Hymenopterous parasites of some bruchid beetles of North and Central America

J. stored Prod. Res. Vol. 24, No. 3, pp. 131-143, 1988 Printedin Great Britain.All rights reserved

Copyright 0

0022-474X/88 $3.00 + 0.00 1988 PergamonPress plc

HYMENOPTEROUS PARASITES OF SOME BRUCHID BEETLES OF NORTH AND CENTRAL AMERICA MARTIN Department

of

Biological

HETZ

and

CLARENCE

D.

JOHNSON

Sciences, Northern Arizona University, Flagstaff, AZ 86011, U.S.A. (Received 8

February 1988)

Abstract-Five genera containing 22 species of hymenopterous parasites associated with 14 genera containing 123 species of seed-infesting bruchid beetles, and 195 species of host plants are listed. Most of the host plants are Leguminosae with some in the Malvaceae and Tiliaceae. The records represent about 20 years of gathering data on bruchids and their host plants. The interactions between the parasites, the bruchids, and the host plants are discussed. Results indicate that the parasites are not narrowly specific to a species of host bruchid but rather have many hosts over a broad geographic area. Most likely the parasites search for seed pods rather than for the kind of insects that feed inside the seeds.

INTRODUCTION A large amount of research has been done on the economic and noneconomic species of the beetle family Bruchidae over the last 25 years, so that we have a basic understanding of the systematics, ecology and host plants of the group (e.g. Janzen, 1969, 1971, 1980; Johnson, 1970, 1981a, b, c, 1983a, b; Johnson and Kingsolver, 1973, 1976; Kingsolver, 1986; Kingsolver and Johnson, 1978; Kingsolver and Whitehead, 1974a, b, 1976; Kingsolver et al., 1977; Whitehead and Kingsolver, 1975a, b, c). One of us (CDJ) has done a great deal of specialized field work and has found many host plants for the noneconomic species of these beetles. These host records have been published largely in the papers by Johnson and by Kingsolver. Although many parasitic Hymenoptera were reared from seeds by Johnson along with bruchids, few papers have been published which included the records of parasites. In this paper we report on the parasitic Hymenoptera which were reared from these seeds and are presumed to be parasites of bruchids. METHODS

Seed pods and seeds were removed from plants and placed in brown paper sacks in the field. A collection number was assigned to each sack and to a corresponding plant sample taken for identification. In the laboratory, pods were allowed to dry in the paper sacks to prevent the growth of mold. They were then transferred to wide mouth mason jars (pint or quart) for emergence of insects. The jars were checked at weekly intervals and all insects that emerged were removed with an aspirator, killed, and mounted. To prevent infestation by pyemotid mites, the sealing lids of the jars were replaced by 1% Kelthane impregnated paper towels and secured with the screw-on ring. The jars were stored on similarly-treated paper towels. We included only those Hymenoptera recorded as parasites of Bruchidae (Krombein et al., 1979) in the list. Parasites identified to genus only were included since they were taxonomically similar to verified bruchid parasites and were reared from seed collections from which bruchids were also reared. These represent probable associations. Because names were not available, either letters or numbers were given to these entities by the specialists who identified them. We used a Honeywell CP6 computer using a sort program. Egg parasites were not included. RESULTS Relatively little is known completed one of the earliest

AND

DISCUSSION

about the Hymenoptera parasitizing bruchids. Cushman (1911) lists devoted to the parasites of numerous bruchid species followed 131

132

MARTIN HETZ and CLARENCE D. JOHNSON

Table 1. Parasites of Bruchidae associated with specific host plants, host beetles, and geographic locations are included in the table. Host plants are those from which the parasites were reared. Host beetles are those that emerged from the same host plant as a parasite. Geographic locations, given in standard abbreviated form, are from where the plant samples were taken that yielded the parasites. Authors of species names are listed only when the name is first cited. Most records of rearing of bruchids and parasites from seeds are not single collections but represent many samples from the same species of plants Parasitoid host plant

Host Bruchidae

Location

BRACONIDAE Heterospilus bruchi Viereck Acacia cymbispina Sprague & Riley A. picachensis Brandegee

A. riparia H.B.K. vel. sp. aff A. willardiana Rose Albizia occidentalis Brandegee Eauhinia ungulata L. Calliandra hottstoniana (Mill.) Standl. Calopogonium mucwtolides Dew. Cassia emarginata L. Leucaena guatemalensis (Britt. & Rose) Mimosa albida Humb. & Bonpl. Pachyrhizus erosus (L.) Phaseohts vulgaris L. Pitheceilobium mongenre Macbride Tephrosia cinerea Pets. Heterospilus prosopidis Viereck Acacia angustissimo (Mill.) Ku&e A. constricta Bentham

A. cymbispina A. greggii Gray Aeschynomene americana L. Calopogonium mucunoides

Mimosestes mimosae (Fabricius) Acanlhoscelides schrankiae (Horn) Merobruchus sp. Stator limbatw (Horn) S. vittatithorax (F’ic) Merobruchus sp. Stator oittotithorax Stator limbatur A4erobruchu.v insolitus (Sharp) Gibbobruchus sp. Stator limbatus Acanthoscelides puellus (Sharp) Amblycerus sp. Sennius militaris (Sharp) Merobruchus sp. Stator limbatus Acanthoscelides cordifer (Sharp) A. d@cilis (Sharp) Acanthoscelides sanfordi Johnson Merobruchus sp. Acanthoscelides flauescens (Fiihraeus)

Merobruchw placidus (Horn) Stator limbatus Mimosestes acaciestes Kingsolver & Johnson M. amicw (Horn) Stator pruininus (Horn) Mimosestes mimosae Stator vachelliae Bottimer Acanthoscelides pertinax (Sharp) Meibomeur sp. Acanthoscelides pue1ht.v

Cassia nicaraguensis Bentham C. ObtusiJolia L.

