334 Florida Entomologist 80(3) September, 1997

FEEDING RECORDS OF COSTA RICAN LEAF (COLEOPTERA: CHRYSOMELIDAE)

R. WILLS FLOWERS1 AND DANIEL H. JANZEN2 1Agricultural Research Programs, Florida A&M University Tallahassee, FL 32307-4100, rfl[email protected]

2Department of Biology, University of Pennsylvania, Philadelphia, PA 19104 [email protected]

ABSTRACT

Host associations are given for 137 species representing 7 subfamilies and 92 genera of Costa Rican Chrysomelidae. A numeric score is introduced to objectively describe confidence that a field observation of an interaction between a chrysomelid and a plant represents true herbivory. Literature host plant records, if they exist, are given for included chrysomelid taxa.

Key Words: herbivory, Criocerinae, Chrysomelinae, Cryptocephalinae, , , Hispinae, Lamprosominae, host

RESUMEN Se presentan asociaciones de plantas hospederas para 137 especies de Chrysome- lidae de Costa Rica, representando 7 subfamilias y 92 géneros de escarabajos. Se in- troduce una calificación numérica para describir objetivamente la confianza en que una observación de campo de una interacción entre un escarabajo y una planta repre- senta un caso verdadero de herbivoría. Se presentan datos de plantas hospederas de la literatura, si existen, para los taxa de escarabajos incluidos.

In recent years, there has been a surge of interest in relationships between tropi- cal plants and . The interest is driven by the related agendas of studying them for their intrinsic scientific interest, and protecting tropical biodiversity through find- ing practical and non-destructive ways to use it. The latter agenda is exemplified by the biochemical prospecting programs recently started in several areas of the world (Reid et al. 1993). Most plant- research begins with a basic event: an observation that a specific plant is somehow important in the life cycle of a specific insect. Unfortunately, huge

Flowers & Janzen: Chrysomelid Feeding Records 335 sections of the tropical insect fauna are still unusable as subjects of insect-plant re- search because that first step of linking plant and insect taxa has been largely ne- glected. In-depth studies of plant-insect interactions have focused on temperate zone insects and on a few relatively well known tropical groups (e.g., Lepidoptera). Only a small percentage of the fauna of tropical herbivores has been similarly studied. The family Chrysomelidae (Coleoptera), or leaf beetles, is a natural subject for studying plant-insect and inter-herbivore interactions (Strauss 1988). Of the esti- mated 37,000 species, world-wide, in this family, almost all, as far as we know, are herbivores or seed predators. However, for about 70% of the described species, we do not have records of host plants. Most of the known host plant records are Holarctic (Jolivet 1988b). For Neotropical Chrysomelidae other than Bruchinae, the most spe- cific information treats economically important species (e.g., King & Saunders 1984, Ostmark 1975, Jolivet 1979, Hilje et al. 1991). However, a review of known host plants of the tortoise beetles (Cassidinae) of Panama was recently published by Windsor et al. (1992); Moldenke (1971) listed host plants for some Mexican Chrysomelidae, and Anaya (1989) reviewed the known host plants of North and Central American Chry- somelinae. Jolivet, in a series of papers (1977, 1978, 1982, 1987a, 1987b, 1988a, 1991; Jolivet et al. 1986) and in a recent book (Jolivet & Hawkeswood 1995) summarized current host plant data on a world level for the Chrysomelidae. However, in much of this literature, species are usually identified only to and their plant hosts only to family. A few field studies have documented significant attacks by chry- somelids on plants in Central American ecosystems (e.g., Rockwood 1974, Memmott et al. 1993), and some detailed field and laboratory studies have been undertaken for several Neotropical species (Bach 1986, Begossi & Benson 1988, Buzzi & Winder 1986, Hsiao 1988, Strong 1977a,b). Apart from these ecological studies of specific chrysomelids, many of the published host plant records are of dubious value, stating merely that beetle X was taken on plant Y (or, all too often, “genus X feeds on plant genus Y”). A further problem, also noted by Furth (1985), is that a large proportion of such records are buried in taxonomic monographs and regional studies (e.g., Bechyné & Bechyné 1975) and accessible only by reading these studies in their entirety. Much more data on a much broader spectrum of chrysomelid taxa will have to be accumu- lated and made available before any credible generalizations about the nature of -plant interactions can be made. In this paper, we present feeding records of adults and larvae for 137 species of Costa Rican Chrysomelidae, representing 7 subfamilies and 92 genera. The majority of these observations were made by the senior author during a six-month sabbatical at Costa Rica’s Instituto Nacional de Biodiversidad (INBio) in 1991, and by the junior author during the years 1978 to 1995 as a byproduct of an on-going intensive study of the caterpillars of the dry forests of Sector Santa Rosa of the Guanacaste Conserva- tion Area (Janzen 1993, Janzen & Gauld 1996). Our records include results from di- rect observations of free-living feeding, feeding tests, and field associations. We have omitted many records where a single beetle was seen or collected on a plant, except for a few cases where the beetle was seen actively feeding. Beetles were identified by the senior author (Criocerinae, Cryptocephalinae, Lam- prosominae, Eumolpinae) and the following specialists: Catherine N. Duckett (Uni- versity of Puerto Rico, Alticini), Vilma Savini P. (Universidad Central de Venezuela, Alticini), David G. Furth (U.S. Natural History Museum, Alticini), Shawn M. Clark (West Virginia Department of Agriculture, ), Charles L. Staines (Maryland Department of Plant Protection, Hispini), and Edward G. Riley (Texas A&M Univer- sity, Cassidini). Plants were identified by the authors and Quirico Jiménez (INBio), Nelson Zamora (INBio), and Pablo Sanchez (Museo Nacional de Costa Rica).

336 Florida Entomologist 80(3) September, 1997

Our data are organized into a table with three supplementary appendices. Table 1 lists observations by chrysomelid taxon, gives field data in summary form, and lists voucher specimens. Appendix 1 is a key to plant family name abbreviations. Appendix 2 gives the full localities for locality codes used in Table 1. Appendix 3 gives miscella- neous field observations, as well as relevant literature citations for many of the chry- somelid taxa. In Table 1 we have followed the higher classification of Reid (1995) which reduces several well-known subfamilies to tribal status and confirms earlier opinions (eg. Crowson 1955, Lawrence 1982) that Bruchidae, or seed weevils, are a subfamily of Chrysomelidae. Bruchinae are not included in this report; for informa- tion on their host associations, see Janzen (1980a), Johnson (1990), and literature ci- tations therein. While not all workers fully agree with all aspects of Reid’s classification, it represents the latest and most comprehensive phylogenetic arrange- ment of the Chrysomelidae. For differing views, see Kingsolver (1995), Verma & Sax- ena (1996), and Reid (1996).

Explanation of Table 1 Leaf Beetle

Scientific names follow Wilcox (1983) and Flowers (1996). In a few cases, approxi- mate species identifications are indicated by “nr.” before the species name: e.g., Pla- giodera nr. uniformis. In some cases only generic identifications were possible, and distinct morphospecies are numbered as such.

Plant

Names follow current usage in the Costa Rica National Herbarium and in the bot- any department at INBio. In cases where species identification is approximate, the term “cf.” is used (e.g., Solanum cf. torvum).

Plant Family

Classification follows the listings of the Flora of Costa Rica by the Missouri Botan- ical Garden and INBio, viewable on the World Wide Web at http://cissus.mobot.org/ manual.plantas/lista.html. Families are coded by initial letters of their family names. See Appendix 1 for full listing.

Stage

A, adult; L, larva; P, pupa

Locality

See Appendix 2 for full locality data.

Date

Date of initial collection is given in cases where beetles were reared from larvae or held for testing.

Flowers & Janzen: Chrysomelid Feeding Records 337

Collectors

DHJ&WH: Daniel H. Janzen & Winnie Hallwachs RWF: R. W. Flowers Names of other collectors are given as they appear on voucher data labels.

Score

This is an attempt to objectively communicate our level of confidence that an ob- served association involved actual feeding by the chrysomelid. 6 Chrysomelids were observed in the field actually eating plant material. 5 Chrysomelids fed on plant when confined. 4 10 or more chrysomelids were collected from a plant and feeding damage that could reasonably be attributed to the beetles was present. 3 Five to nine chrysomelids were collected from a plant and feeding damage that could reasonably be attributed to the beetles was present, or 10 or more chry- somelids were collected from a plant but obvious feeding damage attributable to the beetles was not present. 2 Two to four chrysomelids were collected from a plant and feeding damage that could reasonably be attributed to the beetles was present, or five to nine chry- somelids were collected from a plant but obvious feeding damage attributable to the beetles was not present. 1 Two to four chrysomelids were collected from a plant but no noticeable feeding damage was observed.

Number (No.)

