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PAN-PACIFIC ENTOMOLOGIST

74(1): 1-11, (1998)

LARVAL FOODPLANTS OF EULINI

(LEPIDOPTERA: TORTRICIDAE):

FROM ABIES TO VITIS

JOHN W. BROWN! AND STEVEN PASSOA2

1Systematic Entomology Laboratory, PSI

Agricultural Research Service

U.S. Department of Agriculture

% National Museum of Natural History, MRC 168

Washington, DC 20560

U.S. Department of Agriculture

Animal and Plant Health Inspection Service, PPQ

The Ohio State University

Museum of Biological Diversity

Columbus, OH 43212

Abstract—A list of larval foodplants for members of the tortricid tribe Euliini was compiled

based on a literature search, label data of museum specimens, and the U.S. Department of

Agriculture pest interception database. Of the 70 genera included in the tribe, larval foodplants

are known for one or more species of only 11 genera. Several polyphagous genera (e.g., Proeulia,

Chileula, Bonagota) include pests of cultivated plants, particularly in Chile, Argentina, and

Uruguay. Foodplant records for most Euliini indicate polyphagous feeding habits at both the

generic and species levels. Only a few genera deviate from this pattern: Apolychrosis and Apo-

tomops are restricted to Coniferae, and Anopina and Dorithia appear to be leaf-litter feeders.

Although most Tortricinae are leaf-rollers, many Euliini feed externally on the surface of fruit

and sometimes bore into foodplant tissue.

Key Words.—lInsecta, Lepidoptera, Tortricidae, Euliini, pest, foodplant, Neotropical, Proeulia,

Citrus, Vitis, Prunus.

The tortricid tribe Euliini includes more than 340 described species distributed

primarily in the New World tropics. Among its members are pests of citrus,

grapes, kiwi, stone fruits, pines, and numerous other cultivated and ornamental

plants. As is true of most neotropical tortricids, knowledge of the life histories of

most Euliini is lacking.

We compiled records of larval foodplants of Euliini from three sources. 1) Six

literature databases were searched, including AGRICOLA (1970-—June 1997),

CRIS/ICA (March 1997), Biological Abstracts (1980—June 1997), CAB Abstracts

(1972—April 1997), Zoological Record (1978—June 1997), and AGRIS (1975—May

1997). 2) Specimens were examined for foodplant data at the National Museum

of Natural History, Smithsonian Institution (USNM) and the Essig Museum of

Entomology, University of California, Berkeley (UCB). 3) The U.S. Department

of Agriculture (USDA), Animal and Plant Health Inspection Service (APHIS)

pest interception database was queried. The latter includes records of taxa inter-

cepted at U.S. ports and/or sent to taxonomists (e.g., ARS specialists or APHIS

identifiers) for determination. These efforts represent a fairly comprehensive re-

view of available information and resulted in records of larval foodplants for one

or more species of 11 of the 70 genera included in the tribe.

The term ‘“foodplant’” refers to any plant that larvae of Euliini have been

2 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

discovered feeding upon in the field. The first mention of each common name of

a plant species is accompanied by its scientific name, author, and family; there-

after, only the common name is given. Latin and common names of plants and

familial assignments follow Brako, Rossman & Farr (1995). Table 1 lists all food-

plants we found arranged in alphabetical order by genus and then by species,

along with their associated euliine hervibore(s).

Ano pina Obraztsov

Data from rearing studies (J. A. Powell, unpubl.) suggest that larvae of Anopina

triangulana (Kearfott) feed on leaf litter. This conclusion is based on thorough

collections of larval microlepidoptera in California in which larvae of Anopina

are absent on living plant material at locations where adults are relatively com-

mon. Powell (1964) successfully reared this species from egghatch to maturity on

willow (Salix sp.; Salicaceae), and more recently on dead leaves and synthetic

medium (codling moth diet). The only record of a field collected larva is found

on a genitalic slide of A. ednana (Kearfott) from Bar Harbor, Maine, bearing the

data “‘specimens reared from white birch” (Betula populifolia Marsh.; Betulaceae)

(USNM).

Apolychrosis Amsel

In his description of A. schwardtfegeri, Amsel (1962) listed Pinus ayacahuite

Ehrenb. and P. tenuifolia Benth. (which is a synonym of P. maximinoi H. E.

Moore) (Pinaceae) as larval foodplants of the species in Guatemala. Pogue (1986)

cited Pseudotsuga macrolepis Flous and Abies religiosa Schl. & Cham. (Pinaceae)

as foodplants of A. ferruginus Pogue in Mexico; Pinus leiophylla Schiede &

Deppe and Cupressus benthamii (Klotzsch) Maters (Cupressaceae) as foodplants

of A. ambogonium Pogue in Mexico; Pinus rudis Endl. and P. chiapensis (Mar-

tinez) Andresen as foodplants of A. candidus Pogue in Mexico; and Abies reli-

giosa, Pinus montezumae Lamb., P. ayacahuite, P. rudis, P. hartwegii Lindl., and

Abies sp. as foodplants of A. synchisis Pogue in Mexico. Larvae were reared

primarily from second-year cones.

Apotomops Powell

Prentice (1965) indicates that Apotomops wellingtoniana (Kearfott) is a “‘soli-

tary defoliator’’ in southern Canada (British Columbia to Nova Scotia). He lists

silver fir (Abies amabilis Douglas ex Forbes; Pinaceae) (4 records), western hem-

lock (T'suga heterophylla (Raph.) Sarg.; Pinaceae) (4 records), mountain hemlock

(Tsuga mertensiana (Bong.) Carriére; Pinaceae) (1 record), and spruce (Picea sp.;

Pinaceae) (1 record) as foodplants for this species.

Bonagota Razowski

Label data of the holotype of Bonagota cranaodes (Meyrick) from Argentina

indicate that it was reared from Mimosa (Fabaceae) (Clarke 1963). Specimen label

data (USNM) indicate that B. cranaodes is a general feeder in Uruguay, having

been reared from acacia (Acacia longifolia (Andr.) Willd.; Fabaceae), alfalfa

(Medicago sativa L.; Fabaceae), pear (Pyrus communis L.; Rosaceae), Allophyllus

edulis (Camb.) Radl. (Sapindaceae), Japanese honeysuckle (Lonicera japonica

Thunb.; Caprifoliaceae), pepper-tree (Schinus longifolia (Lindl.) Speg.; Anacar-

Table 1. Eulitine Species by Host Plant. Plants arranged in alphabetic order (USNM = National Museum of Natural History, UCB = University of California,

Berkeley).

Plant species

Abies sp.

Abies religiosa

Abies amabilis

Acacia longifolia

Acer pseudoplatanus

Actinidia deliciosa

Allophyllus edulis

Alnus sp.

Alnus rubra

Alvoradoa sp.

Aristolochia chilensis

Austrocedrus chilensis

Baccharis spicata

Betula sp.

Betula populifolia

Caesalpinia paraguariensis

Citrus sp.

Citrus limon

Citrus paradisi

Citrus sinensis

Corylus sp.

Cupressus benthami

Moth species

Apolychrosis synchisis

Apolychrosis ferruginus

Apolychrosis synchisis

Apotomo ps wellingtoniana

Bonagota cranaodes

Proeulia chrysopteris

Proeulia chryso pteris

Proeulia auraria

Bonagota cranaodes

Eulia ministrana

Cuproxena minimana

Proeulia auraria

Chileulia stalactitis

Clarkeulia nr. virga

Eulia ministrana

Anopina ednana

Proeulia chryso pteris

Deltinea costalimai

Bonagota cranaodes/salubricola

Bonagota cranaodes

Chileulia stalactitis

Clarkeulia bourquini

Clarkeulia dimor pha

Chileulia stalactitis

Proeulia auraria

Proeulia chrysopteris

Eulia ministrana

Apolychrosis ambogonium

Moth distribution

Mexico

Canada/W. United States

Argentina/Chile

Holarctic

Central American/Florida

Argentina/Chile

Uruguay

Holarctic

SE Canada, NE United States

Argentina/Chile

Argentina

Argentina/Chile/Uruguay

Argentina/Chile

Argentina/Brazil

Argentina/Chile

Holarctic

Mexico

Source

Pogue 1986

Prentice 1965

USNM specimen

Gonzales 1990

Gonzales 1986, 1989, 1990; Pucat 1994

Pucat 1994

USNM specimen

Bradley, Tremewan & Smith 1973,

USNM specimen, Ferguson 1975

Prentice 1965

Brown & Powell 1991

Obraztsov 1964

Pastrana 1997

USNM specimen

Bradley, Tremewan & Smith 1973, Pren-

tice 1965

USNM specimen

Pastrana 1997

Pastrana 1961

Biezanko et al. 1974, Pastrana 1997

USNM specimen

Powell 1986

Pastrana 1997

Powell 1986

Gonzales 1990, Pucat 1994

Bradley, Tremewan & Smith 1973

Pogue 1986

8661

SINV'Td LSOH INITTINA :-VOSSVd 2 NMOUT

Table 1. Continued.

Plant species

Diospyros sp.

Epilobium sp.

Fagus sp.

Fraxinus sp.

Geoffroea decorticans

Gladiolus communis

Ipomoea mutabils

Ligustrum lucidum

Ligustrum sinense

Linum usitatissimum

Lonicera japonica

Malus domestica

Medicago sativa

Mentha rotundafolia

Mimosa sp.

Myrciaria jaboticaba

Myrtaceae shrubs

Phytolacca dioica

Picea sp.

Pinus spp.

Pinus ayacahuite

Pinus chiapensis

Pinus hartwegii

Pinus leiophylla

Pinus montezumae

Moth species

Proeulia chryso pteris

Eulia ministrana

Bonagota cranaodes/salubricola

Clarkeulia excer ptana

Bonagota cranaodes/salubricola

Clarkeulia bourquini

Clarkeulia excer ptana

Bonagota cranaodes

Bonagota cranaodes/salubricola

Clarkeulia bourquini

Proeulia chryso pteris

Proeulia auraria

Bonagota cranaodes

Clarkeulia excer ptana

Clarkeulia bourquini

Bonagota cranaodes

Clarkeulia dimor pha

Proeulia auraria

Clarkeulia dimor pha

Apotomo ps wellingtoniana

Proeulia chryso pteris

Apolychrosis schwardtfegeri

Apolychrosis synchisis

Apolychrosis candidus

Apolychrosis synchisis

Apolychrosis ambogonium

Apolychrosis synchisis

Moth distribution

Chile

Holarctic

Argentina/Chile

Argentina/Brazil

Argentina/Chile

Argentina/Brazil

Argentina/Chile

Argentina/Brazil

Argentina/Chile

Argentina/Brazil

Argentina/Chile

Argentina/Brazil

Chile

Argentina/Brazil

Canada/W. United States

Argentina

Guatemala

Mexico

Source

Pucat 1994

Benander 1929

Bradley, Tremewan & Smith 1973

Pastrana 1997

USNM specimen

Pastrana 1997

Gonzales 1990, Pucat 1994

Gonzales 1990

USNM specimen

Pastrana 1997

UCB specimen

Clarke 1963

Pastrana 1997

Pucat 1994

Pastrana 1997

Prentice 1965

Pastrana 1997

Amsel 1962

Pogue 1986

LSIDO'TOWOLNA OIIOVd-NVd AHL

(rl TOA

Table 1. Continued.

Plant species

Pinus radiata

Pinus rudis

Pinus tenuifolia

Platanus orientalis

Polygonum sp.

Prosopis sp.

Prunus sp.

Prunus armeniaca

Prunum avium

Prunus domestica

Prunus persica

Pseudotsuga macrolepis

Pyrus communis

Quercus sp.

Rapanea lorentziana

Rhamnus sp.

Moth species

Proeulia chryso pteris

Apolychrosis candidus

Apolychrosis synchisis

Apolychrosis schwardtfegeri

Bonagota cranaodes/salubricola

Proeulia chryso pteris

Bonagota cranaodes/salubricola

Chileulia stalactitis

Eulia ministrana

Proeulia chryso pteris

Proeulia auraria

Chileulia stalactitis

Proeulia auraria

Eulia ministrana

Bonagota cranaodes/salubricola

Chileulia stalactitis

Clarkeulia bourquini

Proeulia auraria

Proeulia chryso pteris

Apolychrosis ferruginus

Proeulia chryso pteris

Proeulia auraria

Bonagota cranaodes

Dorithia semicirculana

Dorithia trigonana

Dorithia occidentana

Eulia ministrana

Bonagota cranaodes/salubricola

Clarkeulia dimor pha

Eulia ministrana

Moth distribution

Chile

Mexico

Guatemala

Argentina/Chile

Holarctic

Argentina/Chile

Chile

Argentina/Chile

Chile

Holarctic

Argentina/Chile

Argentina/Brazil

Argentina/Chile

Mexico

Argentina/Chile

Rocky Mountains, USA

Arizona to Durango, Mexico

Durango, Mexico

Holarctic

Argentina/Chile

Argentina/Brazil

Holarctic

Source

Parra & Cerda 1992, Pucat 1994

Pogue 1986

Amsel 1962

Pastrana 1997

Gonzales 1990, Pucat 1994

Pastrana 1997

Powell 1986

Bradley, Tremewan & Smith 1973

Obrazstov 1964, Pucat 1994

Pucat 1994

Powell 1986

Pucat 1994

Ferguson 1975

Pastrana 1997

Powell 1986

Pastrana 1997

Gonzales 1990, Pucat 1994

Pogue 1986

Gonzales 1990, Pucat 1994

USNM specimen

Brown & Powell 1991

Bradley, Tremewan & Smith 1973

Pastrana 1997

Bradley, Tremewan & Smith 1973

8661

SINV'TId LSOH INITINA :VOSSVd 9 NMOU

Table 1. Continued.

Plant species

Rosa sp.

Rubus sp.

Rubus chamaemorus

Rubus idaeus

Rubus ulmifolius

Salix sp.

Salix humboltiana

Salix vitellina

Schinus longifolia

Schinus terebinthinfolius

Senecio bonariensis

Simmondsia chinensis

Solanum bonariense

Sorbus sp.

Taxodium distichum

Tilia sp.

Trifolium repens

Tsuga heterophylla

Tsuga mertensiana

Vaccinium myrtillus

Verbenaceae

Viburnum sp.

Vitis spp.