Sennius gultij2r (Sharp) Sennius instabilis (Sharp) S. Iebasi (FPhraeus)

C. reticulata Willdenow Cercis occidentalis Torr. ex Gray Chaetocalyx latisiliqua (Poir.) Benth. ex Hemsl. Desmanthus covillei (Britt. & Rose) Wiggins ex Turner D. fruticosus Rose D. uirgatus (L.) Willdenow

Sennius lebasi Gibbobrwhus minus (Say) Acanlhoscelides mundulw (Sharp) Acanthoscelides desmanrhi Johnson SIator pruininus Acanthoscelides compressicornis (Schaeffer) A. desmanthi Stator pruininw Acanthoscelides ctternauaca Johnson A. desmoditus Johnson Acanrhoscelides kingsolveri Johnson

MEX, OAX MEX, OAX

MEX, COL MEX, MEX, GUAT MEX, GUAT MEX, MEX,

SON BCS NAY MICH OAX

MEX, COL MEX, OAX MEX, MEX, MEX, MEX,

CAMP QR GRO COL

MEX, SIN USA, AZ

MEX, SON USA, AZ MEX, CAMP MEX, MEX, MEX, MEX,

GRO MICH JAL OAX

S. rnoro~u~ (Sharp)

Desmodium distorturn (Aublet) Macbride Indigofera suffruticosa Mill. Leucaena aff. collinsii Britt. & Rose L. esculenta (Mot. & Sesd) Bentham

L. lanceolatn Wats. L. leucocephnla (Lam.) de Wit. Mimosa albida M. invisa Mart. M. langlassei Mich.

M. palmeri Rose Piptadenia flaun (Spreng.) Benth. Prosopis jtdi~7ora (SW.) DC.

Acanthoscelides mocrophthalmus (Schaeffer) Acanfhoscelides boned Johnson A. leucaenicola Johnson A. mankinsi Johnson Acanthoscelides mankinsi Acanthoscelides mocrophthalmus Acanthoscelides mexicanus (Sharp) A. schrankiae Aconthoscelides piceoapicalis (Pit) Acanrhoscelides cordifr A. d@iciiis A. mimosicola Johnson Stator pruinintu Acanthoscelides mexicanus Stator pruininus Acanthoscelides clireliarius (Ftiaeus) Algarobius prosopis (LeConte) Mimosestes amicus

PAN USA, PAN MEX, MEX, MEX, MEX,

AZ SON BCS OAX SIN

MEX, OAX MEX. COL MEX; SON GUAT GUAT MEX, VER MEX, MEX, MEX, MEX,

NAY SIN WC JAL

PAN MEX, OR0

MEX, SIN

MEX, GRO MEX, JAL continued

Parasites

of Brttchidae

of North

and Central

133

America

Table I--contimted Parasitoid host plant

Host Bmchidae

Rhynchosia minima (L.) DC.

senna ~~~~a~~

L.

S pallida (Vahl.) var gaumeri (Britt. & Rose) Irwin & Bameby S. pa&da var pai& (Vah1.f Irwin & Bameby Sertna uilIasa (Mill.) Irwin & Bameby Heterospilw sp. A Ajbizia lebbek (L.) Bentham

Heterospiias sp. B Caesalpinia platyloba S. Wats.

Acanthosceiides jiavescens Sennius lebasi S. morosus S. nrfomaculatus (Mots.) Acanthoscelides obrienonrm Johnson

Se&us auricomw Johnson & Kingsolver Sew&s discolor (Horn] Sennius moroswx Merobrwhus ins&us Stator iimbatus -

Location MEX, VER MEX, GRO

MEX, YUC MEX, GRO MEX, CAMP

MEX, BCS

MEX, SIN

~eterosp~us sp. C

Caiapoganiu~

caeraktan

(Benth.) Sauv.

Pachyrhisus erosus HeterospiI~ sp. D Acacia angastissima Heterospilus sp. E Cassia bicapsuiaris L.

Desmanthus virgatw Leueaena ~c~ep~a Mimosa albidn Stenocorse bruchivora (Crawford) Acacia angustissima

A. constricta

A. coaiteri Bentham A. cymbisptita

A. golobnanii (B&t. 8c Rose) Wiggins A. gaumeri Blake A. mcmurphyi Wiggins A. occidentalh Rose A. picachensis

Albizia sinaloensis Britt. & Rose

&whit& angulata Calopogonil4m caendeum

C. mwwoides

Cassia bicapsularis

Cassia bipora L.

C. ieptocarpa Bentham C. obtustfolia

AcanthosceEdes hectori Kingsolver Caryedes paradisensis Kings&et & Whitehead Acanthoscelides sanfordi

PAN

Merobruchus sp.

MEX, OAX

Senntus celatus (Sharp) S. fallax (Boheman) S. gattijer S. leucostauros Johnson & Kingsolver Acanthoscelides desmanthi Stator pruininus Acanthosce~~es macropht~l~s Aeaathos~el~s mexicanur

MEX, JAL

Merobruehus placidus Merobraeh~ sp. Stator praininas Mimosestes acaciestes M. amicus Stator pruininus Merob~h~ sp. Mimosestes janzeni Kingsolver & Johnson M. mimosae

MEX, GRO USA, AZ

Stator Iimbatus Stator ligate S. vittatithorax Statar limbatus Merobruchus sp. Stator limbatus A~~th~~e~ides schraakiae Merobruchus sp. Stator iimbatus S. vittatithorax Merobruchus insolitus Stator limbalas Gibbobrachus sp. Acanthoscelides hectori A. taboga Johnson Caryedes paradisensb Ac~th~celtdes pueiias

Sennius celatus S. fallax S. guttijer s. letKostawos Acanthoscelides obrienorwn Sennius atripectus Johnson & Kingsolver Sennius celatus S. stasis S. morosus Sennius instabih3 S. lebasi S. morosus Sennius cetatus s. fallax

MEX, CAMP

MEX, SIN MEX, YUC MEX, JAL

USA, AZ

MEX, MEX, MEX, MEX, MEX, MEX,

SON JAL SIN SON BCS YUC

MEX, BCS MEX, SON GUAT MEX, OAX

MEX, SON PAN PAN

MEX, MEX, PAN MEX, MEX,

GRO NAY COL SIN

MEX, OAX MEX, JAL MEX, NAY MEX, OAX

MEX, NAY continued

134

MARTIN HETZ and CLARENCE D. JOHNSON

Table I-continued Parasitoid host plant

c. torn L. C. wislizenii Gray Cercidiwn @ridurn Bcntham

C. microphylium (Torr.) Rose & Johnston C. praecox (Ruiz & Pav.) Harms Desmanthus covillei 1). fruticosuf indigofera sphaerocarpo Gray I. suflruticosa Ipomoea afl simulata I. simulans Hanbury Leucaena glauca (L.) Bentham L. g~te~le~is L. leucoeephaia Lysiloma divaricata (Jacq.) MacBride

Mimosa albida hf. bahamensis Bcntham M. easiir L. M. dysocarpa Bcntham M. langlassei

M. laxipoa Bcntham

M. palmeri

M. pigra L.

M. platycarpa Bet&am M. pudica L. M. setigera Britt. & Rose

M. ~~10s~ Mart. Phaseol~ ltatatus L. Piptadenia Java Piscidia mollis Rose Pitbecellobium sonorae S. Wats Pitryoearpa fZava (Sprcng.) Be&h. Rhynchosia Iongeracemosa (Mart. & Gal.)