Number of vouchered specimens. In general, one feeding record equals one voucher; the few exceptions are mentioned in the Note column.

Voucher

Specimens collected by the senior author have voucher codes in the form “(Collec- tion No.)-RWF(Year)” and are deposited in INBio. Those collected by the junior author have codes in the form “(Year)-SRNP-(Number)” and are nominally specimens of IN- Bio but are on temporary loan to the University of Pennsylvania.

Note

These are numbered consecutively and appear in Appendix 3.

Appendix 2. Localities

Localities cited in Table 1 are listed on an approximate north-south gradient. The first letter of each locality code corresponds to the first letter of its province. Localities in the Area de Conservación Guanacaste also include Lambert Coordinates in paren- theses. Lambert Coordinates are used in Costa Rica in preference to latitude-longi- tude because the 1:50,000 topo sheets are gridded with Lambert Coordinates and, being metric, Lambert positions are easier to use.

338 Florida Entomologist 80(3) September, 1997

DISCUSSION

The data presented in these tables represent only the beginnings of the task of working out host plant relationships for the Central American Chrysomelidae. Our data cover less than 7% of the estimated 2000 chrysomelid species present in Costa Rica alone (Flowers, unpublished data). In some cases, our data confirmed previously published relationships between chrysomelid genera and host plant families (summa- rized in Jolivet & Hawkeswood 1995); 30 of our records represent host plant family range extensions, and 19 records are for chrysomelid genera in which, apparently, no host plants have been recorded previously. Most previously published host plant studies for the Neotropical Chrysomelidae (aside from focused studies on specific taxonomic groups, (e.g., Bach 1986; Begossi & Benson 1988; Windsor 1986) make no distinctions between accidental or casual asso- ciations of plant and beetle and true host relationships. The dangers in not making these distinctions have been demonstrated to us on several occasions when we found chrysomelid species that move off their food plants for resting or defecating. An exam- ple is Omophoeta simulans (Alticini, see Table 1), a group of which was first observed sitting on leaves of a Luehea sapling (Tiliaceae). Although large numbers of beetles were on the Luehea, and their frass was also evident on these leaves, closer inspection revealed that no feeding was taking place on the Luehea and that the true food plant (Evolvulus nummularis; Convolvulaceae) was growing beneath the . Similar warnings about possible confusion of Alticini food plants due to the beetle’s mobility have been given by Hawkeswood and Furth (1994). Nevertheless, collection records can still provide useful information—for many taxa opportunistic collecting has pro- vided the only information we have on possible host plants—if their limitations are clearly acknowledged. For our data we have included a “reliability scale” to roughly measure our confidence that a given association represents a true chrysomelid-host plant relationship. While ecological studies of narrow groups of chrysomelids or plants will always provide the most unambiguous data on feeding requirements, re- cent emphasis on and support for inventory collecting can rapidly increase knowledge of the feeding habits of a broad range of chrysomelids, if observations are qualified in some manner. We intend to continue expanding on the present work, and we encourage other col- lectors of Chrysomelidae to record, categorize and publish the plant associations they observe. Rapidly expanding our knowledge of chrysomelid-plant interactions is im- portant for two reasons. On the practical side, knowing host plants for more chry- somelid species will facilitate programs in chemical prospecting which are currently focused on plants. When a family of plants is being surveyed for active chemicals, the insects feeding on those plants represent another level of chemical derivatives avail- able for screening. The phytophagous insect may produce novel chemical varieties which cannot be synthesized directly from the host plant. A second area where more host plant data are needed is in the testing of hypothe- ses of the evolution of host plant selection. At present there are two competing theo- ries of what chiefly influences this evolution: phylogenetic and ecological mediation. Phylogenetic mediation (cospeciation) postulates that most cases of herbivory arise from cospeciation or parallel descent. This theory has become a popular explanation of host plant selection, under the name “coevolution” (though we caution the reader that this is not the original meaning of the word, see Janzen 1980b). Phylogenetic me- diation has been demonstrated in the Chrysomelidae for Phyllobrotica species (Gal- erucinae) and their hosts in the (Farrell and Mitter 1990). However, their study represents one of the few documented examples of coevolution (Anderson 1993).

Flowers & Janzen: Chrysomelid Feeding Records 339 1 4 116-RWF91 6 2 50-RWF91 2 1 3 80-RWF91 . COLUMNS

zondo OF C. Chavez C.

zondo, L. Rose L. zondo, EXPLANATION

FOR

TEXT

EE . S BEETLES

HIP A G10 25/V/1991 RWF 5 3 22-RWF91 3 DIO A,L G11 3/VII/1983 DHJ&WH 3 10 83-SRNP-707 1 SOL A G23 15/VIII/1991 RWF, MLP A G14 25/VI/1980 DHJ&WH 6 1 80-SRNP-239 4 Plant LEAF Family Stage Locality Date Collectors Score No. Voucher Note

ICAN R L. STE A G7 25/II/1990 RWF 6 3 9-RWF94 5

OSTA (Miers) C sp. sp. DIO DIO A A G2 20/VIII/1991 G2 Eli- R. RWF, 28/V/1991 Eli- R. RWF, OF

Schltdl. & Cham. Schltdl. Dioscorea convolvulaceae Dioscorea Dioscorea Solanum argentium Semialarium mexicanum indica Waltheria (L.) Kunth Byrsonima crassifolia Byrsonima Duval ex. Poiret Duval ex. Mennega

RECORDS

nr. nr. EEDING (Jac.) atricornis subapicalis 1. F nr. nr. sp.

ABLE Suffrian Lac. (Baly) Lema Chev. Lema Lema Metopoceris gemmans Griburius albilabris Chlamisius insignis Jac. Pseudochlamys megalostomoides Leaf Beetle Plant CRYPTOCEPHALINAE s.s. CRYPTOCEPHALINAE s.s. CRYPTOCEPHALINAE: CHLAMISINI CRYPTOCEPHALINAE: CRIOCERINAE T

340 Florida Entomologist 80(3) September, 1997 . COLUMNS 2 6 81-RWF91 10 6 13 74-RWF91 8

OF

Chavez Elizondo EXPLANATION

FOR

TEXT

EE . S BEETLES

LEAF

ICAN R SPI A G19 12/V/1991 RWF 6 11 67-RWF91 6 AST A G8 05/VII/1982 DHJ&WH 1 2 82-SRNP-471 FAB A G17 29/V/1991 RWF 2 6 105-RWF91 9 COC A G2 20/VIII/1991 R. RWF, MLP A G7 15/V/1978 DHJ&WH 6 2 78-SRNP-3.1 7 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

Kunth PIP A G23 15/VIII/1991 C. RWF, RECORDS

(Willd.) (Willd.) sp. AST A G1 18/V/1991 RWF 1 4 16-RWF91 sp. PIP A G1 17/V/1991 RWF 1 2 73-RWF91 EEDING ) F (Jacq.) Thouinidium Thouinidium dicandrum (Humb.&Bonpl.) Radlk. crassifolia Byrsonima Cochlospermum vitifolium Spreng Gliricidia sepium Piper auritum Piper Eupatorium albicaule Vernonia DHJ

Jac. nr. sp. ONTINUED sp. Jac. Har. 1. (C Jac. Har. ABLE cincta CRYPTOCEPHALINAE: CLYTRINI CRYPTOCEPHALINAE: Babia parvula Lamprosoma Oomorphus godmani Antitypona Chalcophana dis- color Chalcophana muta- bilis Chalcophana LAMPROSOMINAE Leaf Beetle Plant EUMOLPINAE T

Flowers & Janzen: Chrysomelid Feeding Records 341 . COLUMNS 1 2 48-RWF91

OF

EXPLANATION J. Saunders J.

FOR

TEXT

EE . S BEETLES

LEAF

ICAN R VIT AVIT G14 12/VI/1989VIT A DHJ&WH A G14 6 12/VI/1980 G10 DHJ&WH 05/VII/1980 1 89-SRNP-182 DHJ&WH 4 6 1 80-SRNP-107 1 80-SRNP-331 VIT A G19 12/VI/1991Tiffer R. RWF, 4 4 96-RWF91 AST A P3 14/V/1995 RWF 4 13 6-RWF95 AST A C1 5/IX/1991 Coto, D. RWF, ASC A G14 22/VI/1980 DHJ&WH 6 1 80-SRNP-190 ASC A G22 02/VII/1991 DHJ&WH 6 5 91-SRNP-1351 CAE A H1 22/I/1989 RWF 6 5 22-RWF94 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

(L.)

