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334

Florida Entomologist

80(3) September, 1997

FEEDING RECORDS OF COSTA RICAN LEAF BEETLES (COLEOPTERA: CHRYSOMELIDAE)

R. W

ILLS

F

LOWERS

1

AND

D

ANIEL

H. J

ANZEN

2

1

Agricultural Research Programs, Florida A&M UniversityTallahassee, FL 32307-4100, [email protected]

2

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

A

BSTRACT

Host plant associations are given for 137 species representing 7 subfamilies and92 genera of Costa Rican Chrysomelidae. A numeric score is introduced to objectivelydescribe confidence that a field observation of an interaction between a chrysomelidand a plant represents true herbivory. Literature host plant records, if they exist, aregiven for included chrysomelid taxa.

Key Words: herbivory, Criocerinae, Chrysomelinae, Cryptocephalinae, Eumolpinae,Galerucinae, Hispinae, Lamprosominae, host plants

R

ESUMEN

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 queuna 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 insects. The interest is driven by the related agendas of studying themfor their intrinsic scientific interest, and protecting tropical biodiversity through find-ing practical and non-destructive ways to use it. The latter agenda is exemplified bythe biochemical prospecting programs recently started in several areas of the world(Reid et al. 1993).

Most plant-insect research begins with a basic event: an observation that a specificplant 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 zoneinsects and on a few relatively well known tropical groups (e.g., Lepidoptera). Only asmall percentage of the fauna of tropical herbivores has been similarly studied.

The family Chrysomelidae (Coleoptera), or leaf beetles, is a natural subject forstudying 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, areherbivores or seed predators. However, for about 70% of the described species, we donot 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 plantsof the tortoise beetles (Cassidinae) of Panama was recently published by Windsor etal. (1992); Moldenke (1971) listed host plants for some Mexican Chrysomelidae, andAnaya (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) summarizedcurrent host plant data on a world level for the Chrysomelidae. However, in much ofthis literature, beetle species are usually identified only to genus and their plant hostsonly to family. A few field studies have documented significant attacks by chry-somelids on plants in Central American ecosystems (e.g., Rockwood 1974, Memmottet al. 1993), and some detailed field and laboratory studies have been undertaken forseveral Neotropical species (Bach 1986, Begossi & Benson 1988, Buzzi & Winder1986, Hsiao 1988, Strong 1977a,b). Apart from these ecological studies of specificchrysomelids, many of the published host plant records are of dubious value, statingmerely that beetle X was taken on plant Y (or, all too often, “genus X feeds on plantgenus Y”). A further problem, also noted by Furth (1985), is that a large proportion ofsuch records are buried in taxonomic monographs and regional studies (e.g., Bechyné& Bechyné 1975) and accessible only by reading these studies in their entirety. Muchmore 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 leafbeetle-plant interactions can be made.

In this paper, we present feeding records of adults and larvae for 137 species ofCosta Rican Chrysomelidae, representing 7 subfamilies and 92 genera. The majorityof these observations were made by the senior author during a six-month sabbaticalat Costa Rica’s Instituto Nacional de Biodiversidad (INBio) in 1991, and by the juniorauthor during the years 1978 to 1995 as a byproduct of an on-going intensive study ofthe 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 haveomitted many records where a single beetle was seen or collected on a plant, except fora 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, Galerucini), Charles L. Staines (MarylandDepartment 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



Our data are organized into a table with three supplementary appendices. Table 1lists observations by chrysomelid taxon, gives field data in summary form, and listsvoucher specimens. Appendix 1 is a key to plant family name abbreviations. Appendix2 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 earlieropinions (eg. Crowson 1955, Lawrence 1982) that Bruchidae, or seed weevils, are asubfamily 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’sclassification, 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 1Leaf 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, anddistinct 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, theterm “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 familynames. 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 orheld for testing.


337

Collectors

DHJ&WH: Daniel H. Janzen & Winnie HallwachsRWF: R. W. FlowersNames 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 tothe beetles was not present.

2 Two to four chrysomelids were collected from a plant and feeding damage thatcould reasonably be attributed to the beetles was present,

or

five to nine chry-somelids were collected from a plant but obvious feeding damage attributable tothe beetles was not present.

1 Two to four chrysomelids were collected from a plant but no noticeable feedingdamage was observed.

Number (No.)

Number of vouchered specimens. In general, one feeding record equals onevoucher; 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 authorhave 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. Thefirst letter of each locality code corresponds to the first letter of its province. Localitiesin 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



D

ISCUSSION

The data presented in these tables represent only the beginnings of the task ofworking out host plant relationships for the Central American Chrysomelidae. Ourdata cover less than 7% of the estimated 2000 chrysomelid species present in CostaRica alone (Flowers, unpublished data). In some cases, our data confirmed previouslypublished relationships between chrysomelid genera and host plant families (summa-rized in Jolivet & Hawkeswood 1995); 30 of our records represent host plant familyrange extensions, and 19 records are for chrysomelid genera in which, apparently, nohost 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 makingthese distinctions have been demonstrated to us on several occasions when we foundchrysomelid 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 observedsitting on leaves of a

Luehea

sapling (Tiliaceae). Although large numbers of beetleswere on the

Luehea

, and their frass was also evident on these leaves, closer inspectionrevealed that no feeding was taking place on the