R. minima

Senna cobanensis (B&t. & Rose) Irwin & Bamcby S. foetidissima (S. & M.) var grandtxora (Bcnth) Irwin & Bameby S. obtusifolia S. pahida (Vahl.) Irwin & Bameby S. villosa Vrosigalphus bruchi Crawford Acacia cochliacantha Numb. & Bonpl. cx. Willd. A. c.vmb~pi~ A. farnesiana (L.) Willd. A. greggii

Host Bmchidae Sennius celatus S. instabilis Acanthoscelides obrienorum Mimosestes amicus M. ufkei (Horn) Stator Iimbatus Mimosesres arm&s Stator limbatus Acanthoscelides derififdi Johnson A. desmanthi Stator pruininus Stator pruininus Acanthosceiides kingsoberi Acanthoscelides kingsohreri Megacerus callirhipis (Sharp) Megacerus sp. Acanthoscelides ~cropht~i~ A~~th~ee~~s ~~opht~~ AcanthosceRdes macrophthaImus Merobruchus insohtus Stator hmbatus S. sordidus (Horn) Acanthoscelides difficilis Ae~thosceIi~s sehrankiae Ae~t~scei~s piceo~~~~ Acanthoscelides chiricahuae (Pall) Acanthoscelides cordtfer A. dtL%ilis A. mimosicola Stator pruininus Acanthoscelides chirhahuae A. mexicanus Stator pruininus Acanthosceiides chiricahuae A. mexicanus Stator pruininus Acanthoscelides di@icilis A. pigricoia Kingsolver A. q~~i~ntat~ (Schacffer) Acanthoscelides mexicanus Acanthoscelides piceoapicalis Acanthoscelides mexicanus A. schrankiae Stator pruininus Aeanth~ceii~s cordiier Acanthoscelides argillareus (Sham) Acanthoscelides clitellarius Ctenocolum janzeni Kingsolver & Whitchead Merobruchus sp. Stator limbatus Aeanthosceli~s ~iitel~ar~s Acanthoscel~s campeche Johnson A. jJaveseens A. pabnasola Johnson Acanthoscelides jhtvescens A. zeteki Kingsolver

Sennius iebasi S. rufomaculatus Sennius fallan Senniu.9morost4.s S. ~foma~Iai~

LXX&OR

MEX, COL USA, AZ USA, AZ

USA? AZ MEX, SIN MEX, SON

MEX, BCS HOND GUAT MEX, COL MEX, NAY MEX, BCS GUAT MEX, YUC MEX, BC MEX, SON MEX, MEX, PAN MEX, MEX,

CHIS YUC SON GRO

MEX, BCS MEX, OAX MEX, SON MEX, SON

HOND MEX, GRO GUAT PAN MEX, JAL

MEX, GRO HOND MEX. MICH MEX, JAL MEX, SON MEX, SON MEX, OAX MEX, CAMP MEX, QR MEX, MEX, MEX, PAN MEX.

GRO JAL MlCH OAX

MEX, CHIS MEX, GRO

Aeant~s~e~~~s obrieno~ Sennius discolor -

MEX, MICH

Mimosestes mimosae M. nubigens (Mot~hulsky) M~mosestes jaeni hf. mimosae Mimosestes nubigens

MEX, SIN

-

MEX, BCS

MEX, SIN MEX, PUB MEX, SON USA, AZ continued

Parasites

of Bruchidae

of North

Table Parasitoid

host plant

Caesalpinia cacalaco Humb. Cassia bicapsularis

& Bonpl.

PiUteceNobium dulce (Roxb.) Prosopis julijora Vitis arizonica Engelm.

Benth.

Cassia leptocarpa Condalia spathularn Gray Ipomoea simulans Lysiloma diuoricara Mimosa biuncifera Bentham M. dysocarpa M. setigera Nissolia schorlii (Torr.) Pip&den&a J%ZW Pirhecellobium sonorae

Gray

P. undulatum (Britt. & Rose) Gentry Pitryocarpo /lava Prosopis pubescens Bentham

A. coulreri A. cymbispina A. farnesiana A. greggii A. occidenralis A. pennatula (Schleetendal & Chamisso) A. schaffneri (S. Wats.) Herman

C. microphyllum

divaricara

Mimosa la.x~~oro Nissolia schotrii

MEX, JAL MEX, MEX,

COL SIN

MEX,

SON

EL SALV USA, AZ USA. AZ

USA, AZ USA, AZ

MEX,

SON

USA, USA, MEX, MEX, USA,

AZ AZ NAY SON AZ

MEX, SON MEX. JAL USA, AZ MEX, GRO MEX, SON

MEX,

SON

MEX, GRO USA. CA

Bentham

Sennius inrrabilis S. lebasi S. morow

MEX,

Mimosestes acaciestes M. amicus Stator pruininus Merobruchus sp. Mimosesles mimosae Mimosesres nubigens

USA,

Merobruchus sp. Slator limbatw Mimosestes humeralis (Gyllenhal) Mimosestes amicus M. nubigenr Merobruchus insolitus Staror limbarus Gibbobruchus sp. Amblycerus sp. Sennius celatu.3 s. fallax S. gulti/er Mimosesres amicus M. ulkei Stator Iimbarus Mimosesles omicus Srntor limb&us

C. praecox Lysiloma

Location

Gibson

Urosigalphus neobruchi Gibson Acacia constricta

Cercidium jloridum

oilis (Schaeffer)

Swor Iimbotus Mimosesles acaciesres M. amicus Staror pruinirw Merobruchus sp. Staror Iimba~w Sennius medialis (Sharp) Neltumius texanus (Schaeffer) Megacerw sp. Stator sordidus Acon~hoscelides chiricahuae A. mexicanus Acanthoscelides chiricahuoe Acan~hoscelides mexicanus SIalor pruininus Acanlhoscelides mundulus Acanrhoscelides clireliarius Merobruchus insolitus Merobrwhus sp. Stator limbarus Merobruchus boucheri Kingsolver M. insolitus Acanthoscelides clirellarius

A. occidentalis

Bauhinia ungulata Cawia bicapsularis

l-continued

Amblycerus

Urosigalphus bruchivorus Crawford Acacia angusrissimo A. consrricm

135

America

Mimosesres mimosae M. viduarua (Sharp) Amblycenrs ocapulcensis Kingsolver Amblycerus sp. Sennius celarus s. fallax S. gurri/er Mimosesles amicus Stator limbarus -

Cercidium microphyllum

Albizia sinoloensis

Central

Host Bruchidae

A. hindrii Bentham

Urosigalphus mexicanus Cassia obtusfolia

and

Merobruchus insolirus St&or limbarus S. sordidus Acanrhoscelides chiricahuae Acanrhoscelides mundu1u.v

OAX

AZ

MEX. MEX; GUAT MEX, USA, MEX,

SON AZ SON

MEX, MEX,

OAX SON

MEX,

SON

MEX, MEX.