L. CAE A G17 29/V/1991 RWF 2 5 103-RWF91 moriflora RECORDS

cf. cf. (Schltdl.) (Schltdl.) sp. MYR A G1 18/V/1991 RWF 5 1 27-RWF91 11 Hook.&Arn. Croat EEDING ) F DC. unident. sp.unident. Vahl. MLP A G5 11/VI/1991 RWFDecne 6 2 101-RWF91 Critonia Cissus rhombifolia Cissus pseudosicy- oides Cissus pseudosicy- oides Cissus rhombifolia Salmea scandens Sarcostemma bi- lobum Blepharodon mucr- onatum Ardisia Cassia biflora Cassia reticulata Willd. (Miller) R. M. King & M. (Miller) R. Rob. H.

hy- ONTINUED Lef. nr. nr. 1. (C Lef. Jac. (Lef.) Jac. ABLE Colaspis Colaspis impressa Colaspis incon- stans Colaspis melanchol- ica Colaspoides uni- color Deuteronoda sutur- alis Lef. Leaf Beetle Plant pochlora T

342 Florida Entomologist 80(3) September, 1997 80-SRNP-191 80-SRNP-290 . 1 COLUMNS 6 2 63-RWF91 6 4 19-RWF94 6 5 15-RWF94

OF

RWF Aguilar DHJ&WH 6 1 EXPLANATION Gonzalez, T. T. Gonzalez,

torga, J. Solis J. torga, FOR

TEXT

EE 2/VII/1980 . S BEETLES

LEAF

ICAN R ASC A G14 22/VI/1980 ASC A G10 30/V/1981 DHJ&WH 6 2 81-SRNP-49 POL A A2 8/III/1990 RWF 3 5 20-RWF94 13 CAE A G11 23/VI/1989 DHJ&WH 6 4 89-SRNP-288 12 VER A G21 20/V/1991 RWF 2 5 111-RWF91 RUB A G1 15/V/1991 RWF 3 5 15-RWF91 14 RUB A G1 15/V/1991 RWF 2 4 8-RWF91 MIM A G29 17/II/1989 M. RWF, MIM A4 18/II/1994 As- Y. RWF, OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

L. COM A G21 20/V/1991 RWF 1 3 112-RWF91

Standley RECORDS

sp. ASC A G23 17/VIII/1991 Chavez, C. Willd. MIM A G8 22/IV/1983 DHJ&WH 6 1 83-SRNP-104 (Donn. (Donn. (Humb. (Humb. H.B.K. EEDING ) F Pithecellobium longi- folium &Bonpl.) Standl. (L.) L. Gonolobus Sarcostemma bilobum Sarcostemma glau- cum Swartzia cubensis germinans Avicennia Conocarpus erecta Coccoloba liportizii &N. Gómez-Laur. Zamora (Britt.&Wils.) Standl. (Britt.&Wils.) Pithecellobium longi- folium Inga vera Gonzalagunia bracteosa Smith) Robinson Ladenbergia sericophylla

hy- nr. nr. ONTINUED sp. (Lef.) Blake 1. (C ABLE (Jac.) Percolaspis sculpta Percolaspis Eumolpus robustus Horn Megascelis Metachroma clarkei Percolaspis poxantha Leaf Beetle Plant Phanaeta ruficollis Lef. T Flowers & Janzen: Chrysomelid Feeding Records 343 79-SRNP-14 80-SRNP-251 . 1 1 COLUMNS 1 3 43-RWF91 4 13 3-RWF94 16 6

OF

Weiler varría, H. H. varría, DHJ&WH 6 EXPLANATION J. Saunders J.

FOR

TEXT

EE 27/VI/1980 . S BEETLES

LEAF

ICAN R STESTE A A L, G10 G14 25/V/1991 16/V/1979 RWF 6 2 113-RWF91 SPO A G10 31/VII/1983 DHJ&WH 6 4 83-SRNP-908 LAU A G8 2/VII/1982 DHJ&WH 6 10 82-SRNP-426 15 CNV A G20 12/V/1991Tiffer R. RWF, 2 3 65-RWF91 RUBRUB ARUB A H1 A 22/I/1989 G4 C1 8/V/1991 RWF 5/IX/1991 RWF Coto, D. RWF, 4 11 3 21-RWF94 5 17-RWF91 MEL A G1 12/V/1991 RWF 2 4 5-RWF91 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

(L.) (L.) Roem. Roem. RECORDS

Standl. Benth. EEDING ) F Ipomoea pes-capra Br. R. Guazuma ulmifolia Lam. Guazuma ulmifolia Rondeletia buddle- oides Sommera donnell- smithii Sabacea villosa & Schult. Ocotea veraguensis Conostygia xalapensis (Bonpl.) D.Don (Meissn.) Mez. Manilkara chicle van Royan sp.unident. MEL A C2 5/II/1994 Cha- M. RWF,

sp. 1 sp. sp 2 sp. 3 sp. sp. 1 sp. ONTINUED sp. 1 sp. 2 sp. 1. (C Jac. ABLE CHRYSOMELINAE Calligrapha argus Stål Phanaeta Phanaeta Typophorus Rhabdopterus Typophorus Rhabdopterus &3 Typophorus vari- Typophorus abilis Leaf Beetle Plant T 344 Florida Entomologist 80(3) September, 1997 18 80-SRNP-69 82-SRNP-225 82-SRNP-533 83-SRNP-1398 83-SRNP-1453 . 2 7 1 1 COLUMNS 6 8 89-RWF91 6

OF

DHJ&WH 6 1 DHJ&WH 6 EXPLANATION

rales, A. Solís A. rales, FOR

TEXT

EE 14/VI/198 3/XII/1983 13/XII/1983 . S BEETLES

LEAF

ICAN R STE A G11 30/V/1981 DHJ&WH 6 2 81-SRNP-50 STE A L, G9 2/VII/1983 DHJ&WH 6 6 83-SRNP-691 AST A G23 16/VIII/1991 RWF 6 5 41-RWF91 19 FLAFLA A L, FLA A L, G9 A L, FLA G26 2/VII/1983 G15 A L, 20/VI/1980 DHJ&WH 15/VI/1983 DHJ&WH G15 DHJ&WH 6 30/V/1991 6 6 DHJ&WH 1 1 83-SRNP-698 9 6 80-SRNP-165 83-SRNP-428 12 91-SRNP-538 SOL A L, G1 16/V/1991 RWF 6 1 16-RWF94 17 APOAPO A A G8 24/VI/1983 G8 DHJ&WH 9/VII/1982 6 1 83-SRNP-564 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

L. MLV A S2 24/VI/1991 Cor- J. RWF, L. FLAA P, G10 6/VI/1980 RECORDS

(Dun.) Fern. EEDING ) F Byttneria aculeata Jacq. (Jacq.) Sida rhombifolia Ayenia micrantha Ayenia Standl. Solanum ochraceofer- rugineum crucis Prockia (HBK) Hemsley Arg. Müll. (Jacq.) Prockia crucis Prockia crucis Prockia Xylosoma flexuosum Xylosoma horrida Rose Koanophyllon pittieri & H. M. (Klatt) R. Robinson trifida Mesechites Prestonia allenii Woodson

. uni- . ONTINUED nr sp. Stål Stål Jac. 1. (C Stål (Rogers) Stål ABLE Calligrapha fulvi- pes Leaf Beetle Plant Calligrapha serpen- tina undec- Leptinotarsa imlineata Plagiodera cerea atritarsis Plagiodera formis Plagiodera Platyphora bicolor Jac. Platyphora petu- lans T Flowers & Janzen: Chrysomelid Feeding Records 345 . COLUMNS

OF

EXPLANATION

FOR

TEXT

EE . S BEETLES

LEAF

ICAN R FAB A G19 12/VI/1991 RWF 3 6 97-RWF91 25 APO A G10 7/VII/1989 DHJ&WH 1 2 89-SRNP-560 CLU A L, G1 17/VI/1991 RWF 6 19 40-RWF91 23 MLP A L, MLP G7 A L, MLP 20/VI/1978 G14 A L, DHJ&WH 2/VII/1980MLP G4MLP DHJ&WH 6 A 12/V/1991MLP A L, 6 3 G27 A G11 RWF 78-SRNP-83 2 4/VII/1983 16/VI/1989 G17 80-SRNP-292 20 DHJ&WH DHJ&WH 6 16/VI/1991 6 6 5 RWF 18 3 31-RWF91 89-SRNP-213 83-SRNP-732 6 21 1 115-RWF91 22 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

Jacq. MLP A G8 14/VI/1984 DHJ&WH 6 2 84-SRNP-557 (L.) L. CEC A G10 31/VII/1983 DHJ&WH 6 2 83-SRNP-903 Kunth. AMA A A1 23/V/1991 RWF 4 9 19-RWF91 24 sp. SOL A G23 20/III/1990 RWF 3 10 17-RWF94

