seasonalpatterns in some Drosophila species near Mysore, India.Rutledge et al. (1976) found large between-site differences inseasonality in species of Panamanian sand flies. Sevastopulo(1976) noted strong differences in seasonality of Charaxesbutterflies in Kenya between coastal gardens and inlandsavannahs. In temperate areas between-site variations inseasonal patterns are commonplace, especially along latitudinalgradients. Similarly, between-year variations in phenology arewell studied in many insects. However, because a cold seasonusually limits the time in which insects can be active, the rangeof variation tends to be relatively small, usually restricted to avariation in the number of generations per year. Information onseasonal patterns of species of weevils is rare, especially fortropical ones.

The present paper analyzes diversity and seasonal patterns inabundance, as shown by light-trap catches, for a large numberof weevil species in seven localities (six for seasonality) in theRepublic of Panama. These sites range in altitudes from 0 to2200 meters and cover a variety of patterns of climaticseasonality. Species richness at each of seven sites, asdemonstrated by the light traps, is presented, and thedistribution of seasonal patterns at the six sites with at least onefull year of data is discussed. Because we could find nosatisfactory method in the literature to describe and summarizeseasonal abundance patterns, elements of circular statistics areproposed here as descriptors of seasonality. From four sitesbetween-year variation in seasonality, rather large in somespecies, is described, as is between-site variation in seasonality

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

for those species that occurred in reasonable numbers at morethan one locality.

ACKNOWLEDGM ENTS

The study on Barro Colorado Island was partly funded bygrants from the Smithsonian Environmental Science program.We thank all those who helped in collecting the daily samples,especially Bonifacio de Leon. The study in Fortuna was madepossible thanks to the generous cooperation of Alcides Salas ofthe Instituto de Recursos Hidraulicos y Electrificacion (IRHE).Personnel of the IRHE collected the daily samples from thelight trap. The project in Boquete was carried out on theproperty of Alberto Sandberg and with the cooperation ofvarious people of the Ministerio de Desarrollo Agropecuario inBoquete, especially Eleuterio Delgado. Rogelio Preto kindly letus use his house and property for the project at GuadalupeArriba. Thanks are due to the untiring support of Cecilio Estribiof the Instituto de Recursos Naturales Renovables in theprovince of Chiriqui who, together with Lionel Quiros, steppedin immediately whenever there were problems either inFortuna, Guadalupe Arriba, or Boquete. The project inCorriente Grande was partially funded and given logisticalsupport by the IRHE and also helped by Abdiel Adames and hisassistants of the Gorgas Memorial Laboratory in Panama City.The project in Miramar was partly funded by a ScholarlyStudies grant from the Smithsonian Institution and received thegenerous practical help and logistical support of YessimYishui. Saturnino Martinez sorted the weevils out of all insectsamples as they came in. We thank Tina Wolda, wife of thesenior author, for her help in collecting the samples in LasCumbres, and Angel Aguirre of the STRI Library in Panama aswell as Zbigniew Witkowski (Krakow, Poland) for their help inlocating weevil literature. Constructive comments by fourreviewers are gratefully acknowledged. Horace R. Burke'sdetailed comments especially are deeply appreciated. The workby CWOB was supported in part by a grant, FLAX 850006,from the Cooperative State Research Service, U.S. Departmentof Agriculture.

Materials and Methods

STUDY SITES

Weevil composition, abundance, and seasonality wereanalyzed from light-trap samples collected at seven localities inPanama, Barro Colorado Island, Las Cumbres, Miramar,Corriente Grande, Boquete, Fortuna, and Guadalupe Arriba.

Barro Colorado Island (BCI), 9°9'19"N, 79°45'19"W, is thelargest of the islands formed in the man-made Gatun Lakewhen this was flooded between 1911 and 1914 to form part ofthe Panama Canal. Light traps were located on a ridge at 120meters above sea level in reasonably undisturbed forest. Fordetails of the forest and its history see Croat (1978), Leigh et al.

(1982), Foster and Brokaw (1982), and Piperno (1990). Therewere two traps, one at two to three meters above ground leveland one in the canopy, 28 meters above the forest floor. For 20years the traps operated virtually every night, all night, and thesamples were collected in the morning and transported to theoffices of the senior author for initial sorting. In early March1977, the glass jar with CC14, used to collect and kill the insectscollected by the trap, was replaced by a stainless steelreceptacle with Kahle's solution. In October 1978, the trapswere moved about 10 meters to a tree nearby. Weevils collectedby these traps were sorted during three years, from March 1976through March 1979.

Las Cumbres, 9°5'36"N, 79°31'54"W, 150 m above sealevel, is a residential area with gardens and some secondgrowth forest, 16 km north of Panama City. A single light trapwas operated behind the house of one of the authors (HW) fora total of seven years on a ridge overlooking a wooded area.The trap operated virtually every night, all night, and weevilswere sorted for three years, from March 1974 through March1976 (see also Wolda, 1980).

Miramar, 9°0'N, 82°15'W, is at sea level, in the province ofBocas del Toro in NW Panama, on the SW corner of theLaguna de Chiriqui. The light trap was located on the edge ofa pasture at the bottom of a very disturbed forested slope, withold cacao trees in the undergrowth. The trap operated everyevening from dusk until 10 P.M. during one year, fromNovember 1978 through November 1979. Due to logisticproblems, data are missing for 12-30 June, 12-25 July, and28-31 July (see Wolda and Flowers, 1985).

Corriente Grande, 9°17'30"N, 82°32'41"W, altitude 100 m,is a forested area along the Changuinola River, near a workcamp of the Instituto de Recursos Hidraulicos e Electrificacion(IRHE). For details of the area see Adames (1980). A light trapwas operated here from dusk to 10 P.M. from mid-Januarythrough mid-May of 1980.

Boquete, 8°48'N, 82°26'W, altitude 1350 m, is in themountains of Western Chiriqui province. The light trap waslocated in a forest remnant surrounded by coffee plantations onthe property of Alberto Sandberg, in the community of AltoLino, and was operated every night, all night, for three years.Weevils were sorted from the samples during two years, fromJuly 1976 through July 1978.

Fortuna, 8°44'N, 82°16'W, altitude 1050 m, is a very wetforested mountain valley along the Rio Chiriqui, some 20 kmeast of Boquete (at present the study area is an artificial lake).A light trap was operated here daily from dusk to 10 P.M. in old,relatively undisturbed forest outside a work camp of theInstituto de Recursos Hidraulicos e Electrificacion (IRHE),from late September 1976 to July 1979, the first nine months atcanopy level, the subsequent two years a few meters above theforest floor (see Adames, 1977).

Guadalupe Arriba, 8°52'27"N, 82°33'12"W, altitude 2200 mis in a very wet cloud forest on the northwestern slopes of theVolcan Bani. A light trap was operated here daily from dusk to

NUMBER 590

TABLE 1.—Annual rainfall and temperature data for seven Panamanian localities as far as available. For Miramarand Corriente Grande, no data are available, so rainfall data from nearby Punta Robalo and Changuinola,respectively, are substituted. Temperature data from Almirante serves as a guideline for temperatures at thesesites. No temperature data are available for Las Cumbres, but the BCI clearing data should be roughly applicable.

Locality

BCI, ClearingBCI, Forest FloorLas CumbresVliramarAlmiranteCorriente GrandeBoqueteFortunaGuadalupe Arriba

N

62-

159_

5814142

Annual rainfallMean ± St.Dev.

2614.0 ±456.0-

2134.2 ±521.71814.7 ±633.1

_

2519.1 ±517.92481.5 ±736.44569.4 ± 839.23874.1 ± 1623

N

18-

148_

814142

Rainy daysMean ± St.Dev.

185.1±20.8-

169.1121.3146.9 ± 25.6

_

214.5137.3192.6126.6326.7 ± 12.9247.6130.3

N

1715--7-842

TemperatureMean

27.123.7

--

25.7-

20.319.114.5

Max

30.924.5

_-

30.2-

25.622.216.5

Min

23.322.8

--

21.2-

15.316.312.6

10 P.M. Weevil data were analyzed from April 1983 throughNovember 1984.

Rainfall data are available, though not necessarily for theyears of our study, for all sites except Miramar and CorrienteGrande. The data are from Caballero (1978, and other annualvolumes in the same series) and Windsor (1990, and pers.comm.). The data from Punta Robalo, a few km north ofMiramar, also on the coast of the Laguna de Chiriqui, shouldprovide a reasonable estimate of the rainfall at Miramar. If thereis a difference, the precipitation at Miramar would likely beslightly higher because of its closer proximity to the mountains.For Corriente Grande the rainfall data from Changuinola are theclosest that are available and are used herein despite the factthat Changuinola is on the coastal plain and Corriente Grandeis south of there in a valley in the foothills between someridges. Temperature data for these two sites also are notavailable. The only data available are from Almirante, north ofMiramar and east of Corriente Grande. We would expectMiramar to have roughly the same temperatures as Almiranteand Corriente Grande, perhaps to be slightly cooler. Notemperature information is available for Las Cumbres. Meanannual rainfall, summarized in Table 1, varied from a little over1800 mm in Miramar to over 4500 mm in Fortuna, a differenceemphasized by the number of rainy days, varying from 147 inMiramar to 327 in Fortuna. In the lowlands mean temperaturesdid not vary much among sites, but, as expected, in themountains the temperatures were lower (Table 1).

At BCI, Las Cumbres, and Boquete there was a clearalternation of an 8-month rainy season and a 4-month dryseason, the latter usually occurring from mid-Decemberthrough mid-April (Figure 1). In the other three sites there wasno distinct dry season. In Fortuna there was a slight decrease inmonthly rainfall in March, and in Changuinola rainfall wasbimodal, with less rain both in February/March and inSeptember. Other sites in the province of Bocas del Toro, suchas Chiriqui Grande, also on the coast of the Laguna de Chiriquieast of Miramar, showed this same bimodal rainfall, which

makes it rather surprising that Miramar (assumed the same asPunta Robalo) did not show any seasonal changes. The dryseason at Guadalupe Arriba was not very pronounced and thereis a suggestion of bimodality in the rainfall. There was nodetectable seasonal variation in mean temperature (Figure 2)anywhere except, perhaps, in Almirante and Guadalupe Arriba,with a minimum around December/January, but even here theannual variation is very small, with at most three degreesCelsius difference between June and December.

All these climatic data were from rain gauges and thermome-ters set up in regular meteorological cabinets installed in theopen, whereas several of the light traps operated inside a forest(BCI, Corriente Grande, Boquete, and Fortuna) where condi-tions are likely to have been different. For temperature, suchdifferences are shown by data from BCI (Table 1; Figure 2).Mean maximum temperature inside the forest, at 1 m above theforest floor, was 6.4°C lower than that outside the forest,whereas the mean nightly minimum showed only an 0.5°Cdifference. We have no rainfall data from inside the forest, butone should keep in mind that part of the rain falling on thecanopy of the trees evaporates or is absorbed by the foliage andthus never reaches the forest floor, whereas another part comesdown as stemflow, a flow of water along the branches and treetrunks, rather than as "rain." For further details on the climateon BCI see Windsor (1990).

COLLECTING METHODS

All weevils (Curculionoidea) reported in the present paperwere collected by HW by means of modified Pennsylvanialight traps as described by Smythe (1982). However, in alllocalities except Las Cumbres, Boquete, and the first year onBarro Colorado Island, the glass collecting jar containingcarbon tetrachloride (CC14) as a killing agent was replaced bya stainless steel receptacle filled with Kahle's solution, amixture of alcohol, formaldehyde, and glacial acetic acid(Borror and DeLong, 1971). Kahle's solution killed the insects

500

0500

B.C.I. (N = 62)100 m

noo Brtfl

600

0500

Las Cumbres (N = 15)150 m

0500

Boquete (N = 14)1350 m

JSSL

0500

J F M A M J J A S O N D600 -T


Fortuna (N = 14)1050 m

Miramar(Punta Robalo, N = 8)0 m

Corriente Grande(Changuinola, N = 12)100 m

J F M A M J J A S O N DGuadalupe Arriba (N=2)2200 m

J F M A M J J A S O N DFIGURE 1.—Seasonal distribution of monthly rainfall in seven Panamanian localities.

much more quickly and generally preserved them in bettercondition.

Possible effects of the differences in the chemical environ-ment of the trap and killing methods were briefly studiedduring one month in January/February 1980 in Las Cumbreswhen two traps were set up six meters apart on a ridge behindthe house of HW, at the same spot the regular light trap hadbeen operating. One trap was operated with a glass jarcontaining CC14 as a killing agent (the "dry" trap), the otherwith a stainless steel receptacle with Kahle's solution (the"wet" trap). In one month the wet trap (Kahle's solution)

collected somewhat fewer weevils (99) than the dry (CC14) trap(125), but the difference was not significant statistically (x2(ld.fr.) = 3.02, p = 0.08), suggesting that an effect of the changefrom CC14 to Kahle's, if any, was fairly small.

Samples collected during the first two years at Las Cumbreswere sorted to species by HPS, and the specimens are depositedin his collection at the Smithsonian Tropical Research Institutein Panama. All other samples were sorted to species by CWOB,and the specimens are in his collection in Tallahassee, Florida.The many unidentified and undescribed species were givenreference codes. Special efforts were made to reconcile the

NUMBER 590

40

30 -

20 -

10 -

040

040

30 -

20 -

10 -

30 -

20 -

10 -

BCI - Clearing N=15

BCI - Forest FloorN = 17

Almirante N=7

J F M A M J J A S O N D

40

30 -

20 -i

10 -

BoqueteN«8


FIGURE 2.—Seasonal distribution of monthly means of maximum, mean, and minimum temperature in sixPanamanian localities.

codes used by CWOB and HPS, but in order to ensurecomparability of the data in the diversity section of the presentpaper, only the samples sorted by CWOB will be used, i.e., allsamples available except the first two years at Las Cumbres. Acomplete list of the species identified by CWOB is given in theappendix.

Every effort was made to identify the weevils to species, andwe are convinced that the vast majority, if not all, of the taxarecognized are good species. The exceptions were six taxa, oneeach in the genera Apion (Apionidae), Notiodes, Phyllotrox,and Terires (Curculionidae: Erirrhininae), and Paratrachelizusand Stereodermus (Brentidae). Each of these taxa may or maynot contain more than one species and are listed as Apioncomplex, Phyllotrox complex, etc. In these genera several

species were identified, but the remainder are referred to asbelonging to a "complex." These complexes, with localitiesand numbers collected, are listed in Table 2. In spite of thesemixed taxa, throughout this paper the designation "species"will be used when referring to individual taxa.

DATA ANALYSIS

At BCI and Las Cumbres, each collecting year started inearly March. For example, the year designated as 1976,extended from March 1976 to March 1977, etc. Yeardesignations in this paper do not necessarily coincide withcalendar years.

DIVERSITY.—In order to compare diversity in different sites


TABLE 2.—List of the six weevil "taxa" recognized that probably are not single species (BCI = Barro ColoradoIsland, LC = Las Cumbres 1976, MIR = Miramar, CGR = Corriente Grande, BOQ = Boquete, FORT = Fortuna,GUA = Guadalupe Arriba.)

ApionidaeBrentidaeBrentidaeCurculionidae

CurculioninaeErirrhininaeErirrhininae

Taxon

Apion complexParatrachelizus complexStereodermus filum complex

Terires complexNotiodes aeratus complexPhyllotrox complex

TOTAL

BCI

509-

173

8821-

4312152624

LC

_

--

-3

474477

MIR

620

1

3-

25642594

CGR

42-

7-

115128

BOQ

9-

12

--

2950

FORT

18-6

--

204228

GUA

_

--

--33

TOTAL

54622

191

88313

4651056103

with different numbers of individuals, as well as differentnumbers of species, diversity indices can be useful. In spite ofsome problems, we prefer the "alpha" (a) parameter of thelogseries (Fisher et al., 1943) as a diversity index over otherpopular indices, such as the Shannon-Weaver index (Taylor etal., 1976; Wolda, 1983a, 1984). If the basic assumptions for theuse of the index are fulfilled, the diversity at different sites canbe compared directly in spite of differences in the number ofindividuals. Moreover, it seems that the usefulness of a is fairlyrobust to departures from the assumptions, especially depar-tures from the logseries distribution.

Comparisons between collections of weevils from differentsites or different years can be made best by means of similarityindices. Most such indices present problems of some kind(Wolda, 1981), but the NESS index (Grassle and Smith, 1976;Smith et al., 1979), a generalization of the Morisita index(Morisita, 1959), is better than most in that its expected valuefor two samples from the same population is 1, its value doesnot depend on the diversity of each of the samples beingcompared and does not depend on sample size, except possiblyfor very small samples. The advantage of the NESS index overthe Morisita index is that it takes into account the rarer speciesand that it has a variance estimate attached so that the indicescan be used for statistical testing. The "m" parameter of theNESS index was taken as 20 wherever possible.

SEASONALITY.—Two phenomena are involved here. Withina year the abundance of an insect species may vary significantlyand the pattern of this variation may or may not repeat itselfover successive years. If it does, and thus is cyclical over time,it is referred to as truly seasonal; if not, it is not truly seasonal.One might argue that the term seasonal(ity) should not be usedunless the observed pattern is proven to repeat itself from yearto year. However, we find that avoiding the term seasonal(ity)confuses more than it helps. We will use seasonal andseasonality throughout this paper when referring to changes inabundance within a year, even if those changes were differentin different years.

For seasonality analysis, only species for which at least 10individuals were collected, i.e., with at least 10 data points onthe annual cycle, were considered. It is impractical to present

graphs of the hundreds of seasonal patterns that we observed.Therefore, to provide some semblance of order to the plethoraof seasonal patterns, we decided to use circular statistics(Batschelet, 1972, 1981; Mardia, 1972), the weeks and monthsbeing basically circular rather than linear variables. The lengthof the "Mean Vector" r is an indication of the concentration ofthe individuals in some part of the year. Throughout the paper,"Mean Vector" is used as a shorthand for "length of the MeanVector." The time of occurrence of each individual is calculatedas an angle (<))) in degrees around the annual circle. Forinstance, for an individual found in week number 25 the angle<|> is 25/52*360 = 173 degrees. If x is the average of the cosinesof all <|>-values in a year and y is the average sine, the MeanVector r is given by

The Rayleigh test (Batschelet, 1981) applied to this MeanVector shows whether or not the distribution of the individualsover the year is significantly different from uniform orsymmetrical. The direction of the mean vector over the yeargives the "Mean Week," which is given by transforming themean angle <J) back to weeks

•={arc tan (y/x) ifx > 0180° + arc tan (y/x) ifx < 0

and the Jupp-Mardia correlation coefficient r2 (which can belarger than unity) correlates two circular variables (Batschelet,1981:190). The "Angular Deviation" s is the circular equivalentof the standard deviation and is given by

5 = V2( l - r )

To compare seasonality patterns in different years, for eachspecies the average Mean Vector with its standard deviationwas calculated, as well as the (circular) average Mean Weekand its angular deviation, the Mean Week itself also being acircular variable. The relations between the two standarddeviations and Mean Vector, as well as Mean Week, provide auseful summary of the between-year or between-site variationin seasonality patterns.

NUMBER 590

In addition to these circular measures of seasonality, the"Peak Week" was used, which gives the mode of the seasonalabundance pattern. It is calculated by taking the running4-week average of the number of individuals, wrapping aroundthe end and the beginning of the year because of the circularnature of the data, and finding the maximum of these runningaverages. The running averages rather than the original datawere used in order to eliminate possible effects of the phases ofthe moon on the catches.

Other measures also were tried. The "Seasonal Maximum"(SM) (Wolda, 1979) is obtained by dividing the maximumnumber of individuals observed in a 4-week period by the totalnumber of individuals observed over the year and multiplyingit by 13, the number of 4-week periods in a year. SM runsbetween 1 (distribution over the year entirely uniform) to 13(all individuals found in one period of four weeks). Asexpected, SM correlates with the Mean Vector. Using theresiduals of the regression of SM on Mean Vector did notprovide any useful extra insight into the nature of the seasonalpatterns; therefore the relevant calculations are not included inthe present paper.

Morisita's "index of seasonal diversity" (1/I6) (Morisita,1967; Yamamoto, 1974) is calculated as

TABLE 3.—Between-year changes in abundance of weevils on BCI in somefamilies and subfamilies classified either as woodborers or as non-woodborers.

q N(N- 1)

where nx = number of individuals in class i

q = number of classes

If the classes are months, q = 12, if weeks, q = 52, etc. Thehigher the value of 1/I6, the more homogeneous the seasonaldistribution. As will be shown (Figure 17), 1/I8 correlates withthe Mean Vector. Although useful when used by itself, it, or itsresiduals of the regression of it on Mean Vector, do not addanything useful to the information already available; thereforethe relevant calculations are not included in the present paper.

Circular statistics, as far as we know, deal basically with neatsymmetrical unimodal seasonal distributions, ideally conform-ing to a Von Mises distribution (Batschelet, 1981), the circularequivalent of a normal distribution. Actual seasonal patterns ofinsects, however, rarely fit this description, which is why thePeak Week, the mode of the seasonal pattern, rarely coincidesprecisely with the mean of this pattern, the Mean Week. Thedifference between the two measures gives an indication ofskewness of the seasonal distribution. This was found more

IndividualsSpeciesWoodborers:

Brentidae (less Ulocerus)Ulocerus sordidus

CossoninaeCryptorhynchinaeDryophthorinaeMagdalidinaeZygopinae

TotalIncrease/Decrease

Total less UlocerusIncrease/Decrease

Non Woodborers:AnthonominaePolydrosinaeCamarotinaeCeratopodinaeCeutorhynchinaeErirrhininae (less Phyllotrox)

Phyllotrox complexHyperinaeEntiminaePrionomerinaeTychiinae

TotalIncrease/Decrease

Total less PhyllotroxIncrease/Decrease

1976

18293586

269113575

156440

227

2752

2639

7618

131

2671

96172032

10423

806

1977

29522872

548315

22523220

30

433

6771146.0%

6456144.6%

361580

1070

182312781

3317

11

1518245.7%

2401197.9%

1978

47518906

1016622433303572

232

383

14550114.9%

832629.0%

49510

1450

114420723

2613

15

2255348.6%

1830-33.8%

illustrative of the patterns occurring than the formal measure-ment of circular skewness.

Results

GENERAL REMARKS

EFFECT OF TRAP DESIGN AND OF THE DYING Tachygalia

TREE.—The difference in killing methods and thus in thechemical environment of the light traps between Las Cumbres,Boquete, and the first year on Barro Colorado Island on onehand, and the other localities and years on the other, may ormay not have affected the efficiency of the traps. The LasCumbres experiment (see "Collecting Methods") suggestedthat the effect, if any, was minimal. However, an analysis ofcollections made before and after the change from CC14 toKahle's solution between 1976 and 1977 on BCI also may beinformative.