CAMP SIN

USA,

SON SON

AZ

MEX, SON USA, AZ MEX, MEX, MEX,

SIN SON SON

MEX, SON USA, AZ

136

MARTIN HETZ and CLARENCED. JOHNSON Table 1-conrinued

Parasitoid host plant Prosopis jtdliporn P. laevigoto (SW.) DC P. pabneri S. Wats. Ziziphus obtusi&Xia Hooker

Host Bruchidae

-

Algarobiw sp. Mimosesres amicus Acanthoscelides prosopoides (S&&%r)

Location USA, TX MEX, DGO

MEX, BCS usa,

AZ

EULOPHIDAE Horismenw bruchophagus Barks Albizia occident& A. sinaloensis Desmanthus fruticosus D. subularw (Britt. & Rose) Wiggins ex. Turner

Horismenus missouriensis (Ashmead) Abulilon ealifornicwn Bentham A. incanum (Link) Sweet A. pringfei Hochr. Acacia angustissima A. eonstricra

Merobruchus sp. ~erob~ch~ ~soiit~ Stator limbatus Slotor pruininus Aconthoscelides derifiedi Smtor pnrininus

Aconthoscelkies suboequalis Johnson Aconthosceiides aequaZ& (Sharp)

A. ~~~~ Aconthoscelides subaequalis Merobruchw placidus Stotor limbaw Mimosesres ncaciesres

Merobruchus sp. Mimosestes mimosae

A. faraesiana

Mintofestes mimosae 84. nubigens

A. penraatulo A. riparioides (Britt. & Rose) Standley A. schaffneri

Mimosestes mimosae Merobruchus sp. Mimosesres amicus M. nubigens Merobruchus SQ. Storor limbatw Acanthoscelides megocornb Kingsolver A. perlinax Meibomeus sp.

Aeschynomene americana

A. rudis Be&ham R. xnsitiva SW. Albizia lebbek A. sinaloensis Amorpha fwticosa L. Asiragalus bisulcatus var bisulcams (Hook) Gray A. lentiginosus Douglas A. thurberi Gray

Bouhinia divoricato (Hart.) ex Steud. B. ungulate Caesalpinia cacaiaco Ca~~~~a eriophylia Bcntham Colopogonium coeruleum

C. mucunoides Canavaiia oxyphylla Standley & L. D. Williams Cossia bicapsuloris

C. btQ?ora C. Ieptocarpa

C. moschato H.B.K. C. occidentalis C. skinneri Bentham

BCS

MEX. SON MEX, BCS MEX, BCS

MEX, BCS MEX, SON MEX. ML MEX; YUC USA, AZ MEX, SIN MEX, SON USA, AZ

M. zzn&w Stotor pruininus

A. coulteri A. cymbispino

A. ~enu~Z~a (L.) Willd.

MIX,

~C~t~s~eI~smdestws(Sharp) Acanrhoscelides mod&us Merobruchus SQ. Stator limbatus A~anthosca~i~s coihtsw (FalI) Acan~hoscelt’des aureohw (Horn) A. fraarercuius (Horn) AconrhosceUs oureolus camp Acanthosceltdes aureolus A. m&us (Horn) A. puilus (Fall)

Gibbobruchus sp. -Amblycerus sp. Stotor pruininus Aconthoscelt’des hectori A. tabogo Caryedes paradisensis Acanthoseeiides puelius A. tabogo Caryerks sp. Sennius celotus S. fallax S. guttifer S. l~~ostauros Sennius auricomus S. insrabilis Sennius ceiams S. insmbilis S. media& S. morosus Pygiopachytnerus SQ. Sennius celarus -

MEX, SON MEX, SIN MEX, SON GUAT MEX, JAL MEX, PUE MEX, SIN MEX, SON MEX, YUC MEX, MICH MEX, SON MEX, CAMP MEX, MICH PAN MEX, MEX, MEX, MEX,

JAL OAX VER SON

USA, AZ USA, UT USA, AZ USA, AZ

MEX, MEX, MEX, USA, PAN

VER CAMP SIN AZ

HOND MEX, MICH PAN MEX, COL MEX, JAL

MEX, SON MEX, NAY USA. AZ PAN MEX, NAY MEX, OAX

137

Parasites of Bruchidae of North and Central America Table Parasitoid

host plant

Host Bruchidae

Chaetocalyx lattsiliqua Condalia globosa Johnston Cracca sp. Dalea aff submontana Rose D. tomentosa (Cav.) Willd. var tomentosa Desmanthus covillei D. virgatus

D. distortum D. incanum DC. D. nicaraguense Oersted. D. tortuowm (SW.) DC. Glycyrrhiza lepidota Pursh. Guaruma ulmijblio Lam. Heliocarpus palmeri S. Wats. Heriwantia crtspa (L.) Briz. Indigofera densiflora M. & G.

I. suffruticosa

Leucaena aff collinsii L. esculenta

L. glauca L. leucocephala L. shannoni Dorm. & Sm. Lotus sp. Lysiloma acapulcensis Bentham L. diuaricata L. thornberi Britt. & Rose

Mimosa albida

M. bahamensis M. biuncifea M. brandegei Robinson M. dysocarpa M. invisa M. Iaxipora M. palmeri

M. pigra M. pudica M. vellosiana Nissolia fiuticosa Jacq. N. schottii Ocimum micranthum Willd. Parryella filifolia Tom. & Gray Piptadenia Java Pithecellobium dulce P. palmeri Hems]. P. undulatum Pitryocarpa j7ova

Location

Sennius auricomus -

C. xiphoidea Bertol. Cerciditun ~oridum C. microphyilum

Desmodium cinereum (H.B.K.)

I--continued

DC.

MEX, SIN USA, AZ MEX, SON

Mimosestes amicus Stator Iimbatus Acanthoscelides mundulur Neltumius texanus Meibomeus? Acanthoscelides Acanthoscelides Acanfhoscelides Stator pruininus Acanthoscelides A. howdenorum A. megaeornis Acanthoscelides A. desmoditus Acanthoscelides

pusillimus (Sharp) desmanthi a’esmanthi

PAN USA, AZ MEX, QR MEX, COL MEX, NAY MEX, SON MEX, SIN

desmoditus Johnson

MEX, MICH MEX, OAX

Johnson

cuernavaca

MEX,

schubertae Johnson

MEX, QR MEX. YUC GUAT MEX, SON

Acanthoscelides desmoditus Acanthoscelides biustulus (Fall) A. cuernavaca Acanthoscelides aureoles Acanthoscelides guazumae Johnson & Acanthoscelides cornis Johnson A. johnsoni Kingsolver Acanthoscelides herissantitus Johnson Acanthoscelides kingsoheri A. ruj?coxis (Sharp) Meibomeus sp. Acanthoscelides ktngsolveri