Vitis vinifera

Moth species

Eulia ministrana

Proeulia sp.

Eulia ministrana

Proeulia sp.

Ano pina triangulana

Eulia ministrana

Clarkeulia sp.

Bonagota cranaodes

Clarkeulia vir ga

Cuproxena auga

Deltinea costalima

Proeulia chrysopteris

Deltinea costalima

Eulia ministrana

Bonagota cranaodes/salubricola

Eulia ministrana

Clarkeulia bourquini

Clarkeulia nr. virga

Apotomo ps wellingtoniana

Eulia ministrana

Clarkeulia bourquini

Eulia ministrana

Proeulia chrysopteris

Bonagota cranaodes/salubricola

Chileulia stalactitis

Proeulia auraria

Proeulia triquetra

Moth distribution

Holarctic

Chile

Holarctic

Chile

California, USA

Holarctic

Uruguay

Argentina/Chile

Argentina/Brazil

Brazil

Argentina

Chile

Arentina

Holarctic

Argentina/Chile

Holarctic

Argentina/Brazil

Uruguay

Canada/W. United States

Holarctic

Argentina

Holarctic

Chile

Argentina/Chile

Source

Bradley, Tremewan & Smith 1973

Pucat 1994, USDA/APHIS database

Benander 1929

USDA/APHIS database

Powell 1964

Prentice 1965

USNM specimen

Brown & Powell 1991

Pastrana 1997

Quiroga et al. 1989, Pucat 1994

Pastrana 1961

Bradley, Tremewan & Smith 1973

Pastrana 1997

Bradley, Tremewan & Smith 1973

USNM specimen

Prentice 1965

Bradley, Tremewan & Smith 1973

Pastrana 1997

Ferguson 1975

Pucat 1994

Pastrana 1997

Powell 1986

Campos et al. 1981; Gonzales 1983,

1990; Pucat 1994

Gonzales 1983, 1990; Pucat 1994

LSIDO'TIONOLNA OIIOVd-NVd AHL

(Dl ‘TOA

1998 BROWN & PASSOA: EULIINI HOST PLANTS 7

diaceae), and grapefruit (Citrus paradisi Macfad.; Rutaceae). Biezanko et al.

(1974) record B. salubricola from lemon (Citrus limon (L.) Burm.; Rutaceae) in

Uruguay.

Pastrana (1997) lists the following as foodplants of Bonagota cranaodes/salu-

bricola (Meyrick) in Argentina: Japanese honeysuckle, morning-glory (Ipomoea

mutabilis Ker; Convolvulaceae), Geoffroea decorticans (Gill. ex Hook. & Am.)

Burkart (Fabaceae), Rapanea lorentziana Mez. (Myrsinaceae), privet (Ligustrum

lucidum Aiton; Oleaceae), Oriental plane-tree (Platanus orientalis L.; Platana-

ceae), knotweed (Polygonum sp.; Polygonaceae), apple (Malus domestica Borkh.;

Rosaceae), plum (Prunus domestica L.; Rosaceae), lemon, bald cypress (Taxo-

dium distichum (L.) Rich.; Taxodiaceae), and grapes (Vitis vinifera L.; Vitaceae).

Although considered distinct species by Razowski (1986), Bonagota salubricola

and B. cranaodes were treated as synonyms by Pastrana (1997), with no expla-

nation for this decision. Hence, it is uncertain to which species his foodplant

records refer. It is possible that the two names represent a single species—B.

salubricola (Meyrick) was described from Buenos Aires and the very similar B.

cranaodes (Meyrick) from Tigre, both in Argentina. However, according to Ra-

zowski (1990), the male genitalia on the slide of the type of B. salubricola are

incorrectly associated, belonging to a Eucosmini and not to the holotype, so com-

parisons of male genitalia with the type are meaningless.

Chileulia Powell

Powell (1986) indicated that larvae of Chileulia stalactitis (Meyrick) feed on

various fruit crops in Chile, including grapes, grapefruit, orange (Citrus sinensis

(L.) Osbeck; Rutaceae), plum leaves, apricot leaves (Prunus armeniaca Marsh.;

Rosaceae), and fruit of mesquite (Prosopis tamarungo Phil.; Fabaceae). Pastrana

(1997) added Austrocedrus chilensis (D. Don) Pic. Serm. & Bizzarri (Cupressa-

ceae) to the list of foodplants in Argentina.

Clarkeulia Razowski

Clarkeulia bourquini (Clarke) has been reared from white clover (Trifolium

repens L.; Fabaceae) (USNM) and Mentha rotundafolia (L.) Huds. (Lamiaceae)

(UCB) in Uruguay, and from alfalfa, Chinese privet (Ligustrum sinense Lout.;

Oleaceae), apple, plum, orange, and various Verbenaceae in Argentina (Pastrana

1997).

According to Pastrana (1997), Clarkeulia dimorpha (Clarke) is a pest of or-

namentals and citrus in Argentina, feeding on Phytolacca dioica L. (Phytolacca-

ceae), Rapanea lorentziana, Myrciaria (Myrtaceae), and orange. Many of the

same species are reported as foodplants in Brazil (d’ Aratijo et al. 1968).

Clarkeulia excerptana (Walker) is recorded from Gladiolus (Iridaceae), alfalfa,

and flax (Linum usitatissimum L.; Linaceae) (Pastrana 1997). Clarkeulia virga

(Clarke) has been reared from flowers of Brazilian pepper-tree (Schinus terebin-

thifolius Raddi; Anacardiaceae) (USNM). An apparently undescribed species (nr.

virga) from Uruguay (USNM) has been found on Baccharis spicata (Lam.) Bail-

lon (Asteraceae) and white clover; and an undetermined species from Uruguay

(USNM) was reared on willows (Salix vitellina Willd. and S. humboltiana Willd.;

Salicaceae).

8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Cuproxena Powell & Brown

Two species of Cuproxena have been reared from field-collected larvae: C.

auga (Razowski and Becker) from Brazilian pepper-tree in Minas Gerais, Brazil,

and C. minimana Brown from Alvoradoa sp. (Simaroubaceae) in southern Florida

(Brown & Powell 1991).

Deltinea Pastrana

Deltinea costalimai Pastrana is reported to bore in the stems or shoots of night-

shade (Solanum bonariense L.; Solanaceae) and to feed on the leaves and flowers

of Citrus sp. (Rutaceae) in Argentina (Pastrana 1961). Pastrana (1997) also cites

Senecio bonariensis Hook. & Am. (Asteraceae) as a foodplant in Argentina.

Dorithia Powell

As with Anopina, circumstantial evidence suggests that Dorithia larvae may

feed on leaf litter (Brown & Powell 1991). Despite extensive searches of potential

host foliage, no larvae have been discovered in the field. Larvae of three species

(i.e., D. semicirculana (Fernald), D. trigonana Brown & Obraztsov, & D. occi-

dentana Brown) have been reared successfully in the laboratory from egghatch

to adults on fresh leaves of Quercus spp. (Fagaceae); the overall distribution of

Dorithia corresponds to the distribution of Quercus from the Rocky Mountains

south to Costa Rica. Although the distribution of Dorithia semicirculana closely

parallels that of Quercus gambellii Nutt., and larvae were successfully reared in

the laboratory on fresh leaves of Quercus lobata Née, which is in the same section

as Q. gambellii (i.e., the white oaks), larvae were never discovered on Q. gam-

bellii in the field.

Eulia Hiibner

Larvae of the holarctic E. ministrana (L.) are polyphagous leaf rollers, recorded

from numerous plant families. Bradley, Tremewan & Smith (1973) list Fraxinus

(Oleaceae), Rhamnus (Rhamnaceae), Sorbus, Prunus (both Rosaceae), Betula, Al-

nus, Corylus (all Betulaceae), and Vaccinium myrtillus L. (Ericaceae) as hosts in

Great Britain, and Rosa (Rosaceae), Fagus, Quercus (both Fagaceae), and Tilia

(Tiliaceae) from the European continent. Benander (1929) reported Epilobium

(Onagraceae) and Rubus chamaemorus L. (Rosaceae) also from Europe.

In Canada, Prentice (1965) cites rearings of Eulia ministrana on red alder

(Alnus rubra Bong.; Betulaceae) (7 records), birch (Betula sp.; Betulaceae) (1

record), and willow (Salix sp.; Salicaceae) (1 record); and Ferguson (1975) cites

cherry (Prunus avium L.) (1 record), alder (Alnus sp.) (2 records), and Viburnum

sp. (Caprifoliaceae) (1 record).

Proeulia Clarke

The genus Proeulia includes 22 described species, nearly all of which are

confined to Chile (Razowski 1995). Several are pests of agricultural plants; con-

sequently, there is considerable economic literature on the genus. Larvae of

Proeulia are polyphagous leaf rollers, also feeding on the surface of fruit (Gon-

zales 1981la, 1981b, 1990); the larvae of some species spend the winter protected

under loose bark (Gonzéles 1972).

1998 BROWN & PASSOA: EULIINI HOST PLANTS 9

Pear, orange, and grape are the main foodplants of P. auraria (Clarke) (Pucat

1994). The life history of this species on grape was detailed by Campos et al.

(1981) and on pear by Alvarez & Gonzales (1982). Other recorded foodplants

include plum, apricot, cherry, apple, Prunus, Platanus (Platanaceae), kiwi (Actin-

idia deliciosa (A. Chev.) C. S. Liang & A. R. Ferguson; Actinidiaceae), and

shrubs of Myrtaceae and Aristolochiaceae (Pucat 1994, Zhang 1994), including

Aristolochia chilensis Bridges ex Lindl. (Obraztsov 1964).

The primary foodplants of P. chrysopteris (Butler) in Chile are apricot, apple,

plum, peach (Prunus persica (L.) Batsch; Rosaceae), nectarine (Prunus persica

var. nucipersica (Suckow) C. K. Schneid.; Rosaceae), pear, and kiwi. Damage

and management of P. chrysopteris on kiwi are discussed in detail by Gonzales

(1986, 1989) and Gonzales & Curkovic (1994). Secondary foodplants of P. chry-

sopteris include Acer pseudoplatanus L. (Aceraceae), Oriental plane-tree, orange,

Diospyros sp. (Ebenaceae), and Vitis spp. (Pucat 1994). Potentially severe damage

to jojoba (Simmondsia chinensis (Link) C. K. Schneid.; Simmondsiaceae) is de-

scribed by Quiroga, Arretz & Araya (1989); and Parra & Cerda (1992) report

damage to Monterey pine (Pinus radiata D. Don; Pinaceae) by P. chrysopteris.

Pucat (1994) cites raspberry (Rubus sp.; Rosaceae) as a “‘rare’’ host; the USDA/

APHIS database includes three records of interceptions of Proeulia on Rubus,

and these records may refer to P. chysopteris. Pastrana (1997) records many of

the same hosts for this species in Argentina, with one additional foodplant: par-

adise-flower (Caesalpina paraguariensis (D. Parodi) Burkart; Fabaceae).

The only recorded foodplant for Proeulia triquetra Obraztsov is grape (Gon-

zAales 1983, 1990, Pucat 1994, Pastrana 1997), and this month is of secondary

economic importance as a grape pest (Gonzales 1990).

The genus Proeulia has been treated in a variety of unpublished USDA/APHIS

circulars, keys, and bulletins, and this body of literature is reponsible for at least

three mistakes that require rectification. 1) In early accounts of pests from Chile,

most species of Proeulia were treated as Eulia. 2) A record of Proeulia from

China is almost certainly based on a misdetermination. 3) A reference to Proeulia

chilense is an error—there is no such species. It is possible that this mistake

resulted from Obraztsov’s (1964) review of the genus in which he identified A.

[Aristolochia] chilense as a larval foodplant of Proeulia in Chile.

SUMMARY AND CONCLUSIONS

Host plant records indicate polyphagous feeding habits for most Euliini at both

the generic and species levels. Only a few genera deviate from this pattern: Apo-

lychrosis and Apotomops are restricted to Pinaceae, and Anopina and Dorithia

appear to be leaf-litter feeders. In contrast to the larvae of most Tortricinae, which

are leaf-rollers on living plant material (MacKay 1962, Powell 1980), many Eu-

liini feed externally on the surface of fruit and sometimes bore into host plant

tissue; species of two genera are believed to feed in leaf-litter.

Larval foodplants are known for one or more representatives of 11 genera in

the tribe; several genera include pests of cultivated and ornamental plants. Doc-

umented hosts encompass 36 vascular plant families: Aceraceae (1 euliine genus),

Actinidiaceae (1), Anacardiaceae (3), Aristolochiaceae (1), Asteraceae (2), Betu-

laceae (3), Caprifoliaceae (2), Convolvulaceae (1), Cupressaceae (2), Ebenaceae

(1), Ericaceae (1), Fabaceae (4), Fagaceae (2), Iridaceae (1), Lamiaceae (1), Lin-

10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

aceae (1), Myrsinaceae (2), Myrtaceae (2), Oleaceae (3), Onagraceae (1), Phyto-

laccaceae (1), Pinaceae (3), Platanaceae (2), Polygonaceae (1), Rhamnaceae (1),

Rosaceae (5), Rutaceae (5), Salicaceae (3), Sapindaceae (1), Simaroubaceae (1),

Simmondsiaceae (1), Solanaceae (1), Taxodiaceae (1) Tiliaceae (1), Verbenaceae

(1), and Vitaceae (3). Reflecting a sampling bias, Rosaceae and Rutaceae each

support five genera of euliine herbivores, more than any other plant family. These

two plant families include a large percentage of the commercially cultivated, fruit-

bearing trees in the New World tropics (e.g., Prunus and Citrus). With the ex-

ception of the use of conifers by Apolychrosis and Apotomops, and rarely by

Proeulia and Bonagota, and the infrequent use of Iridaceae by Clarkeulia, most

euliine genera appear to be restricted to dicotylendous plants; few other trends

are evident.

ACKNOWLEDGMENT

We thank the following for helpful comments on the manuscript and/or iden-

tification of pertinent literature: Jerry Powell, University of California, Berkeley;

Richard Brown, Mississippi State University; James Pakaluk and David Smith,

USDA, Systematic Entomology Laboratory, Washington D.C.; and two anony-

mous reviewers, and Susan Chapman, National Agricultural Library, Beltsville,

Maryland, for the database searches.