RECORDS

(Cav.) Cuatr. (Cav.) EEDING ) F Hiraea reclinata Banisteriopsis muri- cata Banisteriopsis muri- cata Hiraea reclinata Hiraea reclinata Hiraea reclinata Witheringia Prestonia allenii Banisteriopsis muri- cata baccifera Vismia Pl. Tr.& Cecropia peltata Iresine diffusa Lonchocarpus acuminatus (Schecht.) Sousa (Schecht.) Stål ONTINUED sp. Jac. 1. (C sp. ABLE Stilodes neptis s.s. GALERUCINAE: Biblitea jansoni (Jac.) Stilodes modesta Jac. Caraguata pallida (Jac.) Coelomera Isotes Luperosoma vittatum Leaf Beetle Plant T 346 Florida Entomologist 80(3) September, 1997 28 82-SRNP-221 83-SRNP-802 83-SRNP-1479 . 1 2 COLUMNS

OF

DHJ&WH 6 1 EXPLANATION

FOR

TEXT

EE 19/XII/1983 14/VII/1983 . S G14 13/VI/1982 BEETLES

LEAF

A L, A L, ICAN R SOL A G23 16/VIII/1991 RWF 4 14 70-RWF91 RUT L RUT A L, G16 18/VI/1983 DHJ&WH 6 1 83-SRNP-454 BOR A S1 14/II/1990 RWF 6 25 18-RWF94 30 RUB A G2 20/VIII/1991 RWF 4 2 54-RWF91 29 MIM AMIM G4 A 9/V/1991 A4 18/II/1994 RWF RWF 6 2 6 1-RWF94 2 26 4-RWF94 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

L. VER A G14 28/VI/1995 DHJ&WH 6 1 93-SRNP-2995 Kunth. CON A G8 9/VII/1982 DHJ&WH 6 15 82-SRNP-512 RECORDS Standl.

sp. CNV A P3 10/V/1995 RWF 6 1 7-RWF95 31 sp. EUP A G23 15/VIII/1991 RWF 4 11 61-RWF91 27 P. Wilson P. EEDING ) F Pithecellobium palmanum Pithicellobium longi- folium Croton Solanum acerosum Sendt. Rourea glabra Zanthoxylum setulo- sum (Jacq.) Cordia eriostigma Pittier Ipomoea Essenbeckia littoralis Essenbeckia Smith Donn. Coutarea hexandra Lantana camara

ONTINUED sp. sp. (Fabr.) (Jac.) spp. n. sp. n. 1. (C ABLE Malacorhinus decempunctatus Jac. Masurius Monolepta Nestinus viridis Jac. Leaf Beetle Plant Yingaresca ALTICINI GALERUCINAE: Alagoasa seriata Asphaera nobilitata Nestinus (Jac.) Paranapiacaba ru- Paranapiacaba fofasciata T Flowers & Janzen: Chrysomelid Feeding Records 347 . COLUMNS 6 28 72-RWF91 35 4 5 5-RWF94 1 4 88-RWF91

OF

Weiler Chavez varría, H. H. varría, EXPLANATION

zondo, L. Rose L. zondo, FOR

TEXT

EE . S BEETLES

LEAF

ICAN R AST A G1 16/V/1991 RWF 6 2 4-RWF91 32 FAB A G11 12/VIII/1991 RWF 4 17 85-RWF91 SOL A A1 21/V/1991 RWF 3 4 37-RWF91 SOL A C3 5/II/1994 Cha- M. RWF, ERYERY AERY A G19 12/V/1991 A G11 12/VIII/1991 G9 RWF RWF 23/VI/1992 6 DHJ&WH 6 14 6 6 68-RWF91 8 84-RWF91 34 92-SRNP-2424 EUP A G23 14/VIII/1991 C. RWF, EUP A G23 4/V/1995 Ulate E. RWF, 6 19 8-RWF95 MLP A G17 19/V/1991 RWF 6 1 76-RWF91 33 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

m M. M.

sp. MLP A G2 28/V/1991 Eli- R. RWF, arboreum RECORDS

cf. cf. sp. MLV A G1 17/V/1991 RWF 2 5 45-RWF91 Jacq. EEDING ) F Euphorbia elata Brand Euphorbia elata R.&P. Solanum Pavonia Cestrum racemonsu Erythroxylum havan- ense Erythroxylum havan- ense Coursetia elliptica Coursetia Sousa & Rudd Humb. & Bonpl. ex & Bonpl. Humb. Duval Banisteriopsis Vernonia patens Vernonia H.B.K. crassifolia Byrsonima Erythroxylum havan- ense

sp.

ONTINUED sp. 2 sp. sp. 1 sp. 1. (C Bech. &Bech. Bech. lessmanni ABLE Bech.&Bech. Diphaltica Centralaphthona nr. Chaetocnema Diphaltica Diphaulaca aulica (Ol.) Asphaera reichei Har. minor Ayalaia &Bech. Bech. salvador- Ayalaia ense Leaf Beetle Plant T 348 Florida Entomologist 80(3) September, 1997 . COLUMNS 3 9 4 RWF 94 2 8 18-RWF91 4 25 94-RWF91 2 7 109-RWF91 38

OF

lar Rosa Weiler varría, H. H. varría, EXPLANATION

T. A. Ketchem A. T. FOR

TEXT

EE . S BEETLES

LEAF

ICAN R PAS A G9 27/VII/1992 DHJ&WH 6 8 92-SRNP-4026 STESTE LSTE A G9 A 21/XI/1987 G11 11/VIII/1995 H1 DHJ&WH DHJ&WH 11/I/1995 6 6 2 RWF 1 87-SRNP-1343 95-SRNP-7848 36 6 12 1-RWF95 SOL A A2 12/III/1990 RWF 3 26 11-RWF94 SOL A A1 14/VI/1991 RWF 3 6 56-RWF91 39 VER A P2 15/IX/1991Agui- R. RWF, LOG A C4 6/VIII/1991 & M. L. RWF, RUB A C2 5/II/1994 Cha- M. RWF, MIM A A2 12/III/1990 RWF 2 7 13-RWF94 MIM A A2 12/III/1990 RWF 1 2 12-RWF94 37 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

L. SIM A G2 6/V/1991 de la T. RWF, torvum sp FAB A G2 24/VIII/1991 RWF 3 5 59-RWF91 RECORDS

cf. cf. EEDING ) F Desmodium pulchella Passiflora Kunth Byttneria aculeata Byttneria aculeata Byttneria aculeata Solanum Sw. Quassia amara Inga sapindioides Willd. salicifolia Palicourea Standl. Lantana camara Inga sapindioides nitida Benth. Brugmansia candida Pers.

sp. sp. sp. sp. haagi ONTINUED sp. (Jac.) nr. nr. sp. 1 sp. sp. 2 sp. (Lat.) 1. (C sp. Clark ABLE Disonycha quinque- Disonycha lineata trifasci- Disonycha ata Epitrix Leaf Beetle Plant Glenidion Epitrix Genaphthona trans- versicollis Gioia Heikertingerella 2 Har. Heikertingerella 1 Hydmosyne T Flowers & Janzen: Chrysomelid Feeding Records 349 . COLUMNS 2 6 92-RWF91 42 3 12 93-RWF91 2 7 91-RWF91

OF

EXPLANATION

T. A. Ketchem A. T. T. A. Ketchem A. T. T. A. Ketchem A. T. FOR

TEXT

EE . S BEETLES

LEAF

ICAN R AST A C4 6/VIII/1991 & M. L. RWF, SOL A A1 23/V/1991 RWF 3 6 38-RWF91 CAPCAP ACAP A G1 A 12/V/1991 G1 G21 13/V/1991 RWF 23/VIII/1994 RWF RWF 1 4 2 2 25 28-RWF91 11 95-RWF91 26-RWF94 40 41 URT ACLU P1 A 17/II/1990 C4 RWF 6/VIII/1991 &. M. L. RWF, 4 15 14-RWF94 ONA A G28 19/VIII/1991 RWF 4 15 52-RWF91 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

L. BOR A G4 09/V/1991 RWF 2 6 11-RWF91

(L.) sp. SOL A P3 14/V/1995 RWF 2 1 5-RWF95 RECORDS sp. AST A A2 25/III/1996 RWF 6 6 1-RWF96

sp. AST A C4 6/VIII/1991 & M. L. RWF, Jacq. EEDING ) F Lycianthes multiflora Lycianthes Bitt. Witheringia Capparis frondosa Jac. Cleome parviflora Kunth Capparis odoratis- sima glabra Tournefortia Ageratina Senecio andicola Turcz. Urcra caracasana Griseb. (Jacq.) Clibadium Hypericum irazuense Kuntze Ludwigia erecta Hara lit- bor- sp. 1 sp. viola- sp. 1 sp. 2 sp. ONTINUED nr. nr. sp. nr. nr. sp. Bech. & Bech. 1. (C Bech. & Bech. Bech. ABLE Hypolampsis Leptophysa doni Leptophysa toralis Bech. Longitarsus Longitarsus Leaf Beetle Plant Lupraea violacea Jac. Lupraea cea Lupraea Lysathia T 350 Florida Entomologist 80(3) September, 1997 . COLUMNS 4 17 110-RWF91 1 2 108-RWF91 4 10 6-RWF94 45