The total number of individuals and species of weevilscaught each year on Barro Colorado Island is given in the firsttwo lines of Table 3. At first sight these data suggest that thechange from CC14 to Kahle as killing agents directly after 1976

8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

resulted in a substantial increase in both the number of speciesand the number of individuals. There was, however, animportant event that occurred in 1978 that clouds the issue. Thelight traps were suspended from a large tree, Tachygaliaversicolor Standl. and Wms. (Leguminosae: Caesalpinoideae).This tree species is monocarpic, i.e., it flowers once during itsentire life and then dies immediately (Foster, 1977). It is notquite clear whether the tree produces flowers because it is in theprocess of dying or whether flowering kills the tree. The tree towhich the light traps were attached flowered in May 1978,attracting a large number of insects to its flowers (Wolda andRoubik, 1986) and to its apparently already dying wood. Thecondition of the tree deteriorated so rapidly that in October ofthat year branches were already falling off and the trap had tobe moved to a tree nearby. Many weevils were among theinsects aggregating at the dying tree. The large increase in thenumber of individuals between 1977 and 1978 (Table 3) waslikely, at least in part, to have been associated with the death ofthe tree. The increase in the number of species, however, wasminor compared with that in the previous year. Very little isknown about the physiology of Tachygalia trees before andduring flowering and its effects on visiting weevils. Nor is itclear whether the attraction to weevils started well before theflowering process began. Too little is known also about the lifehistories of weevils (Anderson, 1993), especially neotropicalones, to look specifically at species that might be attracted todying Tachygalia trees. However, based on what is known inthe literature and on the personal experience of two of us (COBand HPS), we classified families and subfamilies as either"woodborers" or "nonwoodborers," the latter including speciesthat are leaf-miners, seed-, flowerbud-, or fruit-predators, stemborers of herbaceous plants, leaf-feeders, etc. A number ofweevil families or subfamilies that could not easily be soassigned to either of these groups are ignored herein.

Many of the species classified as woodborers may not havebeen attracted to dying Tachygalia trees, but species that wereare most likely to belong to this category and much less likelyto be nonwoodborers. From 1977 to 1978 there were largeincreases in the number of individuals of both categories (Table3), but this was much more evident among the woodborers.Much of the increase in individuals in 1978, when the treeflowered, was due to only a few species, especially Ulocerussordidus Sharp, a woodborer (Brentidae), and Phyllotroxcomplex (Erirrhininae), a non-woodborer flower visitor, espe-cially palm flowers. The latter taxon was by far the mostcommon and was possibly attracted to the flowers of theTachygalia tree. These two taxa alone contributed 13851individuals to the overall increase of 17996 weevils between1977 and 1978. Spectacular increases between these two yearsalso were found in a number of other species, such asCryptorhynchinae sp. #66 (from 2 to 527), Conotrachelus sp.#6 (from 2 to 73) and Cryptorhynchinae sp. #C155 (from 2 to57). The increase in abundance among the woodborersassociated with the flowering tree, not counting Ulocerus

sordidus, was still 29%, whereas the increase from 1976 to1977 remained at 145% (Table 3). Among the nonwoodborers,excluding the Phyllotrox complex, the numbers decreased from1977 to 1978 by 34%, whereas from 1976 to 1977 the totalnumber of nonwoodborers almost tripled (Table 3). There areobviously large differences in population behavior betweenwoodborers and nonwoodborers, but these do not necessarilysolve the problem. The increase among the woodborers from1977 to 1978 compared with a decrease in the nonwoodborersfits the hypothesis of a strong attractant at the dying of theTachygalia tree. The large increase in Phyllotrox complexdemonstrates that other factors not associated with the dyingtree may be operating. Some of the abundant palms in the traparea may have flowered in 1978, which would account for thelarge numbers of Phyllotrox.

The large to very large increase at this locality among boththe woodborers and the nonwoodborers from 1976 to 1977does not fit the hypothesis of an early attraction by woodborersto the Tachygalia tree. The problem is also that the species thatshowed a rather spectacular increase from 1977 to 1978 werenot necessarily the ones that increased in the previous year, asone would expect if the hypothesis of an early attraction weretrue. In fact, for those species that had at least 20 individuals inthe three years combined, the ratios of abundances in 1977 overthose in 1976 versus these ratios of 1978 over 1977 had ahighly significant negative correlation (r = - 0.267, n = 265).This shows that increases tended to be followed by decreasesand vice versa. Of the species that increased between 1977 and

1978 (47.5% of the total), only 40.3% showed an increase in1976/1977, whereas 15.1% decreased and 44.6% remaining thesame. The vast majority (90.6%) of the species that decreasedbetween 1977 and 1978 (37.3% of the total) had increased inabundance in the previous year. All these points argue againstthe idea of a general early attraction of weevils to the dyingTachygalia tree. The fact that almost 60% of the speciesincreased between 1976 and 1977, whereas only 17%decreased, strongly suggests that some other factor did improvethe trap catches in 1977 and the change in chemicalenvironment of the trap might provide the explanation. If the1976/1977 increase was due to the chemical change, theKahle's solution produced better weevil catches than CC14,which would run against the (nonsignificant) trend observed inthe experiment in Las Cumbres. Numbers of insects dofluctuate from year to year, and in 1977 obviously many moreweevils were caught than in 1976. Whether this was caused bythe chemical changes in the trap or by some other unknownfactor remains an open question. The Tachygalia tree did attractlarge numbers of woodborers when it flowered in 1978, but itprobably did not do so in the year before flowering.

DIVERSITY

PER YEAR.—A total of 113,712 weevils were collectedrepresenting 2030 species (Table 4; Appendix), not counting

NUMBER 590

TABLE 4.—Collecting information, number of individuals, and number of species in seven sites in Panama asrevealed by light-traps, plus the alpha diversity index based on the logseries. (Altitude is given in meters.HPS = H.P. Stockwell; CWOB = C.W. O'Brien.)

Locality

Barro Colorado IslandLas Cumbres (HPS)Las Cumbres (CWOB)MiramarCorriente GrandeBoqueteFortuna (Canopy)Fortuna (low level)Guadalupe Arriba

Total (excl. Las Cumbres HPS)

Altitude

120150150

0100

1350105010502200

Yrs

32'/2

11

'/3

23/4

2l3/4

Period

III 1976-IH 1979X 1973-III1976

III 1976-III1977IX 1978-IX 1979

I-V 1980VII1976-VII1978IX1976-VII1977

VII 1977-VII1979IV 1983-IX 1984

Individ

9533369154011399222292256

8604640

391

113712

Spp.

1239315357170259267195367

51

2030

Spp/Yr

788231360172

_

183_

24836

Alpha

201.0 ±5.768.0 ± 3.894.7 ± 5.036.1 ±2.875.9 ± 4.778.8 ±4.878.7 ± 5.693.9 ±4.915.7 ±2.2

351.6 ±7.8

Alpha/Yr

148.556.597.236.6_

62.9_

70.613.0

the 6915 individuals in 315 species from Las Cumbresanalyzed by HPS. By far the largest collections were made onBarro Colorado Island. Weevils were collected here for threeyears, but were collected for two years or less at the other sites(Table 4). However, the average number of species per year onBCI (788) was much higher than at any other site. The numbersof individuals in the three years on BCI were 18293, 29522,and 47518, respectively, whereas the largest numbers capturedin any year at any of the other sites were only 3992 in Miramarin 1979 and 4011 in Las Cumbres in 1976. On BCI two trapswere used as compared to only one trap each at all other sites.On BCI the traps were operated all night, as they were in LasCumbres and Boquete, but not at the other sites. However,these differences alone seem insufficient to explain the largebetween-site differences in species diversity. Obviously, BCI isan area much richer in weevils than any of the other sites. Thesecond richest area may be Corriente Grande, where 259species were collected in only four months. The area with thelowest number of species, on the other hand, is undoubtedlyGuadalupe Arriba, at 2200 m altitude, with only 51 speciescollected in 20 months. The values for the diversity index a,with their standard deviations, and the mean a per year, whereappropriate, are given in Table 4. The fact that the a for a sitewith the years combined was higher than the average a per yeardemonstrates a between-year heterogeneity in species composi-

tion, and the huge value of a (351.6) found when combining allsites, points to an enormous between-site heterogeneity inspecies composition (see below).

The distribution of species abundances at each site issummarized in Table 5. Not many species were common, asillustrated by the fact that 28% (BCI) to 51% (CorrienteGrande) of all species collected were represented by only onespecimen. On BCI only 31% of the species were represented by10 or more individuals, and at the other sites only 11.8%-15.8% equaled or exceeded this number. Nevertheless, sometaxa were very common, especially the Phyllotrox complex atBCI and Miramar. The three most common taxa at each site hadbetween 25% and 75% of all individuals collected, and at BCIand Miramar the single commonest taxon was represented by45% and 65% of all individuals. To test the formal validity ofthe a diversity index, the distribution of species abundanceswas tested against the expected distributions according to thelogseries. The abundances were classified in "x3-classes," thatis, class boundaries were multiplied by three, starting with thelowest boundary at 0.5. Class 1, then, is between boundaries 0.5and 1.5 (1 individual), class 2 between 1.5 and 4.5 (2-4individuals), class 3 between 4.5 and 13.5 (5-13 individuals),etc. The actual and the logseries distributions for the weevilsfrom BCI are compared in Figure 3, demonstrating the largedifferences that are highly significant (X2(7) = 337.0,

TABLE 5.—The number of species with only 1, at least 5, and at least 10 individuals as percentages of the totalnumber of species. The number of individuals in the combined three most common species at each site isexpressed as a percentage of the total number of individuals, as is the number of the single most common taxon.

Locality

Barro Colorado IslandLas Cumbres (CWOB)MiramarCorriente GrandeBoqueteFortunaGuadalupe Arriba

% Spp.1 indiv.

28.038.949.451.047.645.043.1

% Spp.>5 ind.

42.026.921.821.224.324.425.5

% Spp.>10 ind.

31.015.111.812.714.215.815.7

% Indiv.3 Common

61.4627.9079.2225.7122.2129.5347.31

% Indiv.Commonest

45.3012.3565.2314.447.54

11.8328.39

Commonest taxon

Phyllotrox complexPhyllotrox complexPhyllotrox complexTerioltes sp. #1Micralcinus sp. #1Micromimus continuusCossonus sp. #5


BCI-Weevils

6wz>oLUCC

1000g

100=

10=

1; 1258 Species

95333 Individuals

Alpha = 204.7 ±5.8

1 3 4 5 6 7 8 9X3-CLASSES IN ABUNDANCE

12

Weevil Data Logseries

FIGURE 3.—Distribution of abundances of weevils (Curculionoidea) collected in light traps on Barro ColoradoIsland, as compared with the logseries distribution.

coUJaa.05

1000

900-

800-

700-

600-

400-

300-

200-

100-

y = 22.128 + 4.360 V5T

0.94

10

Guad.Arrib.

joquete

100i i 11 in 1—

1000INDIVIDUALS

11ii 1—i—i 111

10000 100000

FIGURE 4.—Relationship between number of species and number of individuals of weevils (Curculionoidea)caught in light traps in each year at seven Panamanian localities.

NUMBER 590 11

1000

wLUOLU

a.CO

800-

600^

400-

200-

B.C.I.

• Las Cumbres

• Corr.Grande

Miramar

Fortuna• Boquete

Guad.Anib.

500 1000 1500ALTITUDE (m)

2000 2500

FIGURE 5.—Relationship between altitude and number of species of weevils caught by light traps in each year atseven Panamanian localities.

p « 10"10). Compared with the logseries distribution, far toomany rare species, represented by only one or two individuals,and far too few species of intermediate abundance were found.This means that interpretation of the actual values of thea-indices found must be done carefully. However, thedeviation from the logseries at all sites was in the samedirection as for BCI and was highly significant, except forGuadalupe Arriba where no statistical significance could beexpected with the low numbers available. The a-index,therefore, can still be used to compare years and sites. Theactual relationship between the number of species and thenumber of individuals in each year at each site is plotted inFigure 4, showing again the richness of the weevil fauna onBCI and the relative poverty of the weevils from Miramar and,especially, from Guadalupe Arriba. In terms of both numbers ofindividuals and species, Corriente Grande was at least as rich asFortuna, in spite of the fact that insects were collected there foronly four months. The lowest value at Fortuna was for ninemonths in the canopy, the other two points for this site eachrepresent a full year of collecting at a near-ground level. Afive-year study on Homoptera at BCI (Wolda, 1987) showedthat canopy traps tended to collect more individuals but fewerspecies than did traps in the understory. The same was true forweevils at all the sites (see below), except at Fortuna. Thelowest numbers for BCI was for the first year with a differentchemical trap environment and well before the Tachygalia tree,from which the traps were suspended, started flowering (see

above). Part of the differences between sites may have been dueto differences in altitude (cf. Wolda, 1987). There was asignificant decrease in species richness with increasing altitude(Figure 5, p = 0.009), but this was due mostly to the pointsrepresenting the rich fauna at BCI. Without the BCI data therelationship was still significant (p = 0.014) but was almostexclusively due to the data from Guadalupe Arriba, suggestingthat an altitude effect for weevils was not found from lowlandsto intermediate altitudes, but was found only when higherelevations were included. Essentially the same picture wasobtained with the relationship between individuals or a againstaltitude (data not shown). Part of the decrease in diversity withincreasing altitude in these light-trap samples is undoubtedlydue to a decreasing diversity of the fauna that was sampled, butpart of it also may be related to the lower temperatures (Figure2), and an associated decreased flight activity, at higheraltitudes.

PER WEEK AND PER DAY.—All of the above discussion wasabout the diversity of samples representing entire years.However, within each year, diversity, whether expressed asspecies richness or as the diversity index a, is far from constant.This is illustrated for BCI in Figure 6. For some weeks thevalue of a was omitted because it was meaningless in suchsmall samples. For instance, with four individuals and fourspecies in a particular week, a reached the ridiculously highvalue of 264. In one instance (early May, 1977), a also reachedan extremely high value of 275, which was based on a sample


4000 -

2000 -

= Full Moon

300

200 -

100 1

Alpha

1976 1977 1978FIGURE 6.—Diversity (individuals, species, and alpha) per week over three years in light-trap samples collectedon Barro Colorado Island, Panama (timing of full moon also indicated).

of 56 individuals and 51 species. We were inclined to omit thisvalue too, but with this sample size such an omission would notbe justified, and it is included in Figure 6. Both species richnessand the diversity index a had a strong maximum in May-June,at the beginning of the rainy season, and then the number of

species gradually tapered off toward a low in the dry season,especially January through March. There were weeks duringwhich about 300 species were collected, as well as weeks whennone or almost none were taken. There is a highly significantcurvilinear relationship between the number of individuals and

NUMBER 590 13

TABLE 6.—Number of species and number of individuals in each family and subfamily of weevils at sevenPanamanian localities. (The Las Cumbres data are for 1976 only.)

T&xon

ANTHRIB1DAEAPIONIDAEATTELABIDAEBRENTIDAECURCULIONIDAE

ANTHONOMINAEBARIDINAECAMAROTINAECERATOPODINAECEUTORHYNCHINAECOSSONINAECURCULIONINAECRYPTORHYNCHINAEDRYOPHTHORINAEENTIMINAEERIRRHININAEEUGNOM1NAEHYPERiNAEMAGDALIDINAEMOLYTINAEOTIDOCEPHALINAEPETALOCHILINAEPOLYDROSINAEPRIONOMER1NAERHYNCHAENINAERHYTIRRHININAERHYNCHOPHORINAETYCHIINAEZYGOPINAE

TOTAL

Spp.

8040

968

3119

113

147

2440

31

33221

24416

115700

114

148

1239

BCIIndiv.

7751444

1278485

932137

1283

2615788898356

302

467591361

29202

583

771300

138028

2117

95333

Las CumbresSpp.

9131

11

73010

102

16002

11320

101212101329

357

Indiv.

17181

73

93020

461138740

08

lie1470

14672

238620

57564

47

4011

MiramarSpp.

213

110

51050

161

57207210

30102201209

170

Indiv.

426

157

610

120

4863

162150

3044710

11610

1440

1380

11

3992

Corr. GrandeSpp.

162

14

112011

441

78409100

3340000070

40

259

Indiv.

1112

31

157

011

8467

307210

0579

100

67400000

240

79

2229

BoqueteSpp.

3125

11

56070

150

75009010

7700520041

29

267

Indiv.

4399

31

1460

250

4000

55000

104010

90500

13200

581

94

2256

FortunaSpp.

393

12

314070

210

12120

14000

16210

11210

110

39

436

Indiv.

5594

96

524

025

02198

0844525

0398

000

101710

45310

1420

108

5500

Guad.Spp.

0010

06000209106000

17000010017

51

Arr.Indiv

0020

073

000

1140

1230

75000

8800001006

17

391

the number of species (r2 = 0.676, n = 155) and a significantlinear correlation between the number of species and a(r2 = 0.301, n = 150). In fact, there were times when about 200species were collected in a single day (see below, Figure 12),whereas on other days no weevils were caught. The phases ofthe moon often have a strong effect on light-trap catches ofinsects, the numbers of individuals and species being reducedat full moon. The weevils as a group, however, seem at bestonly to be weakly sensitive to the phases of the moon (Figures6,11, 12), and many species may not be subject to effects of themoon at all. Based on these data, diversity varied stronglyseasonally, but not at all or very little with the moon cycle.

FAMILIES AND SUBFAMILIES.—The weevils collected be-longed to five families and, within the Curculionidae, to 24subfamilies (Table 6). The large number of individuals in theErirrhininae belonged mostly to the composite taxon "Phyllo-trox complex" (Table 2), which was the most abundant taxon atBCI, Las Cumbres, and Miramar (Table 5). The Cryptorhyn-chinae and Molytinae were by far the most species-rich groups

at all sites. The next most species-rich groups were theZygopinae at BCI, Boquete, and Fortuna, and the Cossoninae atMiramar and Corriente Grande. At the other extreme, Magdal-inidae were represented by only one species at one site. On BCIthere were relatively many species of both Anthribidae andBrentidae, far more than at any of the other seven localities. Thelow numbers of Camarotinae were in all probability mostly dueto a failure to recognize these flat leaf-miners as weevils whenthese light-trap samples were sorted, as they were found ingreater numbers in samples from later years from these sametraps on BCI (Barria, pers. comm.).

BETWEEN-SITE DIVERSITY.—Between 30% and 69% (aver-

age 50%) of all species caught at a site were not collected at anyof the other sites (Table 7), 18%-47% of the species wereshared with only one other site, and only 1.9% (BCI) to 20%(Miramar) of the species found at one site were found also inthree or more other localities. Only the Phyllotrox "complex"was found at all sites, and these probably were different speciesin different sites in most cases. Table 8 contains the number of


TABLE 7.—Percent of species from each locality occurring at one or more of the other sites.

Locality

Barro Colorado IslandLas Cumbres (CWOB)MiramarCorriente GrandeBoqueteFortunaGuadalupe Arriba

Totalspecies

1239357170259267436

51

Restrictedto site

64.330.532.440.249.458.768.6

Sharedwith only

1 site

25.247.330.632.125.523.517.7

Sharedwith only

2 sites

4.515.417.114.717.211.67.8

Sharedwith only

3 sites

1.44.2

13.59.34.53.63.9

Sharedwith only

4 sites

0.250.842.941.51.11.10

Sharedwith only

5 sites

0.251.402.941.931.871.110

Sharedwith all6 sites

0.050.280.590.390.370.231.96

TABLE 8.—Number of species shared between seven Panamanian localities, asshown by light-trap samples. (BCI = Barro Colorado Island, LC = LasCumbres, 1976, MIR = Miramar, CGR = Corriente Grande, BOQ = Boquete,FORT = Fortuna, GUA = Guadalupe Arriba.)

Total speciesBCILCMIRCGRBOQFORT

BCI LC

1239 357239

_

Number of species

MIR

1708745_

CGR

2591033350-

BOQ

26788261929-

FORT

43611023275873-

GUA

5141254

13

species shared between each pair of sites. All sites butGuadalupe Arriba shared more species with BCI than with anyother site; Guadalupe Arriba shared more species with Fortuna.A better indication of the between-site diversity, however, isgiven by the matrix of NESS similarity indices (Table 9). The

between-year, within-site indices were between 0.90 and 0.95.These values were high, but still significantly different fromunity, pointing to real between-year differences in speciescomposition. The between-site comparisons, however, yieldedmuch lower index values, emphasizing again the high degree ofendemism referred to above (Table 7). The highest among theselow values were between BCI, Las Cumbres, and Miramar, thetwo Atlantic sites and the one Pacific site that does not have ahighland barrier with the Atlantic side of Panama. The thirdAtlantic site, Corriente Grande, also tended to be more similarto these three sites than to the remaining sites with theexception of Fortuna. Fortuna had the highest similarity withBoquete, another highland site only 20 kilometers away, albeitin a very different ecological and climatological situation, andwith Corriente Grande. Guadalupe Arriba had extremely lowsimilarity values with all other sites, which in part may havebeen an artifact caused by the low numbers found at this site.Thirteen of the 52 species found there also occurred at Fortuna,suggesting a higher affinity between these sites than the NESSindex suggests.

TABLE 9.—Within- and between-site diversity in Panamanian weevils as expressed by the NESS-similarity indexand its standard deviation. For Las Cumbres, the within-site comparison is between the first two years, identifiedby H.P. Stockwell, and the between-site comparison is for the last year, identified by C.W. O'Brien. For Fortuna,only the two years with the trap at a low level were considered; for Guadalupe Arriba, the second year was only8 months long.

Within Site:yr. 1 vs. yr. 2yr.2 vs. yr.3yr. 1 vs. yr.3

Between Sites:BCILas CumbresMiramarCorr. GrandeBoqueteFortuna

BCI

0.917 ±0.0030.902 ± 0.0030.892 ± 0.003

-

Las Cumbres

0.919 ±0.007--

0.378 ± 0.005-

Miramar

_-

-

0.469 ± 0.0040.384 ± 0.008

-

Corr. Grande

__

-

0.281 ±0.0100.236 ± 0.0090.359 ±0.012

-

Boquete

0.907 ±0.015_

-

0.148 ±0.0100.172 ±0.0120.161 ±0.0130.152 ±0.010

_

Fortuna

0.899 ± 0.009_

-

0.168 ±0.0060.116 ±0.0050.194 ±0.0080.246 ± 0.0110.240 ± 0.011

-

Guadalupe Arriba

0.946 ± 0.034_

-

0.035 ± 0.0190.027 ±0.0150.038 ± 0.0200.026 ± 0.0110.009 ± 0.0040.058 ±0.017

NUMBER 590 15

1976 1977YEARS

1978

FIGURE 7.—Vertical distribution of weevils in forest on Barro Colorado Island as revealed by light traps: A,percentages of individuals found in canopy trap in each of three years; B, percentage of all species found in trapat ground level; C, percentage of all species found in trap in canopy.

CANOPY VS. GROUND LEVEL.—One BCI trap was locatedtwo to three meters above the forest floor and the other was inthe canopy of a tall tree. The weevil faunas occurring at thesetwo places conceivably could be very different, so that ananalysis of differences in catches between the two traps mightbe rewarding. However, the canopy trap was visible from a fairdistance and from below, whereas the low trap was less visiblefrom the canopy. Thus, some possible habitat differencesbetween canopy and ground level may be less evident in thetrap catches.