LITERATURE CITED

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(Clarke) [Pyrus communis]. Revista Fruticola, 3: 75—80.

Amsel, H. G. 1962. Neue microlepidoptera aus Guatemala. Sond. Z. Angew. Entomol. Bd., 49: 392—

398.

Benander, P. 1929. Zur Biologie einiger Kleinschmetterlinge. Entomol. Tidskr., 50: 123-145.

Biezanko, C. M., A. Ruffinelli & D. Link. 1974. Host-plants and any other foods of the lepidopterous

larvae of Uruguay. Rev. Centro Cienc. Rurais, 4: 107-148.

Bradley, J. PR, W. G. Tremewan & A. Smith. 1973. British tortricoid moths. Royal Entomol. Soc.

London.

Brako, L., A. Y. Rossman & D. F Farr. 1995. Scientific and common names of 7000 vascular plants

in the United States. APS Press, St. Paul, MN.

Brown, J. W. & J. A. Powell. 1991. Systematics of the Chrysoxena group of genera (Lepidoptera:

Tortricidae: Euliini). Univ. Calif. Publ. Entomol., 11: 1-87.

Campos, S. L., M. A. Faccin, L. N. Echerverria & R. L. Sazo. 1981. Agric. Tenica, 41: 249-256.

Clarke, J. EF G. 1963. Catalogue of the Type Specimens of Microlepidoptera in the British Museum

(Natural History) described by Edward Meyrick, Vol 4. Trustees of the British Museum, Lon-

don.

d’Aratijo, A. G., C. R. Gongalves, D. M. Galvao, A. J. L. Gongaves, J. Gomes, M. N. Silva & L.

Simoni. 1968. Quarto cat4logo dos insetos que vivem nas plantas do Brasil. Parte II-1° Tomo.

Insetos, hospedeiros e inimigos naturais. Dept. Defesa Inspec. Agropec., Lab. Centr. Patolog.

Veg., Rio de Janeiro, Brasil.

Ferguson, D. 1975. Host records for Lepidoptera reared in eastern North America. USDA, ARS,

Technical Bull., No. 1521: 1-49.

Gonzales, R. H. 1972. Outbreaks and new records. FAO Plant Protection Bull., 20: 89-91.

Gonzales, R.H. 1981a. Situacion fitosanitaria del manzano y peral. Temporada 1980-1981: Insectos

y acaros. Revista Fruticola, 2: 35-43.

Gonzales, R. H. 1981b. Dos nuevas plagas del peral en la VII Region: Proeulia auraria (Clarke) y

Psylla piricola Forst. Revista Fruticola, 2: 15-18.

Gonzales, R. H. 1983. Manejo de plagas de la vid. Univ. Chile, Publ. Cienc. Agric., 13: 1-115.

Gonzales, R. H. 1986. Plagas del kiwi en Chile. Revista Fruticola, 7: 13-27.

1998 BROWN & PASSOA: EULIINI HOST PLANTS 11

Gonzales, R. H. 1989. Manejo de plagas del kiwi en Chile: 1. Degradacion de residuos de los

insecticidas chlorpyrifos y phosmet. Revista Fruticola, 10: 35-43.

Gonzales, R. H. 1990. Insectos y acaros de importancia agricola y cuarentenaria en Chile. Univ.

Chile, Santiago.

Gonzales, R. H. & S. T: Curkovic. 1994. Manejo de plagas y degradacion de residuos de los pesticidas

en kiwi. Revista Fruticola, 15: 5—20.

MacKay, M. R. 1962. Larvae of North American Tortricinae. Canad. Entomol. Suppl., 28: 1-182.

Obraztsov, N. S. 1964. Neotropical microlepidoptera, V. Synopsis of the species of the genus Proeulia

from central Chile (Lepidoptera: Tortricidae). Proc. U.S. Natl. Mus., 16: 183-196.

Parra, L. E. & L. A. Cerda. 1992. Presencia de Proeulia chrysopteris (Butler) (Lepidoptera: Tortri-

cidae) sobre Pinus radiata D. Don. Bosque, 12: 61-63.

Pastana, J. A. 1961. Una nueva Tortricidae (Lep.) plaga potencial de cultivos citricos. Revista Invest.

Agric., 15: 343-348.

Pastrana, J. A. 1997. Los lepidopteros Argentinos, sus plantas hospedadoras y otros sustratos ali-

menticios (Bachmann, A. O., ed.). Univ. Chile, Santiago. In press.

Pogue, M. 1986. A revision of the Neotropical genus Apolychrosis Amsel with descriptions of new

species, pp. 19-28. Jn Cibrian-Tovar, D., B. Ebel, H. Yates & J. Méndez-Montiel (eds.), Cone

and seed insects of the Mexican conifers. Southeastern For. Exp. Sta., USDA.

Powell, J. A. 1964. Biological and taxonomic studies on tortricine moths, with reference to the species

in California. Univ. Calif. Publ. Entomol., 32: 1-307.

Powell, J. A. 1980. Evolution of larval food preferences in Microlepidoptera. Ann. Rev. Entomol.,

25: 133-159.

Powell, J. A. 1986. Synopsis of the classification of Neotropical Tortricinae, with descriptions of new

genera and species. Pan-Pacif. Entomol., 62: 372-398.

Prentice, R. M. 1965. Forest Lepidoptera of Canada recorded by the Forest Insect Survey, vol. 4:

546-840. Dept. For. Canada.

Pucat, A. M. 1994. Fruit leaf folders, Proeulia spp. Agriculture and Agri-food Canada, Plant Health

Risk Assessment Unit.

Quiroga, D., V. Arretz & J. E. Araya. 1989. Chewing insects on jojoba, Simmondsia chinensis (Link)

Schneider, in the north-central and central regions of Chile, and characterization of damage.

FAO Plant Protection Bull., 3: 121-124.

Razowski, J. 1986. Descriptions of new Neotropical genera of Archipini and rectification of the

Deltinea problem (Lepidoptera: Tortricidae). Sci. Nat, 52:21-25.

Razowksi, J. 1990. Comments on the catalogue of Meyrick types of Tortricidae (Lepidoptera) in the

Museum of Vienna with descriptions of new genera. Polska Akad. Nauk, Annls. Zool., 43:

395-404.

Razowksi, J. 1995. Proeulia Clarke, 1962, the western Neotropical Tortricidae genus (Lepidoptera),

with descriptions of five new species and two allied genera. Acta Zool. Cracov., 38: 271-238.

Zhang, B. C. 1994. Index of economically important Lepidoptera. CAB International.

Received 29 Jul 1997; Accepted 13 Nov 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 12-26, (1998)

REVIEW OF THE GENUS SPHAENOTHECUS DUPONT

(COLEOPTERA: CERAMBYCIDAE)

JOHN A. CHEMSAK! AND FELIPE A. NOGUERA?2

‘Essig Museum of Entomology, University of California, Berkeley

201 Wellman Hall, Berkeley, California 94720-3112, USA

*Estacion de Biologia Chamela, IBUNAM

Apartado Postal 21, San Patricio, Jalisco 48980, MEXICO

Abstract.—The trachyderine genus Sphaenothecus Dupont is reviewed. Sphaenothecus funebris

Bates is transferred to the genus Zalophia Casey and Z. spissicornis Casey is synonymized with

Z. funebris, NEW COMBINATION, Sphaenothecus cribellatus Bates and S. luteicollis Bates are

placed into the genus Ischnocnemis Thomson, NEW COMBINATION. Taranomis Casey is syn-

onymized with Sphaenothecus and Taranomis pallida Schaeffer is placed into Perarthrus

LeConte, NEW COMBINATION. Taranomis cylindricollis Casey is synonymized under Sphaen-

othecus bivittatus Dupont.

As here defined, Sphaenothecus consists of the following seven species: S. argenteus Bates,

Mexico to Guatemala; S. bivittatus Dupont, United States to Honduras; S. picticornis Bates,

Mexico; S. maccartyi NEW SPECIES, Mexico; S. toledoi NEW SPECIES, Mexico-Honduras;

S. facetus NEW SPECIES Guatemala to Costa Rica; S. trilineatus Dupont, Texas and Mexico.

Key Words——Cerambycidae, Sphaenothecus, taxonomic revision, new species, key, distribution,

activity period, hosts.

The trachyderine genus Sphaenothecus Dupont is currently comprised of seven

Mexican and Central American species. Examination of this group indicates that

the presently included species are not congeneric. Zalophia spissicornis Casey 1s

synonymous with S. funebris Bates (NEW SYNONYMY) and funebris enters into

the genus Zalophia Casey (NEW COMBINATION). S. cribellatus Bates and S.

luteicollis Bates are placed into the genus Jschnocnemis Thomson (NEW COM-

BINATION) for the time being. A study of the genus will be necessary to deter-

mine the validity of the reassignments.

Generically we can find no differences between Sphaenothecus and Taranomis

Casey. The species pallida Schaeffer from Baja California is placed into Perar-

thrus LeConte (NEW COMBINATION) and appears to be distinct from the other

two species in that genus.

The genus Sphaenothecus as presently defined consists of seven species, three

of which are previously undescribed.

GENUS SPHAENOTHECUS DUPONT

Sphaenothecus Dupont, 1838: 55; Thomson, 1860: 209; Lacordaire, 1869: 184;

LeConte, 1873: 316; LeConte and Horn, 1883: 301; Leng, 1886: 60; Casey,

1912: 333; Bradley, 1930: 241.

Sphoenothecus; Guerin, 1839: 250; Thomson, 1864: 205.

Sphenothecus; White, 1853: 85; Gemminger and Harold, 1873: 2972; Bates, 1880:

84.

Taranomis Casey, 1912: 333; Linsley, 1962: 95., Type species: Sphaenothecus

bivittatus Dupont, by original designation. NEW SYNONYM Y.

Page charges partially offset by a grant from the C. P. Alexander fund.

1998 CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS 13

Ischnocnemis; LeConte, 1873: 316; LeConte and Horn, 1883: 301; Leng, 1886: 61.

Form moderate sized to large, tapering posteriorly. Head small; palpi short, apical segments short;

mandibles arcuate, acute at apices; eyes finely faceted, deeply emarginate, upper lobes widely sepa-

rated; antennal tubercles divergent, often prominently elevated; antennae slender, often robust in fe-

males, often 12-segmented, much longer then body in males, slightly longer than body and often

subserrate in females, scape usually robust, shorter than segment 3, 4 subequal to 3, segment 3 often

stout, densely clothed with coarse setae. Pronotum convex, sides rounded or with small lateral tuber-

cles; apex narrower than base, disk lacking distinct calluses; prosternum with intercoxal process not

protuberant, abruptly declivous behind, coxal cavities wide open behind; mesosternum with intercoxal

process broad, prominently protruding above coxae, abruptly declivous; metasternum with episternum

broad, subparallel. Elytra tapering posteriorly; disk with longitudinal eburneous costae or longitudinal

pubescent vittae; apices sinuate truncate to bi-emarginate. Legs slender; hind femora often linear,

arcuate; middle femora carinate apically; posterior tarsi slender, rather short, apical segment cleft

almost to base. Abdomen normally segmented.

Type species.—Sphaenothecus tomentosus Dupont, Thomson designation, 1864

(= Sphaenothecus trilineatus Dupont).

This genus may be recognized by the tapering form of the pronotum and elytra,

longitudinally costate or vittate elytra, non protuberant, abruptly declivous pro-

sternal process and by the elevated, declivous mesosternal process.

Seven species are presently known.

KEY TO THE SPECIES OF SPHAENOTHECUS

i. Elytra with elevated, longitudinal, eburneous or dark costae, at least

Heh tad ymMiGdiles:, |, 2 ae eR eM eee, Lets hata ee gS 2:

1h Elytra non-costate, longitudinal pale bands consisting of rows of ap-

pressedapubescemcer 2 e LN A Bee eens Kooks thet ie eaten dol 6

2(1). Pronotum with sides rounded, non-tuberculate ..................... 3

De Pronotum with small lateral tubercles; antennae of males 11-segmented;

elytra with vitta densely, minutely punctate, subopaque; pronotum

with two broad longitudinal bands of golden appressed pubescence.

Length, 14.5—22 mm. Western Mexico from Sinaloa to Guatemala

Be ta Aa Wig ING APR EIT AAS Daa een AEB ye argenteus Bates

3(2). Antennae stout, segment 3 almost as thick as scape, densely clothed

with long, coarse, subdepressed, blue-black setae; scutellum longer

CHAN DRO A Uh ees Fe eee ie a et Bh Se odes Neh RTRs Posen dod 4

Su Antennae slender, segment 3 distinctly narrower than scape, segments

not appreciably apically expanded in either sex; scutellum as broad

as long; longitudinal yellowish vittae of elytra broad; appendages

often partially or entirely reddish. Length, 9-15.5 mm. Southern Cal-

ifornia to Texas, Honduras and Baja California ..... bivittatus Dupont

4(3). Elytra feebly bilobed basally, lobes not extending onto sides of basal

margin of pronotum, yellowish longitudinal vittae broad, apices fee-

bly sinuate truncate or strongly dentate at outer angles ........... 5

4’. Elytra strongly lobed on each side of scutellum, longitudinal, yellowish

vittae, if present, narrow, apices strongly bi-emarginate. Antennae of

males 12-segmented, segment 11 of females almost divided. Form

rather slender. Length, 12—22 mm. Central Mexico .. picticornis Bates

5(4). Elytra with median dark vittae with punctures behind middle separated,

not opaque appearing, apices strongly dentate at outer angles. Hind

14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

tibiae moderately sinuate, dark suberect setae rather sparse. Mesoster-

nal process broadly rounded at apex. Antennae of males 12-seg-

mented. Length, 10-15 mm. Sinaloa, Jalisco and Chiapas ........

DEES PoE e esto date to ETRE EEE th eg pe eee maccartyti NEW SPECIES

oa Elytra with median dark vittae very finely, contiguously punctate be-

hind middle, appearing subopaque, apices with angles unarmed. Hind

tibiae strongly sinuate, dark, suberect setae dense. Mesosternal pro-

cess narrowly rounded at apex. Antennae of males 11-segmented.