Luehea

and that the true food plant(

Evolvulus nummularis

; Convolvulaceae) was growing beneath the shrub. Similarwarnings about possible confusion of Alticini food plants due to the beetle’s mobilityhave been given by Hawkeswood and Furth (1994). Nevertheless, collection recordscan still provide useful information—for many taxa opportunistic collecting has pro-vided the only information we have on possible host plants—if their limitations areclearly acknowledged. For our data we have included a “reliability scale” to roughlymeasure our confidence that a given association represents a true chrysomelid-hostplant relationship. While ecological studies of narrow groups of chrysomelids orplants will always provide the most unambiguous data on feeding requirements, re-cent emphasis on and support for inventory collecting can rapidly increase knowledgeof the feeding habits of a broad range of chrysomelids, if observations are qualified insome 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 theyobserve. 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 currentlyfocused on plants. When a family of plants is being surveyed for active chemicals, theinsects feeding on those plants represent another level of chemical derivatives avail-able for screening. The phytophagous insect may produce novel chemical varietieswhich 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 arisefrom cospeciation or parallel descent. This theory has become a popular explanationof host plant selection, under the name “coevolution” (though we caution the readerthat 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 Lamiales (Farrell and Mitter 1990). However, theirstudy represents one of the few documented examples of coevolution (Anderson 1993).


339

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1R

WF

35

15-R

WF

9114

Lad

enbe

rgia

se

rico

phyl

la S

tan

dley

RU

BA

G1

15/V

/199

1R

WF

24

8-R

WF

91

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote

Flowers & Janzen: Chrysomelid Feeding Records 343

Sab

acea

vil

losa

Roe

m.

& S

chu

lt.

RU

BA

H1

22/I

/198

9R

WF

411

21-R

WF

94

Ph

anae

ta s

p. 1

Ron

del

etia

bu

dd

le-

oid

es B

enth

.R

UB

AG

48/

V/1

991

RW

F3

517

-RW

F91

Ph

anae

ta s

p. 2

Som

mer

a d

onn

ell-

smit

hii

Sta

ndl

.R

UB

AC

15/

IX/1

991

RW

F, D

. Cot

o,

J. S

aun

ders

13

43-R

WF

91

Rh

abd

opte

rus

sp. 1

Oco

tea

vera

guen

sis

(Mei

ssn

.) M

ez.

LA

UA

G8

2/V

II/1

982

DH

J&W

H6

1082

-SR

NP

-426

15

Rh

abd

opte

rus

sp 2

&

3M

anil

kara

ch

icle

(L

.)

van

Roy

anS

PO

AG

1031

/VII

/198

3D

HJ&

WH

64

83-S

RN

P-9

08

Typ

oph

oru

s va

ri-

abil

is J

ac.

un

iden

t. s

p.M

EL

AC

25/

II/1

994

RW

F, M

. Ch

a-va

rría

, H.

Wei

ler

413

3-R

WF

9416

Typ

oph

oru

s sp

. 1C

onos

tygi

a xa

lape

nsi

s (B

onpl

.) D

.Don

ME

LA

G1

12/V

/199

1R

WF

24

5-R

WF

91

Typ

oph

oru

s sp

. 3Ip

omoe

a pe

s-ca

pra

(L.)

R

. Br.

CN

VA

G20

12/V

/199

1R

WF,

R. T

iffe

r2

365

-RW

F91

CH

RY

SO

ME

LIN

AE

Cal

ligr

aph

a ar

gus

Stå

lG

uaz

um

a u

lmif

olia

L

am.

ST

EA

G10

25/V

/199

1R

WF

62

113-

RW

F91

Gu

azu

ma

ulm

ifol

iaS

TE

L, A

G14

16/V

/197

927

/VI/

1980

DH

J&W

H6 6

1 179

-SR

NP

-14

80-S

RN

P-2

51

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote

344 Florida Entomologist 80(3) September, 1997

Byt

tner

ia a

cule

ata

(Jac

q.)

Jacq

.S

TE

AG

1130

/V/1

981

DH

J&W

H6

281

-SR

NP

-50

Cal

ligr

aph

a fu

lvi-

pes

Stå

lS

ida

rhom

bifo

lia

L.

MLV

AS

224

/VI/

1991

RW

F, J

. Cor

-ra

les,

A. S

olís

68

89-R

WF

91

Cal

ligr

aph

a se

rpen

-ti

na

(Rog

ers)

Aye

nia

mic

ran

tha

Sta

ndl

.S

TE

L, A

G9

2/V

II/1

983

DH

J&W

H6

683

-SR

NP

-691

Lep

tin

otar

sa u

nd

ec-

imli

nea

ta S

tål

Sol

anu

m o

chra

ceof

er-

rugi

neu

m (D

un

.) F

ern

.S

OL

L, A

G1

16/V

/199

1R

WF

61

16-R

WF

9417

Pla

giod

era

cere

a at

rita

rsis

Stå

lP

rock

ia c

ruci

s L

.F

LA

P, A

G10

6/V

I/19

8014

/VI/

198

DH

J&W

H6 6

2 780

-SR

NP

-69

82-S

RN

P-2

2518

Pro

ckia

cru

cis

FL

AL

, AG

92/

VII

/198

3D

HJ&

WH

61

83-S

RN

P-6

98P

rock

ia c

ruci

sF

LA

L, A

G26

20/V

I/19

80D

HJ&

WH

61

80-S

RN

P-1

65P

lagi

oder

a n

r. u

ni-

form

is J

ac.

Xyl

osom

a fl

exu

osu

m

(HB

K)

Hem

sley

FL

AL

, AG

1515

/VI/

1983

DH

J&W

H6

983

-SR

NP

-428

Pla

giod

era

sp.