In each of the three years on BCI, between 55% and 59% ofthe individuals were caught in the canopy (Figure 7). Fifty-sixpercent to 64% of the species were collected in the canopy trap,whereas 73% to 77% were found in the trap at ground level.Because of the large number of species common to both traps,the percentages recorded for the two traps totalled more than100%. The distribution of the species, according to thepercentages found in the canopy trap for all three yearscombined, is given in Figure 8. Species that had more than 10,more than 50, or more than 100 individuals are indicatedseparately. There obviously is a large variation among thespecies in vertical distribution, in common species as well as inrarer species. The NESS-similarity index, comparing thecatches at the two levels for all three years combined, was0.884 ± 0.002. This figure is high but is still significantly lowerthan the between-year values (Table 9), showing that between-

level differences are larger than those between years. Of the106 species in 1976 represented by more than 10 individuals,16 species had none and five had all individuals in the canopytrap. Of the 213 such species in 1977, these numbers were 14and 15, respectively; of the 231 such species in 1978, thenumbers were 21 and 26, respectively. This suggests sometendency for some species to be restricted to one forest stratumor the other. For the 23 species in 1976 represented by at least50 individuals, three species had zero and none had allindividuals in the canopy; of 58 such species in 1977, five hadnone and three had all individuals in the canopy; in 1978 thesenumbers among 61 species were two and two, respectively. Soeven in these common species, only a few were restricted toeither the canopy or to a low level in the forest. There aredifferences between what was caught at the two levels, butthese are not spectacular and there is nothing in the present datathat points to a mythical special fauna in the canopy. If there isa special fauna of weevils in the canopy, the species involvedeither do not come to light or they come to both the canopy andthe low trap.

For most species the vertical distribution in the forest wasconsistent from year to year. The fraction in the canopy is givenin Figure 9 for all species that had at least 50 individuals bothin 1977 and in 1978. Graphs made for the other between-yearcomparisons were similar. The vast majority of the speciesclustered around the diagonal showing between-year consis-


30 40 50 60 70PERCENT IN CANOPY

90 100

N>100 50<N<100 10<N<50

FIGURE 8.—Distribution of percentages of all individuals of common species of weevils found in canopy trap onBarro Colorado Island, all three years combined.

tency, but a few did not. For these there was a much higherfraction in the canopy in 1978 than in 1977. Three of thesespecies also were represented by more than SO individuals in1976. For two of these, the percentages in 1976 were identicalto those in 1977 (Paratrachelizus complex (1) 0% (N = 50 and82) both in 1976 and 1977 and 37.8% (N= 189) in 1978,Isotrachelus tibialis (3) 51.8% (N=139) in 1976, 51.7%(N = 209) in 1977 and 100% (N = 159) in 1978), whereas forthe third species (Cryptorhynchinae sp. #C-66 (4)) 1976 wassimilar to 1978 (80.3% (N = 369) in 1976, 6.8% (N = 148) in1977, and 73.1% (N = 725) in 1978).

DISCUSSION OF DIVERSITY

SPECIES RICHNESS IN PANAMA COMPARED WITH NONTROPI-

CAL AREAS.—Weevil collectors usually do not consider lighttraps to be an effective tool and overwhelmingly prefer othermethods, such as beating or sweeping. For instance, even whenthe light source is optimized with respect to the spectralsensitivity of the eyes of the weevil concerned, the light trap isnot considered useful as a survey tool for Diaprepes abbre-viates (L.) (Beavers et al., 1979). The ineffectiveness of thelight trap also is recognized by weevil collectors in theneotropics (CWOB, HPS, unpublished data). A comparison oflight-trap data in the present study with the results from othercollecting methods shows the strong selectivity of light-trap

samples. Locally rich and diverse groups, such as the Baridinaeand Polydrosinae, are poorly represented in the light traps. Forinstance, a little over one-half of the 343 known Panamanianspecies of Cryptorhynchinae (CWOB, unpublished data) arerepresented in our light-trap samples, as compared with aboutone-third for Molytinae and Zygopinae. On the other hand, inthe subfamilies Polydrosinae and Baridinae the percentagesrepresented are 12.1 and 5.1, respectively. In forest andwoodland habitats, light traps seem to be much more effectivein collecting woodborers than in collecting other groups. Thesewoodboring weevils attack dead or dying wood. Species of thefamily Cerambycidae that attack dead wood do come to light,whereas species of Cerambycidae that attack live wood do not.It is not clear why there is a correlation between attraction tolight and the type of substrate attacked. It is also unknownwhether such a correlation exists among weevils. Overall, it isestimated that only 25% to 40% of the weevil species presentwere collected at light. Of the 1666 species in the list of knownPanamanian species (CWOB, unpublished data), 612 (36.7%)were represented in our samples (the other species we collectedare unidentified, see "Appendix"). The light-trap projectreported on herein was actually designed to collect insects otherthan weevils. Yet in the tropical environment in Panama, thecatches of the weevils were surprisingly large, with thousandsof individuals being collected per year in all lowland sites andeven in most years at intermediate altitudes in the mountains.

NUMBER 590 17

0.00.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9

Fraction in Canopy 1978

FIGURE 9.—For all species of weevils with at least 50 individuals in each of the two years 1977 and 1978, fractionof those individuals found in canopy in 1977 plotted against that fraction in 1978. Six species that were ratherdifferent in the two years were (1) Paratrachelizus complex, (2) Pseudoantkonomus tachyon Clark, (3)hotrachelus tibialis (Champion), (4) Cryptorhynchinae sp. #C-66, (5) Caulophilus sp. #7a, (6) Conotracheluspunctiventris Champion.

Only at higher altitudes did sample size decrease dramatically.Also, the average number of species per year varied from 36 atGuadalupe to 788 at BCI.

We are not aware of any analysis of light-trap samples ofweevils from temperate areas. Most colleagues who work withlight traps in the temperate zone tell us that their traps collectvery few weevils (Witkowski, pers. comm.; Spitzer, pers.comm.). However, samples taken by other methods, such assweep-nets, are reported in the literature, especially fromeastern Europe. The contrast in species richness between thosesamples and the Panamanian light-trap samples is striking. In aDanish beech forest, the total number of species obtained overa period of up to three years, using a variety of collectingmethods, was only 29, with only 11 to 14 species per yearobtained with any one method (Overgaard Nielsen, 1974a). Inthe understory of a woodland in England (Phillips, 1992), thetotal number of species collected was 28, varying from one to23, whereas in southern England Owen (1993) found 34species of weevils. The Polish studies yielded some-what more species, with a total of 128 species taken over aperiod of three years at 13 sites near Zabierzow (Witkowski,1975) and varying per year per site from three to 35 (average21). In five sites in two nature reserves south of Katowice, thenumber of species varied between 57 and 84 (Kuska, 1982),and among 13 sites in the western Tatra Mountains of Poland

(Knutelski, 1993), the total number of species was 83, varyingper site between nine and 41. In the Ojcow National Park inPoland, the number per collecting method per year was 25(13-39) (Witkowski, 1969). Other Polish sites had similarspecies numbers (Cmoluch, 1962, 1971; Cmoluch and Kow-alik, 1963; Cmoluch et al., 1975; Jankowska and Witkowski,1978; Witkowski, 1978, 1979; Petryszak, 1981, 1987, 1988;Witkowski and Mazur, 1983; Knutelski, 1988,1991; Petryszakand Kaczmarczyk, 1992). Among nine sites and eight treespecies in Slovakia, a total of 108 species was found, rangingfrom 13 to 50 species per site (Holecova, 1991a, 1991b, 1991c,1993a, 1993b, 1993c), whereas sweeping four sites inoak-hornbeam forests in the same area produced 44 to 73species (Holecova, 1993d). The number of species collected byshaking in seven forests near Moscow (Roginskaya, 1992)varied from 10 to 25, with a total of 30 species. The sweepsamples taken in 54 Finnish sites contained 88 species in total,varying between sites from 0 to 37 (Raatikainen and Iivarinen,1986). Weevils collected in Japan using beating methods variedby location. Only six species were caught in a forest (Isono etal., 1986), whereas on the Japanese Izu Islands the number ofspecies per island varied between 16 and 54 (Morimoto andMiyakawa, 1985). In three years of pitfall-trapping, the totalnumber of weevil species collected in the Orange Free State,South Africa, was 52, at the rate of 33 to 38 per year (Louw,


1987). The poorest Panamanian site, Guadalupe Arriba at 2200m altitude, still compares favorably with the richer of thesetemperate areas. The total number of species collected in theentire Krakow uplands in south Poland, since 1866, was 574(Mazur, 1983), and the total number of weevil species recordedfor the British Islands is 520 (Owen, 1993). The total number ofspecies collected in Panama, on the other hand, far exceeds the2030 species (Table 4) discussed in the present paper (CWOB,unpublished data; HPS, unpublished data). Only 30.1% of thespecies from our traps are known (described) species, and thetotal number of described species from Panama is presentlyestimated as 1666 (CWOB, unpublished data), which suggeststhat the total number of weevil species present in Panama maybe over 5000. In contrast, in 1971 a total of only 2388 weevilspecies were known from the entire Nearctic (O'Brien andWibmer, 1978).

The number of species found depends on sample size, so adiversity index that is more independent of sample size mayprovide a more useful measure of the richness of a fauna. Thediversity index a for the Danish samples varied between 0.73and 1.79, with a value of 2.70 for all samples combined. For theBritish samples (Owen, 1993), a was 5.60. The Polish sampleshad an average a of about 16, varying between four and 39, andthe three Slovakian sites had an average a of 12 (9.9-13.8).The 51 Finnish samples for which a could be calculated had anaverage of 7.01 (3-14.3), the seven Russian samples had anaverage of 2.07 (1.26-3.26), the nine samples from the IzuIslands (Japan) had an average of 8.06 (4.19-10.35), and the afor the South African sample, three years combined, was 14.41(12.3-14.4 per year). There was some variation in diversitybetween these temperate sites, but they were all far poorer thanthe Panamanian samples (Table 4), again with the exception ofthe low diversity of the high-altitude sample at GuadalupeArriba, which was similar in richness to the richer of theseextra-tropical sites. This extraordinary richness of the Panama-nian samples occurred in spite of the fact that they were takenby light traps and thus represented only a small part of the totalfauna, in contrast with the temperate samples that wereobtained by sweeping and/or beating and thus were probablymuch more representative of the fauna as a whole.

Surprisingly, the weevils as a group did not show any cleardepressing effect of the full moon on catches in the light traps(Figures 6, 11, 12). This is in sharp contrast with generalcollecting experience (CWOB, HPS). In other groups, sucheffects have been found in the tropics (Brown and Taylor,1971), with the effect sometimes being very strong (Ito et al.,1993; Wolda, 1977) and sometimes rather weak (Banerjee etal., 1981; Banerjee et al., 1986), even among some insectgroups from the same light traps on BCI with which the presentweevils were collected (Wolda, 1977). The abundance of someinsects did not show any correlation with the phases of themoon (Bandyopadhyay, 1975). The situation in the presentweevil samples contrasts sharply with that in other insect

groups, collected by the same light traps that obtained theweevil samples, in which strong moon-effects were obvious(Wolda, 1977).

FREQUENCY DISTRIBUTIONS OF SPECIES ABUNDANCES.—A

useful model for the frequency distribution of abundances ofspecies is the logseries on which the a-index is based. Theredoes not seem to be a sound reason why natural communitiesshould be built according to the logseries, but as in manystudies the observed distributions did not differ significantlyfrom the logseries, or were at least close to it, the logseriesmodel provides a useful yardstick with which actual distribu-tions can be compared. The distribution of the abundances ofall Panamanian weevil samples was clearly different from thatof the logseries (Figure 3). All differences were in the samedirection, i.e., the samples contained far too many very rarespecies, too few species of intermediate abundances, and, ifanything, too many very abundant species. This is in clearcontrast with most temperate samples of weevils we knowabout (Witkowski, 1969, 1975; Overgaard Nielsen, 1974a;Petryszak, 1981, 1988; Morimoto and Miyakawa, 1985;Raatikainen and Iivarinen, 1986; Louw, 1987; Petryszak andKaczmarczyk, 1992; Roginskaya, 1992; Knutelski, 1993;Owen, 1993.) Some of the temperate samples showedsignificant difference of one kind or another from the logseries,but in only a minority of these temperate samples was thedeviation as described for the Panamanian samples. Thatminority included the samples discussed by Cmoluch (1971),Kuska (1982), Witkowski and Mazur (1983), Knutelski (1988,1991), Holecova (1991a, 1991b, 1991c, 1992, 1993a, 1993b,1993c, 1993d), and Owen (1993), who all found an excess ascompared with the logseries, often statistically significant, ofspecies represented by only one individual, just as in thepresent Panamanian samples. Although the distribution ofspecies abundances in the majority of temperate samples wasdifferent from the Panamanian samples, several other temperatesamples were similar to the tropical ones, suggesting that theremay be no basic difference in this distribution between tropicaland temperate weevil faunas. At least the available informationdoes not prove there was. Morse et al. (1988) found that asmany as 58% of beetles collected by canopy fogging in atropical lowland rainforest in Brunei were represented by onlyone individual. This was an even higher percentage than wefound among the Panamanian weevils (Table 5) and seemedhigher than would be expected from a logseries distribution. Adistribution with many singletons can be obtained when singleindividuals of vagrant species become part of the sample(Taylor, 1978; Wolda et al., 1994). We believe this to be thecase for the Panamanian weevils. The singletons may havebeen occasional immigrants from elsewhere, but they alsocould have been rare captures of local species that, because oftheir behavior, are not normally caught in light traps. Theobserved distribution of abundances of weevil species mayhave been partly due to the capture technique, but it is possible

NUMBER 590 19

that peculiarities of a tropical weevil fauna were at least partlyresponsible. Many weevils are host specific or oligolectic(Janzen, 1980; Paulay, 1985; Knutelski, 1988; Anderson,1993) and, therefore, may have been thinly distributed in adiverse rainforest with its large number of plant species. Theabundance of rare species might be more common in tropicalthan in temperate faunas. However, Microlepidoptera collectedat light traps in a variety of Brunei forests, ranging frommangrove through lowland to montane, had an abundancedistribution that was a close fit to the logseries (Robinson andTuck, 1993). This suggests that not all tropical insect groups, atleast not at all sites, have this excess of rare species. Moreover,the fact that a number of weevil samples from temperateclimates had the same excess suggests that whatever is thecause, it may be similar in Panama and in at least sometemperate areas.

BETWEEN-YEAR COMPARISONS.—The between-year com-

parisons of the Panamanian weevil samples using the NESSsimilarity index (Table 9) provided a mean NESS value of0.912, varying from 0.892 to 0.946. All these values weresignificantly different from unity, although the one fromGuadalupe Arriba was only marginally so, showing thatsamples from different years do show some variation.Between-year comparisons for the samples from Denmark(Overgaard Nielsen, 1974a), Poland (Witkowski, 1969, 1975;Knutelski, 1991; Petryszak and Kaczmarczyk, 1992) and SouthAfrica (Louw, 1987) gave similar between-year NESS values,reflecting the fact that between-year variations in abundance ofinsect species are similar in the tropics and in the temperatezone (Wolda, 1978, 1983b).

BETWEEN-SITE COMPARISONS.—There was obviously a

high degree of endemism (Table 7), a large between-site (beta-)diversity, in these weevils as evidenced by the low similarityvalues (Table 9). This also had been found for other groups ofPanamanian insects, such as cockroaches (Wolda, 1983a;Wolda et al., 1983), Psocoptera (Broadhead and Wolda, 1985),some Coleoptera (Chandler and Wolda, 1987), and Homoptera(Wolda, unpublished data). Unfortunately, there are not muchdata on weevils from elsewhere available to us to see whetheror not these low values are commonplace or are, somehow,restricted to the tropics. Raatikainen and Iivarinen (1986)compared 54 sweep-samples taken in late June to mid-July forthree years from hay meadows all over Finland. The between-site NESS for the 43 sites having at least 10 species was, on theaverage, 0.674 ±0.155. The area covered was much larger thanthat in Panama, but the environments selected (hay meadows)were much more homogeneous. From southern Poland data areavailable from 11 localities and, in many cases, from differentvegetation types within each locality (Witkowski, 1969, 1975;Petryszak, 1981, 1988; Kuska, 1982; Witkowski and Mazur,1983; Knutelski, 1988, 1991, 1993; Petryszak and Kaczmar-czyk, 1992). The mean NESS for samples from differentvegetation types within each locality varied between 0.244 and

0.619, with a mean of 0.45. The mean NESS value for the 11localities, varying widely in altitude and habitat, with sampleswithin each locality lumped, was 0.303, varying from 0.101 to0.756. The average between-island NESS index for the IzuIslands in Japan (Morimoto and Miyakawa, 1985) was 0.505,varying between 0.305 and 0.727. The mean between-sitesimilarity for Slovakian samples varied from 0.453 for samplesfrom Corylus to 0.683 for samples from Alnus (Holecova,1991a, 1992, 1993a, 1993b, 1993c), whereas between fivebiotopes, different sites combined, similarities varied between0.169 (beech forest vs. xerothermic meadows) and 0.695(mesophilic vs. xerothermic meadows) (Holecova, 1991b). Onthe other hand, the mean NESS value for the Panamaniansamples in Table 9 is 0.186, varying from 0.009 to 0.469. Thislimited information strongly suggests that for weevils thetropical between-site similarities were smaller, i.e., the differ-ences in species composition and relative abundances tended tobe larger, than those observed in temperate areas.

The comparisons made here between the present Panama-nian weevil samples and those from the temperate zone sufferfrom differences in collecting methods. All Panamaniansamples were from light traps, whereas none of the Europeansamples was collected in this way. However, the richness of thepresent tropical samples was not due to the collecting methodemployed. The Panamanian samples are so rich, not becauselight traps are so efficient for weevils, but because theneotropical weevil fauna is so rich. This is in line with thegeographic distribution of the described weevil species(O'Brien and Wibmer, 1978), whereas the number of unde-scribed species in the Neotropics undoubtedly is vastly largerthan those in the Nearctic or in the Palearctic (O'Brien andWibmer, 1979). In fact, the majority of the species reported inthe present paper (69.9%) are still undescribed.

VERTICAL STRATIFICATION.—On BCI more weevil indi-

viduals were collected in the canopy trap than at ground level,whereas the ground-level trap caught more species in each ofthe three years. The same pattern was found for Homopterafrom the same traps (Wolda, 1987). The higher species richnessat a low level in the forest for both insect groups was notexpected. Some of the species, especially the ones rarelycaught, could have been migrants from elsewhere, and onemight expect this migration to have taken place predominantlyat the canopy level, leading to a higher species richness here.The present results suggest, however, that our expectationconcerning migration may have been wrong. There wereclear-cut differences between the faunas caught in the canopyand those caught at ground level. The similarity index betweenthese faunas was high, but still significantly lower than thesimilarity between years, showing that the differences betweenlevels in the forest, though not very large, were larger thanthose between successive years. The vast majority of thespecies occurred at both levels, but some species, even verycommon ones, were found exclusively in the canopy or in the

GUAD

ALUP

E ARRIBA

1976

1977

1978

1983

1984

1985

FIG

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NUMBER 590 21

low trap. The differences between the two levels were clear butnot very large, and they do not suggest the existence of a faunain the canopy that is vastly different from what is seen atground level. Brown (1961) found suggestions of a verticalstratification among beetles in a forest in Uganda, and Hingston(1930) also marvelled at the differences in catches at differentlevels in a forest in Guyana. Most authors found that manymore individuals were observed in the canopy than at lowerlevels in several insect groups, such as sphingid moths (Corbet,1961 a), male tabanid flies (Corbet, 196 lb), or insects in general(Sutton and Hudson, 1980; Sutton et al., 1983; Basset et al.,1992). Our observations that both weevils and Homoptera werericher in species at a low level than in the canopy is, however,in contrast to studies by Corbet (1961a) and Basset et al.(1992), who found more insect species high in the forest, orCorbet (1961b), who found no vertical gradient in speciesrichness in mosquitos. In our study, the vertical distribution ofmost species was consistent from year to year, but there werecurious, as yet unexplainable, exceptions, even among somevery common species (Figure 9).

We do not know the extent of the area from which weevilswere attracted to the traps. We also do not have usefulinformation on the dispersal distances of weevils. Beavers et al.(1979) reported that Diaprepes abbreviates (L.) has amaximum single flight of 45 meters, whereas Solbreck (1980)showed that the pine weevil Hylobius abietis L. may migrateover a distance of many kilometers. Sufficient informationalong these lines is not available to make any kind ofgeneralization, certainly not for tropical weevils.

SEASONALITY

WEEVILS AS A GROUP.—The seasonal distribution of all

weevils summarized for each site, except Corriente Grande, isgiven in Figure 10. Data from Corriente Grande are ignoredhere as they cover only four months, which is insufficient forinformation on seasonality. The most seasonal sites, based onrainfall (BCI, Las Cumbres, and Boquete), also had thestrongest seasonal patterns in terms of weevil abundance asdetermined by the light traps. Especially noticeable were thesharp to very sharp seasonal peaks at the beginning of the rainyseason in May, with a second peak occurring around October,especially on BCI. Peaks in the other climatologically lessseasonal sites were far less obvious, but seasonal patterns werestill apparent. In Fortuna, there was an annual low in abundancein the early part of the year in spite of the absence of a distinctdry season, and a peak in abundance in May at what would bethe beginning of the rainy season in more seasonal sites. InMiramar, there may have been a distinct low in abundance inJune-July, but trap problems during part of that period make aninterpretation of the results difficult. In the high-altitude site ofGuadalupe Arriba, there was a seasonal low in abundance inNovember-December. For Miramar, only one year of datawere available, so it is not known whether the possible low in

June-July 1979 was a generally occurring phenomenon, andthus seasonal, or just an accident of that year. We favor theformer explanation. The lows in abundance at the other sites,however, were repeated in successive years and were thusclearly seasonal. At the seasonal sites, the beginning of therainy season seems to be a strong environmental clue forseasonal abundance patterns of weevils.

How closely weevil activity in general follows the onset ofthe rains is indicated in Figure 11 for individuals and Figure 12for species at BCI by showing daily abundances over thethree-month period that invariably includes the transition fromdry to rainy season. In both 1976 and 1977, the dry seasonswere unusually long and dry, especially in 1977, during amoderate El Nino event. In 1978, periods of rain often werefollowed immediately by large increases in abundance of bothindividuals and species. For weevils the rains on 17 April 1978apparently provided an important seasonal cue. No such clearweevil responses were seen in the other years. In 1976, a heavyshower on 9 April was not accompanied by a change in weevilabundance, but the light showers in late April/early May didinitiate weevil activity, as did the heavier rains in late May. In1977, the first moderate flights of weevils were preceded byone light shower on 5 May and then weevil abundancesincreased in late May during and after some heavier rains. Raindid have a strong effect, but not if it occurred too early in theseason, i.e., 10 April was too early in 1976, but 17 April wasnot too early in 1978. In late April or early May, any amount ofrain seems to have had an effect. The full moon did not have adepressing effect at all on weevil catches in the light traps, atleast during the period March through May.