Acanthoscelides Acanthoscelides A. leucaenicola A. mankinsi Acanthoscelides Acanthoscelides Acanthoscelides A. mankinsi

USA, CO PAN MEX, GRO MEX, JAL MEX, SON MEX, OAX

maerophthalmus boneti

MEX, OAX MEX, SIN MEX, SON GUAT GUAT

macrophthalmus macrophthalmus macrophthalmus

MEX, BCS MEX, CAMP MEX, CHIS

Merobruchus sp. Stator limbatus Merobruchus insolitus Stator limbatus Merobruchus insolitus M. knulli (White) Stator limbatus Acanthoscelides cordi/er A. d@cilis A. mexicanus Acanthoscelides schrankiae Acanthoscelides chirichahuae A. mexicanus Acanthoscelides chiricahuae A. mexicanus Acanthoscelides chiricahuae Acanthoscelides piceoapicalis Acanthoscelides mexicanus Stator pruininus Acanthoscelides chiricahuae A. mexicanus Stator pruininus Acanthoscelides digFcilis A. quadridentatus Acanthoscelides piceoapicalis Acanthoscelides Acanthoscelides Acanthoscelides Acanthoscelides Stator limbo&s

OAX

cordifer mundulus mundulus

clitellarius

Merobruchus sp. Acanthoscelides clitellarius

USA, NM GUAT HOND MEX, SON USA. AZ

HOND MEX, CHIS MEX, JAL MEX, YUC USA, AZ MEX,

BC

USA. AZ PAN MEX, SON MEX, SIN MEX, SON HOND COLOM PAN MEX, CR0 MEX, OAX USA, AZ MEX, YUC USA, AZ MEX, GRO MEX. SIN MEX, DGO MEX, SON MEX, CR0 MEX, MICH

MARTINHETZ and CLARENCE D. JOHNSON

138

Table Parasitoid

host plant

Host Bruchidae

Prosopis jldipora P. palmeri P. uelutina Wooton Pseudosamanea guachepele Harms, Rhynchosia calycosa Hemsl. R. longeracemosa R. minima

I.G.

S. pendula Willd. var oualifolia Mart. & Gal. S. pistacifolia (H. B. & K.) Irwin & Bameby S. racemosa (Mill.) Irwin & Bameby S. skinneri (Benth.) Irwin & Bameby S. villosa Sesbania emerus (Aubl.) Urban S. nr macrocarpa Muhlenberg Siaiz acuta Bum. f. S. rhombifolia L. S. spinosa L. Tephrosia cinerea T. multifolia Rose Triumfetta calderoni Standl.

Fawcett

USA, MEX,

zeteki campeche jlavescens

Sennius fallax Amblycerus sp. Sennius fallax S. gutrife S. inanis (Sharp) S. lebasi S. leucostauros -

S. mutisiana Kunth S. pallida var pdlida

(H.B.K.)

Location

Algarobius prosopis Mimosestes amicus Algarobius sp. Mimosestes amicus Acanthoscelides Acanthoscelides Acanthoscelides A. reteki

Sewn foetidissima var grandtjlora S. indecora H. B. & K.

T. lappula L. Vigna peduncularis

I-continued

& Rendle

Sennius auricomus S. b$orae Whitehead Amblycerus sp. -

AZ BCS

USA, AZ COLOM PAN MEX, VER GUAT HOND MEX, CAMP MEX, GRO MEX, MICH MEX, SIN PAN MEX, CHIS MEX, CAMP MEX, MICH

COLOM GUAT & Kingsolver

Acanthoscelides obrienorum Acanthoscelides obrienorum Acanthoxelides obrienorum Acanthoscelides griseolus (Fall) Acanthoscelides griseolw Stator pruininus Acanthoscelides brevipes (Sharp) Acanthoscelides brevipes Acanthoscelides brevipes Acanthoscelides rufovittatus (Sehaeffer) Acanthoscelides rufovittatus Acanthoscelides johnsoni A. sanblas Johnson A. triumfettae Kingsolver Acanthoscelides triumfettae Acanthoscelides suaveolus (Sharp)

MEX, VER COLOM MEX, YUC GUAT MEX, OAX MEX, SIN MEX, COL MEX. JAL MEX; MICH HOND MEX, GRO MEX, JAL MEX, GRO HOND

HOND PAN

EUPELMIDAE Eupelmus amicus Girault Acacia angustissima A. berlanderi Bentham A. constricta

A. cymbispina A. farnesiana A. picachensis

A. riparioides Albizia occidentalis Bauhinia wgulata Cajanus cajan (L.) Millsp. Calopogonium munutoides Cassia btjlora

C. leptocarpa

C. nicaragwnsis C. obtusijolia

C. occidentalis C. reticulata

Merobruchus placidus Stator limbatur Mimosestes acaciestes M. amicus Stator pruirrinus Mimosestes mimosae Stator uachelliae Mimosestes nubigens Stator vachelliae Acanthoscelides schrankiae Merobruchus sp. Stator limbatus S. uittatithorax Merobruchus sp. Merobruchus sp. Gibbobruchus sp. Acanthoscelides reteki Acanthoscelides puellus A. taboga Sennius auricomus S. celatus S. fallax Sennius celatus S. instabilis s. morosus Sennius guttifer Sennius instabilis S. lebasi S. morosus Sennius lawrencei Johnson

MEX,

SON

MEX, COA USA, AZ

MEX, MEX, MEX, MEX, MEX,

SIN SON SON VER OAX

MEX, MICH MEX, BCS PAN PAN HOND MEX,

NAY

MEX,

NAY

MEX, JAL MEX, OAX MEX, SON MEX, PAN

SIN continued

Parasites

of Bruchidae

of North

and Central

America

139

Table I-continued Parasitoid host plant C. wislizenii Cercidium microphyllum Desmanthus covillei

D. fruticosus Indigofew pIatycarpa Rose

I. sphaerocarpa

1. suffruiicosa Ipomoea aE simufata I. hederifolia L. Leucaena glauca Leuhea seemanii Lysiloma acapulcensis L. divarieata

Mimosa aff laxipora Bentham M. albtda M. ~h~ens~s M. brandegei M. iaxipbra

A4. pigra Piptademiaj%va Pithecellobium aff mangeme (Jacq.) MacBride P. dalce Prosopis palmeri Pseudosamanea guaehepele Rhynchosia minima Senna indecora

Sesbaniu nr macrocarpa Sida aggregata F’resl. Tephrosist cinerea Triumfetta lappula Vigna peduncularis Eupelmus sp. Abutilon inranum Acacia angustissima A. constricta

A. mcmurphyi Albizia occi~ntal~ As&rag&s antiselii Gray A, praelongus var lonchopus Sheldon A. thurberi A. trichopodus Gray Bauhinia ungulata Cassia bipora C. wislirenii Desmanthus covillei