OF

lar Weiler Aguilar EXPLANATION

Chavarría, H. H. Chavarría, FOR

TEXT

EE . S BEETLES

LEAF

ICAN R ERI A C3 5/II/1994 M. RWF, BIGBIG A A G14 21/VIII/1995 G10 DHJ&WH 22/VIII/1995 DHJ&WH 6 6 3 95-SRNP-8293 5 95-SRNP-8473 BURBUR ACNV A L, G16 A G13 18/VI/1983 15/VI/1985 DHJ&WH G14 DHJ&WH 21/VII/1994 6 6 RWF 1 3 83-SRNP-455 85-SRNP-401 6 43 5 27-RWF94 44 RUB A P3 14/V/1995 RWF 4 2 4-RWF95 ONA A P2 15/IX/1991 R. RWF, MELMEL A A G1 P2 12/V/1991 14/IX/1991 RWF RWF 3 2 25 4 5-RWF91 107-RWF91 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

(L.) Lam. PAS A P2 15/IX/1991 Agui- R. RWF, Tri- sp. SOL A G1 14/V/1991 RWF 2 4 3-RWF91 RECORDS

sp. PAS A G23 14/VIII/1991 RWF 3 1 60-RWF91 sp. MIM A G1 17/V/1991 RWF 2 2 44-RWF91 (L.) L. EEDING ) F ana Miconia schlimii Witheringia Conostegia xalapensis ochracea Tabebuia (Cham.) Standl. ochracea Tabebuia Passiflora Passiflora biflora Passiflora Sarg. simaruba Bursera Evolvulus nummu- laris Inga Cavendishia bracteata St.- ex J. (Ruis & Pav. Hil.) Hoerold Bursera simaruba Bursera Ludwigia erecta Gonzalagunia rosea

sp. 1 sp. 2 sp. costa- sp. 2 sp. 4 sp. sp. 1 sp. ONTINUED sp. nr. nr. Blake Jac. 1. (C Jac. (Jac.) ABLE Margaridisa Margaridisa Margaridisa Megistops ricensis Monomacra viola- cea Omophoeta simu- lans Paralactica Parasyphraea Parasyphraea Notozona nicara- guensis Nasigona pallida Jac. Leaf Beetle Plant T Flowers & Janzen: Chrysomelid Feeding Records 351 . COLUMNS 6 14 2-RWF94

OF

Weiler EXPLANATION

Chavarría, H. H. Chavarría, FOR

TEXT

EE . S BEETLES

LEAF

ICAN R STE A G11 11/VIII/1995 DHJ&WH 6 2 95-SRNP-7846 FAB A G13 18/VI/1991 RWF 4 7 39-RWF91 EUP A G11 12/VIII/1991 RWF 6 2 83-RWF91 47 RUB A G2 24/VIII/1991 RWF 1 2 53-RWF91 MIM A G17 22/VII/1994 RWF 4 52 23-RWF94 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

Standl. VER A A1 18/VIII/1991 RWF 2 9 55-RWF91 L. MIM A G17 25/V/1991 RWF 4 33 64-RWF91 46 sp. EUP A G8 21/V/1991 RWF 4 7 86-RWF91 sp. ERI A C5 5/II/1994 M. RWF, sp. RUB A A3 26/VII/1994 RWF 5 1 24-RWF94 RECORDS

caribaeum sp. PAS A G23 14/VIII/1991 RWF 3 3 60-RWF91 sp. sp. FLA FLA A A G23 16/VIII/1991 G23 RWF 4/V/1995 Ulate E. RWF, 6 2 4 5 114-RWF91 9-RWF95

Standl. EEDING ) F Cavendishia Byttneria aculeata Canavalia brasilensis Mart. ex Brent. Mart. Dalechampia Mimosa pigra Bernardia nicara- guensis Xylosoma Xylosoma Spermacoce Mimosa pigra Lippia torresii Passiflora Exostema Roem & Schult (Jacq.)

clarki austri- Blake . stra- . ONTINUED nr. nr. nr nr. nr. (Jac.) Har. 1. (C (Jac.) (Fab.) (Schauf.) ABLE Phenrica aca Platyprosopus pal- lens Plectotetra Leaf Beetle Plant Syphrea parvula Jac. Resistenciana pana- mensis Strabala acuminata costaricensis Syphrea bibiana Bech. Baly Resistenciana ob- scura Ptocadica minea T 352 Florida Entomologist 80(3) September, 1997 . COLUMNS 2 4 87-RWF91 3 11 87-RWF91 51

OF

Rose RWF, R. R. RWF, EXPLANATION Elizondo, L. L. Elizondo,

zondo, L. Rose L. zondo, FOR

TEXT

EE . S 24/VIII/1991 BEETLES

LEAF

ICAN R SPI A G11 5/VII/1980 DHJ&WH 6 1 80-SRNP-340 49 BIG A G18 11/VIII/1989 DHJ&WH 6 6 89-SRNP-863 50 STE A H1 11/I/95 RWF 6 14 1-RWF95 CAE A G17 29/I/1989 RWF 1 2 10-RWF94 EUP AEUP G1EUP AEUP 18/V/1991 A G31 A 30/I/1994 RWF G20 13/VIII/1991 G4Tiffer R. RWF, RWF 2 4 10/V/1991 6 7 3 RWF 25-RWF91 49-RWF91 16 4 1-RWF 94 8 7-RWF91 RUB A G2 24/VIII/1991RUB Eli- R. RWF, A G2 28/V/1991 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C

OF

RECORDS

sp. MLV A G1 17/V/1991 RWF 1 2 45-RWF91 48 Pax & K. & K. Pax EEDING ) F Pavonia hetero- Dalechampia morpha Hoffm. O.Will- & L. Standl. iams Serjania schiedeana Schltdl. unident sp. alata H.B.K. ACA A G2 20/VIII/1991 RWF 2 4 51-RWF91 Caperonia palustris Caperonia palustris apodanthes Acalypha Byttneria aculeata Cassia biflora Lindenia rivalis Benth. Lindenia rivalis (L.) A. St.-Hil. (L.) A.

sp. ONTINUED spp. (Schauf.) (Fab.) 1. (C (Jac.) ABLE Syphrea Systena sulphurea Jac. tenui- Walterianella cinta ve- Walterianella nustula Walterianella hu- Walterianella meralis Walterianella Leaf Beetle Plant T Flowers & Janzen: Chrysomelid Feeding Records 353 . COLUMNS 4 14 8-RWF94 54

OF

EXPLANATION

torga, J. Solis J. torga, FOR

TEXT

EE . S BEETLES

LEAF

ICAN R BIGBIG A A L, G10 G9 5/VI/1982 30/VI/1989 DHJ&WH DHJ&WH 6 6 1 2 82-SRNP-169 89-SRNP-448 55 STE A G2 6/V/1991 RWF 1 2 12&13-RWF91 FAB L G17 20/VII/1992 DHJ&WH 6 3 92-SRNP-3595 POA A G30 29/I/1994 RWF 4 26 7-RWF94 53 MLP A G22 27/I/1992 DHJ&WH 6 1 92-SRNP-294 MIM A G9 24/V/1991 RWF 3 6 21-RWF91 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

L. MLP A A5 18/II/1994 As- Y. RWF, sp. RUB A A3 26/VII/1994 RWF 5 1 25-RWF94 RECORDS

sp. MLP L G23 5/III/1991 DHJ&WH 6 1 91-SRNP-74 sp. ZIN A A1 14/VI/1991 RWF 4 1 79-RWF91 52 Benth. EEDING ) F Costus Poaceae Bunchosia Malpighia glabra Spermacoce Centrosema macrocar- pum ochracea Tabebuia Guazuma ulmifolia crassifolia Byrsonima ochracea Tabebuia Centrosema macrocarpum

ONTINUED sp. Baly (Crotch) Uh. 1. (C Pic (Baly) ABLE HISPINAE: s.s. HISPINAE: Cephaloleia suturalis Chalepus bellula (Chapuis) Carinispa never- manni Leaf Beetle Plant Demotispa strandi Uh. Oxychalepus alie- Oxychalepus nus Sumitrosis Uroplata varicos- tata CASSIDINI HISPINAE: Akantaka insidiosa Boh. Xenochalepus Xenochalepus omogera T 354 Florida Entomologist 80(3) September, 1997 . COLUMNS

OF

EXPLANATION

FOR

TEXT

EE . S BEETLES

LEAF

ICAN R BIGBIG A L, A G24 16/VII/1989 G10 DHJ&WH 27/III/1984 6 DHJ&WH 2 6 89-SRNP-701 3 84-SRNP-42 BIGBIG ABIG A L, BIG G11 A L, G10 18/VI/1991 A L, G10 22/VI/1989 G10 3/VI/1992 RWF DHJ&WH 19/V/1994 DHJ&WH 6 DHJ&WH 1 6 6 6 2 89-SRNP-296 5 9 66-RWF91 92-SRNP-1511 94-SRNP-3006 BORBOR ABOR A G10 A L, 30/V/1982 G13 G25 DHJ&WH 16/VI/1989 16/V/1979 DHJ&WH 6 DHJ&WH 6 1 6 82-SRNP-145 3 4 89-SRNP-218 79-SRNP-17 RUBCNV A A L, G15 G17 13/VIII/1982 DHJ&WH 18/I/1989RUB 6 DHJ&WH A 1 6 82-SRNP-721 G11 6 57 4/V/1980 89-SRNP-11 DHJ&WH 6 8 80-SRNP-37 58 MAR A G23 14/VIII/1991 RWF 4 4 75-RWF91 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

S.