BETWEEN-SPECIES COMPARISONS.—Mean Vector:

General seasonal patterns of groups of species are onlymarginally informative, depending as they do on the kinds ofspecies present at each site and their relative abundances. Astudy of the component species at each site is much moreuseful. Data for all complete years of collecting werecombined, and only species represented by at least 10individuals in those years were considered. For the Fortuna site,this meant that only the two years with the trap at ground levelwere included in the analysis, for Las Cumbres the data fromOctober 1973 to March 1974 were excluded, and for GuadalupeArriba the year chosen ran from April 1983 to April 1984.There was a wide variety of seasonal patterns among thespecies at each site, and presenting all those patternsindividually is impractical. The data are best presented insummary form. The distribution of the degree of seasonality asexpressed by the Mean Vector (r) is presented in Figure 13,where zero stands for a uniform distribution over the year and1 means that all individuals were found in the same week. Alarge variation in the value of r was obvious at all sites.Whether or not the seasonal pattern of a species wassignificantly different from a uniform distribution depends bothon the value of r and the number of datapoints (individuals).Nonsignificant cases are represented by the white columns, and


800

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Individuals 1976

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MARCH APRIL MAY

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NUMBER 590

FIGURE 11 (left).—Daily catches of weevils over period March-May in threeyears on BC1, together with daily rainfall over those same periods. Commonesttaxon Phyllotrox complex given separately from the rest of the weevils (timingof full moon also indicated).

significant seasonalities are represented by dark columns inFigure 13. The percentages of species that were significantwere 84.5%, 87.5%, and 81.0% for BCI, Las Cumbres, andBoquete, respectively, whereas for the climatologically lessseasonal sites, Miramar, Fortuna, and Guadalupe Arriba, thesepercentages were only 60%, 58.6%, and 62.5%, respectively.Las Cumbres appeared to be the most seasonal site with 24% ofthe species having r-values over 0.9. For BCI and Boquete,these percentages were 11.3% and 7.1%, whereas at the otherthree sites such high r-values did not occur. Mean values for r,given in Figure 13 with standard deviation and standard error,also illustrate these differences between the sites. Despite thecorrelation between the distribution of the mean vectors and thegeneral climate, it is clear that even in the most aseasonal sitessome species with short seasons (high mean vector values) dooccur.

Mean Week and Peak Week: The Mean Week, i.e., themean of the seasonal distribution, also was calculated for theseweevil species, and the results are plotted in Figure 14, togetherwith the mean and angular deviation for these plots. Only thespecies that were significantly different from uniformlydistributed were included in the calculations of these meanvectors and angular deviations because the concept of MeanWeek is not very meaningful for species whose distribution isnot significantly different from uniform. The distribution of theMean Week for those nonuniform species was not significantlydifferent from uniform in Miramar and Guadalupe, but it washighly significant in the four other sites. In other words, in theformer two sites some species may have had their mean weekat any time of the year, whereas at the other four sites the meanweeks tended to be clustered in a certain season, which in allthese cases was May to July. As expected, the nonsignificantspecies in these plots were distributed throughout the year withno apparent concentration. In spite of the distinct clustering, atvirtually any time of the year some species could be found tohave reached maximum abundance.The average of the signifi-cant Mean Weeks tended to occur in June (BCI, Las Cumbres,Fortuna) or July (Boquete), about a month later than theaverage Peak Week, the actual mode of the seasonal abun-dance, suggesting a skewness to the right in the seasonal patternof many species. In order to properly study this skewness, the(principal axis) regression of Mean Week (y) on Peak Week (x)was calculated for all weevil species represented by at least 10individuals and that had a seasonal distribution significantlydifferent from uniform. The regression was, for all sites takentogether, y = 4.56 + 0.85x and the Jupp-Mardia circularcorrelation was r2 = 1.335 with N = 523 (p<101 0). Thisregression was very close to the diagonal where PeakWeek = Mean Week, so that Peak Week residuals were

23

calculated by simply subtracting Mean Week from Peak Week,while making sure that no residual is larger than 26 weeks inthis seasonal circular world. These residuals were plotted inFigure 15, for each site separately. No particular pattern wasevident among the few species in either Miramar or GuadalupeArriba, but in the other four sites many points occurred in adownward band, starting at a residual value of zero in May andbecoming more and more negative as the points occurred closerto August. These species all had their time of maximumoccurrence in May but had their Mean Week value later in theyear, skewed to the right of the seasonal pattern. This isillustrated in Figure 16A-C. AS is clear from these examples,this "skewness" often reflected bimodality. On BCI a secondband of points was evident, starting in September with residualvalues around zero and going up and to the left, becoming morepositive the closer the points are to June. These points representspecies that had their maximum abundance late in the year butwhose abundance patterns are skewed to the left, often with asecondary peak earlier in the season (Figure 16D,E). In fact, theseasonal pattern of the species in the top band tended to besimilar to that of the species in the lower band, except that itwas the later rather than the earlier peak that was the largest.

The Morisita index of seasonal diversity (Morisita, 1967;Yamamoto, 1974) was calculated for each species from BCIwith at least 10 individuals over three years and was plottedagainst the Mean Vector for those species in Figure 17. Ahigher degree of seasonality is indicated by a higher value forthe Mean Vector and by a lower value for the index of seasonaldiversity, so that the relationship between the two is expectedto be negative. The correlation coefficient between the twomeasures of degree of seasonality is highly significant(r =-0.624, p « 0.001) but is far from perfect, showingthat the two measures are far from identical. We prefer the useof the Mean Vector.

BETWEEN-YEAR WITHIN-SITE COMPARISONS.—As was

shown above, many species at all sites had patterns of variationin abundance over the year that were significantly differentfrom uniform. For most sites these patterns were based on thecombined abundance data of more than one year. Similarly,within each year, the majority of species showed significantabundance variation, but the question is whether or not speciesin different years at the same site showed the same variation inabundance over the year. In other words, was the variation inobserved abundance cyclical and did it reflect true seasonality,or was it just variation that had little or nothing to do withclimatic seasonality, which was shown for mosquitos innorthwestern Panama (Wolda and Galindo, 1981)? In four ofthe sites with between-year information on weevils (BCI, LasCumbres, Boquete, and Fortuna), comparisons were madeusing the Mean Vector, i.e., the indicator of the degree ofseasonality, and the Mean Week, i.e., the mean of the annualdistribution. For each species of which at least 10 individualswere collected in each of the years, these measures werecalculated per year and then the average and standard deviation


100 -

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Species

Rainfall

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idRainfall (mm)

1 l U . i l IMARCH APRIL

O Full Moon

MAY

NUMBER 590 25

FIGURE 12 (left).—Number of species caught per day on BCI in periodMarch-May in three years, together with daily rainfall over same periods(timing of full moon also indicated).

for the Mean Vector and the circular average and the angularstandard deviation for the Mean Week were determined.

For BCI the results for all species of which at least 10individuals were collected in each of the three years of thestudy are presented in Figure 18 A. The standard deviation of theMean Week tends to decrease with increasing values of theMean Vector (Figure 1 8B, r = -0.546, p < 0.001), which meansthat the variation in the mean of the seasonal distribution tendsto be larger when the degree of seasonality is lower. This ishardly surprising, but it is interesting to see that some species

with a high Mean Vector value show a high between-yearvariability in Mean Week, whereas in other species the MeanWeek does not change much between years in spite of a lowdegree of seasonality. Similar increases in the standarddeviation of Mean Week with decreasing Mean Vector werefound for the weevils from Las Cumbres (r = -0.896, p < 0.01),Boquete (r =-0.929, p< 0.001), and Fortuna (r =-0.989,p < 0.001). For the BCI data, the variation inMean Vector decreased with increasing values for the averageMean Vector (r =-0.465, p< 0.001). For Las Cumbres thiscorrelation was just significant (p<0.05) but it was not atBoquete or Fortuna. The variation in Mean Week increasedsignificantly (p < 0.05) at BCI as the season progressed (Figure18c), but no such seasonal change in variability was observed

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FIGURE 13.—Frequency distribution of degree of seasonality among Panamanian weevils as measured by MeanVector (r) in six localities. Species with seasonal pattern significantly different from uniform, i.e., species that aresignificantly seasonal, are indicated separately from those where no significance was observed. Mean, standarddeviation, and standard error of each distribution also given for significantly seasonal species.



4 H

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II

p-0.15 GUAD.ARR.


MEAN WEEKI p<0.05 \ZZ\ N.S.

FIGURE 14.—Frequency distribution of mean week of seasonal pattern among Panamanian weevils as measuredby Mean Week. Species whose seasonal pattern differ significantly from uniform, i.e., significantly seasonalspecies, are indicated separately from those with no significance observed. Mean, standard deviation, andstandard error of each distribution also given for significantly seasonal species.

at any of the other sites. No significant seasonal change invariation in Mean Vector was observed at BCI or anywhereelse.

Some examples illustrate observed between-year differencesin seasonal abundance patterns. Figure 19 gives seasonalpatterns for two species from BCI in two different years. ForPseudapotrepus macrophthalmus Champion the actual abun-dance patterns are similar in the two years. The much lowervalue for Mean Vector in 1977 is due to the relatively lowerpeak in May-June yielding a relatively larger effect of the lowbut broad peak in the fall. In 1977, 88 of the 194 individuals,45%, occurred in the second peak, whereas in 1978, 87 out of672, 13%, were found in the second peak. For Conotrachelus

semirufus Champion the difference between the years wasmuch larger. In 1978, there was a major peak in abundance inthe beginning of the rainy season followed by a broad and lowpeak in the second half of the rainy season. The May-July peakin 1977, on the other hand, was so low that the fall peakdominated the picture. Moreover, the early peak in 1978occurred later than the one in 1977, in spite of the fact that therains in 1978 started earlier. In both species, there werepronounced differences between the two years, but most ofthose involved the relative height of the two peaks ofabundance. There is no reason to call this variation anything butcyclical, i.e., seasonal, in spite of the differences.

Between-year differences in Mean Week are illustrated by

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FIGURE 16.—Six examples of differences between Peak Week and Mean Week, i.e., between mean and mode ofseasonal pattern (vertical lines indicate position of Mean Week (solid line) and Peak Week (dashed line); in F bothlines cover each other): A, Acamptoides sp. #1; B, Apion cretaceicollis Sharp; C, Cossoninae gen. #2, sp. #1; D,Rhinostomus barbirostris (Fabricius); E, Andranthobius sp. #1; F, Apteromechus sp. #8.

two species from BCI, Rhinostomus barbirostris (Fabricius)and Micromimus sp. #17 (Figure 20). For Rhinostomus theseasonal patterns in the three years were not as different as thelarge discrepancies in Mean Week suggest. In 1977 theseasonal low in June-July was particularly low, so that thecircular Mean Week shifted far to the right to January, betweenthe two main peaks of September and April. In 1978, on theother hand, the low in November was particularly pronounced,shifting the Mean Week to June, between the April andSeptember peaks. In 1976 the second low extended fromNovember through March, putting the Mean Week in lateAugust. It was the relative heights of the highs and lows, ratherthan their presence and their timing, that are different betweenyears. A similar explanation may hold for the other species,

Micromimus sp. #17, that had two seasonal peaks, one inMay-July and the other in November-March. The differencebetween the two years was in the seasonal low in September-October, during which period there were several individuals in1978 and none in 1977.

In most species, the general pattern of seasonality is similarbetween years, and between-year differences in measures, suchas Mean Vector and Mean Week, were generated by the relativeheights of the seasonal peaks rather than by a fundamentaldifference in seasonal pattern. In some cases, however,between-year differences are large. For instance, Andrantho-bius sp. #1 on BCI had two major peaks in each year. However,those peaks were in August and October in 1976, in May/Juneand August in 1977, and in April-June and September in 1978.

NUMBER 590 29

Weevils, BCI, 1976-1978

0.1 0.2 0.3 0.4 0.5 0.6 0.7Morisita Seasonal Index

0.8 0.9 1.0

Significant Not Significant

FIGURE 17.—Relationship between Mean Vector and Morisita's Index of Seasonal Diversity for weevils fromBCI, all three years combined. Only species of which at least 10 individuals were collected are included. Linearcorrelation between all 407 data points is -0.681 (p < 0.001).

Similarly, Isotrachelus sp. #3 had its Mean Week in earlyNovember in 1976, in mid-June in 1977, and in mid-April in1978. In 1976, however, only 11 individuals were caught andseven of these occurred in one week in November, with onlytwo in June and two in October. The numbers are small, but thedistribution was very unlike those in 1977 and 1978 when ahigh and sharp peak in abundance occurred early in the rainyseason with low abundances toward the end of the year. It is notclear how much weight should be given to this between-yeardiscrepancy, but it should serve as a warning that the data fromone year may give an erroneous impression of data in otheryears, even when dealing with one species at one site. Twospecies from other sites, Micralcinus sp. #1 from Boquete andCossoninae sp. #6 from Fortuna, emphasize this point (Figure21). The pattern for Micralcinus may be explained by abimodal distribution with the peaks occurring earlier in thesecond year, but the case of Cossoninae sp. #6 is different. Thespecies occurred year-round in both years, with a pronounced

peak around March 1978 and a very broad peak in October1978 through February 1979. This very well may be a case ofpronounced fluctuations without reference to the calendarseasons. A rejection of the null hypothesis of a uniformdistribution does not necessarily mean that the fluctuations athand are cyclical and thus truly seasonal.

One environmental factor affecting the seasonality of severalspecies is the timing of the beginning of the rainy season. AtBCI, Mean Weeks in 1978 tended to be a month earlier than in1976 or 1977, which undoubtedly relates to the earlier start ofthe rains in 1978 (Figure 11).

WITHIN-SPECIES BETWEEN-SITE COMPARISONS.—There are

clear differences in overall seasonality patterns between thesites (Figures 10, 13-15), but between-site comparisons usingonly species shared by these sites would be much moreinformative. Unfortunately, between-site variation in speciescomposition was so large that very few species were generally


o

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0 4 8 12 16 20 24 28 32 36 40 44 48 52

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Frequency

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0 4 8 12 16 20 24 28 32 36 40 44 48 52

10xSD.-Mean Vector

- 6

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Average Mean Week

NUMBER 590 31

FIGURE 18 (left).—Between-year variation in Mean Vector and Mean Week forall species collected at BCI with at least 10 individuals in each of three years:A, Average Mean Vector plotted against average Mean Week with standarddeviations along both axes; B, frequency of classes of average Mean Vector andmeans and standard errors of standard deviations per class of Mean Week andof 10x Mean Vector; C, frequency of classes of average Mean Week and meansand standard errors per class of standard deviations of Mean Week and of 10xMean Vector.

common enough at several sites (collected with at least 10individuals) to make possible between-site comparisons inseasonality of individual species. For each site the data fromdifferent years were pooled. Guadalupe Arriba had to be left outof these comparisons entirely because the only taxon it sharedwith the other sites was the Phyllotrox complex. For the BCI vs.Las Cumbres comparison, there were 52 species available, butthe next most species-rich comparisons were those between

BCI and Fortuna (12 species), BCI and Boquete (11 species),BCI and Miramar, as well as Boquete and Fortuna (10 species),and Las Cumbres and Miramar (nine species). For the othercomparisons, Las Cumbres and Boquete, Las Cumbres andFortuna, Miramar and Boquete, and Miramar and Fortuna, onlyfive, one, two, and two species, respectively, were available.

The between-site variation in seasonal patterns is illustratedhere by classifying the species in classes according to the MeanVector values, as in Figure 18B, and by plotting the mean(angular) standard deviation of the Mean Week (Figure 22) andthe standard deviation of the Mean Vector (Figure 23) againstthe Mean Vector classes (closed symbols, ignoring the fourdata points connected by a line). The within-site between-yeardata, which include the data from Figure 18B, also are given inFigures 22 and 23 (open symbols). For the Panamanian data,the connecting lines for each comparison, as given in Figure18B, are omitted for reasons of clarity. For the

6

4

2

0

40

20 -\

Conotrachelussemirufus

Conotrachelussemirufus

1978r=0.696

\ v \ i i I i r20 -

10 -

Pseudapotrepus macrophthalmus

Pseudapotrepus macrophthalmus1978

r=0.775

M A M J J A S O N D J FFIGURE 19.—Seasonal patterns of two species from BCI with rather extreme between-year differences in MeanVector (graphs represent four-week moving averages of weekly totals).


1 1

Rhinostomusbarbirostris

0 -12 -

8 -

040

20 -

i i i i i i r i i i n

Rhinostomus 1 9 7 7barbirostris

Rhinostomusbarbirostris

i i

Micromimus sp.#17 1977

Micromimus sp.#17 1978

M ' A ' M ' J ' J ' A ' S ' O ' N I ' J ' FFIGURE 20.—Seasonal patterns of two species from BCI with rather extreme between-year differences in MeanWeek (graphs represent four-week moving averages of weekly totals; arrows indicate position of Mean Week).

between-year data points for the Mean Week (Figure 22, opensymbols), there is a clear and highly significant decrease instandard deviation (r =-0.875, p < 0.001). For the between-site data points (closed symbols), this decrease is not

significant (r =-0.360, p>0.10), and this seems to be duealmost entirely to the two left-hand points. When those pointsare deleted, the decrease is significant (r =-0.650, p<0.01).However this may be, it is clear that for moderate Mean Vector

NUMBER 590 33

40

20 -

0

4 -

2 -

0

8

4 H

0

6 -

4 -

2 -

Cossoninae sp.#6

FORTUNA

1977/8

i I i i r

Cossoninae sp.#6

FORTUNA

1978/9

Micralcinus sp.#1

BOQUETE

1976/7

Micralcinus sp.#1

BOQUETE

1977/8

J A S O N D J F M A M JFIGURE 21.—Seasonal patterns of two species with pronounced between-year differences in Mean Week, onefrom Fortuna, and one from Boquete (graphs represent four-week moving averages of weekly totals; arrowsindicate position of Mean Week).

values, those between 0.25 and 0.65, the between-site variationin Mean Week tends to be much larger than that between yearswithin each site. Highly seasonal species tend to have the samevariation in Mean Week between sites as within sites. Foraseasonal species, one would expect an unpredictable and largevariation in Mean Week between years as well as between sites,but the few data points on the left of Figure 22 do not confirmthat expectation. The between-site standard deviation of theMean Vector (Figure 23) is roughly the same as that betweenyears. The degree of seasonality between sites varies by thesame amount as that within each site between years.

For four selected species collected both at BCI and Las

Cumbres, between-site variation in seasonal patterns isillustrated in Figure 24. The differences can obviously be quitelarge even in two climatically similar sites, such as BCI and LasCumbres. One extreme example of such differences is given byMicromimus minimus (Boheman), which will be discussedbelow (Figure 27). Another such example is Metriophilusminimus Champion (data not shown). At both sites this specieswas found between December and August, but at BCI there wasa major peak in May-June, with low abundances betweenDecember and April, whereas at Las Cumbres there was abroad peak from December through March with low abun-dances afterward.


Weevils

0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9Average Mean Vector

Between-Years Between Sites Poland

FIGURE 22.—Between-year and between-site variation in mean of seasonal distribution, as expressed by angulardeviation of Mean Week, in relation to average Mean Vector (cf. Figure 18B). Between-year data for BCI, LasCumbres, Boquete, and Fortuna; between-site data for BCI vs. Las Cumbres, Boquete, and Fortuna, Las Cumbresvs. Miramar, and Boquete vs. Fortuna. (Four between-site points connected by line represent data from LublinPlateau in Poland (Cmoluch, 1962).)

At climatically very different sites the differences can beeven larger. This is illustrated for six species in the BCI vs.Miramar comparison, the seasonal patterns of which are givenin Figure 25. Especially in Cossoninae gen. #2, sp. #1, but alsoin Conotrachelus turbatus Faust and Heilus bioculatus(Boheman), the differences are spectacular. Similarly,Conotrachelus cristatus Fahraeus at Las Cumbres (data notshown) occurred throughout the rainy season, with a maximumin October-November, and was absent during the dry season(January-April), but at Miramar had a sharp peak in April.Fortuna also has an aseasonal climate, but the differencesbetween Fortuna and BCI in the seasonal patterns of all 12shared useable species (Figure 26) are not very spectacular inmost instances. For many species the seasonal pattern wasrather similar (Figure 26A,D,H,J,L), in others there was a secondpeak at Fortuna that was absent or nearly absent at BCI (Figure26B,G,K), whereas in only a few species there was a suggestionof the pattern being less seasonal at Fortuna (Figure 26c,E). Thesimilarity in many species between Fortuna and more seasonalsites may be related to the proximity of Fortuna to the seasonalPacific side of Panama. For example, Boquete and Fortuna,which are only 20 kilometers apart, have very different weatherpatterns, yet eight of their ten usable species have similarseasonal patterns. Conotrachelus Julvescens Champion was

bimodal in Boquete, with pronounced peaks in May and inNovember-December, but had just one sharp peak in Februaryin Fortuna. On the other hand, Conotrachelus crenatusChampion had one sharp peak in April-May in Boquete andwas bimodal in Fortuna, with peaks in April-May and inJanuary-February.

Unfortunately, species that occurred in reasonable numbersat more than two sites were very rare. Apart from the Phyllotroxcomplex, none of the species had at least 10 individuals in allsix sites or even at five sites. Only three species were"common" at four sites. The seasonal patterns of these threespecies are shown in Figure 27. Hemiliopsis nudicollis(Chevrolat) occurred year-round in at least three, possibly all,of the four sites. On BCI it had a major peak in April-June, atthe beginning of the rainy season, with the abundance taperingoff toward the end of the year. In January-February, the earlydry season, there was a secondary peak in seasonal abundance.In Fortuna, in spite of its much less seasonal climate, the patternwas almost identical to that on BCI, except for the absence ofthe January-February peak. In Miramar the numbers were toolow to make definite statements about the seasonal patterns, butthe absence of any individuals in May-July is curious. In LasCumbres there was a broad peak in abundance from Novemberto March, with lower abundances the rest of the year. Only at

NUMBER 590 35

Weevils

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1Average Mean Vector

<=> Between-Years • Between Sites Poland

FIGURE 23.—Between-year and between-site variation in degree of seasonality, as expressed by standarddeviation of Mean Vector, in relation to average Mean Vector (cf. Figure 18B; for further details see Figure 22).

BCI and Fortuna was the seasonal distribution of abundancesignificantly different from uniform. Pseudapotrepns macro-phthalmus Champion had a major peak in April-June in BCI,Boquete, and Fortuna, which is the early rainy season at thefirst two sites but not in Fortuna, with a low abundance duringthe rest of the rainy season and a near-absence during the dryseason. The pattern in Miramar was clearly different in that thespecies occurred year-round with peaks that seem unlikely tobe seasonal. Miramar was a relative aseasonal site, but so wasFortuna. The seasonal variation in abundance was significantlydifferent from uniform in all sites but Miramar. Micromimusminimus (Boheman) was even more curious. The fairly sharpseasonal peak on BCI occurred in April-June, but in LasCumbres, as well as Boquete, the peak occurred in Novemberthrough January, whereas the pattern at Miramar was a hybridbetween these two patterns (see below for the taxonomy of thisspecies in Boquete). Again, the pattern at Miramar was the onlyone not different from uniform.