Xyl

osom

a h

orri

da

Ros

eF

LA

L, A

G15

30/V

/199

1D

HJ&

WH

612

91-S

RN

P-5

38

Pla

typh

ora

bico

lor

Jac.

Koa

nop

hyl

lon

pit

tier

i (K

latt

) R

. M. &

H.

Rob

inso

n

AS

TA

G23

16/V

III/

1991

RW

F6

541

-RW

F91

19

Pla

typh

ora

petu

-la

ns

Stå

lM

esec

hit

es t

rifi

da

(Jac

q.)

Mü

ll. A

rg.

AP

OA

G8

24/V

I/19

83D

HJ&

WH

61

83-S

RN

P-5

64

Pre

ston

ia a

llen

ii

Woo

dson

AP

OA

G8

9/V

II/1

982

3/X

II/1

983

13/X

II/1

983

DH

J&W

H6

1 1 1

82-S

RN

P-5

3383

-SR

NP

-139

883

-SR

NP

-145

3

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Pre

ston

ia a

llen

iiA

PO

AG

107/

VII

/198

9D

HJ&

WH

12

89-S

RN

P-5

60S

tilo

des

mod

esta

Ja

c.B

anis

teri

opsi

s m

uri

-ca

ta (

Cav

.) C

uat

r.M

LP

L, A

G7

20/V

I/19

78D

HJ&

WH

63

78-S

RN

P-8

320

Ban

iste

riop

sis

mu

ri-

cata

ML

PL

, AG

142/

VII

/198

0D

HJ&

WH

62

80-S

RN

P-2

92

Ban

iste

riop

sis

mu

ri-

cata

ML

PL

, AG

412

/V/1

991

RW

F6

531

-RW

F91

Sti

lod

es n

epti

s S

tål

Hir

aea

recl

inat

a Ja

cq.

ML

PA

G8

14/V

I/19

84D

HJ&

WH

62

84-S

RN

P-5

57H

irae

a re

clin

ata

ML

PA

G27

4/V

II/1

983

DH

J&W

H6

383

-SR

NP

-732

Hir

aea

recl

inat

aM

LP

L, A

G11

16/V

I/19

89D

HJ&

WH

618

89-S

RN

P-2

1321

Hir

aea

recl

inat

aM

LP

AG

1716

/VI/

1991

RW

F6

111

5-R

WF

9122

GA

LE

RU

CIN

AE

: s.s

.

Bib

lite

a ja

nso

ni

(Jac

.)W

ith

erin

gia

sp.

SO

LA

G23

20/I

II/1

990

RW

F3

1017

-RW

F94

Car

agu

ata

pall

ida

(Jac

.)V

ism

ia b

acci

fera

(L

.)

Tr.

& P

l.C

LU

L, A

G1

17/V

I/19

91R

WF

619

40-R

WF

9123

Coe

lom

era

sp.

Cec

ropi

a pe

ltat

a L

.C

EC

AG

1031

/VII

/198

3D

HJ&

WH

62

83-S

RN

P-9

03Is

otes

sp.

Ires

ine

dif

fusa

Ku

nth

.A

MA

AA

123

/V/1

991

RW

F4

919

-RW

F91

24L

upe

roso

ma

vitt

atu

m J

ac.

Lon

choc

arpu

s ac

um

inat

us

(Sch

ech

t.)

Sou

sa

FA

BA

G19

12/V

I/19

91R

WF

36

97-R

WF

9125

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Mal

acor

hin

us

dec

empu

nct

atu

s Ja

c.

Pit

hec

ello

biu

m

palm

anu

m S

tan

dl.

MIM

AG

49/

V/1

991

RW

F6

21-

RW

F94

26

Pit

hic

ello

biu

m l

ongi

-fo

liu

mM

IMA

A4

18/I

I/19

94R

WF

62

4-R

WF

94

Mas

uri

us

spp.

Cro

ton

sp.

EU

PA

G23

15/V

III/

1991

RW

F4

1161

-RW

F91

27S

olan

um

ace

rosu

m

Sen

dt.

SO

LA

G23

16/V

III/

1991

RW

F4

1470

-RW

F91

Mon

olep

ta s

p.R

oure

a gl

abra

Ku

nth

.C

ON

AG

89/

VII

/198

2D

HJ&

WH

615

82-S

RN

P-5

12N

esti

nu

s vi

rid

is

Jac.

Zan

thox

ylu

m s

etu

lo-

sum

P. W

ilso

nR

UT

L L, A A

G14

13/V

I/19

8214

/VII

/198

319

/XII

/198

3

DH

J&W

H6

1 1 2

82-S

RN

P-2

2183

-SR

NP

-802

83-S

RN

P-1

479

28

Nes

tin

us

n. s

p.E

ssen

beck

ia l

itto

rali

s D

onn

. Sm

ith

RU

TL

, AG

1618

/VI/

1983

DH

J&W

H6

183

-SR

NP

-454

Par

anap

iaca

ba r

u-

fofa

scia

ta (

Jac.

)C

outa

rea

hex

and

ra

(Jac

q.)

RU

BA

G2

20/V

III/

1991

RW

F4

254

-RW

F91

29

Yin

gare

sca

sp.

Cor

dia

eri

osti

gma

Pit

tier

BO

RA

S1

14/I

I/19

90R

WF

625

18-R

WF

9430

GA

LE

RU

CIN

AE

: ALT

ICIN

I

Ala

goas

a se

riat

a (J

ac.)