(R. & (R. (L. (L. (Ver- RECORDS

Donn. Smith Donn. sp. sp. CNV CNV A L G8 10/VI/1989 G8 DHJ&WH 25/VII/1995 DHY&WH 6 6 2 89-SRNP-162 1 56 95-SRNP-7192 EEDING ) F Tabebuia ochracea Tabebuia Watson Tabebuia rosea Tabebuia Calathea crotalifera Ipomoea Ipomoea Ipomoea trifida Don (Kunth) G. Cordia alliodora Cordia alliodora Cydista diversifolia Cydista diversifolia Cydista diversifolia tol.) DC. Guettarda mac- rosperma Cordia alliodora Oken P.) Alibertia edulis A. Rich. Rich.) Cydista diversifolia Miers (H.B.K.)

sp. nr. nr. sp. ONTINUED (Boh.) 1. (C Spaeth ABLE Aslamidium Charidotella egregia Charidotis costari- cea Chelymorpha Coptocycla leprosa Coptocycla sordida Boh. Boh. Leaf Beetle Plant T Flowers & Janzen: Chrysomelid Feeding Records 355 60 82-SRNP-13 80-SRNP-309 83-SRNP-699 80-SRNP-218 82-SRNP-323 79-SRNP-311 . 1 1 1 1 4 1 COLUMNS 6 4 76 47-RWF91 61 4 27 90-RWF91 1

OF

Ketchem TA & K. I. I. & K. TA DHJ&WH 6 DHJ&WH 6 DHJ&WH 1 EXPLANATION J. Saunders J.

Flowers, LM, LM, Flowers, FOR

TEXT

EE 8/I/1982 2/VII/1983 24/VI/1982 . S G9 4/VII/1980 G8 23/VI/1980 G10 5/XI/1979 BEETLES

LEAF

A L, A L, L, A L, ICAN R BIG A L, G25 16/V/1979 DHJ&WH 6 1 79-SRNP-16 59 BIG P L, BIG G25 A 16/V/1979 DHJ&WH G3 1 9/VII/1991 RW & CA 2 79-SRNP-16B LAU A BOR L G19 4/VII/1983 DHJ&WH 6 24 83-SRNP-714 64 BOR A G11 14/V/1985 DHJ&WH 6 2 85-SRNP-185 BOR A BOR A BUR A G10 9/I/1982 DHJ&WH 1 4 82-SRNP-17 62 OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

L. BOR A C1 5/IX/1991 Coto, D. RWF, L. BORA L, G5 11/VI/1991 RWF 6 1 100-RWF91 (Mart. ex (Mart. RECORDS

11890 BOR A G14 13/V/1980 DHJ&WH 6 1 80-SRNP-56 63 EEDING ) F Hemsl. Bourreria huanita Cordia Cordia inermis Cordia inermis Bursera simaruba Bursera Cordia panamensis Cordia spinescens Cordia panamensis Riley Tabebuia impetigi- Tabebuia nosa Ocotea veraguensis Tabebuia Tabebuia impetiginosa Standl. DC.) Tabebuia impetigi- Tabebuia nosa

ONTINUED nr. nr. (Boh.) 1. (C ABLE alutacea Physonota alutacea Boh. Physonota Omocerus caerule- opunctata Orexita wagneri (Boh.) Ischnocodia annulis Ischnocodia (Fab.) Dorynota aurita (Boh.) Leaf Beetle Plant T 356 Florida Entomologist 80(3) September, 1997 . COLUMNS

OF

EXPLANATION

FOR

TEXT

EE . S BEETLES

LEAF

ICAN R OSTA Plant Family Stage Locality Date Collectors Score No. Voucher Note C OF

RECORDS

sp. CNV A A1 21/V/1991 RWF 3 5 46-RWF91 Mill. SOLA L, G14 30/VII/1986 DHJ&WH 6 7 86-SRNP-477 65 EEDING ) F Asteraceae esculen- Lycopersicon tum Ipomoea AST A G10 11/VII/1982 DHJ&WH 6 2 82-SRNP-545

(Boh.) ONTINUED (Boh.) 1. (C ABLE Plagiometriona crucipennis Plagiometriona testudinaria Xenocassis ambita (Champ.) Leaf Beetle Plant T Flowers & Janzen: Chrysomelid Feeding Records 357

The alternative hypothesis is that ecological mediation (colonization and host transfer) is the primary explanation for current host associations. In cases of ecologi- cal mediation, phylogenies of herbivores and host plants are not congruent, and host shifts are not necessarily between sister taxa of plants Anderson (1993). In a survey of the Curculioninae (Curculionidae), Anderson (1993) found that in taxa where sys- tematics and plant associations were reasonably well known, evidence for cospecia- tion of plant and insect taxa is lacking, and ecological mediation appeared to be the rule. However, like the Chrysomelidae, the majority of curculionine taxa lack any host plant data. Until host plants are known for a much larger proportion of phytophagous insect taxa, speculations on the evolution of host plant selection by insects will con- tinue to be based on small subsets of the phytophagous insect universe.

ACKNOWLEDGMENTS

We sincerely thank the staffs of the Area de Conservación Guanacaste (ACG), and the Instituto Nacional de Biodiversidad (INBio) for their assistance and many kind- nesses during the course of this study. This research was funded in part by a grant (FLAX 91005) from the CSRS, USDA, to Florida A&M University, a National Science Foundation Mid-Career Fellowship (BSR-9003898) to the senior author, and NSF DEB-9400829 to the junior author.

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MOLDENKE, A. R. 1970. A revision of the Clytrinae of North America north of the Isth- mus of Panama (Coleoptera: Chrysomelidae). Stanford University. MOLDENKE, A. R. 1971. Host-plant relations of phytophagous beetles in . Pan- Pacific Entomol. 47: 105-116. MONRÓS, F. A. 1949. Descripción de las metamorfosis de Lamprosoma chorisiae Mon- rós y consideraciones taxonómicas sobre Lamprosominae (Col. Chrysomel- idae). Acta Zoo. Lill. 7: 449-466. OSTMARK, H. E. 1975. Banana pests in the genus Colaspis including description of a new species (Coleoptera: Chrysomelidae). Florida Entomol. 58: 1-8. REID, C. A. M. 1995. A cladistic analysis of subfamilial relationships in the Chry- somelidae sensu lato (Chrysomeloidea), pp. 557-631 in Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson, J. Pakaluk and S. A. Slipinski,´ ´ eds. Muzeum I Instytut Zoologii PAN, Warszawa. REID, C. A. M. 1996. More on the family Bruchidae. Chrysomela Newsletter 31: 3. REID, W. V., S. A. LAIRD, R. GÀMEZ, A. SITTENFELD, D. H. JANZEN, M. A. GOLLIN, AND C. JUMA. 1993. Biodiversity prospecting: using genetic resources for sustain- able development. World Resources Institute. Baltimore, MD. 341+ix pp. ROCKWOOD, L. L. 1974. Seasonal changes in the susceptibility of Crescentia alata leaves to the flea beetle Oedionychus sp. Ecology 55: 142-148. SCHRODER, W. W., B. PUTTLER, S. S. IZHEVSY, AND D. GANDOLFO. 1994. Viviparity and larval development of Platyphora quadrisignata (Germar) in Brazil. Coleopts Bull. 48: 237-243. STRAUSS, S. Y. 1988. The Chrysomelidae: a useful group for investigation of herbi- vore-herbivore interactions, pp. 91-106 in Biology of Chrysomelidae, P. Jolivet, E. Petitpierre and T. H. Hsiao, eds. Kluwer, the Netherlands. 640 pp. STRONG, D. R. 1977a. Rolled-leaf hispine beetles (Chrysomelidae) and their Zingib- erales host plants in Middle America. Biotropica 9: 156-169. STRONG, D. R. 1977b. Insect species richness: hispine beetles of Heliconia latispatha. Ecology 58: 573-582. VERMA, K. K., AND R. SAXENA. 1996. The status of Bruchidae a a family. Chrysomela Newsletter 32: 3. WILCOX, J. A. 1983. Checklist of the beetles of North and Central America and the West Indies. Vol. 8. The Leaf Beetles and the bean weevils. Family 129. Chry- somelidae. E. J. Brill. New York, 166 pp. WINDSOR, D. M. 1986. Natural history of a subsocial tortoise beetle, Acromis sparsa Boheman (Chrysomelidae, Cassidinae) in Panama. Psyche 94: 127-150. WINDSOR, D. M., E. G. RILEY, AND H. P. STOCKWELL. 1992. An introduction to the bi- ology and systematics of Panamanian tortoise beetles (Coleoptera: Chrysomel- idae: Cassidinae), pp. 372-391 in D. Quintero and A. Aiello, eds. Insects of Panama and Mesoamerica. Oxford Univ. Press, Oxford, 692+xxii pp. 360 Florida Entomologist 80(3) September, 1997

APPENDIX 1—ABBREVIATIONS OF PLANT FAMILY NAMES IN TABLE 1.