Length, 16—22 mm. Oaxaca to Honduras .... toledoi NEW SPECIES

6(1'). Antennae with scape 2 X as long as broad, dorsally flattened and im-

pressed at base. Integument reddish-brown. Length 15—22 mm. Gua-

temala.to' Costa Rica™ 4:00.04 eee Se facetus NEW SPECIES

6’. Antennae with scape at least 2.5 X as long as broad, slightly flattened

dorsally at base but not impressed. Integument black. Length, 15—25

mm. Southern Texas to Veracruz, Sinaloa to Oaxaca .............

SPHAENOTHECUS ARGENTEUS BATES

(Pl. 2)

Sphenothecus argenteus Bates, 1880: 84.

Sphaenothecus argenteus; Aurivillius, 1912: 472; Chemsak and Noguera, 1995:

64.

Male.—Form moderate sized, tapering; integument black to reddish, elytra with two longitudinal,

yellowish, eburneous vittae on each side; pubescence dense, golden, appressed, fine, pale, suberect

and fine, appressed and dark. Head small; antennal tubercles prominently elevated, blunt at apices;

vertex with a linear, glabrous callus; pubescence dense, appressed, golden, long, erect, golden hairs

numerous; antennae slender, 11-segmented, extending at least 4 segments beyond elytra, scape cylin-

drical, slender, moderately flattened only slightly beneath at base, basal segments densely clothed with

subdepressed black setae, outer segments densely clothed with very fine, very short pubescence, third

segment only slightly narrower than scape, segment 11 appendiculate at apical one-third. Pronotum

broader than long, sides with a small tubercle behind middle; disk with median area convex, slightly

elevated, broadly impressed at middle at base; basal margin lobed at middle; punctures at middle fine,

arranged in irregular contiguous rows; pubescence dense, appressed, golden, forming 2 broad longi-

tudinal vittae at sides of middle; sides sparsely to densely clothed with long or short erect hairs; lateral

longitudinal dark vittae subopaque, finely, irregularly punctate; prosternum shallowly impressed, usu-

ally densely golden pubescent; meso- and metasternum medially glabrous, very finely, sparsely punc-

tate, sides densely golden pubescent. Elytra a little more than 2 X as long as broad, tapering apically;

basal margin broadly lobed; each elytron with a broad, longitudinal eburneous fascia near suture and

a narrow one near lateral margin, the 2 coalescing near apex; punctures on median black vittae

subcontiguous near base, punctures on outer black vittae very fine, dense, subopaque; pubescence

along suture dense behind scutellum, golden, appressed, arranged transversely, black areas with fine,

dark, appressed pubescence; apices sinuate, sutural angles dentate. Scutellum narrow, elongate, gla-

brous. Legs slender, hind femora not extending to apices of elytra; femora rather sparsely punctate

and pubescent; front tibiae arcuate, densely pubescent beneath, hind tibiae densely clothed with dark,

subdepressed setae; hind tarsi moderately broad, segment 1 shorter than two following together. Ab-

domen medially glabrous, sides densely clothed with golden appressed pubescence, long erect hairs

dense to sparse; last sternite shallowly emarginate at middle. Length, 15-22 mm.

Female.—Form similar, slightly more robust. Antennae slightly longer than body. Abdomen with

last sternite broadly rounded, narrowly, shallowly emarginate at apical margin. Length, 14.5-—21.5 mm.

Diagnosis.—This species is very distinctive by the broad, longitudinal bands

1998 CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS 15

of golden, appressed pubescence of the pronotum and by the eburneous longitu-

dinal vittae of the elytra.

Type Locality.—San Geronimo, Guatemala.

Range.—Western Mexico from Sinaloa to Guatemala.

Flight Period.—June to December.

Remarks.—The northernmost Mexican specimens have short, sparse, erect pu-

bescence on the pronotum while in Guatemala, this is long and dense. The un-

derside of the Guatemala material also has the long erect hairs much more nu-

merous and the pubescence of the middle of the prosternum much denser.

Adults have been collected on flowers of Croton.

Material Examined —MEXICO. CHIAPAS: 21 males, 6 females, 16 km W of Ocozocuatla, ‘‘El

Aguacero”’, 4-5 Aug 1994, 4-30 Sep 1994, 1-3 Oct 1994, 4-16 Nov 1994, V. H. Toledo. JALISCO:

11 males, 6 females, Estacion de Biologia, Chamela, 14—23 Oct 1986, Chemsak; 7 Dec 1983, S. H.

Bullock; 1 Sep 1985, R. Ayala; 13 Sep 1985, 22 Nov 1985, 25 Sep 1986, 27 Oct 1986, 25 Nov 1986,

FE A. Noguera. MICHOACAN: 1 female, La Huacana, 10 Oct 1988, G. Rodriguez. OAXACA: 1 female,

E] Camaron, 32 km (20 mi) E of Oaxaca, 21 Jul 1956, J. W. MacSwain; 3 males, 1 female, 89.6 km

(56 mi) NW of Tehuantepec, 27 Jul 1963, J. Doyen, W. Foster. SINALOA: 2 females, 4 km (2.5 mi)

N of Mazatlan, 10-11 Aug 1970, J. A. Chemsak. GUATEMALA. BAJA VERAPAZ: 5 males, 6 fe-

males, Las Limas, Mpio. Salama, Baja Verapaz, 1150 m, 16 Jun 1986, 8—28 Sep 1986, 16 Oct 1986,

7 Nov 1986, 18 Dec 1986. EL PROGRESO: 1 female, 12.8 km (8 mi) NE of El Progreso, 8 Jul 1965,

A. Raske, C. Slobodchikov; 1 male, Magdalena, Mpio. San Agustin, Acasaguastlan, 16 Jun 1989.

GUATEMALA: 1 male, San Geronimo, Champion. ZACAPA: 1 female, Zacapa, 1 Oct 1929, D. M.

Bates.

SPHAENOTHECUS BIVITTATUS DUPONT

Sphaenothecus bivittatus Dupont, 1838: 58, pl. 220, Fig. 1; Guerin, 1844: 250;

Linsley, 1934: 61; Saalas, 1936: 111; Linsley, 1940: 562.

Sphenothecus bivittatus; White, 1853: 86; Bates, 1880: 84.

Ischnocnemis bivittatus; LeConte, 1873: 316; Leng, 1887: 193; Townsend, 1903:

TA:

Taranomis bivittata; Casey, 1912: 333; Linsley, 1942: 63; Linsley, 1962: 95;

Rogers, 1977: 227; Chemsak, Linsley and Mankins, 1980: 33.

Taranomis bivittatus, Duffy, 1960: 109.

Taranomis bivittata bivittata; Linsley, 1960: 95; Turnbow and Wappes, 1978: 368;

Hovore and Penrose, 1982: 25; MacKay, Zak and Hovore, 1987: 364; Hovore,

Penrose and Neck, 1987: 296.

Leptocera bilineata Gory, 1831: pl. 45, Fig. 9; Castelnau, 1840: 490, pl. 34, Fig. 4.

Taranomis cylindricollis Casey, 1924: 267. NEW SYNONYMY.

Taranomis bivittata cylindricollis, Linsley, 1962: 96.

Taranomis nematocera Casey, 1924: 268.

Male.—Form small, tapering; integument black, legs and often pronotal disk reddish, elytra with 2

longitudinal eburneous vittae on each side; pubescence short, pale or golden, appressed and subde-

pressed. Head small; antennal tubercles not elevated; vertex coarsely, confluently punctate; pubescence

erect, sparse, antennae slender, 12-segmented, 2.5 to 3 X longer than body, scape conical, slender,

dorsally carinate, at least near base, basal segments clothed with short, dark, suberect setae, third

segment longer than scape, fourth subequal to third. Pronotum broader than long, sides broadly round-

ed; disk convex, broadly impressed at middle at base; basal margin shallowly lobed at middle; punc-

tures moderately coarse, sparse to dense, usually transverse on apical one-half, center behind middle

usually with an irregular glabrous area; each side with a broad, longitudinal, densely appressed, pu-

bescent vitta, sides with numerous long, erect hairs; prosternum shallowly impressed, densely pubes-

16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

cent; meso- and metasternum densely clothed with silvery appressed pubescence and longer, erect

hairs, middle glabrous. Elytra about 2.5 X as long as broad; basal margin moderately bilobed; each

elytron with 2 longitudinal eburneous fasciae, one beginning inside of humeri and extending almost

to apex, one lateral usually not reaching lateral margin and extending almost to apex; punctures

between fasciae fine, dense, becoming finer and denser toward apex; pubescence dense, fine, appressed,

long, erect hairs usually numerous basally; apices sinuate truncate, angles usually moderately aentate.

Scutellum about as long as broad, apex narrow, acute, sparsely pubescent. Legs slender; hind femora

extending beyond body, moderately densely punctate and pubescent; hind tibiae moderately densely

clothed with suberect, dark setae; hind tarsi slender, first segment as long as following two together.

Abdomen densely clothed with silvery appressed pubescence, middle glabrous; last sternite lightly

emarginate at middle. Length 9—15.5 mm.

Female.—Form similar. Antennae a little longer than body, 11-segmented. Legs with femora not

extending beyond body. Abdomen with last sternite broadly truncate. Length, 9-14 mm.

Diagnosis.—This species averages smaller in size than other Sphaenothecus.

The slender, 12-segmented antennae and elongated hind femora of males and the

usual dark lateral margins of the elytra also characterize this species. The color

of the pronotum varies from black to reddish and does not appear to be a constant

geographical character.

Discussion.—We have been informed that (Monné 1994) in his recent catalog

has reinstated the name bilineatus Gory for bivittatus. We have not seen this

reference and in view of the confusion regarding publication dates are mantaining

the long accepted synonymy of bilineatus. Unless the name is a homonym, Mon-

né’s usage may be correct.

Type Locality— Of bivittatus, Mexico; bilineata, Mexico; cylindricollis, near

Tucson, Arizona; nematocera, near Tucson, Arizona.

Range.—Southern California to Texas, Honduras and Baja California (Pl. 1).

Flight Period.—April to November.

Host Plants.—Acacia, Prosopis.

Remarks.—There appears to be no justification for segregating this species into

another genus and it is being treated as originally proposed by Dupont, 1838.

SPHAENOTHECUS PICTICORNIS BATES

(Pl. 3)

Sphenothecus picticornis Bates, 1880: 84.

Sphaenothecus picticornis; Aurivillius, 1912: 472.

Male.—Form moderate sized, tapering; integument reddish to black, pronotum and underside often

reddish, antennal segments dark at apices, elytra usually with 2 narrow, eburneous, longitudinal vittae

on each side, legs with tibiae and femora dark at apices, tarsi black; pubescence moderately dense,

short, appressed silvery to golden, pronotum laterally with a longitudinal band of dense, appressed

pubescence. Head small; antennal tubercles prominent, rounded; vertex with an often vague, medially

impressed callus; pubescence around eyes dense; antennae 12-segmented, often 2.5 X longer than

body, scape cylindrical, about 2 X as long as broad, dorsally carinate, basal segments densely clothed

with short, dark, vaguely bluish, suberect setae, outer segments clothed with minute appressed pubes-

cence, segments dark annulate at apices, segment 3 about as broad as scape, segments 4—9 slightly

expanded at apices, segment 3 longer than scape, 4 slightly longer than 3, 5 longer than 4. Pronotum

slightly broader than long, sides broadly rounded; disk convex, transversely impressed near base, basal

margin strongly lobed at middle; punctures fine, denser over apical one-half, middle toward base

usually impunctate; each side with a moderately broad, longitudinal, silver or golden pubescent vitta,

vittae widely separated on disk, sides with numerous long, erect, pale hairs; prosternum barely im-

pressed, densely pubescent; meso- and metasternum densely clothed with silvery or golden, appressed

pubescence, middle glabrous, long, pale, erect hairs numerous, mesosternal process prominently ele-

vated. Elytra about 2.5 X as long as broad, basal margin strongly bilobed; each elytron with 2 ebur-

CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS 17

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THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

1998 CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS 19

neous, longitudinal fasciae extending almost to apex; each side narrowly costate down middle; basal

punctures fine, sparse, becoming very dense toward apex; pubescence fine, short, silvery, densely,

transversely appressed along sutural area; apices sinuate, angles acutely dentate. Scutellum strongly

tapering, about 2 X as long as broad, glabrous. Legs slender; hind femora not extending beyond body,

sparsely punctate and pubescent; hind tibiae moderately densely clothed with fine, suberect setae; hind

tarsi slender, segment 1 slightly shorter than two following together. Abdomen densely pubescent,

narrowly glabrous medially; last sternite shallowly emarginate at middle. Length, 12—22 mm.

Female.—Form similar. Antennae subserrate, slightly longer than body; 11-segmented, segment 11

strongly appendiculate. Abdomen with last sternite broadly truncate at apex. Length, 14-19 mm.

Diagnosis.—This species varies greatly in size and coloration. The eburneous

fasciae of the elytra vary in length and width and occasionally are black. The

pronotum is often totally black and the legs and underside vary in the amount of

infuscation. The thickened, slightly arcuate, densely pubescent third segment of

the antennae, strongly lobed basal pronotal margin and strongly bilobed elytral

base make this species distinctive.

Type Locality.—Mexico.

Range.—Central Mexico from Michoacan to Veracruz and Oaxaca.

Flight Period.—October to January.

Material Examined.—All from MEXICO. GUERRERO: 1 male, 2 km SE of Grutas on Hwy 55, 6

Jan 1989, R. L. Minckley and B. N. Danforth. MEXICO: 1 male, km 20 of Tejupilco-Bejucos, 6 Jun

1964, Barrera, Brailovsky. MICHOACAN: 1 male, 3 km N of Arteaga, 850 m, 1 Nov 1987, R. Ayala.

MORELOS: 16 males, 6 females, 2.5 km W of Huautla, Estacion CEAMISH, 940 m, 20 Nov 1995

to 16 Mar 1996, EF A. Noguera, A. Rodriguez, A. Morales, E. Ramirez, E. Gonzalez; 3 males, 1

female, 2.5 km W of Ajuchitlan, 950 m, 11 Nov 1995 to 17 Jan 1996, R. Ayala, E A. Noguera, E.

Gonzalez. OAXACA: 1 male, 63 km N of Huajuapan, 14 Oct 1978, E. Giesbert; 4 males, 1 female,

Petlalcingo, 25 Nov 1963, A. E. Michelbacher. PUEBLA: 1 male, 1 female, 15 km WNW of Izucar

de Matamoros, 1300 m, 6 Nov 1976, P. H. Sullivan. VERACRUZ: 3 males, 1 female, Cordoba, 28

Oct 1963, A. B. Lau. STATE UNKNOWN: 1 male, 1 female.