Lan

tan

a ca

mar

a L

.V

ER

AG

1428

/VI/

1995

DH

J&W

H6

193

-SR

NP

-299

5

Asp

hae

ra

nob

ilit

ata

(Fab

r.)

Ipom

oea

sp.

CN

VA

P3

10/V

/199

5R

WF

61

7-R

WF

9531

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Asp

hae

ra r

eich

ei

Har

.V

ern

onia

pat

ens

H.B

.K.

AS

TA

G1

16/V

/199

1R

WF

62

4-R

WF

9132

Aya

laia

min

or

Bec

h. &

Bec

h.

Byr

son

ima

cras

sifo

lia

ML

PA

G17

19/V

/199

1R

WF

61

76-R

WF

9133

Ban

iste

riop

sis

sp.

ML

PA

G2

28/V

/199

1R

WF,

R. E

li-

zon

do, L

. Ros

e1

488

-RW

F91

Aya

laia

sal

vad

or-

ense

Bec

h. &

Bec

h.

Ery

thro

xylu

m h

avan

-en

se J

acq.

ER

YA

G19

12/V

/199

1R

WF

614

68-R

WF

9134

Ery

thro

xylu

m h

avan

-en

seE

RY

AG

1112

/VII

I/19

91R

WF

66

84-R

WF

91

Ery

thro

xylu

m h

avan

-en

seE

RY

AG

923

/VI/

1992

DH

J&W

H6

892

-SR

NP

-242

4

Cen

tral

aph

thon

a n

r. le

ssm

ann

i B

ech

.&B

ech

.

Eu

phor

bia

elat

a B

ran

dE

UP

AG

2314

/VII

I/19

91R

WF,

C.

Ch

avez

628

72-R

WF

9135

Eu

phor

bia

elat

aE

UP

AG

234/

V/1

995

RW

F, E

. Ula

te6

198-

RW

F95

Ch

aeto

cnem

a sp

.P

avon

ia s

p.M

LVA

G1

17/V

/199

1R

WF

25

45-R

WF

91D

iph

alti

ca s

p. 1

Ces

tru

m r

acem

onsu

m

R.&

P.S

OL

AA

121

/V/1

991

RW

F3

437

-RW

F91

Dip

hal

tica

sp.

2S

olan

um

cf.

arbo

reu

m

Hu

mb.

& B

onpl

. ex

Du

val

SO

LA

C3

5/II

/199

4R

WF,

M. C

ha-

varr

ía, H

. W

eile

r

45

5-R

WF

94

Dip

hau

laca

au

lica

(O

l.)C

ours

etia

ell

ipti

ca M

. S

ousa

& R

udd

FA

BA

G11

12/V

III/

1991

RW

F4

1785

-RW

F91

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Des

mod

ium

sp

FA

BA

G2

24/V

III/

1991

RW

F3

559

-RW

F91

Dis

onyc

ha

quin

que-

lin

eata

(L

at.)

Pas

sifl

ora

pulc

hel

la

Ku

nth

PA

SA

G9

27/V

II/1

992

DH

J&W

H6

892

-SR

NP

-402

6

Dis

onyc

ha

trif

asci

-at

a C

lark

Byt

tner

ia a

cule

ata

ST

EL

G9

21/X

I/19

87D

HJ&

WH

62

87-S

RN

P-1

343

36

Byt

tner

ia a

cule

ata

ST

EA

G11

11/V

III/

1995

DH

J&W

H6

195

-SR

NP

-784

8B

yttn

eria

acu

leat

aS

TE

AH

111

/I/1

995

RW

F6

121-

RW

F95

Epi

trix

sp.

1S

olan

um

cf.

torv

um

S

w.

SO

LA

A2

12/I

II/1

990

RW

F3

2611

-RW

F94

Epi

trix

sp.

2Q

uas

sia

amar

a L

.S

IMA

G2

6/V

/199

1R

WF,

T. d

e la

R

osa

28

18-R

WF

91

Gen

aph

thon

a tr

ans-

vers

icol

lis

(Jac

.)In

ga s

apin

dio

ides

W

illd

.M

IMA

A2

12/I

II/1

990

RW

F2

713

-RW

F94

Gio

ia s

p.P

alic

oure

a sa

lici

foli

a S

tan

dl.

RU

BA

C2

5/II

/199

4R

WF,

M. C

ha-

varr

ía, H

. W

eile

r

39

4 R

WF

94

Gle

nid

ion

nr.

haa

gi

Har

.In

ga s

apin

dio

ides

MIM

AA

212

/III

/199

0R

WF

12

12-R

WF

9437

Hei

kert

inge

rell

a sp

. 1

Bu

dd

leja

nit

ida

Ben

th.

LO

GA

C4

6/V

III/

1991

RW

F, L

. M. &

T.

A. K

etch

em4

2594

-RW

F91

Hei

kert

inge

rell

a sp

. 2

Lan

tan

a ca

mar

aV

ER

AP

215

/IX

/199

1R

WF,

R. A

gui-

lar

27

109-

RW

F91

38

Hyd

mos

yne

sp.

Bru

gman

sia

can

did

a P

ers.

SO

LA

A1

14/V

I/19

91R

WF

36

56-R

WF

9139

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Lyc

ian

thes

mu

ltifl

ora

Bit

t.S

OL

AA

123

/V/1

991

RW

F3

638

-RW

F91

Wit

her

ingi

a sp

.S

OL

AP

314

/V/1

995

RW

F2

15-

RW

F95

Hyp

olam

psis

sp.