ACA Acanthaceae CNV Convolvulaceae MYR Myrsinaceae AMA Amarantaceae DIO Dioscoreaceae ONA Onagraceae APO Apocynaceae ERY Erythroxylaceae PAS Passifloraceae ASC Asclepiadaceae ERI Ericaceae PIP Piperaceae AST Asteraceae EUP Euphorbiaceae POA Poaceae BIG FAB Fabaceae: POL Polygonaceae BOR Boraginacaea Papilionoidea RUB Rubiaceae BUR Burseraceae FLA Flacourtiaceae RUT Rutaceae CAE Fabaceae: HIP Hippocrateaceae SPI Sapindaceae Caesalpinoidea LAU Lauraceae SPO Sapotaceae CAP Capparidaceae LOG Loganiaceae SIM Simarubaceae CLU Clusiaceae MLP Malpighiaceae SOL Solanaceae CEC Cecropiaceae MLV Malvaceae STE Sterculiaceae COC Cochlospermaceae MAR Marantaceae URT Urticaceae COM Combretacaea MEL Melostomataceae VER Verbenaceae CON Connaraceae MIM Fabaceae: VIT Vitaceae Mimosoidea ZIN Zingiberaceae Flowers & Janzen: Chrysomelid Feeding Records 361

APPENDIX 2— LOCALITIES FROM TABLE 1.

G1 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Pitilla, Estacion Pitilla, 8 km S Santa Cecilia, 700 m (N330000, E380400) G2 Guanacaste Prov., Area de Conservacion Guanacaste, Sector El Hacha, Cerro el Hacha, 10 km SE La Cruz, 300 m (N331700, E365400) G3 Guanacaste Prov., Area de Conservacion Guanacaste, Area Recreativa Junquillal, 3 km N Cuajiniquil, 0 m (N328000, E351700) G4 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Orosi, Esta- cion Maritza, 20 km SE La Cruz (N326500, E372200) G5 Guanacaste Prov., Area de Conservacion Guanacaste, Estacion Pocosol, 20 km S La Cruz, 250 m (N319000, E361100) G6 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Orosí, Esta- cion Maritza, sendero Casa Fran, 21 km SE La Cruz, 600 m (N326000, E373300) G7 Guanacaste Prov., Area de Conservacion Guanacaste, Estacion Santa Rosa, 28 km NNW Liberia, 250 m (N313700, E359000) G8 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Bosque Humedo, 30 km NNW Liberia, 300 m (N314800, E360500) G9 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Cafetal, 31 km NNW Liberia, 300 m (N315500, E360200) G10 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Bosque San Emilio, 29 km NNW Liberia, 300 m (N313800, E359800) G11 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Sendero Natural, 28 km NNW Liberia, 250 m (N313100, E359900) G12 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Finca Rosa Maria, 26 km NNW Liberia, 250 m (N311000, E359500) G13 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Casona, 28 km NNW Liberia, 250 m (N313000, E359900) G14 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Area Administrativa, 29 km NNW Liberia, 250 m (N313500, E358900) G15 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Cliff Top Light, 31 km NNW Liberia, 300 m (N315200, E360200) G16 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Casetilla Entrada, 33 km NNW Liberia, 300 m (N317800, E362600) G17 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Laguna Escondida, 30 km NNW Liberia, 250 m (N314500, E357900) G18 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Llano Guacimal, 32 km NNW Liberia, 300 m (N317000, E361600) G19 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Canyon del Tigre, 18 km NW Irigaray, 200 m (N310000, E356800) G20 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Naranjo, Playa Naranjo, 0 m (N307000, E354500) G21 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Naranjo, Sen- dero Real, 10 m (N309000, E354000) G22 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Cruz de Piedra, 33 km NNW Liberia, 300 m (N317200, E360900) G23 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Cacao, EstacionCacao, 9 km N Quebrada Grande, 1000 m (N323100, E375500) 362 Florida Entomologist 80(3) September, 1997

APPENDIX 2—(CONTINUED) LOCALITIES FROM TABLE 1.

G24 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Finca Jenny, 30 km NNW Liberia, 200 m (N316200, E364200) G25 Guanacaste Prov., Area de Conservation Guanacaste, Sector Santa Rosa,Vado Rio Poza Salada, 17 km NW Irigaray, 10 m (N308900, E355700) G26 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Quebrada Guapote, 27 km NNW Liberia, 200 m (N312700, E361700) G27 Guanacaste Prov., Area de Conservacion Guanacaste, Sector Santa Rosa, Quebrada Costa Rica, 250 m (N312200, E357500) G28 Guanacaste Prov., Potrerillos, Rio Tempisque, 23 km NNW Liberia, 100 m (N310900, E367400) G29 Guanacaste Prov., Finca La Pacifica, 5 km NW Canas. G30 Guanacaste Prov., 12 km NW of Bebedero, Hacienda Horizontes. G31 Guanacaste Prov., Bebedero, Ingenio Taboga. A1 Alajuela Prov., Finca San Gabriel, 2 km SW Dos Ríos, 600 m A2 Alajuela Prov., Reserva Forestal San Ramon, 900 m A3 Alajuela Prov., Bijagua, 20 km S Upala, 500 m A4 Alajuela Prov., Canton La Guacima, Río Segundo, 780 m A5 Alajuela Prov., Canton Ciruelas, Río Ciruelas, 800 m H1 Heredia Prov., Estac. Biol. La Selva, 50 m S1 San José Prov., San Pedro, Univ. Costa Rica S2 San José Prov., El Rodeo, 1.5 km S Ciudad Colon C1 Cartago Prov., Pavones, nr. Turrialba C2 Cartago Prov., Madreselva, nr. Empalme C3 Cartago Prov., Carretera Interamericana, 7 km S. Cartago C4 Cartago Prov., Cerro Asunción, paramo vegetation, 3396 m C5 Cartago Prov., Tapanti, Refugio Vida Silvestre P1 Puntarenas Prov., Reserva Forestal Monteverde P2 Puntarenas Prov., Peninsula de Osa, Est. Boscosa, Reserva Forestal Golfo Dulce P3 Puntarenas Prov., Peninsula de Osa, Cerro de Oro Flowers & Janzen: Chrysomelid Feeding Records 363

Appendix 3—Notes to Table 1.