SPHAENOTHECUS TOLEDOI CHEMSAK AND NOGUERA, NEW SPECIES

(Pl. 5)

Types.—Holotype male, allotype: MEXICO. CHIAPAS: 16 km W of Ocozo-

cuatla, ““El Aguacero’’, 26 Jan 1995, V. H. Toledo; deposited: Instituto de Biolo-

gia, UNAM, Mexico. Paratypes, all with same data as follow: 1 male, 1 female,

2 Jan 1995; 3 males, 1 female, 28 Jan 1995; 1 male, 2 females, 30 Jan 1995; 2

males, 1 female, 1 Feb 1995; 1 male, 28 Feb 1995; 1 male, 1 Mar 1996; 1 male,

2 Mar 1996; 5 males, 3 Mar 1995; 1 male, 2 females, 4 Mar 1995; 1 male, 22

Dec 1994; deposited in EMEC; EBCC and V. H. Toledo collection. Additional

paratypes include: OAXACA: 1 male, 15 km E Tehuantepec, emerged Dec 1980

from dead thorned vine, E. Giesbert (GIES). HONDURAS. LA PAZ: 1 male,

Taladro, 5 Jan 1976, J. V. Mankins (EMEC).

Male.—Form moderate sized, tapering; integument reddish, head pronotum, legs and underside

usually partially infuscated, antennae narrowly black at apices of segments, elytra testaceous with

longitudinal black vittae; pubescence sparse to dense, erect and appressed. Head small; antennal tu-

bercles prominent, obtuse at apices; vertex with a usually raised median line; pubescence fine, pale,

Plates 2-5. Plate 2, Sphaenothecus argenteus Bates; Plate 3, S. picticornis Bates; Plate 4, S.

maccartyi NEW SPECIES, and Plate 5, S$. toledoi NEW SPECIES.

20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

suberect; antennae slender, 11-segmented, often more than 2 X as long as body, scape rather slender,

cylindrical, about 3 X longer than broad, carinate basally, basal segments moderately densely clothed

with short, subdepressed blue-black setae, outer segments minutely pubescent, segments 3-8 slightly

expanded at apices, segment 3 narrower than scape except at apex, segment 3 longer than 1, segments

3-5 equal, 11 long, appendiculate. Pronotum broader than long, sides broadly rounded; disk convex,

impressed at middle near base, basal margin broadly lobed at middle; punctures moderately coarse,

irregular, transversely placed, area before impression usually non-punctate; sides with widely separated,

longitudinal bands of pale, appressed pubescence, long, erect hairs numerous at sides; prosternum

barely impressed, densely pubescent; mesosternal process prominently projecting, meso- and meta-

sternum densely golden pubescent, medially glabrous. Elytra a little more than 2 X longer than broad;

basal margin moderately strongly bilobed; each elytron with two broad, longitudinal, yellowish vittae,

suture, middle of disk and often, lateral margins, dark vittate; pubescence toward apex dense, ap-

pressed; apices sinuate truncate, inner angles very lightly dentate. Scutellum glabrous, black, longer

than broad, acute at apex. Legs slender, stout; hind femora not attaining elytral apices, sparsely punctate

and pubescent; hind tibiae moderately densely clothed along edges with short, suberect setae; hind

tarsi short, moderately broad, segment 1 shorter than two following together. Abdomen densely pu-

bescent, medially glabrous; last sternite subtruncate, shallowly emarginate at middle. Length, 16-21

mm.

Female—Form similar. Antennae subserrate, a little longer than body, segment 11 pointed at apex.

Abdomen with last sternite broadly truncate. Length, 16-19 mm.

Diagnosis.—S. toledoi may be separated from S. bivittatus by the larger size,

11-segmented antennae of males and basally thicker antennal segments. It differs

from S. picticornis by the feebly bilobed base of the elytra, usually broad yellow-

ish elytral vittae, and 11-segmented antennae of males. S. maccartyi may be sep-

arated by the strongly dentate outer angles of the elytral apices and 12-segmented

antennae of males.

Etymology.—We dedicate this species to V. H. Toledo to acknowledge his col-

lecting efforts.

SPHAENOTHECUS MACCARTYI CHEMSAK AND NOGUERA, NEW SPECIES

(Pl. 4)

Sphaenothecus sp.; Chemsak and Noguera, 1993: 64.

Taranomis bivittata bivittata; (not Dupont) Chemsak and Noguera, 1993: 64.

Types.—Holotype male: MEXICO. JALISCO: Chamela, 18 Feb 1986, R. Ay-

ala; allotype, Estacion de Biologia Chamela, emerged ex Stemadenia sp., 7 Oct

1985, E A. Noguera; deposited in Instituto de Biologia, UNAM, Mexico. Para-

types, all from Estacion de Biologia Chamela as follows: 25 males, 10 females,

1-8 Dec 1988, J. A. Chemsak, J. D. MacCarty (EMEC and J. D. MacCarty

collection); 1 male, 7 Dec 1977, H. Brailovsky (UNAM); 3 males, 3 females,

Dec 1986, host Caesalpinia eriostachys Benth, FANM 499, S. H. Bullock; 1

female, Dec 1986, host Caesalpinia sclerocarpa Standl., FANM 501, S. H. Bul-

lock; 1 male, 3 Feb 1987, FANM 449, host Amphipterigium adstringens

(Schlecht.) Schiede, EK A. Noguera; 1 female, 3 Feb 1987, FANM 454, host Com-

ocladia engleriana Loes, EK A. Noguera; 1 male, 2 females, 5 Feb 1987, FANM

462, host Caesalpinia eriostachys, EF A. Noguera; 1 female, 9 Feb 1987, FANM

485, host Spondias purpurea L., E A. Noguera; 1 female, 13 Feb 1987, FANM

521, host Caesalpinia sclerocarpa, KF A. Noguera; 2 males, 13 Feb 1987, FANM

525, host Bursera instabilis McVaugh & Rzed., E A. Noguera; 1 female, 18 Feb

1987, FANM 539, host Delonix regia (Bo}j.), E A. Noguera; 2 females, 20 Feb

1987, FANM 604, host Caesalpinia caladenia Standl., EK A. Noguera; 1 female,

1998 CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS 21

2 Apr 1987, FANM 610, host Pithecellobium mangense (Jacq.) MacBride, EF A.

Noguera; 1 male, ex: Stemadenia, 2 Oct 1985, Noguera; 4 males, 3 females, 7—

9 Dec 1986, on flowers of Caesalpinia eriostachys, F A. Noguera; 3 males, 2

females, 9 Dec 1987, 14 Jan 1987, on Caesalpinia flowers, EF A. Noguera; 1 male,

27 Sep 1987, Noguera; 1 female, 9 Dec 1984, ovipositing on dead branch of

Ipomoea wolcottiana Rose, Noguera; 1 male, 22 Nov 1985, Noguera; 1 male, 18

Feb 1986, R. Ayala (EBCC). One additional male paratype from 8 km (5 mi) N

of Mazatlan, Sinaloa, Mexico, 12 Dec 1988, Chemsak (EMEC), and 2 males, 1

female, Rio San Nicolas, Chamela, 18 Nov 1985, R. Ayala (EBCC).

Male.——Form moderate sized, tapering; integument reddish, head pronotum, apices of antennal seg-

ments, apices of femora, elytra often partially infuscated; pubescence sparse to dense, short, appressed

and long and erect. Head small; antennal tubercles not prominent, apically obtuse; vertex linearly

impressed, vaguely callused; pubescence fine, short, golden, appressed with long, erect hairs numerous

around eyes; antennae slender, 12-segmented, about 2 x as long as body, scape slender, slightly

cylindrical, a little more than 2 X as long as broad, carinate basally, basal segments moderately densely

clothed with short, dark, subdepressed setae, outer segments very finely pubescent, segments 3-8

slightly enlarged at apices, segment 3 as broad as and longer than scape, 4 equal to 3, 5 subequal to

4. Pronotum broader than long, sides broadly rounded; disk convex, impressed at middle near base,

basal margin narrowly lobed at middle; punctures fine, sparse, irregular, transverse; each side with a

slightly curved, broad, golden pubescent, longitudinal vitta, sides with numerous long, erect hairs;

prosternum lightly impressed near apex, densely golden pubescent, medially glabrous. Elytra a little

more than 2 X as long as broad; basal margin moderately strongly bilobed; punctures fine, sparse at

base, dense along suture and between vittae; each elytron with 2 narrow, longitudinal eburneous vittae,

vittae usually broadly yellowish and extending to apex, suture and disk dark vittate; pubescence rather

sparse, short depressed; apices sinuate, outer angles usually strongly dentate, inner angles with small

teeth. Scutellum about 1.5 X longer than broad, glabrous. Legs slender; hind femora not attaining

elytral apices, sparsely punctate and pubescent; hind tibiae moderately densely clothed with fine, dark,

suberect setae; tarsi black, hind pair slender, segment 1 slightly shorter than two following together.

Abdomen densely pubescent, glabrous medially; last sternite narrowly subtruncate. Length, 10-18

mm.

Female.—Form similar. Antennae 11 segmented, slightly longer than body, segments subequal in

diameter, eleventh appendiculate. Abdomen with last sternite broadly subtruncate. Length, 12-15 mm.

Diagnosis.—This species is distinctive by the usually strongly dentate outer

angles of the elytral apices. It averages smaller in size than S. toledoi and also

differs from that species by the 12-segmented antennae of males. S. maccartyi

also differs from S. bivittatus by the broader yellowish vittae of the elytra and by

the shorter hind femora which do not attain the apices of the elytra.

Etymology.—We are pleased to dedicate this species to J. D. McCarty for his

interest, assistance, and collecting efforts.

Remarks.—A series of 12 males, 7 females from 16 km W of Ocozocoautla,

‘““E] Aguacero’’, Chiapas, Mexico, 2 Nov 1994 to 4 Mar 1995, V. H. Toledo,

appear assignable to this species. These are identical in all respects to the Jalisco

and Sinaloa populations except for the outer angles of the elytral apices which

are only lightly dentate.

SPHAENOTHECUS FACETUS CHEMSAK AND NOGUERA, NEW SPECIES

(Pl. 6)

Types.—Holotype male: COSTA RICA. GUANACASTE: Santa Cruz, 28 Nov

1985; allotype: Comelco, 8 km NW of Bagaces, 10 Apr 1971, P. Opler; deposited:

Essig Museum of Entomology, University of California, Berkeley. Paratypes, all

ae THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Plates 6-7. Plate 6, Sphaenothecus facetus NEW SPECIES; Plate 7, S. trilineatus Dupont.

from same Province as follow: 3 males, 5 females, Comelco, 8 km NW of Ba-

gaces, 10 Apr 1971, 24—25 Jan 1972, 3 Feb 1972, P. Opler; 1 male, 1 female,

Comelco, 4 km NW of Bagaces, Area A, on flowers of Bullthorn (Acacia sp.),

P. Opler; 2 males, 1 female, Cafias, 5 Jan 1972, E Cordero; 2 males, 1 female,

Hacienda La Pacifica, 5 km NW of Cafias, 1—10 Jan 1974, 1 Feb 1972, G. Frankie,

17 Feb 1972; 1 male, 1 female, Santa Rosa National Park, 7—9 Dec 1979, D. H.

Janzen, 2—11 Mar 1980, Janzen and Hallwachs. Two additional male paratypes

from: GUATEMALA. ZACAPA: 1 male, Gualan, 22 Jan 1905, J. A. Hine; 1

female, 2 km E of Santa Cruz, 14 Jun 1991, E. Giesbert. All the paratypes de-

posited in EMEC except 2 specimens in EBCC and 1 in Giesbert Collection.

Male.—Form moderate sized, tapering; integument reddish-brown, antennae darker; pubescence

white to creamy, appressed, forming 3 longitudinal vittae on elytra and 2 on pronotum. Head small;

antennal tubercles prominent, apices acutely elevated; front short, deeply impressed; vertex linearly

impressed; pubescence sparse, denser on front; antennae slender, 12-segmented, extending 5—6 seg-

ments beyond apices of elytra, scape short, about 2 X as long as broad, flattened basally dorsally and

ventrally, dorsally carinate near base, segments 3-8 slightly enlarged at apices, segments 5—10 dorsally

vaguely carinate, basal segments clothed with short, subdepressed, blue-black setae, segment 3 longer

than 1, 4 subequal to 3, 5 longer than 4. Pronotum slightly broader than long, sides ocassionally with

a small lateral tubercle; base broadly impressed laterally; disk convex, impressed medially near base;

punctures moderately sparse, transversely placed; pubescence sparse, each side with a broad densely

pubescent longitudinal vitta, basal impression with a rounded pubescent spot, long, fine, erect hairs

numerous at sides below vittae; prosternum narrowly impressed at apex, densely clothed with sub-

depressed pubescence; meso- and metasternum medially glabrous, sides clothed with dense, appressed

1998 CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS 23

pubescence. Elytra a little more than 2 X as long as broad, tapering apically; basal margin moderately

sinuate; each elytron with a narrow pubescent fascia along suture and a slightly broader one on disk

ending just before apex; punctures fine, well separated; apices sinuate. Scutellum narrow, elongate,

densely pubescent. Legs slender; femora gradually enlarged toward apex, not extending to elytral

apices, finely, moderately densely punctate, pubescence short, depressed, moderately dense; hind tibiae

clothed with short, dark, subdepressed setae; hind tarsi moderately broad, segment 1 shorter than two

following together. Abdomen medially glabrous, sides densely clothed with appressed pubescence;

last sternite very shallowly, narrowly emarginate truncate at apex. Length, 15-22 mm.

Female.—Form similar, more robust. Antennae about as long as body. Abdomen with last sternite

broadly rounded. Length, 15-22 mm.

Diagnosis.—The broad, basally flattened antennal scape immediately identifies

this species. Additionally the dark reddish-brown integument and usually creamy

colored pubescent vittae make it distinctive.

Etymology.—The name, facetus, alludes to the elegant appearance of this species.