These between-site differences in seasonal patterns ofindividual species do not seem to be clearly related todifferences in climate. Miramar did have the least seasonalabundance patterns in both Micromimus and Pseudapotrepus,but Fortuna did not show any hint of an effect of its relativelyaseasonal climate. This brings us to a difficult problem. Howsure are we that we are indeed dealing with the same species atall these sites? After careful re-examination of the specimens ofMicromimus minimus, it was found that the specimens from

Boquete had slightly different male genitalia; thus theyprobably represent a new species, referred to here as "nr.minimus." However, no such differences were found betweenthe specimens from BCI, Las Cumbres, and Miramar. So thelarge difference in seasonal pattern between BCI and LasCumbres remains, and we may have to accept that the almostidentical patterns of Las Cumbres and Boquete were producedby different species. No other between-site morphologicaldifferences have been found thus far in any of the other species,so that to the best of our knowledge we are dealing with thesame species in each case. We will assume that ouridentifications were correct, although we realize that thetaxonomy of these tropical weevils is still in its infancy. Manyof the species we collected remain to be described, and most ofwhat we know is based on morphology, with virtually nothingknown about weevil behavior or genetics. Sibling species, wellknown in many groups in temperate areas and thus likely tooccur in the tropics as well, are unknown at this moment amongPanamanian weevils. We must proceed under the assumptionthat the specimens, which on morphological grounds areconsidered to belong to the same species, are indeedconspecific.

DISCUSSION OF SEASONALITY

In discussions of seasonal abundance patterns, the data areusually presented in graphical form, which works well for a


10

10

0

100

50

0

2

1 -

Conotrachelus turbatus

BCI

Conotrachelus turbatus

LAS CUMBRES

Cossoninae n.gen.jjh ,sp.#1

BCI

Cossoninae n.gen.f"! ,sp.#1

LAS CUMBRES


20 •

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1.0 •

0.5 -

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50 -

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Cossoninae n

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Cossoninae n

LAS CUMBRES

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CUMBRES•

J F M A M J J A S O N DFIGURE 24.—Seasonal patterns (four-week moving averages) of four selected species of weevils in Las Cumbresand on BCI.

limited number of species. However, with a large number ofspecies presenting graphs for all species becomes impracticaland some form of data reduction becomes necessary. Attemptshave been made to provide summary seasonality measures,such as the Morisita Index of Seasonal Diversity (Morisita,1967; Yamamoto, 1974) and Seasonal Range and SeasonalMaximum (Wolda, 1979). However, we found those measuresunsatisfactory. As the above chapters show, circular statisticsdo provide useful descriptors of seasonal patterns, especiallywhen dealing with large numbers of species. The measures,Mean Vector, an indicator of the degree of seasonality, MeanWeek, the mean of the seasonal distribution, and the differencebetween mode (Peak Week) and mean (Mean Week) of theseasonal distribution, turned out to be very useful.

Information on seasonal patterns of temperate weevils as agroup were given by Petryszak (1981, 1987, 1988, 1991),Kuska (1982), Knutelski (1988, 1991), Petryszak and

Kaczmarczyk (1992), and Holecova (1993a, 1993b), but littleinformation is available on seasonality in individual weevilspecies. A number of papers described the generally shortseason of adult activity in many species (Harris and Coppel,1967; Silver, 1968; Fye and Bonham, 1970; OvergaardNielsen, 1974b, 1974c; Hansen, 1987; Stachowiak, 1991;Holecova, 1991a, 1991b, 1991c, 1992, 1993c, 1993d; Stacho-wiak, 1991; Knutelski, 1993). Although the season of somespecies may extend over most of the summer (Attah andLawton, 1984; Isono et al., 1986; Holecova, 1991a, 1991b,1991c, 1992, 1993c, 1993d), no species seem to be active mostof the year. The active season is often interrupted byhibernation in winter (Davey, 1956; DeSteven, 1981; Menu,

FIGURE 25 (right).—Seasonal patterns (four-week moving averages) of sixspecies of weevils from BCI and Miramar.

NUMBER 590 37

10

5 -

Conotrachelusturbatus

Cossoninaeng.#2,nsp#1

3

0

2

1

1.0

0.5

/. Polydacrys depressifrons

Stenommatus sp.#1

/TV

1.0

0.5

Cossoninaeng.#1,nsp#1

Heilusbioculatus

- 1

- 0.4

0.0J F M A M J J A S O N D

0 0


3 3 i i i i i i • i i i i i i 3

2000 -

1000 -

J F M A M J J A S O N D J F M A M J J A S O N D•FORTUNA (Right ordinate)


•B.C.I. (Left ordinate)FIGURE 26.—Seasonality patterns (four-week moving averages) of 12 species of weevils with at least 10individuals both at Barro Colorado Island and at Fortuna (data from different years are pooled; Mean Vectorvalues and their significance indicated on left for BCI and on right for Fortuna): A, Apion "complex";B, Conotrachelus sp. #77; C: Conotrachelus brevisetis Champion; D, Hemiliopsis nudicollis (Chevrolat); E,Micromimus sp. #2; F, Micromimus sp. #3; G, Paratrachelizus sp. #2; H, Phyllotrox "complex"; I, Pisaeus variousChampion; J, Pseudanchonus occultus (Champion); K, Pseudapotrepus macrophthalmus Champion; L,Stereodermus calvus Sharp (Mean Week indicated by vertical lines).

1993; Menu and Debouzie, 1993). In very few cases does thepublished information on temperate weevils allow calculationof seasonality measures, such as the Mean Vector. Exceptionsare studies in Poland by Cmoluch (1962) at the Lublin Plateauand by Cmoluch et al. (1975) at the Sandomierz area. In thesestudies, the percentages of species with a Mean Vector largerthan 0.80 were 90.6% and 62.7%, respectively, whereas forPanamanian weevils these percentages ranged from 3.5% atFortuna and 8.0% at Miramar to 26.3% at BCI and 38.5% atLas Cumbres (Figure 13). Mean Vector values smaller than0.60 occurred in only one percent of the species at the Lublin

Plateau and two percent at Sandomierz. For all Panamaniansites, these percentages were much higher (Figure 13), rangingfrom 32.7% at Las Cumbres and 48.2% at BCI to 79.3% atFortuna and 80.0% at Miramar. The Mean Week for all speciesat the two Polish sites occurred between May and September,whereas at all Panamanian sites the Mean Week was found tooccur at any time of the year (Figure 14). Some of this mayhave been due to the fact that no samples were taken at thePolish sites from October to April, but we have been told thatvery few, if any, weevils are active in eastern Poland at thattime of the year. This summary illustrates, for the first time in

NUMBER 590 39

10

Hemiliopsisnudicollis (Chevrolat)

8

6

4 •

2 •

0

2

0.8

0.4

0.0

BCI

Las Cumbres

Miramar

Fortuna

100

Pseudapotrepusmacrophthalmus Champion

100

Micromimusminimus (Boheman)


FIGURE 27.—Seasonal patterns (foursites.

J F M A M J J A S O N D J F M A M J J A S O N D

week moving averages) of three species with at least 10 individuals at four

quantitative terms, the differences in seasonality patternsbetween weevils from a tropical area and two sites in thetemperate zone. In broad lines, these data may in fact representdifferences between the temperate zone and the tropics ingeneral, although details will have to be filled out when moredata become available.

Little is known about between-year variation in seasonalityin most temperate weevils. Some reported no differencesbetween years (Fye and Bonham, 1970; Hansen, 1987),whereas others discussed a between-year variation that ex-tended at most a few weeks (Overgaard Nielsen, 1974c; Vittumand Tashiro, 1987; Phillips, 1992; Knutelski, 1993). We knowof no published information on between-year variation inseasonality of temperate weevils that can be used for ananalysis similar to the one given in Figure 18. For between-sitevariation in seasonality, the only useable information we foundwas on weevils at the Lublin Plateau in Poland (Cmoluch,1962). Those data are from six sites, and of 25 weevils species

collected, at least 10 individuals were taken from at at least twoof those sites. The between-site variation in Mean Week isindicated for these data in Figure 22. Similarly, the between-site variation in Mean Vector in Poland is not much differentfrom that in Panama (Figure 23). If anything, the between-sitevariation in both Mean Vector and Mean Week in Panama issomewhat larger than that in Poland, but if such differences invariability exist, they are not very pronounced, at least not inthe present data.

Almost no published information is available on seasonalityof tropical weevil species and its between-year and between-site variability. It is known that weevils as a group occurredyear-round, walking up and down tree trunks, in a seasonallyflooded forest near Manaos, Brazil, but no information onindividual species was available (Adis, 1979, 1981). Leon(1980) reported that the boll weevil in Nicaragua is active for alonger period than at temperate latitudes, e.g., North Carolina(Fye and Bonham, 1970).


The Panamanian weevils as a group (Figures 6, 10) had astrong tendency toward bimodality, at least at the moreseasonal sites, with one peak occurring near the beginning ofthe rainy season in May/June and one later in that season,around October. A similar bimodality was found in manygroups of temperate weevils (Kuska, 1982; Petryszak, 1987,1988; Knutelski, 1988, 1991, 1993; Petryszak and Kaczmarc-zyk, 1992; Holecova, 1992), with some exceptions (Holecova,1991a, 1993a, 1993b, 1993c). A major difference, however,between temperate and tropical weevils is that in the tropics atleast some weevils are active at any time throughout the year.Among tropical weevils there are pronounced peaks inabundance, but the annual lows for total weevil abundance arerarely zero.

At the level of individual Panamanian species a very largevariation in seasonal patterns was observed. Some speciesoccurred throughout the year, sometimes without any clearseasonal variation, whereas at the other extreme some specieshad very short activity periods, with a sharp seasonal peak(Figure 13), just as do many temperate species. Many species,however, were bimodal in distribution, similar to manytemperate species (Stachowiak, 1991; Knutelski, 1993). Thedifference, again, is that species that are active the year round,with or without clear seasonal peaks, have not been observed inthe temperate zone, at least in areas with a cold winter. Forthose few temperate data sets where we have been able tocalculate the same seasonality measures used for the Panama-nian weevils, we found an absence of low values of the MeanVector, no species with nonsignificant values of Mean Vector,and a preponderance of high Mean Vector values. Notsurprisingly, the temperate data showed a much higher degreeof seasonality than the tropical ones, but that differences isquantified here for the first time. The means of the seasonaldistribution (Mean Week) for temperate weevils are concen-trated in the summer months only, with none occurring fromfall through spring. However, in a warm temperate area, such asthe Orange Free State in South Africa (Louw, 1987), theseasonal distributions of weevil species seems remarkablysimilar to the one observed for weevils in Panama. There too,some species were found active at any time of the year,although often with clearcut seasonal peaks. For other groupsof insects in Panama, a similar large variation in seasonalpatterns has been observed (Wolda, 1980; Wolda and Fisk,1981; McElravy et at., 1982; Wolda and Broadhead, 1985;Wolda and Flowers, 1985; Cwikla and Wolda, 1986; Woldaand Ramos, 1992; Wolda, unpublished data).

For most Panamanian species, seasonal patterns are rathersimilar in successive years, in degree of seasonality (MeanVector), and in the mean of the seasonal distribution (MeanWeek) (Figure 18), although there may be a shift with a later orearlier onset of the rainy season (Figures 11, 12). However,there are strong exceptions to this general rule (Figures 19-21).In a number of these instances, the between-year differencesobserved were so large that additional years of data would be

needed to determine whether we were dealing with a variablebut seasonal pattern or with a variation in abundance that wasunrelated to seasons. In other cases, it was clear that anyrelationship between abundance and climatic seasons wastenuous, if it existed at all. This is very different from thesituation among temperate weevils based on the small amountof information we have been able to glean from the literature.Between-year differences in seasonal patterns among thesetemperate weevils seems to be restricted to shifts of a fewweeks at best, often related to spring temperatures (Silver,1968; Fye and Bonham, 1970; Overgaard Nielsen. 1974c;Holecova, 1991a). Large between-year differences in seasonalpatterns in tropical insects are not restricted to weevils but alsohave been found in nocturnal bees (Wolda and Roubik, 1986),and Homoptera and Blattaria (Wolda, 1989; unpublished data).Trichoptera from Puerto Rico showed large between-yeardifferences in emergence patterns (Masteller and Flint,1992).

At intermediate degrees of seasonality, between-site varia-tion in Mean Week tends to be considerably larger thanvariation between years at each site (Figure 22), but for moreseasonal species, the ones with higher values for Mean Vector,no such difference was found. The variation in the degree ofseasonality was essentially the same between sites as between-years (Figure 23). Seasonal patterns of several weevil specieswere generally rather similar in different Panamanian sites, inspite of differences in altitude or climate, but in a number ofspecies the between-site differences were striking (Figures24-27), even when the climatic conditions at the sites weresimilar, such as on Barro Colorado Island and in Las Cumbres.The three species illustrated in Figure 27 clearly demonstratesuch large between-site differences in seasonality. In somecases, the different patterns were clearly related to differencesin climate. Pseudapotrepus macrophthalmus Champion (Fig-ure 27) was much less seasonal in Miramar, a site that isclimatically also relatively nonseasonal. On the other hand, inFortuna many clearly seasonal patterns were observed, al-though it also is rather nonseasonal in rainfall (Figure 26).Among other Panamanian insects, similar between-site differ-ences were observed (Wolda, 1982, unpublished data; Rutledgeet al., 1976). Examples of such between-site differences inseasonality in temperate weevils are rare in the literature (Fyeand Bonham, 1970), although they probably are common alonglarge latitudinal gradients. For smaller areas, however, thebetween-site differences seem to be small. Unfortunately, mostpublished data only allow qualitative comparisons as they donot permit calculations of measures, such as Mean Vector orMean Week. Stachowiak (1991) observed very similar seasonalpatterns in two different Polish sites. Knutelski (1993) foundthat the activity seasons at higher altitudes (± 1500 m) tendedto occur a little later than those at lower levels (± 900 m) in theTatra Mountains in southern Poland, but even there thedifferences seemed to be small relative to many of those foundamong the Panamanian weevils. The only data set known to us

NUMBER 590 41

that can be used for quantitative between-site comparisons(Cmoluch, 1962) suggests that, if anything, Panamanianweevils show higher between-site variability in both MeanWeek (Figure 22) and Mean Vector (Figure 23) than do weevilsfrom eastern Poland. However, the result is far from convinc-ing, and more data from different areas both in the temperatezone and in the tropics are needed to answer questions abouttropical-temperate differences in seasonality patterns.

The richness in seasonality patterns among tropical insectspecies is spectacular, as are the sometimes large between-yearand between-site differences in those patterns, as illustratedhere for Panamanian weevils. This is a first large-scaledescriptive study of those patterns, and analyses and explana-tions about the causes of these patterns and the between-species, between-year, and between-site differences thereinwill have to await future studies.

Appendix

List of Species of Weevils Collected by Light Trapsin Seven Localities in Panama

(BCI = Barro Colorado Island, LCC = Las Cumbres, MIR = Miramar, CGR = Corriente Grande, BOQ = Bo-quete, FOR = Fortuna, GUA = Guadalupe Arriba.)

Taxon BCI LCC MIR CGR BOQ FOR GUA

ANTHRIBIDAE

1980aAnthribidae2071 Anthribidae2688 Anthribidae2881 Anthribidae2883 Anthribidae2896 Anthribidae2941 Anthribidae2943 Anthribidae2956 Anthribidae2961 Anthribidae2962 Anthribidae2972 Anthribidae2987 Anthribidae2994 Anthribidae3021 Anthribidae3022 Anthribidae3044 Anthribidae3047 Anthribidae3048 Anthribidae3109 Anthribidae3112 Anthribidae3119 Anthribidae3126 Anthribidae3181 Anthribidae3185 Anthribidae3231 Anthribidae3239 Anthribidae3265 Anthribidae3271 Anthribidae3279 Anthribidae3283 Anthribidae3288 Anthribidae3293 Anthribidae3309 Anthribidae3327 AnthribidaeAnthribidae Gen.?Anthribidae n. gen. nr. AllandrusAnthribidae n. gen. nr. BrevibarraAnthribidae sp. #2Anthribidae sp. #3Anthribidae sp. #4Anthribidae sp. #7Anthribidae sp. #14Anthribidae sp. #18Corrhecerus mixtus JordonDomaptolis championi JordonEugonus decorus JordonEugonus n. sp. nr. particolor JordonEugonus subcylindricus Fahraeus

1811

11

11151154314417121111111111

1213111

14111121

541

1443

2

42

NUMBER 590 43


Euparius nigritarsis JordonEuparius polius JordonEuparius sp. #1Euparius sp. #5Euparius sp. #8Euparius sp. #9Euparius sp. #19Euparius sp. #20Euparius sp. #21Euparius stratus JordonEuparius suturalis JordonEuparius torquatus JekelGoniocloeus fractus JordonGymnognathus scalaris JordonGymnognathus sp. n. #1Homocloeus sp. near xanthopus JordonHypselotropis allatus JordonIschnocerus aeneus Jordon[schnocerus sp. #1Lagopezus inversus JordonLagopezus tenuicornis FabriciusMeanthribus apicalis JordonNeanthribus championi JordonMeanthribus pistor JordonOrmiscus sp. #1Phaenithon curvipes (Germar)Phaenithon discifer Jordon

722

21144134

21--23-1--13--2-52

Phaenithon sp.Piezocorynus dimidiatus JordonPiezocorynus plagifer JordonPiezocorynus sp. #1Piezocorynus sp. #2Piezocorynus sp. #10Piezocorynus sp. #12Piezocorynus sp. #17Ptychoderes brevis JordonPtychoderes rugicollis JordonPtychoderes tricostifrons FahraeusStenocerus angulicollis JekelStenocerus longulus Jekel

APIONIDAE

Apion bicolor GerstaeckerApion "complex"Apion costaricense WagnerApion cretaceicolle SharpApion darlingtoni KissingerApion grallarium SharpApion hastifer SharpApion inflatipenne SharpApion lebasii GyllenhalApion nodicorne SharpApion peculiare WagnerApion sp. #1Apion sp. #2Apion sp. #3Apion sp. #4Apion sp. #6Apion sp. #7Apion sp. #8Apion sp. #9Apion sp. #10Apion sp. #12

32641211

1182

2926

10

509

91821

20291792

6

18

6 6

2



Apion sp. #13Apion sp. #14Apion sp. # 15Apion sp. #16Apion sp. #17Apion sp. #18Apion sp. #19Apion sp. #20Apion sp. #21Apion sp. #22Apion sp. #23Apion sp. #24Apion sp. #25Apion sp. #26Apion sp. #27Apion sp. #28Apion sp. #29Apion sp. #30Apion sp. #31Apion sp. #33Apion sp. #34Apion sp. #35Apion sp. #36Apion sp. #37Apion sp. #38Apion sp. #40Apion sp. #41Apion sp. #42Apion sp. #43Apion sp. #44Apion sp. #45Apion sp. #46Apion sp. #47Apion sp. #48Apion sp. #49Apion sp. #50Apion sp. #51Apion sp. #52Apion sp. #53Apion sp. #54Apion sp. #55Apion sp. #56Apion sp. #57Apion sp. #58Chrysapion auctum SharpChrysapion chrysocomum (Gerstaecker)

ATTELABIDAE

Attelabus sp. # 1Auletobius optatus SharpAuletobius sp. #1Eugnamptus divisus SharpEugnamptus godmani SharpEugnamptus sp. # 1Eugnamptus sp. #2Eugnamptus sp. #3Eugnamptus sp. #4Eugnamptus sp. #7Eugnamptus sp. #8Eugnamptus sp. #10Eugnamptus sp. # 11Eugnamptus sp. #12Eugnamptus sp. #13

168065

211

1322

71

192

221

150211

247182

1016431

164

3961

21613

--

11241211

31

NUMBER 590 45


Euscelus sp. near breviceps (Sharp)Hemilypus sp. #1Pselaphorhynchites chiriquensis (Sharp)Pseudauletes inermis (Sharp)Rhynchites basalis Sharp

BRENTIDAE

2154 Brentidae2201 Brentidae2287 Brentidae2288 Brentidae2332 BrentidaeAbactrus championi SharpAcratussp. #1Acratus sp. #2Acratus sp. #3Arrhenodes belti (Sharp)Arrhenodes goudoti KirschArrhenodini belti (Sharp)Arrhenodini sp. #4Brentidae gen. #1; sp. #1Brentus clavipes SharpClaeoderes bivittata KirschClaeoderes sp. #1Ephebocerus mexicanus SharpHyperephanus hirtellus ErichsonHyperephanus sp. #1Nemobrenthus aeneipennis SharpNemocephalus femoratus SharpNemocephalus guatemalensis SharpNemocephalus puncticeps SharpNemocephalus sp. #1Nemocoryna godmani SharpNemocoryna sericata SharpParatrachelizus adustus (Boheman)Paratrachelizus aureopilosus (Senna)Paratrachelizus cognatus (Sharp)Paratrachelizus "complex"Paratrachelizus elevatus (Sharp)Paratrachelizus fracticornus (Sharp)Paratrachelizus frontalis (Sharp)Paratrachelizus robustus (Sharp)Paratrachelizus sp. #1Paratrachelizus sp. #2Paratrachelizus sp. #3Paratrachelizus sp. #4Paratrachelizus turgidirostris BohemanProteramocerus sp. #1Rhaphirhynchus sp. #1Rhaphirhynchus sp. #2Rhaphirhynchus sp. #3Schoenfeldtia impressicollis SennaStereobates pedator SharpStereobatinus efferus KleineStereodermus barbirostris SharpStereodermus breviceps SharpStereodermus calvus SharpStereodermus dentipennis SharpStereodermus dentipes SharpStereodermus filum complexStereodermus latirostris SharpStereodermus pygmaeus GyllenhalStereodermus sp. #3

3412621291_2717011017612711413117

QO

11516995431_41201519423561035121221211118_4417327504

-_____1-_1_------_--_---

___5----2_34_-1_---------6___-614

-___-_-----------_1-_1--

_-_--20--------------5---7_-193_

--____-4-----------3----

_--1-2-------------11--32_--3-_

-_____-----------2------

_-1----1--11--------2----3-12--_

-____1-----------7--2---

_-16--------16--------13--222

68



Stereodermus sp. #10Stereodermus sp. #11Stereodermus sp. #12Stereodermus sp. #13Stereodermus sp. #14Stereodermus sp. #15Stereodermus sp. #16Stereodermus sp. #17Stereodermus sp. #18Stereodermus sp. nr. dentipes SharpStereodermus sp. nr. puncticollis SharpTaphroderes apicalis SharpTaphroderes beltianus SharpTaphroderes oscillator SharpTaphroderes ventralis SharpTeramocerus belti SharpTrachelizini sp. #5Trachelizus turgidirostris (Boh.)Tychaeus myrmecophagus HerbstUlocerus sordidus SharpUlocerussp. #1