1. These beetles were sitting on heavily eaten leaves, 1.5 m above ground. 2. Additional specimens were observed at time of collection, and C. Chavez reported seeing this species frequently on the same host plant. 3. This species was tested on the host plant. Jolivet (1978) gave Asteraceae, Mimo- saceae, Ericaceae and Fagaceae as other host plant families of this genus. 4. This species was found feeding at shoot tips of its host plant. 5. The host plant is an abundant roadside weed on the entrance road in Sector Santa Rosa and elsewhere in this sector. Beetles have been collected both in the rainy and dry seasons. The larvae make cone-shaped cases, apparently utilizing hairs of the host’s leaves. Moldenke (1971) listed both Malvaceae and Convolvu- laceae as host plant families for this species. 6. The vouchers were collected from a swarm of this species feeding on the low bush in dense dry forest. The intense feeding and mating activity was similar to that observed in other Clytrinae (Flowers et al. 1994, Moldenke 1971). Jolivet (1978) lists Mimosaceae as the predominant host for this genus. 7. The beetle was seen eating bark of new stems. Monrós (1949) and Jolivet (1978) described bark feeding by other members of this genus. 8. This species was very abundant on the leaves of its host at several regenerating pasture sites in 1991. This cosmopolitan genus has been recorded from Arali- aceae from the Palearctic and from Myrtaceae from Puerto Rico (Jolivet 1978). 9. In 1991 this species was very abundant in the pastures and open areas after the onset of the summer rains. Individuals were also collected on other pasture . The collection of Jan Bechyné in Maracay Venezuela contains several specimens of this species collected in and bearing the (apparently) manuscript name "saltator". The only other host record for this genus is Theo- broma cacao L. (Sterculeaceae) for an unidentified species (Jolivet 1987b). 10. Jolivet (1987b) stated that all host observations of the genus Chalcophana have been Asteraceae. 11. Although only one voucher was preserved, numerous adults were observed, and several were tested on the leaves of the plant host. Jolivet (1987b) noted that this genus is both cosmopolitan and polyphagous. 12. Adults were feeding at night on very new expanding leaves of a 1.5 m shoot at base of . Jolivet (1987b) stated that the only reliable feeding records for this genus are from Fabaceae. 13. The only host records in the literature for Percolaspis are from Poaceae and Theo- broma cacao (Jolivet 1987b). 14. This species has been found feeding on several species of Rubiaceae. Adults are agile leapers when disturbed. Jolivet (1987b) gave a single record for this genus: Persea (Lauraceae) for a Cuban Phanaeta. 15. Adults of this genus were found on new foliage and in some years defoliated their hosts. 16. This species was very common feeding on various species of Melastomataceae. Jolivet (1987b) described Typophorus as polyphagous but does not list any Melas- tomataceae among its host plants. 17. The voucher is one of many collected, seen and reared at Estación Pitilla and San Gabriel on various species of Solanum. 18. Larvae skeletonize host plant leaves. This species extensively defoliates its host during some years. Literature records for New World Plagiodera are limited to Salix, Populus (Salicaceae), Croton (Euphorbiaceae), and Lueha (Tiliaceae); how- 364 Florida Entomologist 80(3) September, 1997

ever, species in the Philippines and India have been reported on Xylosoma and Flacourtia (Flacourtiaceae) (Jolivet & Hawkeswood 1995). 19. This is the most commonly collected of the Costa Rican species of Platyphora. During one feeding test, two very small larvae were observed in the plastic bag which up till then held a single female, suggesting that Platyphora bicolor is vi- viparous. Schroder et al. (1994) described the biology of the viviparous Platy- phora quadrisignata (Germar) from southern Brazil. 20. Adults and larvae were frequently found feeding on host plant throughout the 1991 rainy season. Apparently, our observations represent the only known host plant data for Stilodes. 21. A group was followed from egg to adult. Larvae feed and rest on underside of leaves. Pupation takes place in leaf litter. 22. Larvae are sooty black, covered with branched hair-like projections, and with red heads. Pupae are yellow. This chrysomelid was parasitized by Myopharous (Ta- chinidae: Diptera). 23. In 1991 this beetle caused a major defoliation of its host plant, a pioneer species in cleared pastures. 24. The host plant of this galerucine was found growing along the edge of a small patch of forest. 25. The host plant was a low understory tree in tropical dry forest. 26. This galerucine was seen on several occasions feeding on young leaves of its host plant. This genus has been recorded from Acacia (Fabaceae) in the USA (Jolivet 1987a). 27. A large group of these Masurius (which may represent more than one species) was found feeding on the two host plants growing within a few yards or each other along a trail in montane forest. 28. This and the following species were reared to adult. 29. In addition to the voucher specimens, other specimens were collected two years earlier on the same host plant. 30. RWF has observed adults of this species every year since 1989 defoliating basal shoots of a tree growing in front of the main administration building at the Uni- versity of Costa Rica. Jolivet (1987a) listed Cordia and Lantana (Verbenaceae) as hosts of this genus. 31. RWF observed on individual at night eating a hole in the middle of a leaf of the Ipomoea host plant. 32. In both cases, beetles were observed feeding on the host plant. Jolivet (1991) listed Labiaceae and Verbenaceae as probable hosts for this genus and noted other citations of Lauraceae, Buddlejaceae, Asteraceae, Umbelliferae, Sterculi- aceae, and Fabaceae. 33. In addition to the vouchered specimen from Byrsonima crassifolia, this species was abundant on this host plant at Estacion Maritza (G4) in 1991. 34. Unlike many other chrysomelids which were found associated only with young foliage, A. salvadorense was found actively feeding late in the rainy season on older leaves. 35. A large group of these beetles was found on a broken stalk of the host plant, feed- ing on sap and milky latex. The host plant was growing in the shaded understory of montane forest. 36. Adults were reared from larvae feeding on the host plant. 37. Jolivet (1991) listed Samanea (Fabaceae/pap.) as a host of this genus. 38. Jolivet (1991) cited Theobroma and Tecoma (Bignoniaceae) as other known host plants of this alticine genus. Flowers & Janzen: Chrysomelid Feeding Records 365

39. Both this and the following host plant were growing close together in a mixed stand next to a road. 40. The host plant, growing in a wet depression in a cleared area, sustained heavy feeding damage from this alticine in 1991. Jolivet (1991) listed Cleome, Solanum, Beta (Chenopodiaceae), Labiaceae, Cordia, and Adiantum (Adiantaceae) as host plants of Leptophysa. 41. These beetles were swept from a tree that showed heavy feeding damage to the leaves. No active feeding was observed, but this collection was made during an abnormal dry spell during what was supposed to be the wet season. 42. This Longitarsus is a flightless species. 43. Field observations by DHJ indicate that the adult appears on the host plant to oviposit; larvae are free living and cut islands out of leaf margin. 44. The host plant is a small prostrate weed. The beetles were first observed resting and defecating on a shrub of Luehea (Tiliaceae) which grew over the Evolvulus. When no feeding damage on the Luehea was seen, despite the beetle activity, a wider search revealed the true host plant. 45. These small pinkish-orange flea beetles were observed feeding on newly expand- ing leaves (which are also reddish to pinkish orange) of their ericaceous hosts. 46. This alticine was collected abundantly from a very dense stand of its host plant. In 1994 it was found equally abundantly in the same stand of plants. 47. RWF has observed this species over several years, actively feeding on Euphorbi- aceae even during the dry season in quite arid habitats. 48. These represent five different morphospecies of Syphrea collected on various plants. 49. This species feeds by scraping pits in the expanding leaves of this host plant. The following plant record may be an alternate dry season food source. 50. Huge numbers of this species were found defoliating the host plant during the voucher year. In 1991, on the other hand, no specimens were found and no dam- age to the host was apparent. This is the species called Oedionychis sp. in Rock- wood (1974). Bechyné (1955) restricted the definition of true Oedionychis to a small group of flightless Mediterranean flea beetles. New World species formerly in Oedionychis are now placed in Walterianella, Alagoasa and other genera. 51. Jolivet (1991) listed Venezuelan records of Gardinia (Rubiaceae) and Tabebuia (Bignoniaceae) for this genus. 52. The genus Cephaloleia is well known from various species of Heliconia and other Zingiberales (Strong 1977a,b). This species was regularly encountered in rolled- up terminal leaves of Costus at this and other localities. 53. This hispine was very abundant in a dense stand of grass growing on a river sand bar. 54. A large number of these hispines were feeding on and heavily damaging leaves of a shrub of its host growing along the bank of a river in deep shade. 55. These cassids have black larvae with long black caudal brushes; the pupae have a creamy white thorax. Adults were reared. 56. Windsor et al. (1992) gave Ipomoea lindenii Mart. & Gal. as host plant for true C. egregia. 57. This species periodically defoliates its host. 58. Feeding on young leaves of Alibertia was seen; some feeding damage was also seen on the two bignoniaceous plants as well. 59. The cassid caused a major defoliation in 1979, but has been rare since. The 1991 record was from a single tree growing by the seashore and heavily damaged by a group of the cassids. Jolivet (1988a) also listed Tabebuia and other Bignoniaceae as hosts for this genus. 366 Florida Entomologist 80(3) September, 1997

60. These records are of beetles aestivating in the dry season; see Flowers (1991) for more details on this behavior. Windsor et al. (1992) listed several species of Cor- dia as the true host plants of this species. 61. The host plant was an understory plant in a pine plantation. Jolivet (1988a) also listed Hyptis (Labiaceae) as a host plant for this genus. 62. The record from Bursera simaruba is for beetles hiding under bark plates during the dry season. Jolivet (1988a) listed Phaseolus (Fabaceae) and Passiflora (Pas- sifloraceae) for this genus. 63. In 1991 this species was common during the rainy season. A colony at the Ad- ministration Area in Sector Santa Rosa (G14) was followed for several months, during which time predatory pentatomids were observed resting on foliage above the cassids, and occasionally descending to feed on them. 64. Windsor et al. (1992) listed Cordia spinescens for a P. nr. alutacea from Panama. 65. Windsor et al. (1992) also list Solanum seaforthianum Andr. and Physalis cor- data Mill (Solanaceae).

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