1C

appa

ris

fron

dos

a Ja

c.C

AP

AG

112

/V/1

991

RW

F1

228

-RW

F91

Lep

toph

ysa

nr.

bor-

don

i B

ech

. & B

ech

.C

leom

e pa

rvifl

ora

Ku

nth

CA

PA

G1

13/V

/199

1R

WF

425

95-R

WF

9140

Lep

toph

ysa

nr.

lit-

tora

lis

Bec

h. &

B

ech

.

Cap

pari

s od

orat

is-

sim

a Ja

cq.

CA

PA

G21

23/V

III/

1994

RW

F2

1126

-RW

F94

41

Lon

gita

rsu

s sp

. 1T

ourn

efor

tia

glab

ra L

.B

OR

AG

409

/V/1

991

RW

F2

611

-RW

F91

Lon

gita

rsu

s sp

. 2A

gera

tin

a sp

.A

ST

AC

46/

VII

I/19

91R

WF,

L. M

. &

T. A

. Ket

chem

26

92-R

WF

9142

Sen

ecio

an

dic

ola

Tu

rcz.

AS

TA

C4

6/V

III/

1991

RW

F, L

. M. &

T.

A. K

etch

em3

1293

-RW

F91

Lu

prae

a vi

olac

ea

Jac.

Urc

ra c

arac

asan

a (J

acq.

) G

rise

b.U

RT

AP

117

/II/

1990

RW

F4

1514

-RW

F94

Lu

prae

a n

r. vi

ola-

cea

Cli

bad

ium

sp.

AS

TA

A2

25/I

II/1

996

RW

F6

61-

RW

F96

Lu

prae

a sp

.H

yper

icu

m i

razu

ense

K

un

tze

CL

UA

C4

6/V

III/

1991

RW

F, L

. M. &

. T.

A. K

etch

em2

791

-RW

F91

Lys

ath

ia s

p.L

ud

wig

ia e

rect

a (

L.)

H

ara

ON

AA

G28

19/V

III/

1991

RW

F4

1552

-RW

F91

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Lu

dw

igia

ere

cta

ON

AA

P2

15/I

X/1

991

RW

F, R

. A

guil

ar4

1711

0-R

WF

91

Mar

gari

dis

a sp

. 1C

onos

tegi

a xa

lape

nsi

sM

EL

AG

112

/V/1

991

RW

F3

255-

RW

F91

Mar

gari

dis

a sp

. 2M

icon

ia s

chli

mii

Tri

-an

aM

EL

AP

214

/IX

/199

1R

WF

24

107-

RW

F91

Mar

gari

dis

a sp

. 4W

ith

erin

gia

sp.

SO

LA

G1

14/V

/199

1R

WF

24

3-R

WF

91M

egis

tops

nr.

cost

a-ri

cen

sis

Bla

keT

abeb

uia

och

race

a (C

ham

.) S

tan

dl.

BIG

AG

1421

/VII

I/19

95D

HJ&

WH

63

95-S

RN

P-8

293

Tab

ebu

ia o

chra

cea

BIG

AG

1022

/VII

I/19

95D

HJ&

WH

65

95-S

RN

P-8

473

Mon

omac

ra v

iola

-ce

a (J

ac.)

Pas

sifl

ora

bifl

ora

Lam

.P

AS

AP

215

/IX

/199

1R

WF,

R. A

gui-

lar

12

108-

RW

F91

Nas

igon

a pa

llid

a Ja

c.G

onza

lagu

nia

ros

eaR

UB

AP

314

/V/1

995

RW

F4

24-

RW

F95

Not

ozon

a n

icar

a-gu

ensi

s Ja

c.B

urs

era

sim

aru

ba (L

.)

Sar

g.B

UR

AG

1618

/VI/

1983

DH

J&W

H6

183

-SR

NP

-455

43

Bu

rser

a si

mar

uba

BU

RL

, AG

1315

/VI/

1985

DH

J&W

H6

385

-SR

NP

-401

Om

oph

oeta

sim

u-

lan

s Ja

c.E

volv

ulu

s n

um

mu

-la

ris

(L.)

L.

CN

VA

G14

21/V

II/1

994

RW

F6

527

-RW

F94

44

Par

alac

tica

sp.

Pas

sifl

ora

sp.

PA

SA

G23

14/V

III/

1991

RW

F3

160

-RW

F91

Par

asyp

hra

ea s

p. 1

Inga

sp.

MIM

AG

117

/V/1

991

RW

F2

244

-RW

F91

Par

asyp

hra

ea s

p. 2

Cav

end

ish

ia b

ract

eata

(R

uis

& P

av. e

x J.

St.

-H

il.)

Hoe

rold

ER

IA

C3

5/II

/199

4R

WF,

M.

Ch

avar

ría,

H.

Wei

ler

410

6-R

WF

9445

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Cav

end

ish

ia s

p.E

RI

AC

55/

II/1

994

RW

F, M

. C

hav

arrí

a, H

. W

eile

r

614

2-R

WF

94

Ph

enri

ca n

r. au

stri

-ac

a (S

chau

f.)B

yttn

eria

acu

leat

aS

TE

AG

1111

/VII

I/19

95D

HJ&

WH

62

95-S

RN

P-7

846

Pla

typr

osop

us

pal-

len

s (F

ab.)

Can

aval

ia b

rasi

len

sis

Mar

t. e

x B

ren

t.F

AB

AG

1318

/VI/

1991

RW

F4

739

-RW

F91

Ple

ctot

etra

nr.

clar

ki

Bal

yL

ippi

a to

rres

ii S

tan

dl.