CURCULIONIDAE

ANTHONOMINAE

Anthonomus caeruleisquamis ChampionAnthonomus calvescens ChampionAnthonomus excelsus Clark and BurkeAnthonomus flavirostris ChampionAnthonomus fortunatus ClarkAnthonomus marmoratus ChampionAnthonomus monostigma ChampionAnthonomus paleatus ChampionAnthonomus prodigiosus ClarkAnthonomus pruinosus ChampionAnthonomus sextuberculatus ChampionAnthonomus sp. #2Anthonomus sp. #3Anthonomus sp. #5Anthonomus sp. #7Anthonomus sp. #8Anthonomus sp. #9Anthonomus sp. #11Anthonomus sp. #14Anthonomus sp. #17Anthonomus sp. #18Anthonomus sp. #19Anthonomus sp. #20Anthonomus sp. #21Anthonomus sp. #22Anthonomus sp. #24Anthonomus subparallelus ChampionAnthonomus sulcipygus ChampionAnthonomus triangularis ChampionAnthonomus triangulifer ChampionAnthonomus venustus ChampionAnthonomus veraepacis ChampionAtractomerus inaequalis ChampionLoncophorellus nitidus (Champion)Loncophorus fortis (Champion)Loncophorus fusiformis (Champion)Loncophorus obliquus ChevrolatLoncophorus santarosae (Clark)

21

2052

5

216652

38

414

11413

1

141

47861

110

114-471111

31

34122

-

-

_-3-

11

NUMBER 590 47

Taxon

Melexerus hispidus (Champion)Pseudanthonomus nucleon ClarkPseudanthonomus tachyon Clark

BAR1DINAEBaridinae Centrinini sp. #1Baridinae sp. #2Baridinae sp. #3Baridinae sp. #4Baridinae sp. #5Baridinae sp. #6Baridinae sp. #7Baridinae sp. #8Baridinae sp. #9Baridinae sp. # 10Baridinae sp. #12Baridinae sp. #13Baridinae sp. #14Baridinae sp. # 15Baridinae sp. #16Baridinae sp. #17Baridinae sp. # 19Baridinae sp. #21Baridinae sp. #24Baridinae sp. #25Baridinae sp. #26Baridinae sp. #27Baridinae sp. #28Baridinae sp. #29Baridinae sp. #30Buchananius carinifer (Champion)Buchananius sp. #1Buchananius sp. #2Buchananius sp. #3Buchananius sp. #4Buchananius sp. #5Cylindrocerus comma SchoenherrCylindrocerus glabripectus ChampionCylindrocerus subulatus ChampionCyrionyx scapulosus (Boheman)Cyrionyx sp. #1Cyrionyx sp. #2Diorymerus laeviusculus ChampionDiorymerus longirostris ChampionDiorymerus sp. #10Eugeraeus discifer ChampionGeraeus balaninoides ChampionGeraeus sp. #1Geraeus sp. #3Glyptobaris ugata BohemanIasides cincticollis ChampionLamprobaris cucullata ChampionLamprobaris sp. #1Loboderes flavicornis GyllenhalLoboderes sulphureiventris ChampionMadarellus eruptus ChampionParisoschoenus sp. #1Plocamus clavisetis ChampionPseudoccntrinus sp. #1Revena laevipennis Hustache

CAMAROTINAE

Camarotussp. #1

BCI

_-

272

---------

3415112121-------

712----1---2--1---------51-141-

1

LCC MIR CGR

1 11

1

111- - -_- - -_-

1- - --_- - -- - -- - -____-_

112

_43

_1

- - -_____

-_

- - -1

- - -— — —

1- - -

3_ _ —

11

_- - -__

1___

1

_

BOQ

_--

--------

------------------------111~

-

1*"1—1

—-

------—

-

FOR GUA

_--

---5 4321111---------111---

161

120

71

- -- -

— ~

1

11— ~-— •*— ~- -

-



CERATOPODINAE

Catiline sp. #1Ceratopus bisignatus BohemanCeratopus longiclava ChampionCeratopus sp. #1Ceratopus sp. #2Ceratopus sp. #3Ceratopus sp. #5Ceratopus sp. #6Ceratopus sp. #8Ceratopus sp. #9Ceratopus sp. #11Ceratopus sp. #12Ceratopus sp. # 13Ceratopus sp. #14Ceratopus sp. #15Ceratopus sp. #16Ceratopus sp. #17Ceratopus sp. #18Ceratopus sp. #19Ceratopus sp. #20Ceratopus sp. #21Ceratopus sp. #22Ceratopus sp. #23Ceratopus sp. #24Ceratopus tessellatus Champion

CEUTORHYNCHINAE

Hypocoeliodes dietzi ChampionHypocoeliodes sp. #1

COSSONINAE

Acamptini n. gen. # 1, sp. # 1Acamptini n. gen. #1, sp. #2Acamptopsis encausta ChampionAcamptus plurisetosus (Champion)Acamptus sp. # 1Acamptus sp. #2Acamptus sp. #3Acamptus verrucosus VossAnchacamptus mandli VossCaulophilus oryzae (Gyllenhal)Caulophilus rufotestaceus (Champion)Caulophilus sp. #laCaulophilus sp. #2Caulophilus sp. #2aCaulophilus sp. #3Caulophilus sp. #3aCaulophilus sp. #4Caulophilus sp. #4aCaulophilus sp. #5Caulophilus sp. #5aCaulophilus sp. #6aCaulophilus sp. #7aCaulophilus sp. #8aCaulophilus sp. #9Caulophilus sp. #9aCaulophilus sp. #10Caulophilus sp. #10aCaulophilus sp. #11Caulophilus sp. #12Caulophilus sp. #13Choerorrhynchus sp. # 1Cossoninae Rhyncolini sp. #2

26548

10112

776

114

12

27--1-11

--10226__

312018_1153__2____12_3421298_---3_-15-3

57---------_---__--_-------_-_

-3--------------_--632-4--_-27-_

---1-1--

125_2-1-9-2

108-_1527-1-1112_

-------63---1--7-1--_----1___-_

---------

152-5-16-1_----_--2_____

NUMBER 590 49

Taxon

Cossoninae n. gen. #1; sp. #1Cossoninae n. gen. #2; sp. #1Cossoninae n. gen. #3, sp. #1Cossoninae sp. #3Cossoninae sp. #4Cossoninae sp. #5Cossoninae sp. #6Cossoninae sp. #7Cossonus canaliculatus (Fabr.)Cossonus planirostris ChampionCossonus sp. #1Cossonus sp. #2Cossonus sp. #4Cossonus sp. #5Himatiumsp. #1Macrancyloides perlongus ChampionMicromimus continuus ChampionMicromimus dehiscens ChampionMicromimus minimus (Boheman)Micromimus sp. #1Micromimus sp. #2Micromimus sp. #3Micromimus sp. #6Micromimus sp. #7Micromimus sp. #8Micromimus sp. #9Micromimus sp. #10Micromimus sp. #11Micromimus sp. #12Micromimus sp. #14Micromimus sp. #15Micromimus sp. #16Micromimus sp. #17Micromimus sp. #18Micromimus sp. #19Oocorynus corrosus ChampionOocorynus sp. #1Pentarthrum elumbis (Boheman)Prionarthrus sp. #1Prionarthrus sp. #2Prionarthrus sp. #3Prionarthrus sp. #4Prionarthrus sp. #5Pseudapotrepus macrophthalmus ChampionPseudapotrepus sp. #1Pseudapotrepus sp. #2Pseudeucoptus macrocephalus ChampionPseudopentarthrum sp. #1Pseudopentarthrum sp. #2Rhinanisus sp. #1Rhinanisus sp. #3Rhisanisus planatus ChampionRhyncolini sp. #1Stenancylus sp. #1Stenomimus sp. #6Stenomimus sp. #8Stenomimus sp. #9

CRYPTORHYNCHINAEAcamptoides angustus ChampionAcamptoides sp. #1Acamptoides sp. #3Apteromechus deciduus Champion

BCI

1450391

-5

86---------

3359

1-

900131767

4544

31---1-

662237

---

625-17

2039

1945

1--23--7-8--

1964

-2

LCC

3115------1---------

385---------------5---4-------------1---

--5-

MIR

9981

-21

2---------1---

78-2-----7

19---------------

59—

-------

45--

----

CGR

4459

2--

1112--

1811----1

109-6

33-----

172--

3791-

14204-----

60-

48---11--

12187

10

----

BOQ

_

--------1------

233965

--21-------1-----—-——

—~

16529

—1-—--

——

-——-

FOR GUA

_ _

1207

---

450------- I l l--

651213

4022

------ --- -- -11- -- -~ —

— ~

•* •*

197429

1

•• ~

— "••

"* ~

10~* ~6— —— *

— —



Apteromechus flavopuntatus ChampionApteromechus nitidifrons ChampionApteromechus opacifons ChampionApteromechus parvus ChampionApteromechus pigmentatus ChampionApteromechus rugulifrons ChampionApteromechus scabrosus ChampionApteromechus sp. #3Apteromechus sp. #4Apteromechus sp. #5Apteromechus sp. #6Apteromechus sp. #7Apteromechus sp. #8Apteromechus sp. #9Apteromechus sp. #10Apteromechus sp. #11Apteromechus sp. #12Apteromechus sp. #13Apteromechus sp. #14Apteromechus sp. #15Apteromechus sp. #16Apteromechus sp. #19Apteromechus sp. #20Apteromechus sp. #21Apteromechus sp. #22Apteromechus sp. #23Apteromechus sp. #25Apteromechus sp. #26Apteromechus sp. #27Apteromechus sp. #28Arthrocorynus dotatus ChampionAtrichis sp. #1Atrichis sp. #2Atrichis sp. #3

Bothrobatys laticollis BohemanCoelosternus leporinus (Champion)Coelosternus quadnfasciatus (Champion)Coelosternus sp. nr. biolleyi (Champion)Coelosternus variisquamis (Champion)Cophes asperatus ChampionCophes gibbus ChampionCophes hieroglyphicus ChampionCophes longiusculus (Boheman)Cophes sp. #2Cophes sp. #3Cophes sp. #4Cophes sp. #6Cophes sp. #7Cophes sp. #8Cophes sp. #9Cophes sp. #10Cryptorhynchinae sp. #C1Cryptorhynchinae sp. #C2Cryptorhynchinae sp. #C4Cryptorhynchinae sp. #C5Cryptorhynchinae sp. #C7Cryptorhynchinae sp. #C8Cryptorhynchinae sp. #C9Cryptorhynchinae sp. #C10Cryptorhynchinae sp. #C 11Cryptorhynchinae sp. #C 12Cryptorhynchinae sp. #C13Cryptorhynchinae sp. #C14

41608-1_653__25762--21254133312115112

21119

-4-5111-1223151_3121--______----

_-__

26

9165

1138422966-

-

114_3I-3

134488

104

3-2

___---2

46324245389432

6--

_-_----

--_--______

10

NUMBER 590 51


Cryptorhynchinae sp. #C 15Cryptorhynchinae sp. #C 16Cryptorhynchinae sp. #C17Cryptorhynchinae sp. #C18Cryptorhynchinae sp. #C19Cryptorhynchinae sp. #C21Cryptorhynchinae sp. #C22Cryptorhynchinae sp. #C23Cryptorhynchinae sp. #C24Cryptorhynchinae sp. #C25Cryptorhynchinae sp. #C29Cryptorhynchinae sp. #C30Cryptorhynchinae sp. #C31Cryptorhynchinae sp. #C32Cryptorhynchinae sp. #C33Cryptorhynchinae sp. #C34Cryptorhynchinae sp. #C35Cryptorhynchinae sp. #C36Cryptorhynchinae sp. #C37Cryptorhynchinae sp. #C38Cryptorhynchinae sp. #C39Cryptorhynchinae sp. #C40Cryptorhynchinae sp. #C41Cryptorhynchinae sp. #C43Cryptorhynchinae sp. #C44Cryptorhynchinae sp. #C45Cryptorhynchinae sp. #C46Cryptorhynchinae sp. #C47Cryptorhynchinae sp. #C48Cryptorhynchinae sp. #C49Cryptorhynchinae sp. #C50Cryptorhynchinae sp. #C51Cryptorhynchinae sp. #C52Cryptorhynchinae sp. #C53Cryptorhynchinae sp. #C54Cryptorhynchinae sp. #C55Cryptorhynchinae sp. #C57Cryptorhynchinae sp. #C60Cryptorhynchinae sp. #C61Cryptorhynchinae sp. #C62Cryptorhynchinae sp. #C63Cryptorhynchinae sp. #C64Cryptorhynchinae sp. #C65Cryptorhynchinae sp. #C66Cryptorhynchinae sp. #C67Cryptorhynchinae sp. #C68Cryptorhynchinae sp. #C69Cryptorhynchinae sp. #C70Cryptorhynchinae sp. #C72Cryptorhynchinae sp. #C74Cryptorhynchinae sp. #C76Cryptorhynchinae sp. #C77Cryptorhynchinae sp. #C79Cryptorhynchinae sp. #C80Cryptorhynchinae sp. #C155Cryptorhynchinae sp. #CR1Cryptorhynchinae sp. #CR2Cryptorhynchinae sp. #CR3Cryptorhynchinae sp. #CR4Cryptorhynchinae sp. #CR5Cryptorhynchinae sp. #CR6Cryptorhynchinae sp. #CR7Cryptorhynchinae sp. #CR8

104-_10126_2966

1310

8510

22217482

6

14

530

59

7312429083166721252__

------111122

1111

166995


Taxon

Cryptorhynchinae sp. #CR10Cryptorhynchinae sp. #CR12Cryptorhynchinae sp. #CR13Cryptorhynchinae sp. #CR15Cryptorhynchinae sp. #CR17Cryptorhynchinae sp. #CR19Cryptorhynchinae sp. #CR20Cryptorhynchinae sp. #CR22Cryptorhynchinae sp. #CR24Cryptorhynchinae sp. #CR25Cryptorhynchinae sp. #CR26Cryptorhynchinae sp. #CR27Cryptorhynchinae sp. #CR28Cryptorhynchinae sp. #CR29Cryptorhynchinae sp. #CR30Cryptorhynchinae sp. #CR31Cryptorhynchinae sp. #CR32Cryptorhynchinae sp. #CR33Cryptorhynchinae sp. #CR34Cryptorhynchinae sp. #CR35Cryptorhynchinae sp. #CR36Cryptorhynchinae sp. #CR38Cryptorhynchinae sp. #CR39Cryptorhynchinae sp. #CR40Cryptorhynchinae sp. #CR41Cryptorhynchinae sp. #CR43Cryptorhynchinae sp. #CR44Cryptorhynchinae sp. #CR45Cryptorhynchinae sp. #CR46Cryptorhynchinae sp. #CR47Cryptorhynchinae sp. #CR48Cryptorhynchinae sp. #CR49Cryptorhynchinae sp. #CR50Cryptorhynchinae sp. #CR51Cryptorhynchinae sp. #CR52Cryptorhynchinae sp. #CR53Cryptorhynchinae sp. #CR54Cryptorhynchinae sp. #CR55Cryptorhynchinae sp. #CR56Cryptorhynchinae sp. #CR57Cryptorhynchinae sp. #CR58Cryptorhynchinae sp. #CR60Cryptorhynchinae sp. #CR61Cryptorhynchinae sp. #CR62Cryptorhynchinae sp. #CR63Cryptorhynchinae sp. #CR64Cryptorhynchinae sp. #CR66Cryptorhynchinae sp. #CR68Cryptorhynchinae sp. #CR69Cryptorhynchinae sp. #CR70Cryptorhynchinae sp. #CR75Cryptorhynchinae sp. #CR77Cryptorhynchinae sp. #CR79Cryptorhynchinae sp. #CR80Cryptorhynchinae sp. #CR82Cryptorhynchinae sp. #CR84Cryptorhynchinae sp. #CR85Cryptorhynchinae sp. #CR89Cryptorhynchinae sp. #CR90Cryptorhynchinae sp. #CR91Cryptorhynchinae sp. #CR92Cryptorhynchinae sp. #CR93Cryptorhynchinae sp. #CR94

BCI

_-9------------1-------1--3-----------7----2---

6043229------67227

16

LCC MIR CGR

_______________ _ _

2__ _ ______- - -- - -- - -___-___- - -- - ---_1______- - -____2

6143

1 1515

______

BOQ

_----1-----------------------8----------

20123222---------------_

FOR GUA

2018

112

1123115

1861

253211111541111

17122111121

196---1--------------_--__

NUMBER 590 53

Taxon

Cryptorhynchinae sp. #CR95Cryptorhynchinae sp. #CR96Cryptorhynchinae sp. #CR97Cryptorhynchinae sp. #CR98Cryptorhynchinae sp. #CR100Cryptorhynchinae sp. #CR101Cryptorhynchinae sp. #CR102Cryptorhynchinae sp. #CR103Cryptorhynchinae sp. #CR104Cryptorhynchinae sp. #CR105Cryptorhynchinae sp. #CR106Cryptorhynchinae sp. #CR107Cryptorhynchinae sp. #CR108Cryptorhynchinae sp. #CR109Cryptorhynchinae sp. #CR 110Cryptorhynchinae sp. #CR111Cryptorhynchinae sp. #CR112Cryptorhynchinae sp. #CR113Cryptorhynchinae sp. #CR114Cryptorhynchinae sp. #CR116Cryptorhynchinae sp. #CR117Cryptorhynchinae sp. #CR118Cryptorhynchinae sp. #CR119Cryptorhynchinae sp. #CR120Cryptorhynchinae sp. #CR121Cryptorhynchinae sp. #CR122Cryptorhynchinae sp. #CR123Cryptorhynchinae sp. #CR125Cryptorhynchinae sp. #CR126Cryptorhynchinae sp. #CR127Cryptorhynchinae sp. #CR130Cryptorhynchinae sp. #CR131Cryptorhynchinae sp. #CR134Cryptorhynchinae sp. #CR135Cryptorhynchinae sp. #CR136Cryptorhynchinae sp. #CR137Cryptorhynchinae sp. #CR138Cryptorhynchinae sp. #CR139Cryptorhynchinae sp. #CR140Cryptorhynchinae sp. #CR141Cryptorhynchinae sp. #CR142Cryptorhynchinae sp. #CR143Cryptorhynchinae sp. #CR144Cryptorhynchinae sp. #CR145Cryptorhynchinae sp. #CR147Cryptorhynchinae sp. #CR149Cryptorhynchinae sp. #CR150Cryptorhynchinae sp. #CR152Cryptorhynchinae sp. #CR154Cryptorhynchinae sp. #CR155Cryptorhynchinae sp. #CR156Cryptorhynchinae sp. #CR157Cryptorhynchinae sp. #CR158Cryptorhynchinae sp. #CR163Cryptorhynchinae sp. #CR164Cryptorhynchinae sp. #CR165Cryptorhynchinae sp. #CR166Cryptorhynchinae sp. #CR167Cryptorhynchinae sp. #CR169Cryptorhynchinae sp. #CR170Cryptorhynchinae sp. #CR171Cryptorhynchinae sp. #CR172Cryptorhynchinae sp. #CR173

BCI LCC MIR CGR BOQ FOR GUA

11

1 _ _ _ _ _ _4 _ _ _ _ _ _3 _ _ _ _ _ _

16 - - - - - -11 - - - - - -11 _ _ _ _ _

1 _ _ _ _ _ _1 _ _ _ _ _ _2 - - - - - -3 _ _ _ _ _ _2 - - - - - -5 _ _ _ _ - -5 _ _ _ _ _ _1 _ _ _ _ _ _

28 - - - - - -2 - - - - - -1 _ _ _ _ _ _

2 - - - - - -2 - - - - - -

188 - - - - - -5 _ i2 - - - - - -4 - - - - - -2 - - - - - -

96 - - - - - -1 7 - 1 - -

1 _ _ _ _ _ _96 - 1 - - -16 - - - - - -6884 - - - - ... -18 - - - - - -25 - - - - - -33 _ _ _ _ _ _24 - - - - - -33 - - - - - -

9 _ _ _ _ _ -11 - - - - - -11 _ _ _ _ _ -

14 - - - - - -7 _ _ _ _ _ _

14 _ _ _ _ _ _31 - - - - - -

4 _ _ _ _ _ -3 _ _ _ _ _ -7 _ _ _ _ _ _

12 - - - - - -47 - - - - - -34 _ _ _ _ - -16 - - - - - -4 _ _ _ _ _ _

14 _ _ _ _ _ _22 - - - - - -15 _ _ _ _ _ _11 _ _ _ _ _ _5 _ _ _ _ _ _3 _ _ - _ - -3 _ _ _ - - -g _ _ _ _ _ _2 - - - - - -2 - - - - - -



Cryptorhynchinae sp. #CR174Cryptorhynchinae sp. #CR175Cryptorhynchinae sp. #CR176Cryptorhynchinae sp. #CR177Cryptorhynchinae sp. #CR178Cryptorhynchinae sp. #CR179Cryptorhynchinae sp. #CR180Cryptorhynchinae sp. #CR181Cryptorhynchinae sp. #CR182Cryptorhynchinae sp. #CR183Cryptorhynchinae sp. #CR184Cryptorhynchinae sp. #CR185Cryptorhynchinae sp. #CR187Cryptorhynchinae sp. #CR188Cryptorhynchinae sp. #CR189Cryptorhynchinae sp. #CR190Cryptorhynchinae sp. #CR191Cryptorhynchinae sp. #CR192Cryptorhynchinae sp. #CR193Cryptorhynchinae sp. #CR196Cryptorhynchinae sp. #CR197Cryptorhynchinae sp. #CR198Cryptorhynchinae sp. #CR199Cryptorhynchinae sp. #CR200Cryptorhynchinae sp. #CR202Cryptorhynchinae sp. #CR203Cryptorhynchinae sp. #CR204Cryptorhynchinae sp. #CR205Cryptorhynchinae sp. #CR206Cryptorhynchinae sp. #CR207Cryptorhynchinae sp. #CR209Cryptorhynchinae sp. #CR210Cryptorhynchinae sp. #CR211Cryptorhynchinae sp. #CR212Cryptorhynchinae sp. #CR213Cryptorhynchinae sp. #CR214Cryptorhynchinae sp. #CR215Cryptorhynchinae sp. #CR216Cryptorhynchinae sp. #CR217Cryptorhynchinae sp. #CR218Cryptorhynchinae sp. #CR220Cryptorhynchinae sp. #CR221Cryptorhynchinae sp. #CR222Cryptorhynchinae sp. #CR223Cryptorhynchinae sp. #CR224Cryptorhynchinae sp. #CR225Cryptorhynchinae sp. #CR226Cryptorhynchinae sp. #CR227Cryptorhynchinae sp. #CR228Cryptorhynchinae sp. #CR229Cryptorhynchinae sp. #CR230Cryptorhynchinae sp. #CR231Cryptorhynchinae sp. #CR232Cryptorhynchinae sp. #CR233Cryptorhynchinae sp. #CR234Cryptorhynchinae sp. #CR235Cryptorhynchinae sp. #CR236Cryptorhynchinae sp. #CR237Cryptorhynchinae sp. #CR238Cryptorhynchinae sp. #CR239Cryptorhynchinae sp. #CR240Cryptorhynchinae sp. #CR241Cryptorhynchinae sp. #CR242