SPHAENOTHECUS TRILINEATUS DUPONT

(Pl. 7)

Sphaenothecus trilineatus Dupont, 1838: 57, pl. 219, Fig. 3; Casey, 1912: 333;

Linsley, 1935: 100; Chemsak, Linsley and Mankins, 1980: 33; Chemsak and

Noguera, 1993: 64.

Sphenothecus trilineatus,; White, 1853: 85; Bates, 1880: 84; 1885: 329.

Sphaenothecus tomentosus Dupont, 1838: 56, pl. 219, Fig. 1; Casey, 1912: 333.

Sphenothecus tomentosus; White, 1853: 85; Bates, 1880: 84; 1885: 328.

Sphoenothecus tomentosus; Thomson, 1864: 205.

Sphaenothecus trilineatus V. tomentosus,; Blackwelder, 1946: 589.

Sphaenothecus lateralis Bates, 1880: 329.

Sphaenothecus trilineatus v. lateralis; Blackwelder, 1946: 589.

Male.—Form moderately large, tapering; integument black; pubescence short, dense, appressed,

creamy to yellowish, forming 2 longitudinal fasciae on pronotum and 3 prominent fasciae on elytra.

Head small; antennal tubercles prominently elevated, apices obtuse; vertex linearly impressed, irreg-

ularly punctate; pubescence on front, above and below eyes dense, appressed; antennae slender, 12-

segmented, at least 2 X as long as body, scape cylindrical, slightly broader at apex, vaguely carinate

basally, basal segments clothed with short subdepressed, violet-black setae, outer segments densely

clothed with very short, fine, dark, appressed pubescence, segments 4—8 slightly enlarged at apices,

segments 11 and 12 almost fused, segment 3 slightly narrower than scape, segment 3 longer than

scape, 4 subequal to 3, 5 longer than 4. Pronotum slightly broader than long, sides broadly rounded;

disk convex, impressed at middle near base; basal margin broadly arcuate, not lobed; punctures fine,

rather sparse, center often with an impunctate area; each side with a broad, longitudinal, densely

appressed pubescent vitta, middle often with an incomplete vitta or at least a pubescent spot on basal

impression, long, erect, pale hairs numerous at sides; prosternum barely impressed, densely pubescent;

meso- and metasternum densely golden pubescent, middle glabrous. Elytra a little more than 2 X as

long as broad; basal margin slightly arcuate, not bilobed; punctures fine, sparse; each elytron with a

narrow to broad longitudinal, densely appressed pubescent fascia, sutural pair usually narrower, epi-

pleura with a narrow pubescent fascia on basal one-half; apices sinuate truncate, angles unarmed.

Scutellum longer than broad, densely pubescent. Legs slender, robust; hind femora not extending to

elytral apices, rather sparsely punctate, usually with a dorsal patch of appressed pubescence at basal

one-half; hind tibiae densely clothed with short, dark, subdepressed setae; hind tarsi moderately broad,

first segment shorter than following two together. Abdomen densely pubescent, glabrous medially; last

sternite narrow, shallowly emarginate at apex. Length, 16-25 mm.

Female.—Form similar, more robust. Antennae a little longer than body, 11-segmented, segments

subserrate, 11 strongly appendiculate. Abdomen with last sternite broadly rounded at apex. Length,

15-25 mm.

24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Diagnosis.—The large size, black integument and pubescent fascia of the elytra

distinguish this species. It differs from S. facetus by the dark integument and

longer, non-flattened antennal scape.

Type Locality.—Of trilineatus, Mexico; tomentosus, Mexico; lateralis, Mexico.

Range.—Southern Texas to Veracruz, Sinaloa to Oaxaca.

Flight Period.—September to February.

Host Plants—Amphipterigium adstringens, Spondias purpurea, Lonchocarpus

magallanesii, Delonix regia, Acacia angustissima (Mill.) KTze., Caesalpinia er-

iostachys, Caesalpinia platyloba S. Wats, Caesalpinia sp., Ceiba pentandra (L.)

Gaertn.

Remarks—The width of the pubescent elytral fasciae varies considerably.

Specimens from eastern Mexico tend to have the fasciae thin and cream colored.

Those from western Mexico tend to have the fasciae broader and yellow.

There is at hand a single male from Brownsville, Texas, June 1925, G. Linsley,

which represents the first record to the United States fauna.

Adults can be encountered on the flowers of Acacia, Apoplanesia paniculata

Presl., Caesalpinia, Croton, Leucaena, Lonchocarpus and Ipomoea.

Material Examined.—MEXICO. CHIAPAS: 23 males, 8 females, 16 km W of Ocozocuatla, El

Aguacero, 4-16 Nov 1994, 3-10 Dec 1994, 31 Jan 1995, V. H. Toledo. GUERRERO: 3 males, 2

females, 13 km NW of Iguala, 12 Sep 1982, Powell & Chemsak; 1 male, 1 female, 23 km W of

Iguala, 11-16 Sep 1982, Powell & Chemsak; 9 males, 11 females, Zopilote Canyon, 8 km S of

Mezcala, 17 Sep 1982, Chemsak. JALISCO: 65 males, 59 females from Estacion de Biologia Chamela

as follow, 3 Jan 1983, S. H. Bullock; 12 Nov 1983, Bullock; 11 Dec 1983, Bullock; 17 Jan 1985,

THA 104, host Caesalpinia sp., emerged 27 Jul 1985, Atkinson & Noguera; 6 Feb 1985, THA 143,

host Ceiba pentandra, emerged 5 Jul 1985, Atkinson; 23 Feb 1985, FANM 205, host Lonchocarpus

magallanesii, emerged 13—26 Jun 1985, 1-27 Jul 1985, 2—24 Aug 1985, 2-15 Sep 1985, Noguera;

14 Mar 1985, FANM 220, host Caesalpinia platyloba, emerged 5 Jul 1985, Noguera; 1 Apr 1985,

Noguera; 18 Jun 1985, Noguera; 30 Sep 1985, A. Rodriguez; 25 Oct 1985, M. Sanchez; 13 Dec 85,

on flowers of Caesalpinia eriostachys, A. Rodriguez; 2 Jul 1986, Noguera; 22 Jul 1986, AR30a, on

flowers of Acacia sp., A. Rodriguez; 28 Aug 1986, M. Sanchez; 22 Sep 1986, FANM 410a, Noguera;

3 Oct 1986, R. A. Usela; 14/23 Oct 1986, Chemsak; 14 Nov 1986, Noguera; 7 Dec 1986, FANM

433d, on flowers of Caesalpinia eriostachys, Noguera; 9 Dec 1986, FANM 436, Noguera; 5 Feb 1987,

FANM 464, host Spondias purpurea, Noguera; 9 February 1987, FANM 505, host Spondias purpurea,

Noguera; 12 Feb 1987, FANM 510-512, host Delonix regia, Noguera; 18 Feb 1987, FANM 543, host

Delonix regia, Noguera; 13 Mar 1987, FANM 570, host Acacia angustissima, Noguera; 13 Mar 1987,

FANM 574, host Amphipterigium adstringens, Noguera; 25 Sep 1987, FANM 683b, on flowers of

Leucaena sp., Noguera; 25 Sep 1987, FANM 683c, on flowers of Apoplanesia paniculata, Noguera;

7 Oct 1987, FANM 685b, on flowers of Leucaena sp., Noguera; 17 Oct 1987, Chemsak; 20-22 Oct

1987, J. Powell; 7-15 Jul 1987, Chemsak, E. & J. Linsley; 1-8 Dec 1988, Chemsak; 21 Nov 1991,

E Ramirez; 2 males, Jose Maria Morelos, 13 Aug 1985, 5 Sep 1985, M. Sanchez; 5 females, Quemaro,

25 Oct 1985, Atkinson; 6 males, 3 females, 7 km NE of Melaque, 16-23 Oct 1986, Chemsak; 4

males, 2 females, 4 km N of Melaque, 27 Oct 1986, FANM 427a, Noguera; 3 males, 1 female from

Careyes as follow, 17 Oct 1986, on flowers of Lonchocarpus sp., Noguera; 21 Dec 1992, Chemsak

& Katsura; 5 females, 1 male, 8 km N of Autlan, 8 Oct 1993, Carrillo, Rodriguez & Noguera; 3

males, 1 female, El Corcovado, 12 Sep 1994, alt. 940 m, G. Nogueira; 1 female, Sayula, Isla Chica,

B. espinoso, 1366 m, 13 Nov 1994, S. Gallegos. MORELOS: 3 males, 1 female, 9.6 km (6 mi) S of

Amacuzac, 9 Oct 1963, Michelbacher; 1 male, 1 female, Tequesquitengo, 1 Oct 1959, E. Gonzalez;

1 female, Yautepec, 13 Jul 1963, Parker, Stange; 1 male, Cafion de Lobos, 22 Dec 1948, A. C. Smith;

1 female, Progreso, 3 Feb 1949, J. Hernandez; 2 females, Cuernavaca, 5 Nov 1989, 1700 m, A.

Burgos; 24 males, 8 females, 2.5 km N, 4 km W Huatla, Estacion CEAMISH, 940 m, 18°40'.671 N

99°02'.475 W, 16 Nov 1995 to 14 Jan 1996, R. Ayala, E A. Noguera, E. Gonzalez, E. Ramirez, A.

Rodriguez; 6 males, 12 females, 2.5 km W Ajuchitlan, 950 m, 18°28'.065 N 98°59'.548 W, 17 Nov

1995 to 17 Jan 1996, R. Ayala, A. Rodriguez, A. Perez, Noguera. MICHOACAN: 1 female, Laguna

1998 CHEMSAK & NOGUERA: REVIEW OF SPHAENOTHECUS D5

Zicuiran, 4 Nov 1992, Rodriguez & Noguera; 1 female, 6 km W of La Placita, 214 km W Playa Azul,

7 Nov 1987, R. Ayala. OAXACA: 2 males, 1 female, 36.8 km (23 mi) S of Matias Romero, 14 Aug

1963, Paker, Stange; 2 females, 6.4 km (4 mi) E of El Camaron, 13 Oct 63, A. E. Michelbacher; 1

male, 7.2 km (4.5 mi) E of Tehuantepec, 13 Sep 1959, Cantrall, Cohn; 2 females, 12.8 km (8 mi)

NW of Tutla, 6 Oct 1975, Chemsak; 1 female, 15 km S of Rio Grande, 24 Oct 1989, Noguera &

Rodriguez; 5 females, 2.5 km NE of Puerto Angel, 27 Oct 1989, Noguera & Rodriguez. PUEBLA: 2

males, 9.6 km (6 mi) SE of Acatlan, 8 Oct 1975, Chemsak; 1 female, 9.6 km (6 mi) S of Zapotitlan,

6 Oct 1975, Chemsak. SYNALOA: 2 males, Elota, 29 Aug 1960, Westcott; 2 males, 3 females, Gua-

muchil, 27 Oct 1965, G. E. & A. S. Bohart; 2 females, 1.6 km (1 mi) SE of Mazatlan, 25 Dec 1968,

D. L. Briggs. VERACRUZ: 1 female, 43.2 km (27 mi) E of Jalapa, 31 Oct 1973, Williams, Mullinex.

ACKNOWLEDGMENT

We gratefully acknowledge the following institutions and individuals for the

use and loan of material: Essig Museum of Entomology, University of California,

Berkeley (EMEC); Estacion de Biologia Chamela, IBUNAM, Chamela (EBCC);

Instituto de Biologia, UNAM, Mexico (UNAM); American Museum of Natural

History, New York; United States National Museum of Natural History, Wash-

ington, D.C.; Centro de Zoologia, CUCBA-UdeG, Guadalajara; E. Giesbert, V.

H. Toledo, J. D. McCarty, D. Heffern, J. L. Navarrete, A. Burgos, and B. Eya.

LITERATURE CITED

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Bates, H. W. 1879-1885. Biologia Centrali-Americana, Insecta, Coleoptera, 5, Longicornia, 436 p.

Blackwelder, R. E. 1946. Checklist of the coleopterous insects of Mexico, Central America, the West

Indies, and South America. Pt. 4. U.S. Nat. Mus. Bull., 185: 551-763.

Bradley, J. C. 1930. A manual of the genera of beetles of America north of Mexico. 360 pp. Ithaca,

N.Y.

Casey, T. L. 1912. Studies in the Longicornia of North America. Memoirs on the Coleoptera, 3: 215—

376.

Casey, T. L. 1924. Additions to the known Coleoptera of North America. Memoirs on the Coleoptera,

11: 1-347.

Castelnau, F 1840. Histoire naturalle des animaux articules. Insectes, coleoptéres, Vol. 2, 564 pp.

Chemsak, J. A., E. G. Linsley and J. V. Mankins. 1980. Records of some Cerambycidae from Hon-

duras. Pan-Pacific Entomol., 56: 26—37.

Chemsak, J. A. and E A. Noguera. 1993. Annotated checklist of the Cerambycidae of the Estacion

de Biologia Chamela, Jalisco, Mexico, Coleoptera, with descriptions of new genera and species.

Folia Entomol. Mex., 89: 55-102.

Duffy, E. A. J. 1960. A monograph of the immature stages of Neotropical timber beetles. Br. Mus.,

Nat. Hist., London.

Dupont, H. 1838. Monographie des trachyderides de la famille des longicornes. Mag. Zool., 8: i—xili,

1-59.

Gemminger, M. and E. von Harold. 1873. Catalogus Coleopterorum hucusque descriptorum synon-

ymicus et systematicus, 10: 2989-3232. Monachii.

Gory, H. L. 1831. Jn FE E. Guerin-Meneville, Iconographie du régne animal de G. Cuvier . . . insectes,

7. 5-576. Paris.

Guerin-Meneville, EF E. 1844. Iconographie du régne animal. Vol. 1, Insectes, 576 pp. Paris.

Hovore, FE T. and R. L. Penrose. 1982. Notes on Cerambycidae coinhabiting girdles of Oncideres

pustulatus LeConte, Coleoptera: Cerambycidae Southwest. Nat., 27: 23-27.

Hovore, E T., R. L. Penrose and R. W. Neck. 1987. The Cerambycidae, or Longhorned beetles, of

Southern Texas: A faunal survey (Coleoptera). Proc. Calif. Acad. Sci., 44: 283-334, 20 Figs.

Lacordaire, T. 1869. Histoire naturelle des insectes. Genera des coleoptéres, 8: 1-552.