VE

RA

A1

18/V

III/

1991

RW

F2

955

-RW

F91

Pto

cad

ica

nr.

stra

-m

inea

Har

.P

assi

flor

a sp

.P

AS

AG

2314

/VII

I/19

91R

WF

33

60-R

WF

91

Res

iste

nci

ana

ob-

scu

ra (

Jac.

)E

xost

ema

cari

baeu

m

(Jac

q.)

Roe

m &

Sch

ult

RU

BA

G2

24/V

III/

1991

RW

F1

253

-RW

F91

Res

iste

nci

ana

pan

a-m

ensi

s (J

ac.)

Xyl

osom

a sp

.F

LA

AG

2316

/VII

I/19

91R

WF

25

114-

RW

F91

Xyl

osom

a sp

.F

LA

AG

234/

V/1

995

RW

F, E

. Ula

te6

49-

RW

F95

Str

abal

a ac

um

inat

a co

star

icen

sis

Bla

keS

perm

acoc

e sp

.R

UB

AA

326

/VII

/199

4R

WF

51

24-R

WF

94

Syp

hre

a bi

bian

a B

ech

.M

imos

a pi

gra

L.

MIM

AG

1725

/V/1

991

RW

F4

3364

-RW

F91

46

Mim

osa

pigr

aM

IMA

G17

22/V

II/1

994

RW

F4

5223

-RW

F94

Syp

hre

a pa

rvu

la

Jac.

Ber

nar

dia

nic

ara-

guen

sis

Sta

ndl

.E

UP

AG

1112

/VII

I/19

91R

WF

62

83-R

WF

9147

Dal

ech

ampi

a sp

.E

UP

AG

821

/V/1

991

RW

F4

786

-RW

F91

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Syp

hre

a sp

p.P

avon

ia s

p.M

LVA

G1

17/V

/199

1R

WF

12

45-R

WF

9148

Dal

ech

ampi

a h

eter

o-m

orph

a P

ax &

K.

Hof

fm.

EU

PA

G1

18/V

/199

1R

WF

26

25-R

WF

91

Cap

eron

ia p

alu

stri

s (L

.) A

. St.

-Hil

.E

UP

AG

3130

/I/1

994

RW

F3

161-

RW

F 9

4

Cap

eron

ia p

alu

stri

sE

UP

AG

2013

/VII

I/19

91R

WF,

R. T

iffe

r4

749

-RW

F91

Aca

lyph

a ap

odan

thes

S

tan

dl. &

L. O

.Wil

l-ia

ms

EU

PA

G4

10/V

/199

1R

WF

48

7-R

WF

91

Byt

tner

ia a

cule

ata

ST

EA

H1

11/I

/95

RW

F6

141-

RW

F95

Sys

ten

a su

lph

ure

a Ja

c.S

erja

nia

sch

ied

ean

a S

chlt

dl.

SP

IA

G11

5/V

II/1

980

DH

J&W

H6

180

-SR

NP

-340

49

Cas

sia

bifl

ora

CA

EA

G17

29/I

/198

9R

WF

12

10-R

WF

94W

alte

rian

ella

hu

-m

eral

is (

Fab.

)L

ind

enia

riv

alis

B

enth

.R

UB

AG

224

/VII

I/19

91R

WF,

R. E

li-

zon

do, L

. Ros

e2

487

-RW

F91

Wal

teri

anel

la t

enu

i-ci

nta

(Ja

c.)

un

iden

t sp

.A

CA

AG

220

/VII

I/19

91R

WF

24

51-R

WF

91

Wal

teri

anel

la v

e-n

ust

ula

(S

chau

f.)C

resc

enti

a al

ata

H.B

.K.

BIG

AG

1811

/VII

I/19

89D

HJ&

WH

66

89-S

RN

P-8

6350

Wal

teri

anel

la s

p.L

ind

enia

riv

alis

RU

BA

G2

28/V

/199

124

/VII

I/19

91R

WF,

R.

Eli

zon

do, L

. R

ose

311

87-R

WF

9151

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


HIS

PIN

AE

: s.s

.

Cep

hal

olei

a su

tura

lis

Bal

yC

ostu

s sp

.Z

INA

A1

14/V

I/19

91R

WF

41

79-R

WF

9152

Ch

alep

us

bell

ula

(C

hap

uis

)P

oace

aeP

OA

AG

3029

/I/1

994

RW

F4

267-

RW

F94

53

Car

inis

pa n

ever

-m

ann

i U

h.

Bu

nch

osia

sp.

ML

PL

G23

5/II

I/19

91D

HJ&

WH

61

91-S

RN

P-7

4

Mal

pigh

ia g

labr

a L

.M

LP

AA

518

/II/

1994

RW

F, Y

. As-

torg

a, J

. Sol

is4

148-

RW

F94

54

Dem

otis

pa s

tran

di

Uh

.S

perm

acoc

e sp

.R

UB

AA

326

/VII

/199

4R

WF

51

25-R

WF

94

Oxy

chal

epu

s al

ie-

nu

s (B

aly)