32223

212111111211

22111111111111111114423511111

55

NUMBER 590 55


Cryptorhynchinae sp. #CR243Cryptorhynchinae sp. #CR244Cryptorhynchinae sp. #CR245Cryptorhynchinae sp. #CR246Cryptorhynchinae sp. #CR247Cryptorhynchinae sp. #CR248Cryptorhynchinae sp. #CR249Cryptorhynchinae sp. #CR250Cryptorhynchinae sp. #CR251Cryptorhynchinae sp. #CR252Cryptorhynchinae sp. #CR253Cryptorhynchinae sp. #CR254Cryptorhynchinae sp. #CR255Cryptorhynchus aequalis (Champion)Cryptorhynchus albopunctatus (Champion)Cryptorhynchus alutaceus (Champion)Cryptorhynchus atrosignatus (Champion)Cryptorhynchus bifenestratus (Champion)Cryptorhynchus bimaculatus (Champion)Cryptorhynchus bioculatus (Champion)Cryptorhynchus cancellatus (Champion)Cryptorhynchus carinifer (Champion)Cryptorhynchus cinctipes (Champion)Cryptorhynchus cinereus (Champion)Cryptorhynchus collabismoides (Champion)Cryptorhynchus concentricus (Champion)Cryptorhynchus consobrinus RosenschoeldCryptorhynchus contaminatus (Champion)Cryptorhynchus cordubensis (Champion)Cryptorhynchus curtirostris (Champion)Cryptorhynchus decoratus (Champion)Cryptorhynchus degressus (Champion)Cryptorhynchus degressus var. (Champion)Cryptorhynchus disciger (Champion)Cryptorhynchus disjunctus ChampionCryptorhynchus duplaris ChampionCryptorhynchus erraticus (Champion)Cryptorhynchus eruptus (Champion)Cryptorhynchus evanescens (Champion)Cryptorhynchus ferox (Champion)Cryptorhynchus formosus (Champion)Cryptorhynchus foveatus BohemanCryptorhynchus foveifrons (Champion)Cryptorhynchus fraterculus (Champion)Cryptorhynchus furvus (Champion)Cryptorhynchus fuscatus (Champion)Cryptorhynchus humilis (Champion)Cryptorhynchus ignobilis (Champion)Cryptorhynchus infuscatus (Champion)Cryptorhynchus interlitus (Champion)Cryptorhynchus melanophthalmus (Champion)Cryptorhynchus mesomelas (Champion)Cryptorhynchus murinus (Champion)Cryptorhynchus octonotatus (Champion)Cryptorhynchus oculeus (Champion)Cryptorhynchus paleatus (Champion)Cryptorhynchus pallidisetis ChampionCryptorhynchus plagiaticollis (Champion)Cryptorhynchus plumipes (Boheman)Cryptorhynchus propinquus (Champion)Cryptorhynchus quadrisignatus (Champion)Cryptorhynchus sedulus (Champion)Cryptorhynchus simplex (Champion)

23

9

105856

4137

2122

19

5

1049

222

15

281153

1106

3

3

2

2

2

2183

2

4 3

2 1

4

51

611 12

1

2



Cryptorhynchus singularis (Champion)Cryptorhynchus sp. near biollei ChampionCryptorhynchus sp. nr. nigroplagiatus (Champion)Cryptorhynchus stigmaticus ChampionCryptorhynchus stigmatophorus ChampionCryptorhynchus strigatus (Champion)Cryptorhynchus subcaudatus (Champion)Cryptorhynchus tirunculus BohemanCryptorhynchus tortuosus (Champion)Cryptorhynchus uncipes (Champion)Cryptorhynchus undulatus (Champion)Cy lindrocorynus dentipes BohemanDiaporesis distincta PascoeElpinus palmatus ChampionElytrocoptus lemniscatus BohemanEpiscirrus propugnator (Gyllenhal)Eubulomus multicostatus ChampionEubulomus reflexirostris ChampionEubulomus sp. # 1Eubulomus sp. #2Eubulomus sp. #3Eubulomus sp. #4Eubulomus sp. #5Eubulomus sp. #6Eubulomus sp. #7Eubulomus sp. #8Eubulomus sp. #9Eubulomus sp. #10Eubulomus sp. #11Eubulomus sp. #15Eubulomus squamiventris ChampionEubulopsis edentata ChampionEubulus bifasciculatus ChampionEubulus brevis (Rosenschoeld)Eubulus carinitrons ChampionEubulus circumlitus ChampionEubulus coecus (Fabricius)Eubulus crinitus ChampionEubulus crispus ChampionEubulus crispus var. ChampionEubulus densus var. ChampionEubulus discoideus ChampionEubulus fulvodiscus ChampionEubulus gracilicornis ChampionEubulus ignifer ChampionEubulus marcidus ChampionEubulus melanotus ChampionEubulus miser ChampionEubulus moerens ChampionEubulus nigricollis ChampionEubulus nigrosignatus ChampionEubulus niveipectus Hustache

Eubulus ocellatus Champion

Eubulus orthomastius (Germar)

Eubulus punctifrons Champion

Eubulus reticulatus Champion

Eubulus signaticollis Champion

Eubulus sp. #1

Eubulus sp. #3

Eubulus sp. #4

Eubulus sp. #6

Eubuius sp. #7

107

12

1027

5736-31246413221218323723016114374117-1115102

411

17-41431-

151729-

1122423

5-

-

_--131108--___---_---151_11652-

__-

_

221_1-16-476419

NUMBER 590 57


Eubulus sp. #8Eubulus sp. #9Eubulus sp. #10Eubulus sp. #11Eubulus sp. #12Eubulus sp. #13Eubulus sp. #14Eubulus sp. #15Eubulus sp. #16Eubulus sp. #17Eubulus sp. #18Eubulus sp. #19Eubulus sp. #20Eubulus sp. #21Eubulus sp. #22Eubulus sp. #23Eubulus sp. #24Eubulus sp. #25Eubulus sp. #26Eubulus sp. #27Eubulus sp. #28Eubulus sp. #30Eubulus sp. #31Eubulus sp. #32Eubulus sp. #35Eubulus sp. #37Eubulus sp. #38Eubulus sp. #39Eubulus sp. #40Eubulus sp. #41Eubulus sp. #42Eubulus sp. #43Eubulus sp. #44Eubulus sp. #45Eubulus sp. #47Eubulus sp. #49Eubulus sp. #51Eubulus sp. #52Eubulus sp. #53

Eubulus sp. #54Eubulus sp. #56Eubulus sp. #57Eubulus sp. #58Eubulus sp. #59Eubulus sp. #60Eubulus sp. #61Eubulus sp. #62Eubulus sp. #63Eubulus sp. #64Eubulus sp. #65Eubulus sp. #66Eubulus sp. #67Eubulus sp. #69Eubulus sp. #70Eubulus sp. #71Eubulus sp. #72Eubulus sp. #74Eubulus sp. #75Eubulus stipator (Boheman)Eubulus truncatus ChampionEubulus unidentatus ChampionEuscepes longisetis ChampionEutinobothrus pilosellus (Boheman)

852344

1356111

186_

-

212

124

15

24

72111114231132121111112

28

-_---______1_

11

_

---_-

-

_

-------------_-------

-

-

1

2 2



Faustinus apicalis (Faust)Hemiliopsis nudicollis (Chevrolat)Isus mnigrum ChampionLeiomerus glabrirostris (Boheman)Macromeropsis binotata ChampionMacromerus numenius ErichsonMacromerus numenius succinctus ChevrolatMacromerus sp. #1Mantias gracilitarsis ChampionMerocnemus horni FaustMetadupus nodatus BohemanMetraniella nigrolineata ChampionMetraniella sp. #1Metraniella sp. #2Metriophilus cribricollis ChampionMetriophilus definitus (Rosenschoeld)Metriophilus fugax ChampionMetriophilus horridulus ChampionMetriophilus minimus ChampionMetriophilus miscellus ChampionMetriophilus nigroterminatus ChampionMetriophilus nitidus ChampionMetriophilus occultus ChampionMetriophilus ramosus ChampionMetriophilus ramulosus ChampionMetriophilus rugirrons ChampionMetriophilus sp. #2Metriophilus sp. #3Metriophilus sp. #4Metriophilus sp. #5Metriophilus sp. #6Metriophilus sp. #7Metriophilus sp. #8Metriophilus sp. #9Metriophilus sp. #10Metriophilus sp. #11Metriophilus sp. #12Metriophilus sp. #14Metriophilus sp. #15Metriophilus sp. #16Metriophilus sp. #17Metriophilus sp. #21Metriophilus sp. #22Metriophilus sp. #23Metriophilus sp. #24Metriophilus sp. #25Metriophilus sp. #26Metriophilus sp. #27Metriophilus sp. #28Metriophilus sp. #29Metriophilus sp. #30Metriophilus sp. #31Metriophilus v-fulvum ChampionMicroxypterus binotatus ChampionMicroxypterus suturalis ChampionOxypteropsis armata ChampionOxytenopterus asper BohemanOxytenopterus clotho (Kirsch)Oxytenopterus obliquus (Champion)Oxytenopterus sp. #1Oxytenopterus torvidus (Faust.)Phalias laticrus ChampionPhilonis inermis Champion

200147_437-_

26--_---_

17-2_---1

10-__---1

5____1-81

118____1115

22131109200

391

9419-22172

--_5

1289

_

7-

1-7811__

362

13032

131121

1238

15

5

7

100

19

1910

NUMBER 590 59


Phyrdenus divcrgens (Germar)Phyrdenus setiferus (Boheman)Phyrdenus subnotatus (Boheman)Pisaeus complanatus ChampionPisaeus sp. #1Pisaeus sp. #2Pisaeus sp. #3Pisaeus sp. #4Pisaeus sulcatus ChampionPisaeus various ChampionPseudomopsis bicristata ChampionPseudomopsis distigma ChampionPseudomopsis notaticollis ChampionPseudomopsis similis ChampionPseudomopsis sp. # 1Pseudomopsis sp. #3Pseudomopsis sp. #4Pseudomopsis sp. #5Pseudomopsis sp. #6Pseudomopsis sp. #8Pseudomopsis sp. #9Pseudomopsis sp. #10Pseudomopsis sp. # 12Pseudomopsis sp. # 13Pseudomopsis sp. #14Pseudomopsis sp. #15Ptous sp. # 1Ptous sp. #2Rhinochenus stigma (Linnaeus)Rhinochenus transversalis ChevrolatScedasus muricatus ChampionSemnorhynchus fulvopictus (Champion)Semnorhynchus planirostris (Champion)Semnorhynchus sp. #1Semnorhynchus sp. #2Semnorhynchus tristis (Champion)Siron exornatus (Boheman)Staseas granulatus ChampionStaseas granulatus var. ChampionStaseas mexicanus ChampionStaseas sp. #1Staseas sp. #2Staseas sp. #3Staseas sp. #5Staseas sp. #6Staseas sp. #10Staseas sp. #11Sternocoelus acutidens (Champion)Sternocoelus erubescens (Champion)Sternocoelus multidentatus (Fiedler)Sternocoelus sp. #1Sternocoelus sp. #2Sternocoelus sp. #3Sternocoelus sp. #4Sternocoelus sp. #5Sternocoelus tardipes (Boheman)Trachalus micronychus ChampionTyloderma aeneotinctum ChampionTyloderma circumcaribbeum WibmerTyloderma expansum WibmerTyloderma hustachei WibmerTyloderma lepidogramma WibmerTyloderma pilosellum (Chevrolat)

-213

25

29321

6301-

-

_--_

7510313

35

-

-

_

_

_

-

2_101

-2

1_213

_

2-4

-1

1_2-

_--1

20415

4295324

_

17-_

228589316_591133___79439_1

32_12-

5171__274

---57__11---3__---12

1-_---

6--_

-_1

--11-1_----___-----

_-_--1

2-3621--_

---4--2-----__4-1--

_-----

1------_

------------74---

Pi~

1


Taxon

Tyloderma variabile WibmerTyrannion albosignatum ChampionTyrannion breviculum ChampionTyrannion imbelle ChampionTyrannion nigrosellatum ChampionTyrannion scabidum ChampionTyrannion sp. #1Tyrannion sp. #2Tyrannion sp. #3Tyrannion sp. #4Tyrannion sp. #6Tyrannion sp. #9Tyrannion sp. #10Tyrannion sp. #11Tyrannion sp. #12Tyrannion sp. #14Tyrannion sp. #15Tyrannion sp. #16Tyrannion sp. #17Tyrannion sp. #18Tyrannion sp. #19Tyrannion sp. #20Tyrannion sp. #21Tyrannion sp. #22Tyrannion sp. #23Tyrannion sp. #24Tyrannion sp. #25Tyrannion sp. #26Tyrannion sp. #27Tyrannion sp. #28Tyrannion sp. #29Tyrannion sp. #30Tyrannion sp. #31Tyrannion sp. nr. tricnstatum ChampionTyrannion tricnstatum ChampionTyrannion validus ChampionUlosominus sp. #1Ulosomus sp. #1Zascelis brevicollis ChampionZascelis consputa (Boheman)Zascelis rugosa ChampionZascelis sp. #1Zascelis sp. #2Zascelis sp. #3Zascelis sp. #4Zascelis sp. #5

CURCULIONINAE

Terires "complex"Terires pilosus ChampionTerires plurisetosus Champion

DRYOPHTHORINAE

Dryophthorus americanus BedelDryophthorus sp. # 1Stenommatus sp. # 1Stenommatus sp. #2Stenommatus sulcifrons Champion

ENTIMINAE

Hypoptus macularis ChampionPromecops unidentata Champion

ERIRHININAE

Anchylorhynchus bicarinatus O'Brien

BCI

21----------

272-1

31114--------2------

1837171-1--

51

882168

-

131

16-

-

2

-

2

LCC

---------------------------------------2----

17

-

-13

125

----

-

17

-

MIR

----1------------------1---------------

10-----

-

3-

-

--

13-

2

-

-

-

CGR

--22-1-----1--2-------------2------1-3-------

-

7-

-

-25

1262930

-

-

-

BOQ

------142--15--------4

1-----------------1

1911-

-

--

-

----

-

-

-

-

FOR

--2

113----1

22------------2112------1------1---

-

--

-

--1-

524

-

-

-

GUA

---------------------------1

2-1111-----------

-

--

-

----

3

_

-

_

NUMBER 590 61

Taxon

Anchylorhynchus sp. # 1Andranthobius palmarum (Champion)Andranthobius sp. # 1Brachybamus sp. # 1Celetes attaleae (Champion)Celetes sp. #1Celetes sp. #2Cotithenesp. #1Cyrtobagous singularis HustacheDerelomini sp. #1Derelomini sp. #2Derelomini sp. #3Derelomini sp. #4Derelomini sp. #5Derelomini sp. #6Derelomini sp. #7Derelomini sp. #8Derelomini sp. #9Erirrhininae gen. #1; sp. #1Erirrhininae gen. #2; sp. #1Grypidiopsis variegata ChampionHelodytes foveolatus (Duval)Helodytes litus KuschelLissorhoptrus isthmicus KuschelNeochetina eichhorniae WarnerNotiodes sp. #1Notiodes sp. aeratus LeConte complex)Ochetina bruchi HustacheOchetina induta ChampionOnychylis meridionalis ChampionOnychylis setiger ChampionPenestes sp. #1Phyllotrox "complex"Phyllotrox marcidus ChampionPhyllotrox sp. #2Phyllotrox sp. #4Phyllotrox sp. #5Phyllotrox sp. #6Phyllotrox sp. #7Phyllotrox sp. #8Phyllotrox sp. #9Phyllotrox sp. #10Phyllotrox sp. #11Phyllotrox sp. #12Phyllotrox sp. #13Phyllotrox sp. #14Phyllotrox sp. #15Phyllotrox sp. #16Phyllotrox sp. #17Phyllotrox sp. #19Phyllotrox sp. #20Phytotribus sp. #1Pistiacola cretatus (Champion)Scybis pubescens ChampionTerioltes circumdatus ChampionTerioltes sp. #1Terioltes sp. #2Terioltes sp. #3

EUGNOMINAEUdeus eugnomoides ChampionUdeus sp. #1Udeus sp. #2Udeus sp. #3

BCI

64964

125932

7301-

156-

185

1429

----

126-712

2091-

18131-

43121--311-------11----13

434----

1-

12-

LCC

_240

-1-

43--------------1117--3-----

474------------------2-3----

13

10-

MIR

_---------6---31-------

16-----

452--

2564--------------2----------

--52

CGR

_------------6---1--------------

115--------73--

--11-----

322123

~

1---

BOQ

_--------6----------------------

2954

-34

51

-

-

-

-

-

1~—

-----1--

----

FOR GUA

_1-----

8--------2--------------

1204 3

- -1- -- ~

5923114

1

16

- --2 22

6453

25

-

_ _- -


Taxon

HYPERINAE

Larinosomus isthmica (Champion)Phelypera distigma (Boheman)

MAGDALIDINAE

Laemosaccus sculpturatus Champion

MOLYTINAEAeatus costulatus ChampionAeatus ebeninus ChampionAeatus sp. #1Aeatus sp. #2Aeatus sp. #3Aeatus sp. #4Aeatus sp. #5Anchonus sp. #1Anchonus sp. #2Arniticussp. #1Arniticus sp. #2Arniticus sp. #3Arniticus sp. #4Chalcodermus angulicollis FahraeusChalcodermus calidus (Fabricius)Chalcodermus curvipes ChampionChalcodermus dentipes ChampionChalcodermus ovalis FiedlerChalcodermus radiatus ChampionChalcodermus serripes FahraeusChalcodermus sp. #1Chalcodermus sp. #2Chalcodermus sp. #3Chalcodermus variolosus ChampionChalcodermus vittatus ChampionCholus canescens PascoeCleogonus armatus ChampionCleogonus fratellus FiedlerCleogonus rubetra (Fabricius)Conotrachelus albifrons ChampionConotrachelus albolineatus ChampionConotrachelus alborosaceus FiedlerConotrachelus annulipes ChampionConotrachelus arachnoides ChampionConotrachelus aristatus ChampionConotrachelus bilineatus ChampionConotrachelus brevisetis ChampionConotrachelus ciliatus ChampionConotrachelus compressus ChampionConotrachelus constrictus ChampionConotrachelus continuus ChampionConotrachelus crenatus ChampionConotrachelus cristatus FahraeusConotrachelus curtirostris ChampionConotrachelus curvicostatus MarshallConotrachelus dentiferus FahraeusConotrachelus dentimanus ChampionConotrachelus deplanatus ChampionConotrachelus diaconitus (Klug)Conotrachelus divirgatus ChampionConotrachelus divisus ChampionConotrachelus extrusus ChampionConotrachelus fasciculatus ChampionConotrachelus flavangulus ChampionConotrachelus flexuosus ChampionConotrachelus foveicollis Champion

BCI

4021

2

114485622---7---

24131018--

37396

361-

152257

-1

12-

51296

-101

-4-

50--4

429

861676

-----

120

-

LCC

16

-

-1031------3--16-4-22---

221-571-8---

10-5-2-7-

86-3_

95-_--1--63

MIR

-1

-

--------1------------------------5-------_

27-4_-8-----

2-

3

CGR BOQ

1

-

-

-----

---1 4-_---_----------1--

1-------

5---- 1253-_

1_--

74

-_-_

1

FOR GUA

-

-

-

-------1---6

1--11--------1-----12--9

321-

23-

21_-__ __-_--_

4-_

-

NUMBER 590 63

Taxon

Conotrachelus fulvescens ChampionConotrachelus fulvibasis ChampionConotrachelus fulvopictus ChampionConotrachelus gibbirostris ChampionConotrachelus incanus ChampionConotrachelus inexplicatus FaustConotrachelus lateralis ChampionConotrachelus latirostris ChampionConotrachelus leucocephalus ChampionConotrachelus longipennis ChampionConotrachelus longirostris ChampionConotrachelus multituberculatus (Fabr.)Conotrachelus obliquelineatus ChampionConotrachelus parvulus ChampionConotrachelus picticollis ChampionConotrachelus planifrons ChampionConotrachelus punctiventris ChampionConotrachelus quadrinodosus ChampionConotrachelus quadrinotatus FahraeusConotrachelus quadripustulatus ChampionConotrachelus rectirostris ChampionConotrachelus robustus ChampionConotrachelus rubidus ChampionConotrachelus scapularis FahraeusConotrachelus semirufus ChampionConotrachelus serpentinus (Klug)Conotrachelus sextuberculatus ChampionConotrachelus signatus KirschConotrachelus sinuaticollis ChampionConotrachelus sobrinus BohemanConotrachelus sp. #1Conotrachelus sp. #2Conotrachelus sp. #4Conotrachelus sp. #5Conotrachelus sp. #6Conotrachelus sp. #7Conotrachelus sp. #8Conotrachelus sp. #9Conotrachelus sp. #10Conotrachelus sp. #11Conotrachelus sp. #12Conotrachelus sp. #14Conotrachelus sp. #16Conotrachelus sp. #18Conotrachelus sp. #19Conotrachelus sp. #20Conotrachelus sp. #21Conotrachelus sp. #22Conotrachelus sp. #23Conotrachelus sp. #24Conotrachelus sp. #25Conotrachelus sp. #26Conotrachelus sp. #27Conotrachelus sp. #28Conotrachelus sp. #30Conotrachelus sp. #31Conotrachelus sp. #33Conotrachelus sp. #34Conotrachelus sp. #37Conotrachelus sp. #38Conotrachelus sp. #39Conotrachelus sp. #41Conotrachelus sp. #42

BCI

_

----7

2129--1

576-

2071783713---

1531

4202---

1012

1382-

809-

11-

43-383--2

10026

-759

1-----------

LCC

_

-4---2----3-

57-

365132--1--1

18-

15-

1758141921

123133

1262531

127

21212711---------

MIR CGR BOQ

- 117_

49__

1_

3326

27__

3491

- - -- - -5

1- - -- - -

19__

3- - -- - -__

1___- - -_- - -________

1______

1466

23111

15

FOR GUA

289-11

11-1------

55-8-1

15

38--61-----

---------- -- -- --- -- -

-- -

-- -- -5- -- -1-

1-1-- -24



Conotrachelus sp. #43Conotrachelus sp. #44Conotrachelus sp. #46Conotrachelus sp. #47Conotrachelus sp. #48Conotrachelus sp. #49Conotrachelus sp. #50Conotrachelus sp. #51Conotrachelus sp. #52Conotrachelus sp. #53Conotrachelus sp. #53aConotrachelus sp. #54Conotrachelus sp. #56Conotrachelus sp. #57Conotrachelus sp. #58Conotrachelus sp. #59Conotrachelus sp. #60Conotrachelus sp. #61Conotrachelus sp. #62Conotrachelus sp. #63Conotrachelus sp. #65Conotrachelus sp. #67Conotrachelus sp. #69Conotrachelus sp. #71Conotrachelus sp. #72Conotrachelus sp. #73Conotrachelus sp. #75Conotrachelus sp. #77Conotrachelus sp. #79Conotrachelus sp. #81Conotrachelus sp. #82Conotrachelus sp. #83Conotrachelus sp. #85Conotrachelus sp. #87Conotrachelus sp. #88Conotrachelus sp. #90Conotrachelus sp. #91Conotrachelus sp. #92Conotrachelus sp. #93Conotrachelus sp. #94Conotrachelus sp. #96Conotrachelus sp. #97Conotrachelus sp. #98Conotrachelus sp. #99Conotrachelus sp. #100Conotrachelus sp. #101Conotrachelus sp. #102Conotrachelus sp. #103Conotrachelus sp. #104Conotrachelus sp. #105Conotrachelus sp. #106Conotrachelus sp. #107Conotrachelus sp. #108Conotrachelus sp. #109Conotrachelus sp. #110Conotrachelus sp. #111Conotrachelus sp. #112Conotrachelus sp. #113Conotrachelus sp. #114Conotrachelus sp. #115Conotrachelus sp. #116Conotrachelus sp. #117Conotrachelus sp. #118