LeConte, J. L. 1873. Classification of the Coleoptera of North America. Part II. Smithson. Misc.

Collect, 11(265): 279-348.

LeConte, J. L. and G. H. Horn. 1883. Classification of the Coleoptera of North America. 2nd Ed.

Smithson. Misc. Collect, 26(507): 1-567.

26 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Leng, C. W. 1886. Synopses of Cerambycidae. Entomol. Amer., 2: 27-32, 60-63, 81-83, 102-103.

Leng, C. W. 1887. Synopses of Cerambycidae. Entomol. Amer., 3: 4—8, 23-24, 44.

Linsley, E. G. 1934. Studies in the Cerambycidae of Lower California, Coleoptera Pan-Pac. Entomol.,

10: 59-63.

Linsley, E. G. 1935. Studies on the Longicornia of Mexico. Trans. Amer. Entomol. Soc., 61: 61-102.

Linsley, E. G. 1940. Notes on Oncideres twig girdlers. J. Econ. Entomol., 33: 561-563.

Linsley, E. G. 1942. Contributions toward a knowledge of the insect fauna of Lower California. No.

2 Coleoptera, Cerambycidae. Proc. Calif. Acad. Sci., 24: 21-96.

Linsley, E. G. 1962. Cerambycidae of North America. Part. III. Taxonomy and classification of the

subfamily Cerambycinae, tribes Opsimini through Megaderini. Univ. Calif. Publ. Entomol., 21:

1-165.

MacKay, W. P, J. C. Zak and E T. Hovore. 1987. Cerambycid beetles, Coleoptera: Cerambycidae of

the northern Chihuahuan Desert, South Central New Mexico. Coleopt. Bull., 41: 361-369.

Monné, M. A. 1994. Catalogue of the Cerambycidae (Coleoptera) of the Western Hemisphere. Part

XI. Subfamily Cerambycinae: Tribes Torneutini, Trachyderini, Basipterini. Sociedade Brasileira

de Entomlogia, Sao Paulo.

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Kansas Entomol. Soc, 50: 222-228.

Saalas, U. 1936. Uber des Flugelgeader und die Phylogenetische entwicklung der Cerambyciden.

Ann. Zool. Soc. Zool.-Bot. Fenn. Vanamo, 4: 1-198.

Thomson, J. 1860. Essai d’une classification de la famille de cerambycides et materiaux pour servir

a une monographie de cette famille. 404 pp. Paris.

Thomson, J. 1864. Systema cerambycidarum ou expose de tous les genres compris dans la famille

des cerambycides et familles limitrophes. Mem. Soc. R. Sci. Liege, 19: 1-540.

Townsend, C. H. T. 1903. Contributions to a knowledge of the coleopterous fauna of the lower Rio

Grande Valley in Texas and Tamaulipas, with biological notes and special relevance to geo-

graphical distribution. Trans. Texas Acad. Sci., 1902, 5: 49-101.

Turnbow, R.H., Jr. and J. E. Wappes. 1978. Notes on Texas Cerambycidae. Coleopt. Bull., 32: 367—

372.

White, A. 1853. Catalogue of the coleopterous insects in the collection of the British Museum. Part

7. Longicornia, 1: 1-174.

Received 5 Dec 1996; Accepted 12 Nov 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 27-31, (1998)

ESTABLISHMENT AND REDISTRIBUTION OF

SPHENOPTERA JUGOSLAVICA OBENBERGER

(COLEOPTERA: BUPRESTIDAE) FOR BIOLOGICAL

CONTROL OF DIFFUSE KNAPWEED (CENTAUREA

DIFFUSA LAMARCK) IN THE MIDWESTERN AND

WESTERN UNITED STATES

R. F. LANG!, G. L. PIPER’, & E. M. Coomss?

‘United States Department of Agriculture, Animal and Plant Health

Inspection Service

Plant Protection and Quarantine, Forestry Sciences Laboratory

Montana State University, P. O. Box 170278

Bozeman, Montana 59717

*Department of Entomology, Washington State University

P. O. Box 646382, Pullman, Washington 99164

-Oregon Department of Agriculture, 635 Capitol Street NE

Salem, Oregon 97310

Abstract-—S phenoptera jugoslavica Obenberger (Coleoptera: Buprestidae) is a root boring beetle

imported to the United States as a biological control agent against Centaurea diffusa Lamarck

(Asterales: Asteraceae). United States Department of Agriculture, Animal and Plant Health In-

spection Service, Plant Protection and Quarantine personnel and research collaborators conducted

an intensive collection and redistribution program, releasing this biological control agent in 11

states and 61 counties from 1980 to 1996. Establishment and recovery has been confirmed in 8

states and 48 counties.

Key Words.—Insecta, biological control, weed, Sphenoptera, Centaurea.

Diffuse knapweed (Centaurea diffusa Lamarck) (Asterales: Asteraceae), an ac-

cidentally introduced European biennial, or short-lived perennial has invaded and

lowered the productivity of rangeland, pasture, wildlands, and wastelands in North

America. The plant is seed dependent for reproduction, survives on dry disturbed

lands with a northern distributional range in British Columbia at latitude 51° N,

and is found at altitudes ranging from 150 to over 900 m (Watson & Renney

1974). Diffuse knapweed has an elongated taproot and usually forms a rosette in

the spring or fall (Watson & Renney 1974). The plant is adapted to remain in the

rosette state for more than one year if climatic conditions or the stage of growth

are not conducive to bolting and seed production, an adaptation that is controlled

by vernalization (Thompson & Stout 1991).

In western North America diffuse knapweed was first collected in an alfalfa

field in Klickitat County, Washington in 1907 (Howell 1959). No other collections

were reported in Washington until the 1930’s. By 1985, diffuse knapweed was

reported in Washington (11 counties), Montana (10 counties), Idaho (5 counties),

and Oregon (5 counties) (Roché & Talbott 1986). Between 1989 and 1993, diffuse

knapweed infested area in Washington (Piper 1989) increased from 173,150 ha

to 641,993 ha of rangeland (Roché 1994). To diminish the severity and to halt

the rapid spread of C. diffusa, a complex of European phytophages has been

evaluated and released against the weed in North America (Schroeder 1985, Miill-

28 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

er & Schroeder 1989, Rees et al. 1996). Importations have included the capitulum-

attacking insects Bangasternus fausti (Reitter) (Coleoptera: Curculionidae) (1991),

Larinus minutus Gyllenhal (Coleoptera: Curculionidae) (1991), Urophora affinis

Frauenfeld (Diptera: Tephritidae) (1970), and U. quadrifasciata (Meigen) (Dip-

tera: Tephritidae) (1970), and the root-destroying insects Agapeta zoegana L.

(Lepidoptera: Cochylidae (1984), Pelochrista medullana (Staudinger) (Lepidop-

tera: Tortricidae) (1983), Pterolonche inspersa Staudinger (Lepidoptera:

Pterolonchidae) (1986), Cyphocleonus achates (Fahraeus) (Coleoptera: Curcu-

lionidae) (1987), and Sphenoptera jugoslavica Obenberger (Coleoptera: Bupres-

tidae) (1979) (Story & Nowierski 1984).

Spheno ptera jugoslavica adults consume diffuse knapweed foliage and the lar-

vae feed in the roots (Powell & Myers 1988). The adults emerge in early July

and live up to 30 d. Mating begins within 1 to 2 d and oviposition within 5 to

12 d continuing for 7 to 22 d (Zwolfer 1976). The eggs are generally positioned

between the appressed petioles of the rosette leaves and sometimes on the petioles.

Oviposition is synchronized with plant growth, the eggs being laid on inactive

rosettes in order to prevent them from being crushed or displaced during rapid

plant growth (Zwolfer 1976, Powell & Myers 1988). A female S. jugoslavica

produces about 47 eggs during her lifetime, laying 1 to 7 eggs per plant (Zwélfer

1976). Normally only one larva develops in a root (Harris & Shorthouse 1996).

A newly hatched larva feeds externally on basal tissue prior to its first molt after

which it tunnels into the root. The plant forms a spindle-shaped gall around the

larva which feeds upon the cortical tissue lining the gall (Powell & Myers 1988,

Harris & Shorthouse 1996). The entry tunnel leading to the gall and the gall itself

are filled with frass. The beetle overwinters as a larva in the root and resumes

feeding in the spring. Pupation begins in late May, and the pupal period lasts 15

to 21 d (Rees et al. 1996). Because of microclimatic variables that affect larval

and pupal development, there is a staggered emergence of adults from May to

July. Newly formed adults remain in the pupal chamber for 2 to 5 d before

gnawing an exit hole near the root crown and emerging (Zwolfer 1976). The

insect is univoltine.

MATERIALS AND METHODS

Spheno ptera jugoslavica was approved for release as a biological control agent

for diffuse knapweed in the United States on 17 May 1979 by the United States

Department of Agriculture (USDA), Animal and Plant Health Inspection Service

(APHIS), Plant Protection and Quarantine (PPQ). The beetle was first released in

the United States in 1980 in California, Idaho, Oregon, Washington and in Mon-

tana (1983) by USDA, Agricultural Research Service. The release material orig-

inated in Greece (Table 1). Major releases of S. jugoslavica adults, collected at

White Lake, British Columbia, Canada, were made in Idaho, Montana, Oregon,

and Washington between 1987 to 1992 by USDA, APHIS, PPQ and research

collaborators. Caged and uncaged releases of the biocontrol agent were made from

1987-1996 in Arizona, Colorado, Idaho, Montana, Nebraska, Nevada, Oregon,

South Dakota, Utah, Washington, and Wyoming (Table 1). To document estab-

lishment, some cooperators were instructed to use insect nets to sweep in con-

centric circles starting at the point of release and working their way outward to

a distance of approximately 6 m. Sweeping was conducted in mid-July during the

1998 LANG ET AL.: SPHENOPTERA ESTABLISHED 29

Table 1. Sphenoptera jugoslavica releases and status in the midwestern and western United States,

1983-1996.

State County Release Year Status

Arizona Coconino 1993 Not established

Gila 1994 Not established

Colorado Archuleta 1991 Not established

Boulder 1991 Not established

Douglas 1991, 1993-1994 Established?

Jefferson 1992 Not established

Idaho Benewah 1994 Not monitored

Blaine 1987 Not monitored

Camas 1990 Not monitored

Gooding 1990 Established

Jerome 1987-1989 Established

Lincoln 1987 Established

Montana Broadwater 1996 Not monitored

Fergus 1990 Established

Lewis & Clark 1983, 1989-1990 Established

Mineral 1993 Not monitored

Sweet Grass 1994-1995 Recovered

Nebraska Pierce 1993, 1995 Not established

Nevada Washoe 1994 Not established

Oregon Baker 1993 Established

Deschutes 1989 Established

Gilliam 1989 Established

Hood River 1989 Established

Jackson 1983 Established

Jefferson 1989 Established

Lake 1989 Established

Malheur 1989 Established

Morrow 1989 Established

Multnomah 1996 Recovered

Sherman 1988 Established

Umatilla 1989 Established

Union 1989 Established

Wallowa 1989 Established

Wasco 1987-1989 Established

Wheeler 1996 Recovered

South Dakota Shannon 1994, 1995 Established

Tripp 1990 Established

Utah Weber 1993, 1994 Established

Washington Adams 1990 Established

Asotin 1989 Established

Benton 1988-1990 Established

Chelan 1995 Established

Columbia 1995 Established

Ferry 1987 Established

Franklin 1995 Established

Garfield 1989 Established

Grant 1988, 1993 Established

Kittitas 1988 Established

Klickitat 1985, 1989-1990, 1993 Established

Lewis 1992 Established

Lincoln 1989 Established

30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Table 1. Continued.

State . County Release Year Status

Okanogan 1983, 1985-1987,

1991-1993 Established

Skamania 1988 Established

Spokane 1988, 1989, 1990, 1993 Established

Stevens 1985-1989, 1991,

1993-1995 Established

Walla Walla 1985-1986, 1989 Established

Whitman 1984, 1986, 1989 Established

Washington Yakima 1988-1990 Established

Wyoming Johnson 1995 Established

Lincoln 1995 Not established

Natrona 1994 Established

4 Spheno ptera jugoslavica is considered established if larvae or adults are recovered the second year

after release.

latter part of the day from 1700 to 2000 h on calm, warm, dry days. Sites in

Colorado, Idaho, Montana, Nebraska, Utah, Washington, and Wyoming were per-

sonally (R. F. L.) visited during March and 20 plants were collected randomly

within 6 m of the release point and taken to a laboratory where they were dissected

to ascertain larval presence.

The buprestid is considered established if larvae or adults were recovered the

second year after release. Collection of adult S. jugoslavica for redistribution is

recommended when 25% of a 20 root sample is infested with larvae.

RESULTS AND DISCUSSION

Spheno ptera jugoslavica has been released in 11 states and 61 counties (Table

1). Establishment has been confirmed in all states except Arizona, Nebraska, and

Nevada. In Colorado, establishment and redistribution has occurred in Douglas

County from the 1991-1994 releases; no recoveries were made in the other release

counties. In Idaho, confirmed establishment occurred in Jerome and Lincoln

Counties from 1987—1989 releases. In Montana, establishment occurred in Fergus

and Lewis & Clark Counties from 1987-1990 releases. Beetles collected from

Lewis & Clark County were released and later recovered in Sweet Grass County.

In Oregon, establishment occurred in 16 counties from 1987-1990 releases. Adult

S. jugoslavica are strong fliers and have readily colonized uninfested knapweed

stands in Oregon. In South Dakota, establishment occurred in Shannon and Tripp

counties from 1990, 1994, and 1995 releases. In Utah, Weber county establish-

ment occurred from 1993-1994 releases and in Washington county establishment

occurred in 20 counties from 1987-1995 releases. In Wyoming establishment

occurred in Johnson and Natrona counties from 1994-1995 releases (Table 1).

Intrastate collections and redistribution of S. jugoslavica in Montana, Oregon,

Utah, Washington, and Wyoming have been made from established populations.

Sphenoptera jugoslavica is presently well established in the midwestern and

western United States. Extensive collection and redistribution has occurred in 5

states and researchers have noted that the bioagent is redistributing itself naturally.

Sphenoptera jugoslavia will continue to spread unaided to other diffuse knapweed

1998 LANG ET AL.: SPHENOPTERA ESTABLISHED 31

infestations throughout the midwestern and western states. This movement should

be monitored as the beetle invades new states and counties.