Cen

tros

ema

mac

roca

r-pu

m B

enth

.M

IMA

G9

24/V

/199

1R

WF

36

21-R

WF

91

Su

mit

rosi

s sp

.G

uaz

um

a u

lmif

olia

ST

EA

G2

6/V

/199

1R

WF

12

12&

13-R

WF

91U

ropl

ata

vari

cos-

tata

Pic

Byr

son

ima

cras

sifo

lia

ML

PA

G22

27/I

/199

2D

HJ&

WH

61

92-S

RN

P-2

94

Xen

och

alep

us

omog

era

(Cro

tch

)C

entr

osem

a m

acro

carp

um

FA

BL

G17

20/V

II/1

992

DH

J&W

H6

392

-SR

NP

-359

5

HIS

PIN

AE

: CA

SS

IDIN

I

Aka

nta

ka i

nsi

dio

sa

Boh

.T

abeb

uia

och

race

aB

IGA

G10

5/V

I/19

82D

HJ&

WH

61

82-S

RN

P-1

6955

Tab

ebu

ia o

chra

cea

BIG

L, A

G9

30/V

I/19

89D

HJ&

WH

62

89-S

RN

P-4

48

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

reN

o.V

ouch

erN

ote


Tab

ebu

ia o

chra

cea

BIG

L, A

G24

16/V

II/1

989

DH

J&W

H6

289

-SR

NP

-701

Tab

ebu

ia r

osea

(V

er-

tol.)

DC

.B

IGA

G10

27/I

II/1

984

DH

J&W

H6

384

-SR

NP

-42

Asl

amid

ium

sp.

Cal

ath

ea c

rota

life

ra S

. W

atso

nM

AR

AG

2314

/VII

I/19

91R

WF

44

75-R

WF

91

Ch

arid

otel

la n

r. eg

regi

a (B

oh.)

Ipom

oea

sp.

CN

VA

G8

10/V

I/19

89D

HJ&

WH

62

89-S

RN

P-1

6256

Ipom

oea

sp.

CN

VL

G8

25/V

II/1

995

DH

Y&

WH

61

95-S

RN

P-7

192

Ch

arid

otis

cos

tari

-ce

a S

paet

hG

uet

tard

a m

ac-

rosp

erm

a D

onn

. Sm

ith

RU

BA

G15

13/V

III/

1982

DH

J&W

H6

182

-SR

NP

-721

57

Ch

elym

orph

a sp

.Ip

omoe

a tr

ifid

a (K

un

th)

G. D

onC

NV

L, A

G17

18/I

/198

9D

HJ&

WH

66

89-S

RN

P-1

1

Cop

tocy

cla

lepr

osa

Boh

.C

ord

ia a

llio

dor

a (R

. &

P.)

Oke

nB

OR

AG

1030

/V/1

982

DH

J&W

H6

182

-SR

NP

-145

Cor

dia

all

iod

ora

BO

RA

G13

16/V

I/19

89D

HJ&

WH

63

89-S

RN

P-2

18C

ord

ia a

llio

dor

aB

OR

L, A

G25

16/V

/197

9D

HJ&

WH

64

79-S

RN

P-1

7C

opto

cycl

a so

rdid

a B

oh.

Ali

bert

ia e

du

lis

(L.

Ric

h.)

A. R

ich

.R

UB

AG

114/

V/1

980

DH

J&W

H6

880

-SR

NP

-37

58

Cyd

ista

div

ersi

foli

a (H

.B.K

.) M

iers

BIG

AG

1118

/VI/

1991

RW

F1

266

-RW

F91

Cyd

ista

div

ersi

foli

aB

IGL

, AG

1022

/VI/

1989

DH

J&W

H6

689

-SR

NP

-296

Cyd

ista

div

ersi

foli

aB

IGL

, AG

103/

VI/

1992

DH

J&W

H6

592

-SR

NP

-151

1C

ydis

ta d

iver

sifo

lia

BIG

L, A

G10

19/V

/199

4D

HJ&

WH

69

94-S

RN

P-3

006

TA

BL

E 1

.(C

ON

TIN

UE

D)

FE

ED

ING

RE

CO

RD

S O

F C

OS

TA

RIC

AN

LE

AF B

EE

TL

ES. S

EE

TE

XT F

OR

EX

PL

AN

AT

ION

OF C

OL

UM

NS.

Lea

f B

eetl

eP

lan

tP

lan

t Fa

mil

yS

tage

Loc

alit

yD

ate

Col

lect

ors

Sco

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The alternative hypothesis is that ecological mediation (colonization and hosttransfer) is the primary explanation for current host associations. In cases of ecologi-cal mediation, phylogenies of herbivores and host plants are not congruent, and hostshifts are not necessarily between sister taxa of plants Anderson (1993). In a surveyof 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 therule. However, like the Chrysomelidae, the majority of curculionine taxa lack any hostplant data. Until host plants are known for a much larger proportion of phytophagousinsect 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), andthe 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 ScienceFoundation Mid-Career Fellowship (BSR-9003898) to the senior author, and NSFDEB-9400829 to the junior author.