15

623

937

919343778581441

1516

2712

1

3

1636

161014

12

6783

28

1101038

-___1-_---___----__-_--_1_-___

-__-11

21321211

1421111311

142121

144172

292

NUMBER 590 65


Conotrachelus sp. #119Conotrachelus sp. #120Conotrachelus sp. #121Conotrachelus sp. # 122Conotrachelus sp. #123Conotrachelus sp. # 124Conotrachelus sp. #125Conotrachelus sp. #126Conotrachelus sp. # 128Conotrachelus sp. #129Conotrachelus sp. # 131Conotrachelus sp. #134Conotrachelus sp. #135Conotrachelus sp. # 136Conotrachelus sp. #137Conotrachelus sp. #138Conotrachelus sp. #139Conotrachelus sp. #142Conotrachelus sp. #144Conotrachelus sp. #147Conotrachelus sp. #150Conotrachelus sp. #151Conotrachelus sp. #153Conotrachelus sp. #155Conotrachelus sp. #157Conotrachelus sp. #159Conotrachelus sp. #160Conotrachelus sp. #161Conotrachelus sp. #162Conotrachelus sp. #163Conotrachelus sp. #164Conotrachelus sp. #165Conotrachelus sp. #168Conotrachelus sp. #169Conotrachelus sp. # 171Conotrachelus sp. # 172Conotrachelus sp. #173Conotrachelus sp. #174Conotrachelus sp. #175Conotrachelus sp. #176Conotrachelus sp. #177Conotrachelus sp. #179Conotrachelus sp. #180Conotrachelus sp. #181Conotrachelus sp. #182Conotrachelus sp. #183Conotrachelus sp. #184Conotrachelus sp. #185Conotrachelus sp. #186Conotrachelus sp. #187Conotrachelus sp. #188Conotrachelus sp. #189Conotrachelus sp. #190Conotrachelus sp. #191Conotrachelus sp. #192Conotrachelus sp. #193Conotrachelus sp. #194Conotrachelus sp. #196Conotrachelus sp. #197Conotrachelus sp. #198Conotrachelus sp. #199Conotrachelus sp. #200Conotrachelus sp. #201

1

39

163313

13

12122111

61112111

1633

-2---_-_----

_----_-_

2---_-

10288611122___

336311111195113116117118132-------

10_-

155

22



Conotrachelus sp. #202Conotrachelus sp. #203Conotrachelus sp. #204Conotrachelus sp. #205Conotrachelus sp. #206Conotrachelus sp. #207Conotrachelus sp. #208Conotrachelus sp. #209Conotrachelus sp. #210Conotrachelus sp. #211Conotrachelus sp. #212Conotrachelus sp. #213Conotrachelus sp. #214Conotrachelus sp. #215Conotrachelus sp. #216Conotrachelus sp. #217Conotrachelus sp. #218Conotrachelus sp. #219Conotrachelus sp. #220Conotrachelus sp. #221Conotrachelus sp. #222Conotrachelus sp. #223Conotrachelus sp. #224Conotrachelus sp. #225Conotrachelus sp. #226Conotrachelus sp. #228Conotrachelus sp. #229Conotrachelus sp. #230Conotrachelus sp. #231Conotrachelus sp. #232Conotrachelus sp. #233Conotrachelus sp. #235Conotrachelus sp. #236Conotrachelus sp. #237Conotrachelus sp. #238Conotrachelus sp. #239Conotrachelus sp. #240Conotrachelus sp. #241Conotrachelus sp. #242Conotrachelus sp. #243Conotrachelus sp. #244Conotrachelus sp. #245Conotrachelus sp. #246Conotrachelus sp. #247Conotrachelus sp. #248Conotrachelus sp. #249Conotrachelus sp. #250Conotrachelus sp. #251Conotrachelus sp. #253Conotrachelus sp. #254Conotrachelus sp. #255Conotrachelus sp. #256Conotrachelus sp. #257Conotrachelus sp. #258Conotrachelus sp. nr. anaglypticus (Say)Conotrachelus sp. nr. corallifer BohemanConotrachelus sp. nr. uniformis ChampionConotrachelus spinifer ChampionConotrachelus squamulatus ChampionConotrachelus striatirostris ChampionConotrachelus subfasciatus BohemanConotrachelus subulatus ChampionConotrachelus suturalis Champion

---111

------1---------11

11 I1----

211111112

-_ _

4 4

4

36

214 22

31159423

19

NUMBER 590 67

Taxon

Conotrachelus tegulatus FiedlerConotrachelus tetrastigma ChampionConotrachelus tnannulatus ChampionConotrachelus tridens ChampionConotrachelus turbatus FaustConotrachelus unidentatus ChampionConotrachelus venustus ChampionConotrachelus verticalis (Klug)Conotrachelus vittaticollis ChampionHeilipodus atrosignatus (Champion)Heilipodus biplagiatus (Boheman)Heilipodus choicus (Germar)Heilipodus cinctipennis (Champion)Heilipodus cynicus (Pascoe)Heilipodus dorbignyi (Guerin)Heilipodus jocosus (Boheman)Heilipodus lutosus (Pascoe)Heilipodus naevulus (Mannerheim)Heilipodus nigromaculatus (Champion)Heilipodus phrynodes (Pascoe)Heilipodus sp. nr. appendiculatus (Champion)Heilipodus spinipennis (Champion)Heilipodus suspensus (Pascoe)Heilipodus trinotatus (Champion)Heilipodus unifasciatus (Champion)Heilipus areolatus (Champion)Heilipus clathratus (Champion)Heilipus draco (Fabricius)Heilipus elegans GuerinHeilipus ornatus (Champion)Heilipus sp. #1Heilipus sp. #2Heilipus sp. #3Heilipus sp. #5Heilipus sp. #6Heilipus sp. #7Heilipus sp. #8Heilipus sp. #9Heilipus sp. #10Heilipus sp. #11Heilipus sp. #12Heilipus sp. #13Heilipus sp. #15Heilipus sp. #16Heilipus sp. #18Heilipus sp. #20Heilipus sp. #21Heilipus sp. #22Heilipus sp. #23Heilipus sp. #24Heilipus sp. #26Heilipus sp. #27Heilipus sp. #28Heilipus sp. #29Heilipus sp. #31Heilipus sp. #32Heilipus sp. #33Heilipus sp. #34Heilipus sp. #35Heilipus sp. #36Heilipus sp. #37Heilipus sp. #38Heilipus sp. #39

BCI

-54

-7557

72

153-

51-

421--

21199

3-

57--1-

152--

18-

11106

--1

5479

223

9121241

3-------1581

10132315

101

LCC

11--

68-

20130

---------3------7-----

19275---------1-------------------

MIR CGR BOQ

_

3 - 1 226

_10

1_

11

_2

_-_- - 4- - -

4- - -- - -

2 23- - -- - -- - -

3

53

- - 3_

1- - -- - -_

13

__

1__1__

4- - -

1_- - -- - -- - -- - -- — -- - -- - -- - -

11- - -

2-- - -- - -__

-

F O R G U A

_4------93---1

37---2-

322

---2 31-------- --1-- -

- -- -- -— —- -35121— —- -— ~1- -- -- —- -_ _

_ _- -- -


Taxon

Heilipus sp. #41Heilipus sp. #42Heilipus sulcifer ChampionHeilipus trifasciatus (Fabr.)Heilipus unifasciatus ChampionHeilus bioculatus (Boheman)Heilus guttatus (Boheman)Hilipinus latipennis ChampionHilipinus sp. #1Hilipinus sp. #2Hilipinus sp. #3Hilipinus sp. #4Hilipinus sp. #6Hilipinus sp. #7Hilipinus sp. #8Hilipinus sp. #9Hilipinus sp. #10Hilipinus sp. #11Hilipinus sp. #12Hilipinus sp. #13Hilipinus sp. #14Hilipinus sp. #15Hilipinus sp. #16Hilipinus sp. #17Hilipinus sulcicrus (Champion)Homalinotus dorsalis (Kirsch)Hypnideus multimaculatus Rosado NetoIthaura humilis KuschelIthaura nitida PascoeIthaura sp. # 1Ithyporini sp. #2Marshallius chiriquensis (Champion)Marshallius guttatus (Boheman)Marshallius leucostictus (Champion)Marshallius securifer (Champion)Marshallius sp. #1Marshallius sp. nr. securifer (Champion)Micralcinus sp. #1Microhyus erinaceus ChampionMicrohyus hystrix ChampionMicrohyus longisetis ChampionMicrohyus pallidisetis ChampionMicrohyus sp. # 1Microhyus sp. #2Microhyus sp. #6Microhyus sp. #7Microhyus sp. #9Microhyus sp. #10Mitrephorus curvilineatus HustacheOncorhinus latipennis ChampionOncorhinus scabricollis GyllenhalOzoctenus sp. # 1Ozopherus muricatus PascoeParabyzes angulosus (Champion)Pheloconus flavicans (Fiedler)Pheloconus rubicundulus (Boheman)Pheloconus sp. # 1Pheloconus sp. #2Pheloconus sp. #3Pseudanchonus debilis (Champion)Pseudanchonus larvatus KuschelPseudanchonus occultus (Champion)Pseudanchonus sp. # 1

BCI

1----

148-

4928

2162----44-

6833

---

364-

12038

2065-

72-----12-----112

12-

8222

36-41---_

41_

LCC

---1---1----------------21-5

134

72-8---------------2----

2-

10---__

4_

MIR

-11--

112---------------1-----1---2------------------------_-__

6_

CGR

----71------------1----------1--1--1--15-------3----------

11-

4_

BOQ

---------------------1-----------2---

170-----------------

18-_-

34310_

FOR GUA

-----1--4---

1523

14-------

1-----1-3-2

12-

15--6 113

5742----1_-_-

15-_

1-

585321

1

NUMBER 590 69


Pseudanchonus sp. #2Pseudanchonus tuberculifer (Champion)Rhineilipus cuvieri (Boheman)Rhineilipus intensus (Pascoe)Rhineilipus penicillatus (Champion)Rhineilipus sulcifer (Champion)Rhyparonotus sp. #1Rhyssomatus dilaticollis ChampionRhyssomatus nigerrimus FahraeusRhyssomatus nitidus ChampionRhyssomatus sp. #1Rhyssomatus sp. #2Rhyssomatus sp. #3Rhyssomatus sp. #4Rhyssomatus sp. #5Rhyssomatus sp. #6Rhyssomatus sp. #7Rhyssomatus sp. #8Rhyssomatus sp. #9Rhyssomatus sp. #10Rhyssomatus sp. #11Rhyssomatus sp. #12Rhyssomatus sp. #13Rhyssomatus sp. #14Sternechus brevicollis ChampionSternechus nitidus ChampionSternechus sp. #3Sternechus sp. #6Sternechus sp. #8Sternechus sp. #9Sternechus sp. #10Sternechus sp. #11Sternechus subrufus FiedlerThrasyomus sp. #1Thrasyomus sp. #2Thrasyomus rumens PascoeThrasyomus uniformis Champion

OTIDOCEPHALINAE

Hammatostylus argala (Erichson)Ludovix bifasciatus (Champion)Myrmex crassirostris (Champion)Myrmex grandis (Chevrolat)Myrmex laevipennis var. (Champion)Myrmex laevis (Champion)Myrmex sp. #1Myrmex sp. #2Myrmex sp. #3Myrmex sp. #4Myrmex sp. #5Myrmex sp. #6Myrmex sp. #9Pimelerodius sharpi (Sleeper)Pimelerodius sp. #2Prosicoderes bituberculatus (Champion)Sicoderus antilope (Fabricius)Sicoderus laevigatus (Champion)Sicoderus sp. #2Sicoderus sp. #4

PETALOCHILINAE

Hormops sp. #1Spermologus sp. #1

281551

_-31__641

2101323

1463222723610

621

1131_2_2141__5121111111

----

1111146--2

_-----

1271-----5

-

-

-----1--------1---_

--11

_---__---

_-----

_--_-----

-

-

---------1--------_

-11-

_-____---

_-----

_--------

-

-

_

---1--

11

1

23



POLYDROSINAE

Claeoteges obliterata ChampionClaeoteges sp. #1Compsus nigropunctatus ChampionCompsus sp. #1Eustylus sexguttatus ChampionExophthalmus carneipes ChampionExophthalmus jekelianus (White)Exophthalmus sp. #1Exophthalmus sp. #2Exophthalmus sp. #3Exophthalmus sp. #4Exophthalmus sp. #5Exophthalmus sp. #6Exophthalmus sulcicrus ChampionMacrostylus serripes (Champion)Macrostylus sp. #1Macrostylus sp. #2Macrostylus sp. #7Pandeleteius hieroglyphicus ChampionPolydacrys depressifrons BohemanPolydrosinae gen. #2; sp. #1Polydrosinae gen. #2; sp. #2Polydrosinae sp. #1Polydrosinae sp. #2Polydrosinae sp. #3Polydrosinae sp. #4Polydrosinae sp. #5Polydrusus sp. #1

PR1ONOMERINAE

Camptocheirus ornatus PascoeEctyrsus elongatus ChampionOdontopus sp. #2Odontopus sp. #3Odontopus sp. nr. femoralis (Champion)Piazorhinus sp. #1Piazorhinus sp. #2Piazorhinus sp. #3Piazorhinus sp. #4Piazorhinus sp. #5Piazorhinus sp. #6Piazorhinus sp. #7Themeropis binodosa ChampionThemeropis divergens Pascoe

RHYNCHAENINAE

Pedetinus halticoides (Champion)

RHYNCHOPHORINAE

Mesocordylus abditus VaurieMesocordylus bracteolatus (Boheman)Mesocordylus dispersus ChampionMesocordylus pustulosus ChampionMesocordylus secundus VaurieMesocordylus sp. #1Mesocordylus sp. #2Mesocordylus spumosus VaurieMesocordylus striatus (Boheman)Mesocordylus subulatus (Germar)Metamasius cincinnatus ChampionMetamasius dasyurus ChampionMetamasius hebetatus (Gyllenhal)Metamasius hemipterus sericeus (Olivier)Metamasius sp. #1

3111

13224

7 22

85 13

20

10

1310

5657---12238---12_

------_1521----_

6---____-----_

----__3_---351

---23____1--___

-_14515_47__11224

NUMBER 590 71


Metamasius sp. #2Orthognathus subparallelus (Chevrolat)Rhinostomus barbirostris (Fabricius)Rhinostomus thompsoni VaurieSitophilus oryzae (L.)

RHYTIRRHININAE

Listronotus dietrichi (Stockton)

TYCHIINAE

Lignyodes pallidus (Champion)Lignyodes sp. #2Lignyodes sp. #3Lignyodes sp. #4Lignyodes sp. #5Plocetes avertifer ClarkSibinia rotundata Champion

ZYGOPINAE

Arachnomorpha circumlineata ChampionArchocopturus regalis (Boheman)Copturomimus asperatus ChampionCopturomimus caeruleotinctus ChampionCopturomimus cinereus HellerCopturomimus sp. #1Copturomimus sp. #2Copturomimus sp. #3Copturomimus sp. #4Copturomimus sp. #5Copturomimus sp. #6Copturomimus sp. #7Copturomimus sp. #8Copturomimus sp. # 10Copturomimus sp. #11Copturomimus sp. #12Copturomorpha albomaculata ChampionCopturomorpha funerea ChampionCopturomorpha leucosticta ChampionCopturomorpha sp. #1Copturomorpha sp. #2Copturomorpha sp. #3Copturomorpha sp. #4Copturomorpha sp. #5Copturomorpha sp. #6Copturomorpha sp. #7Copturomorpha sp. #8Copturomorpha sp. #9Copturomorpha sp. #10Copturomorpha sp. #11Copturomorpha sp. # 12Copturomorpha sp. #13Copturomorpha sp. #15Copturomorpha sp. # 16Copturomorpha sp. #17Copturomorpha sp. #18Copturomorpha sp. #19Copturomorpha sp. #20Copturus colymbus HellerCopturus fulvomaculatus ChampionCopturus lamprothorax HellerCopturus lynceus ChampionCopturus sp. #1Copturus sp. #2Copturus sp. #3Copturus sp. #4

114842

511

322

1

2

2

20 2

57 13

1 14

3 - 41

1 - 11

13 12I

42



Copturus sp. #5Copturus sp. #7Copturus sp. #8Copturus sp. #10Copturus sp. #11Copturus sp. #12Copturus sp. #13Copturus sp. #15Copturus sp. #16Copturus sp. #17Copturus sp. #18Copturus sp. #19Copturus sp. #20Cratosomus aspersus ChampionCratosomus curassavicus (Voet)Cratosomus lentiginosus (Germar)Cratosomus sp. #1Cratosomus sp. #3Cratosomus sp. #4Eulechriops corusca ChampionEulechriops ductilis ChampionEulechriops sp. #6Eulechriops sp. #7Eulechriops sp. #8Eulechriops sp. #9Eulechriops sp. #10Eulechriops sp. # 11Eulechriops sp. #12Eulechriops sp. #13Eulechriops sp. #14Eulechriops sp. # 15Eulechriops sp. #16Eulechriops sp. #17Eulechriops sp. #18Eulechriops sp. #19Eulechriops sp. #20Eulechriops sp. #21Eulechriops sp. #22Eulechriops sp. #23Eulechriops sp. #24Eulechriops sp. #25Eulechriops sp. #26Eulechriops sp. #27Eulechriops sp. #28Eulechriops sp. #32Eulechriops sp. #33Eulechriops sp. #34Eulechriops sp. #35Eulechriops sp. #36Eulechriops sp. #37Eulechriops sp. #38Eulechriops sp. #39Eulechriops sp. #40Eulechriops sp. #41Eulechriops sp. #42Eulechriops sp. #43Eulechriops sp. #44Eulechriops sp. #45Eulechriops sp. #46Eulechriops sp. #47Eulechriops sp. #48Eulechriops sp. #49Eulechriops sp. #50

5181

1311

101

181222

612214231222113121

11

271212-_

NUMBER 590 73


Eulechriops sp. #51Eulechriops sp. #52Eulechriops sp. #53Eulechriops sp. #54Eulechriops sp. #55Eulechriops sp. #56Eulechriops sp. #57Eulechriops sp. #58Eulechriops sp. #59Eulechriops sp. #60Eulechriops sp. #61Eulechriops sp. #62Eulechriops sp. #63Eulechriops sp. #64Eulechriops sp. #65Eulechriops sp. #66Eulechriops sp. #68Eulechriops sp. #69Eulechriops sp. #70Eulechriops sp. #71Eulechriops sp. #76Eulechriops sp. #77Eulechriops sp. #78Eulechriops tenuirostns ChampionHoplocopturus scintillans ChampionIsotrachelus sp. #1Isotrachelus sp. #2Isotrachelus sp. #3Isotrachelus sp. #4Isotrachelus sp. #5Isotrachelus sp. #6Isotrachelus sp. #7Isotrachelus sp. #8Isotrachelus tibialis (Champion)Lechriops analis ChampionLechriops bicolor ChampionLechriops canescens ChampionLechriops disparilis ChampionLechriops maculiceps ChampionLechriops parilis ChampionLechriops parotica (Pascoe)Lechriops rufomaculata ChampionLechriops rugicollis ChampionLechriops sp. #2Lechriops sp. #3Lechriops sp. #4Lechriops sp. #5Lechriops sp. #6Lechriops sp. #7Lechriops sp. #10Lechriops sp. #11Lechriops sp. # 12Lechriops sp. # 13Lechriops sp. #15Lechriops sp. # 16Lechriops sp. #17Lechriops sp. #18Lechriops sp. #21Lechriops sp. #22Lechriops sp. #23Lechriops sp. #24Lechriops sp. #25Lechriops sp. #26

417921510134

5134

5191

27121

675

11121____

111

----2111-

_

1 12


Taxon

Lechriops sp. #27Lechriops sp. #28Lechriops sp. #30Lechriops sp. #31Lechriops sp. #32Lechriops sp. #33Lechriops sp. #34Lechriops sp. #35Lechriops sp. #36Lechriops sp. #37Lechriops sp. #38Lechriops sp. #41Lechriops sp. #42Lechriops sp. #43Lechriops sp. #44Lechriops sp. #45Lechriops sp. #46Lechriops sp. #47Lechriops sp. #48Lechriops sp. #53Lechriops sp. #54Lechriops vestita (Boheman)Lissoderes subnudus ChampionMacrolechriops sp. #1Microzurus championi HustacheMicrozurus trinotatus ChampionMnemynurus championi HellerMnemynurus poecilideres ChampionPhilenis fuscofemorata ChampionPiazorhinus sp. #4Piazurus alternans KirschPiazurus helleri ChampionPiazurus maculipes GyllenhalPiazurus sp. #1Piazurus sp. #2Piazurus succivus BohemanPiazurus sulphuriventris HellerPseudopiazurus centraliamericanus (Heller)Pseudopinarus quadratus (Champion)Pseudopinarus rana (Heller)Psomus (genus near Ps.) sp. #1Psomus sp. #1Psomus sp. #2Psomus sp. #3Psomus sp. #4Trichodocerus brevilineatus ChampionTrichodocerus brevilineatus var. ChampionTrichodocerus sp. # 1Trichodocerus sp. #3Trichodocerus sp. #4Trichodocerus spinolae ChevrolatZygopinae sp. # 1Zygopinae sp. #2Zygopinae sp. #3Zygopinae sp. #4Zygopinae sp. #5Zygopinae sp. #6Zygopinae sp. #7Zygopinae sp. #8Zygopinae sp. #9Zygopinae sp. #10Zygopinae sp. #11Zygopinae sp. #12

BCI

1111291111215--------1-7-42-2-1-

383--2-3-6

90233

23641

---

1652---____

2___

LCC MIR CGR

_

_____________

1123

_________

612

__

4_

1__

8___ _ _____

22 1 131

32

13____-__

1-

21

- _ _

BOQ

_------------1----------1--------------4---------__

2511111_____

FOR GUA

_---------------2-1

56

12---11---7-1-

1---9---------____________ __ _

2

NUMBER 590 75

Taxon

Zygops centromaculatus DesbrochersZygops maculipes DesbrochersZygops marmoreus DesbrochersZygops mexicanus BohemanZygops sp. #2Zygops sp. #3Zygops tridentatus GyllenhalZygops tripartitus Desbrochers

Subtotal IndividualsSubtotal Species

Total IndividualsTotal Species

BCI

114

120

1-

51

953331239

1137122030

LCC

_

------

-

4011357

MIR

_

------

-

3992170

CGR

_

2----3

-

2229259

BOQ

_

------

-

2256267

FOR

_

--2-1-

-

5500436

GUA

_

------

-

39151

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