D. fruticosus D. subulatus Eysenhardtia adenostyiti Baill. Indigofera suffruticosa

Host Brncbidae Acanthoscelides obrienorum Mimosestes amicus Stator limbatus Acanthoscelia%s compressicornis A. &@eidi A. desmanthi Srator ~~~n~ Stator pruinirms Acanthoscelides baboquivari Johnson A. kingsolveri Meibomeus sp. Acanthoscetides kingsolueri Megaeerus cai~irh~~ Megacerw callirhipis M. eulophus Acanthoscelides macrophthalmus Merobruchus sp. Stator Iimbatus Merob~c~s insoiitus Merobruchus sp. Stator Iimbatus Acanthoscelides chiricahuae A. mexicanus Acanthosceiides cordger Ac~t~scelides schrankhze Acanthosceiirles chiricahuae A. mexicanus Acanthoscelides chiricahuae A. mexicanus Stator pruininus Acanthosceiides pigricola A. ~i~niat~ Acanthoseelides cliteliarius Merobruchus boucheri Acanthoscelkies jiavescens A. reteki Ambfycerus sp. Sennius guttijer S. inanis S. lebasi S. Ieucostauros Acanthoscefides grijeolus Stator paling Acanthoscelides Javescens Acanthoscelides johnsoni Acanthoscelides suaveohts

Aeanthosre~i~s ~ae4ua~is Stator limbatus Mimosestes acaciestes M. amicus Stator pruininw Stator limbatus ~~rob~ch~ &so&us Acanthoscelides aureolus AcanthosceUs mixtus AcanthoscelidPs puliw Gibbobruchus sp. Acanthoscelides obrienorum Senn~s atripectus Acanth~cei~s obrienorum Sennius sp. Acanthoscelidss Lrifieldi A. desmanthi Stator pruininur Stator pruininw A~ant~s~ii~s ~~z~e~i Stator pruininus -

Location MEX, PUE USA, AZ MEX, SON

MEX, BCS MEX, MICH

HOND MEX, SON MEX, COL MEX, JAL MEX, BCS PAN GUAT GUAT MEX, BC MEX, OAX MEX, BCS MEX, MICH MEX, YUC MEX, BC MEX, SON

HOND MEX, GRO MEX, SIN MEX, SIN MEX, BCS COLOM MEX, MICH MEX, QR MEX, CAMP

MEX, COL MEX, SON MEX, COL PAN PAN

MEX, COL MEX, SON USA. AZ

MEX, BCS MEX, BCS USA, CA USA, AZ USA, AZ MEX, BC GUAT MEX, OAX USA. AZ MEX, SON MEX, BCS

MEX, BCS MEX, OAX MEX, SIN MEX, SON continued

140

MARTINHETZ and CLARENCED. JOHNSON Table l-continued

Parasitoid host plant

Host Bmchidae

Let&a seemanii Lysiloma divaricata Mimosa aff laxipora

M. albida

N. biuncifera M. brandegei M. dysocarpa M. laxipora M. paimeri M. pigra

M. polyanthoides M. pudica M. setigera

Robinson

Nissolia schottii Piptadenia flaoa Pithecellobium duke Rhynchosia minima Senna hirsuta L. var hirta Irwin & Bameby

S. pallida Setma skinneri Triumfetta loppula

Location

-

PAN MEX, BC

Merobruchus insolitus Stator limbatus Acanthoscelides chiricahuae A. mexicanus Acanthoscelides cordifer A. difFcilis A. mimosicola Acanthoscelides chiricahuae A. mexicanus Acanthoscelides chiricahuae A. mexicnnus Acanthoscelides chiricahuae

MEX, BCS MEX, COL MEX, OAX USA, AZ MEX, BC MEX, USA, MEX, MEX, MEX,

Acanthoscelides mexicanus Stator pruininus Acanthoscelides mexicanus Stator pruininus Acanthoscelides diffilir A. pigricola A. quadridentatus Acanthoscelides schrankiae Acanthoscelides piceoapicalis Acanthoscelides mexicanus Stator pruininus Acanthoscelides

SON AZ OAX SON SIN

HOND

MEX, GRO PAN MEX, JAL USA, MEX, MEX, MEX, MEX,

clitellarius

Acanthoscelides zeteki Sennius lebasi S. morows S. rttfomaculatus Acanthoscelides obrienorum Sennius discolor Acanthoscelides obrienorum Acanthoscelides johnsoni

AZ GRO SIN GRO OAX

MEX, MICH MEX, MICH PAN

PTEROMALIDAE Lariophagus texamts Crawford Acacia cochliacantha A. collinsii Safford A. cornigera (L.) Willd.

A. cymbispina A. fame&ma

A. greggii A. schaffneri Albizia lebbek Callkmdra

eriophylla

Leucaena shamtoni Lysiloma divaricata

Prosopis palmeri

Zatropis incertus (Ashmead) Abutilon amplexifolium (DC.) G. Don A. calt~ornicum

Mimosestes mimosae M. nubigens Stator sordidus S. vachelliae Acanthoscelides oblongoguttatus A. sousai Johnson Mimosestes mimosae Stator vachelliae Mimosestes nubigens Stator vachelliae

MEX, SIN MEX, CHIS (FBhraeus)

Mimosestes nubigens Merobruchus isnolitus Stator limbatus Stator Chihuahua Johnson & Kingsolver S. limbatus S. pruininus Acanthoscelides macrophthalmus Merobruchus insolitus Mimosestes mimosae Stator limbatus S. sordidus Algarobius sp. Mimosestes amicu.9

A. incnnum

Acanthoscelides tepic Johnson Abutiloneus idoneus Bridwell Acanthoscelides subaequalis Acanthoscelides subaequalis

A. palmeri Gray A. trisulcatum (Jacq.) Urban Acacia angustissima

Acanthoscelides sleeperi Johnson Acanthoscelides subaequalis Merobruchus sp.