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

ACA Acanthaceae CNV Convolvulaceae MYR MyrsinaceaeAMA Amarantaceae DIO Dioscoreaceae ONA OnagraceaeAPO Apocynaceae ERY Erythroxylaceae PAS PassifloraceaeASC Asclepiadaceae ERI Ericaceae PIP PiperaceaeAST Asteraceae EUP Euphorbiaceae POA PoaceaeBIG Bignoniaceae FAB Fabaceae: POL PolygonaceaeBOR Boraginacaea Papilionoidea RUB RubiaceaeBUR Burseraceae FLA Flacourtiaceae RUT RutaceaeCAE Fabaceae: HIP Hippocrateaceae SPI Sapindaceae

Caesalpinoidea LAU Lauraceae SPO SapotaceaeCAP Capparidaceae LOG Loganiaceae SIM SimarubaceaeCLU Clusiaceae MLP Malpighiaceae SOL SolanaceaeCEC Cecropiaceae MLV Malvaceae STE SterculiaceaeCOC Cochlospermaceae MAR Marantaceae URT UrticaceaeCOM Combretacaea MEL Melostomataceae VER VerbenaceaeCON Connaraceae MIM Fabaceae: VIT Vitaceae

Mimosoidea ZIN Zingiberaceae


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 RecreativaJunquillal, 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, 20km 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 SantaRosa, 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)


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 SantaRosa,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 mA2 Alajuela Prov., Reserva Forestal San Ramon, 900 mA3 Alajuela Prov., Bijagua, 20 km S Upala, 500 mA4 Alajuela Prov., Canton La Guacima, Río Segundo, 780 mA5 Alajuela Prov., Canton Ciruelas, Río Ciruelas, 800 mH1 Heredia Prov., Estac. Biol. La Selva, 50 mS1 San José Prov., San Pedro, Univ. Costa RicaS2 San José Prov., El Rodeo, 1.5 km S Ciudad ColonC1 Cartago Prov., Pavones, nr. TurrialbaC2 Cartago Prov., Madreselva, nr. EmpalmeC3 Cartago Prov., Carretera Interamericana, 7 km S. CartagoC4 Cartago Prov., Cerro Asunción, paramo vegetation, 3396 mC5 Cartago Prov., Tapanti, Refugio Vida SilvestreP1 Puntarenas Prov., Reserva Forestal MonteverdeP2 Puntarenas Prov., Peninsula de Osa, Est. Boscosa, Reserva Forestal Golfo

DulceP3 Puntarenas Prov., Peninsula de Osa, Cerro de Oro

APPENDIX 2—(CONTINUED) LOCALITIES FROM TABLE 1.


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 therainy and dry seasons. The larvae make cone-shaped cases, apparently utilizinghairs 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 bushin dense dry forest. The intense feeding and mating activity was similar to thatobserved 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 regeneratingpasture 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 theonset of the summer rains. Individuals were also collected on other pastureshrubs. The collection of Jan Bechyné in Maracay Venezuela contains severalspecimens of this species collected in El Salvador 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 havebeen Asteraceae.

11. Although only one voucher was preserved, numerous adults were observed, andseveral were tested on the leaves of the plant host. Jolivet (1987b) noted that thisgenus is both cosmopolitan and polyphagous.

12. Adults were feeding at night on very new expanding leaves of a 1.5 m shoot atbase of tree. Jolivet (1987b) stated that the only reliable feeding records for thisgenus 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 areagile 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 theirhosts.

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 SanGabriel on various species of Solanum.

18. Larvae skeletonize host plant leaves. This species extensively defoliates its hostduring some years. Literature records for New World Plagiodera are limited toSalix, Populus (Salicaceae), Croton (Euphorbiaceae), and Lueha (Tiliaceae); how-


ever, species in the Philippines and India have been reported on Xylosoma andFlacourtia (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 bagwhich 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 the1991 rainy season. Apparently, our observations represent the only known hostplant data for Stilodes.

21. A group was followed from egg to adult. Larvae feed and rest on underside ofleaves. Pupation takes place in leaf litter.

22. Larvae are sooty black, covered with branched hair-like projections, and with redheads. 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 speciesin cleared pastures.

24. The host plant of this galerucine was found growing along the edge of a smallpatch 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 (Jolivet1987a).

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 eachother 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) ashosts of this genus.

31. RWF observed on individual at night eating a hole in the middle of a leaf of theIpomoea 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 notedother citations of Lauraceae, Buddlejaceae, Asteraceae, Umbelliferae, Sterculi-aceae, and Fabaceae.

33. In addition to the vouchered specimen from Byrsonima crassifolia, this specieswas abundant on this host plant at Estacion Maritza (G4) in 1991.

34. Unlike many other chrysomelids which were found associated only with youngfoliage, A. salvadorense was found actively feeding late in the rainy season onolder 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 understoryof 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.


39. Both this and the following host plant were growing close together in a mixedstand next to a road.

40. The host plant, growing in a wet depression in a cleared area, sustained heavyfeeding damage from this alticine in 1991. Jolivet (1991) listed Cleome, Solanum,Beta (Chenopodiaceae), Labiaceae, Cordia, and Adiantum (Adiantaceae) as hostplants of Leptophysa.

41. These beetles were swept from a tree that showed heavy feeding damage to theleaves. No active feeding was observed, but this collection was made during anabnormal 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, awider 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 variousplants.

49. This species feeds by scraping pits in the expanding leaves of this host plant. Thefollowing plant record may be an alternate dry season food source.

50. Huge numbers of this species were found defoliating the host plant during thevoucher 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 asmall group of flightless Mediterranean flea beetles. New World species formerlyin 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 otherZingiberales (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 agroup of the cassids. Jolivet (1988a) also listed Tabebuia and other Bignoniaceaeas hosts for this genus.


60. These records are of beetles aestivating in the dry season; see Flowers (1991) formore 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) alsolisted Hyptis (Labiaceae) as a host plant for this genus.

62. The record from Bursera simaruba is for beetles hiding under bark plates duringthe 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 abovethe 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-

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data Mill (Solanaceae).

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334 Florida Entomologist - ufdcimages.uflib.ufl.eduufdcimages.uflib.ufl.edu/uf/00/09/88/13/00048/sn... · describe conﬁdence that a ﬁeld observation of an interaction between