MEX, VER MEX, MEX, MEX, MEX, MEX, MEX, USA, MEX, MEX,

SIN SON JAL MICH SIN SON AZ GTO BCS

USA, AZ

GUAT MEX, SON

MEX. BCS

MEX, NAY MEX, BCS MEX, SON MEX, BCS MEX, SIN MEX, SON USA, AZ MEX, SON MEX, GRO GUAT MEX, CAMP continued

141

Parasites of Bruchidae of North and Central America

Tabk l-canrirwd Parasitoid host plant A. constricra

Host Brucbidac

A. greggii

Mimasestes ucaeiestrs N. amicus Slator pruinim4s -

A. hay&i Bentham Aeschynomene americana

Ac~th~~e~~s

Anoda c&rata (L.) Schlccht

hfeibomeus sp. Acanrhoscelides anodilus Johnson

Calopogmtium mucunoiaks

Chamaecrista romndifolia Greene Dolea aK submonlana D. formosa

_ D. ieporina Ait. ex Hem& D. rincloria (T. S. Brandog.) var arenaria M. E. Fonts D. lomenfosa VBTfomentosa Desmanrhw virgatus Desmodiam incarmm D. neomexicammz Gray D. orb&&are Schlccht D. tor~uosum Heliocarpus palmeri

Ittdigofera ~ff~~Ieosa Mimosa aibido M. biuncijea M. dysocarpa M. pigra M. pm&a M. somnians H. & B. cx Willd. M. vellosiana Petalosremum candidum Willdenow PiUwcelIobium mangeme Prosopis wIuri?m Semza hirsata var h&a

S. occidentalis

S. pa&&a war p&da Sida rho~~o~~ Tephrosia cana Brandegee T. multifolio T. rhodanrha Brandegee T. viscioides Schlecht Triumfeet~acalderom’

T. Iappuia

W&a&a

amplissima (L.) R. E. Fries

Zarropis sp. 1 Sidn aggregaia Zalropis sp. 2 Asrragalus pycrtostachyus Gray Zarropis sp. 3 Mimosa biunci/era Rhynchosia longeracemosa R. minima

per&ax

Acanihoscelides puelius A. taboga Semdas ceiatus s. insmb&9 s. morosus Acamhoscelkies pertinax Ac~t~s~l~s per&ax A. pu&imus Acanthoscelides pwillimus Acanrhoscelkies compressicontis A. desmanrhi Acan~hoseeiiuizsp~~~ps~ Johnson A. schuber~ae Acanthoscelides cuernavaca Acanthoscelides desmodicola Johnson Acanthoscelides cornis Acan&wee&&s cordrfer A. dJicilis AcanrhoscelIdes chiricahuae Acanthascelides cbiricahuae Aca&wscelides &#iciHs A. ~~i~~a~ Acanrhoscelkks piceoap&aI~ Acanlhoscelides cordfer Merobruchus sp. Sear&s k&a& S. morasw S. rufomaeuiatas Sennius faIlax S. lebasi S. rufomaculatur Sent& auricoma.5 A~anth~ee~~s bre&pes Acanthosce#&s rufwittaws Acanthoscelides r@vittatus Acanthoscelides rufwiuarw AcanthoscelIdes pertincur Acanihosceti&s johnsoni A. sa&as A. triumfel~~ Acanrhoqiides johnsani A. lriumfertae

Lo&ion USA, AZ

USA, AZ PAN COLOM MEX, MICH PAN MEX, JAL HOND PAN MEX, JAL

MEX, MICH MEX, COL NEX, JAL USA, AZ MBX, JAL MEX, BCS MEX, NAY MEX, OAX MBX, SON MEX. QR PAN USA, AZ MEX, PUE MEX, SON MBX, GRO MEX, JAL MEX, MICH MEX, SON MEX, OAX MEX, VER USA, AZ USA, AZ HOND COLOM PAN MEX, GRO USA, AZ MEX, GRO U=%Az MEX, OAX

MEX, OAX

MEX, MICH HOND MEX, BCS PAN MEX, MICH MEX, MICH HOND

-

MEX. CHIS MEX, OAX PAN MEX, WC

-

MEX, SON

-

USA, CA

Acamhoscelides chiricabuae AcanthosceIIdes campeche A. paWscens A. ~~~a Acanrhoscelides j?mvscens A. sereki

USA, AZ MEX, CAMP

142

MARTIN HETZ and CLARENCE D. JOHNSON Table I-continued Host Bruchidac

Parasitoid host plant TORYMIDAE Microdontomerus anthonomi (Crawford) Astragalus douglasii Gray A. praelongus Sheldon A. praelongus var. lonchopus A. thurberi Calliandra eriophylla Cassia couesii Gray Desmanthtu covillei

D. uirgatur Indigofew st&ticosa

Acanthoscelides oureolus A. pullus Aconthoscelides nureolw Acanthoscelides mix&s Stator limbatus S. prtdnint4s Sennius morosus Acanthoscelides der$eldi A. dewtanthi Stator pndninus Aconthoscelides compressicornis A. desmanthi -

Location

USA, CA USA, AZ USA, AZ USA, AZ USA, AZ MEX, DGO MEX, SON

MEX, OAX MEX, SON

by De Luca (1965), Whitehead (1975) and Center and Johnson (1976). The last edition of the catalogue of Hymenoptera in America north of Mexico (Krombein et al., 1979) provides the most complete account of bruchid parasites and their hosts for that area. The most recent summary of the activities of bruchid parasites was by Steffan (1981). Because of the probability of the occurrence of more than one bruchid species per seed pod, the chances of an absolute parasite/host record are reduced. Thus, the primary concern to readers of data from rearings of field-collected material is the authenticity of host records. But, if one surveys published host records of bruchid parasites an extreme amount of variation in number of hosts per parasite is encountered. This has been noted several times (Cushman, 1911; Whitehead, 1975; Krombein et al., 1979; Steffan, 1981). Steffan (1981) gave a most dramatic account when he stated “In North America E. [Eupelmus] amicus Girlt. not only attacks five species of bruchids, but also parasitizes 7 Lepidoptera, 4 Coleoptera, excluding the bruchids, and 2 Hymenoptera”. He also notes that one species of Chalcididae attacks only one bruchid species. Because of the nature of the seed pod environment and the extreme polyphagy of several bruchid parasites already recorded, the concern over absolute host records in this environment need not be real. The seed pod (or seed) is an entity potentially housing a host for a parasite attuned to that entity. The parasite is not occupied with finding a particular bruchid species but rather finding any host (bruchid, weevil, etc.) within a known environment. Steffan (1981) reflects our findings in his statements, “When looking for a host (ovipositing) many parasitic Hymenoptera seem to respond more to a given environment than to the systematic position of the host “and” . . . there is no more host specificity, but a biotope specificity (the coconut palm frond), and a microbiotope specificity (the gallery bored in the leaf)“. We propose through a comprehensive association of parasites, host plants and host beetles that host specificity in parasites of bruchid beetles is relative to a particular environment, not a particular beetle. The parasite-bruchid-host plant associations in Table 1 are from the seed collections of Johnson during about 20 years of continuous field sampling of bruchid beetles and their host plants from the southwestern United States, Mexico, Central America, and a few collections in northern South America. After analysis of the data on parasites in Table 1, we have concluded that relatively few species of parasitic Hymenoptera exploit a large number of species of seed predators of seed-pod bearing plants. Plants that bear seed pods are a resource which includes desert and lowland scrub forests, mostly species of Leguminosae, but with some species of Malvaceae and Tiliaceae and their seed predators, bruchid beetles. Seeds of economic importance (e.g. Phase&s spp.) also are hosts to several bruchid species and their parasites. There are 22 species of hymenopterous parasites associated with 123 species of bruchids, which in turn, feed in seeds of 195 species of plants. The list of parasitic Hymenoptera known to attack Bruchidae from the southwestern United States to northern South America is presented in Table 1. The Hymenoptera are associated with the plants that provide the environment and the bruchids that are their hosts. This list is of importance because it is a first step in associating parasites with bruchids and their hosts over a broad geographical area, and publication is necessary for ecological studies currently under way.

Parasites of Bruchidae of North and Central America

143

~c~o~ledge~e~t~-We would like to give special thanks to E. E. Grisell, P. M. Marsh, and M. E. Schauff of the Systematic Entomology Laboratory, USDA, for identifications of the parasitoids; and we thank M. Johnson, N. Schmelzle, K. Appleby, and L. Holter for help in various aspects of the gathering and analysis of data. The research could not have been done without the support of USDA Grants 12-loll-9187~33), 12-14-1~~7~33), and NSF Grants DEB78-05962, DEB81-09995, BSR82-11763 to CDJ. The help of all is gratefully acknowledged. REFERENCES Center T. D. and Johnson C. D. (1976) Host plants and parasites of some Arizona seed feeding insects. Ann, ent, Sot. Am. 69, 195-201.

Cushman R. A. (1911) Notes on the host plants and parasites of some North American Bruchidae. J. .&on. Ent. 4,489-510. De Luca Y. (1965) Catalogue des metazoaires parasites et predateurs de bruchides (Coleoptera). J: stored Prod. Res. I, 51-98. Janzen D. H. (1969) Seed-eaters versus seed size, number, toxicity, and dispersal. Evolution 23, l-27. Janzen D. H. (1971) Escape of Cassia grand& L. beans from predators in time and space. &o/ogy 52, 964-979. Janzen D. H. (1980) Specificity of seed-attacking beetles in a Costa Rican deciduous forest. J. Ecol. 68, 929-952. Johnson C. D. (1970) Biosystematics of the Arizona, California, and Oregon Species of the Seed Beetle Genus Acanthoscefkies Schilsky (Coleoptera: Bruchidae). Univ. CuiiJ Pubf. Ent. 59, l-116. Johnson C. D. (198la) Seed beetle host specificity and the systematics of the Leguminosae (pp. 995-1027, f61 pp. microfiche). In Advances in Legume Systematics (Edited by R. M. Polhill and P. H. Raven). xvi + 1050 pp. (2 Parts). Royal Botanic Gardens, Kew. Johnson C. D. (1981b) Relations of Acanfhosceffdes with their plant hosts. In The Ecology ofBruchf& Atfacking Legumes (Pulses) (Edited by Labeyrie V.). Series Entomofogica, Vol. 19, xiv + 233 pp. W. Junk, The Hague. Johnson C. D. (1981~) Interactions between bruchid (Coleoptera) feeding guilds and behavioral patterns of pods of the Leguminosae. Enuir. Entomof. 10, 249-253. Johnson C. D. (1983a) Ecosystematics of Acanthoscefides (Coleoptera: Bruchidae) of Southern Mexico and Central America. Misc. Pubf. ent. Sot. Am. 56, l-370. Johnson C. D. (1983b) Handbook on seed insects of Prosopb Species. Ecology, Control, and Identz~cutfon of seed-infesting Insects of New World Prosopis (Leguminosae), 55 pp. The Food and Agriculture Organization of the United Nations. Johnson C. D. and Kingsolver J. M. (1973) A reoision of the Genus Sennius of North and Central America (Cofeoptera: Bruchidae), 135 pp. U.S. Dept. Agric. Tech. Bull. 1462. Johnson C. D. and Kingsolver J. M. (1976) Systematics of Stator ofNorth and Central America (Cofeoptera: Bruchidae), 101 pp. U.S. Dept. Agric. Tech. Bull. 1537. Kingsolver J. A?. (1986) A taxonomic study of the genus Afgarobius (Coleoptera: Bruchidae). Entomography 4, 109-136. Kingsolver J. M. and Johnson C. D. (1978) Systematics of the Genus Mimosestes (Cofeoptera: Bruchidae). 106 pp. U.S. Dept. Agric. Tech. Bull. 1590. Kingsolver J. M., Johnson C. D., Swier S. R. and Teran A. (1977) Chapter 6, Prosopfs fruits as a resource for invertebrates. Mesquite Its Biofogy in Two Desert Scrub Ecosystems. (Edited by Simpson B. B.) pp. 108-122. US/IBP Synthesis Series 4. Dowden, Hutchinson d Ross, Stroudsburg, Penn. Kingsolver J. M. and ~itehead D. R. (1974a) Biosystematics of Central American species of Ctenocofum, a new genus of seed beetles (Coleoptera: Bruchidae). Froc. Eiof. Sot. Wash. 87, 283-312. Kingsolver J. M. and Whitehead D. R. (1974b) Classification and comparative biology of the seed beetle genus Caryedes Hummel (Coleoptera: Bruchidae). Trans. Am. ent. Sot. 100, 341-436. Kingsolver J. M. and Whitehead D. R. (1976) The North and Central American species of Meibomeur (Coleoptera: Bruchidae: Bruchinae). U.S. Dept. Agric. Tech. Buff. 1523. 54 pp. Krombcin K. V., Hurd P. D. Jr, Smith D. R. and Burks B. D. (1979) Catafogue of ~y~nopteru in America north of Meuieo. 3 Vols. l-2735. Steffan J. R. (1981) The parasites of bruchids, pp. 223-229. In The Ecology ofBruchfdF Attacking Legumes (Pulses). (Edited by Labeyrje V.) Series ~ntomoiogiea Vol. 19, pp. 223-229. W. Junk, The Hague. Whitehead D. R. (1975) Parasitic Hymenoptera associated with infested fruits in Costa Rica. J. Wash. Acad. Sci. 65, 108-l 16.

Whitehead D. R. and Kingsolver J. M. (1975a) Beetles and wasps associated with Cussia bzjfora L. (Cae~lpiniac~e) fruits in Costa Rica, with a new speices of Sennius (Coleoptera: Bruchidae). J. Wash. Acad. Sci. 65, 154-1.57. Whitehead D. R. and Kingsolver J. M. (1975b) Megasennius, a new genus for Acanzhoscelides muricatus (Sharp) (Coleoptera: Bruchidae), a seed predator of Cassti grandis L. (Caes~~nia~ae) in Central America. Proc. ent. Sot. Wash. 77, 460-465.

Whitehead D. R. and Kingsolver J. M. (197.5~)Biosystematics of the North and Central American Species of Gibbobruchus (Coleoptera: Bruchidae: Bruchinae). Trans. Am. ent. Sot. 101, 167-225.