LITERATURE CITED

Harris, P. & J. D. Shorthouse. 1996. Effectiveness of gall inducers in weed biological control.

Can. Entomol., 128: 1021-1055.

Howell, J.T. 1959. Distributional data on weedy thistles in western North America. Leafl. West.

Bot., 9: 17-32.

Miiller, H. & D. Schroeder. 1989. The biological control of diffuse and spotted knapweed in

North America—what did we learn? pp. 151-169. In Fay, P. K. & J. R. Lacey (eds.). Proc.

Knapweed symp. 4-5 April 1989, Bozeman, Montana. Montana State Univ. Coop. Ex. Serv.

EB 45. Bozeman, Montana.

Piper, G. L. 1989. Releases of Sphenoptera jugoslavica on diffuse knapweed infestations bor-

dering highway rights-of-way in Eastern Washington. pp. 175-179. In Lacey, J. R. & P. K.

Fay (eds.). Proc. of the Knapweed Symp., Bulletin 1315, Great Falls, MT. Montana State

University Cooperative Extension Service, Bozeman, MT.

Powell, R. D. & J. H. Myers. 1988. The effect of Sphenoptera jugoslavica Obenb. (Col., Bu-

prestidae) on its host plant Centaurea diffusa Lam. (Compositae). J. Appl. Entomol., 106: 25—

46.

Rees, N. E., D. C. Quimby, Jr., G. L. Piper, E. M. Coombs, C. E. Turner, N. R. Spencer & L. V.

Knutson. 1966. Biological control of weeds in the West. Western Soc. Weed Sci., Boze-

man, Montana.

Roché, B. F., Jr. 1994. Status of knapweeds in Washington. Wash. State Univ. Coop. Ext. Serv.

Knapweed Newsletter, 8: 2-4.

Roché, B. F., Jr. & C.J. Talbott. 1986. The collection history of Centaureas found in Washington

State. Agric. Res. Center Res. Bull. XB 0978. Wash. State Univ. Pullman, Washington.

Schroeder, D. 1985. The search for effective biocontrol agents in Europe. 1. Diffuse and spotted

knapweed. pp. 103-119. Jn Delfosse, E. S. (ed.). Proc. VI. Int. Symp. Biol. Contr. Weeds, 19-

25 August 1984, Vancouver, Canada. Agriculture Canada, Ottawa.

Story, J. M. & R. M. Nowierski. 1984. Status of biocontrol knapweed research in Montana. pp.

65-70. In Lacey, J. R. & P. K. Fay (eds.). Proc. Knapweed Symp., 3-4 April 1984, Great Falls,

Montana. Montana State Univ. Coop. Ext. Serv. Bull. 1315. Bozeman, Montana.

Thompson, D. J. & D. G. Stout. 1991. Duration of the juvenile period in diffuse knapweed

(Centaurea diffusa). Can. J. Bot., 69: 368-371.

Watson, A.K. & A.J. Renney. 1974. The biology of Canadian Weeds. 6. Centaurea diffusa

and C. maculosa. Can. J. Plant Sci., 54: 687-701.

Zwolfer, H. 1976. Investigations on Sphenoptera (Chilostetha) jugoslavica Obenb. (Col., Bu-

prestidae), a possible biocontrol agent of the weed Centaurea diffusa Lam. (Compositae) in

Canada. Z. ang. Entomol., 80: 170-190.

Received 25 Jul 1997; Accepted 30 Oct 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 32-38, (1998)

LACANOBIA SUBJUNCTA (LEPIDOPTERA: NOCTUIDAE)

ON TREE FRUITS IN THE PACIFIC NORTHWEST

PETER J. LANDOLT

United States Department of Agriculture, Agricultural Research Service

Yakima Agricultural Research Laboratory

5230 Konnowac Pass Road, Wapato, Washington, USA 98951

Abstract.—Larvae of the noctuid moth Lacanobia subjuncta (Grote & Robinson) were collected

on commercial apple trees, as well as pear and plum trees, in central Washington and north

central Oregon, during June to September 1996. Larvae were reared to adult on foliage of the

tree species on which they were collected. Adult moths were captured in molasses-baited traps

placed in apple orchards, primarily from mid May to early June and again in August 1996 near

Yakima, Washington. In the laboratory, L. subjuncta were successfully reared from egg to adult

on several herbaceous weed species common in local tree fruit orchards. Multiple generations

were successfully reared from egg to adult on apple seedlings grown in a greenhouse.

Key Words.—Insecta, cutworm, apple, host plant, rearing.

Fruitworms and cutworms in the family Noctuidae are occasional and minor

pests of tree fruits. Howell (1993) reviewed the biology of several species of

Noctuidae occurring on foliage and fruit of tree fruits in the Pacific Northwest

(PNW), referring primarily to apple and pear. Three noctuid species were reported

to use tree fruits as primary hosts and are occasional pests in organic orchards or

on backyard trees (Howell 1993). These are the green fruitworm Lithophane an-

tennata (Walker), the speckled green fruitworm Orthosia hibisci Guenee, and the

pyramidal fruitworm Amphipyra pyramidoides Guenee.

All three species of fruitworm are widespread throughout much of temperate

North America (Covell 1984, Crumb 1956), but their pest status in the PNW is

unknown. Recently, damage to apple fruit in Washington has been linked to cut-

worms without identification of the species causing the problem (Wamer 1996,

Landolt 1997). It is not clear what species of Noctuidae may be causing damage

to apple in the PNW because published accounts of collections and identifications

of noctuid larvae from tree fruits are generally lacking.

The objective of this study was to verify the species of noctuids on apple in

central Washington that may be contributing to crop losses, particularly in organic

orchards and orchards using mating disruption to control codling moth, Cydia

pomonella L. Reported here is the identification of a single species, Lacanobia

subjuncta Grote & Robinson, from larvae collected and reared from apple, pear,

and plum from multiple localities in Washington and Oregon. Additionally, the

pattern of adult L. subjuncta activity through the season (indicating the number

of generations per year) and successful larval development on five species of

herbaceous plants common in orchards are reported.

MATERIALS AND METHODS

In order to obtain identifiable adult specimens, all noctuid larvae collected from

fruit trees during 1996 were reared in the laboratory on foliage of the tree species

on which they were collected. Collections were largely from three sources: 1)

weekly systematic sampling for spiders (from beating tray samples) in three apple

1998 LANDOLT: LACANOBIA SUBJUNCTA BIOLOGY 33

orchards (June through mid-October), 2) nonsystematic sampling for larval leaf-

rollers (Torticidae) in apple, pear and plum orchards, and 3) from pest control

advisors sampling in response to visible damage to foliage and fruit. Larvae were

placed in glass jars with field-collected foliage of the host species on which they

were collected. Foliage was added or exchanged as needed and 3 cm of damp

soil was placed in the bottom of jars as a pupation medium. Emerged moths were

pinned and spread as voucher specimens which are deposited in the collection of

the author, in the M. T. James Entomological Collection, Washington State Uni-

versity, Pullman, WA, and in the W. E Barr Entomological Collection at the

University of Idaho, Moscow, ID.

Lacanobia subjuncta were reared in the laboratory for one generation on apple

sucker growth cut in commercial apple orchards, and then for multiple generations

on apple seedlings grown in a greenhouse. Eggs for this colony were obtained

from females that were captured in a light trap placed in an apple orchard during

August 1996. About 60 females were placed individually in 20 ml polystyrene

snap cap vials. Eighteen of these females laid eggs. Vials containing eggs were

held in an environmentally controlled room (22° C, 70% RH) until eggs hatched

and neonate larvae were moved with a fine camel hair brush to ten, 4-liter glass

jars containing apple foliage. Fifty larvae were placed in each jar. Foliage was

added or exchanged as needed and 3 cm of damp soil was added when larvae

were mature, as a pupation medium. About 250 pupae were removed from the

soil, sorted by sex, and placed into paper cups in screened cages for adult emer-

gence.

Subsequent generations were reared in the laboratory on potted apple seedlings

grown in a greenhouse. Eggs were obtained from lab-reared moths placed in 10

x 15 X 20cm plastic boxes with screened lids. Each box contained water and

sugar-water on cotton balls in plastic petri dishes (4 cm diam) and one or two

pairs of adult moths. Eggs or hatching larvae were transferred to 45 x 45 X 45

cm screened cages containing potted apple seedlings (40—60 cm tall). Seedlings

were added as needed, and 3—5 cm of damp vermiculite was placed in the bottom

of cages as a pupation medium. One hundred to 200 larvae were placed in each

cage. Pupae were removed from vermiculite and potting soil, sorted by sex, and

placed into paper cups in screened cages for adult emergence. This colony was

used to provide larvae for assessments of host plant suitability.

The seasonal pattern of L. subjuncta moth activity was documented in two field

tests of moth attraction to food baits. Both trapping experiments were part of a

study of codling moth attraction to sweet baits. Traps for both tests were set up in

early May and were maintained until the third week of September, 1996. Traps

were checked, and moths removed, on most days and bait was replaced every two

weeks. In test A, eight traps were baited with 10% aqueous solutions of molasses

(Grandma’s Molasses, unsulphured, Motts; USA, Stamford, CT) and eight traps

were baited with aqueous solutions of jaggery, a palm sugar extract (Indian Kol-

hapur Jaggery, House of Spices Inc., Jackson Heights, N.Y.) (Landolt 1995). All

16 traps were glass McPhail traps (Newell 1936) containing 200 ml of bait. The

test was conducted within a Golden Delicious apple orchard near Parker, Yakima

County, Washington. Traps were placed at a height of 2 m and 30 m apart. Data

were transformed (square root [x + %2]) and subjected to ANOVA, and treatment

means were compared by F test (SAS Institute 1985) to determine if captures of

34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Table 1. Collections of noctuid larvae in 1996 from commercial tree fruit orchards. Larvae reared

to adult stage in laboratory for identification.

Date Locale Number Host

VI-1-1996 OR, Umatilla Co., Milton-Freewater 2 Apple

VII-1996 WA, Grant Co., Mattawa 2 Apple, Golden

VII-3-1996 OR, Umatilla Co., Milton-Freewater 1 Apple

VII-11-1996 WA, Mattawa 1 Apple, Fuji

VII-24-1996 WA, Yakima Co., Parker 4 Pear, Bartlett

VII-29-1996 WA, Yakima Co., Yakima 1 Apple

VIII-1996 OR, Umatilla Co., Milton-Freewater Z Apple

VIUI-29-1996 WA, Yakima Co., Parker 12 Plum

IX-6-1996 WA, Benton Co., Prosser 6 Apple

IX-12-1996 WA, Benton Co., Prosser 14 Apple

IX-12-1996 WA, Yakima Co., Parker 3 Apple

IX-23-1996 WA, Yakima Co., Parker 1 Apple

IX-30-1996 OR, Umatilla Co., Milton-Freewater 1 Apple, Red Delicious

X-1-1996 WA, Yakima Co., Parker 1 Apple, Red Delicious

X-7-1996 WA, Yakima Co., Parker 1 Apple

X-8-1996 WA, Yakima Co., Parker 1 Apple

L. subjuncta moths with bait types were not equal. In test B, 4 trap designs were

compared, with all traps were baited with 200 ml of 10% molasses. Trap types

were glass McPhail traps, Multipher traps (BioControl Services Inc.), Trappit Dome

Trap (Agrisense, Palo Alto, CA), and open bowl traps (white plastic disposable

cereal bowls suspended by a wire hanger). Six replicates of these 4 trap designs

were maintained through the field season. This experiment was established in one

orchard, but was moved to a second orchard on July 6. Both orchards were Golden

Delicious apple plantings. Data were transformed (square root [k + -%]) and sub-

jected to ANOVA. Significant differences between treatments were determined us-

ing Tukey’s Studentized Range Test (SAS Institute 1985).

Host suitability data was obtained for L. subjuncta larvae on foliage of black

medic (Medicago lupina L.), dandelion (Taraxacum officinale Weber), sow thistle

(Sonchus oleraceus L), bindweed (Convolvulus arvensis L.), redroot pigweed

(Amaranthus retroflexus L.), mallow (Malva neglecta Wallr), and lambsquarters

(Chenopodium berlandieri Mogq.). Five newly hatched larvae were placed into

100 ml wax coated paper cups containing plant foliage as an assay unit. Foliage

was added or replaced as needed until larvae were dead or neared maturity. Three

cm of damp vermiculite was then added as a pupation medium. Pupae were

removed, sorted by sex, and placed in 500 ml clear plastic cups until adult eclo-

sion. This assay was conducted using five cups for each host plant species or tree

fruit variety. Data were recorded for evidence of feeding, pupation and adult

emergence, date of adult emergence, and sex of adults.

RESULTS

Sixteen samples of noctuid larvae were collected from 6 localities in Washing-

ton (the Parker area SE of Yakima, west Yakima, Moxee, Zillah, Mattawa, and

Prosser) and near Milton-Freewater, Oregon (Table 1). Although most samples

were taken from commercial apple orchards, one was from a commercial pear

orchard and one from a commercial plum orchard. Fifty-three larvae were reared

1998 LANDOLT: LACANOBIA SUBJUNCTA BIOLOGY 35

to adult and were determined to be L. subjuncta. Two additional specimens taken

in an apple orchard near Prosser, WA on 12 Sep 1996, were parasitized by an

ichneumonid wasp but are considered to have been L. subjuncta based on col-

oration and pattern. There were no other species of Noctuidae in these samples.

One thousand nine hundred and ninety-eight L. subjuncta moths were captured

in traps baited with sweet baits from May through mid September 1996 (Fig. 1).

Two distinct peaks of moth catches were evident in both plots, from mid May

through mid June, and again in August into early September. Very few moths

were captured during July. The sex ratio of moths captured in traps baited with

sweet baits was 2:3 females to males. In both plots, numbers of moths captured

during the first flight period were higher than in the second (Fig. 1). Numbers of

L. subjuncta males and females captured in traps baited with molasses (2.8 + 0.5

females, 2.6 + 0.5 males) and traps baited with jaggery (2.4 + 0.6 females, 1.5

+ 0.4 males) were not significantly different (F = 3.78, P > 0.05). In the com-

parison of trap types, captures of L. subjuncta moths were significantly reduced

in bowl traps (5.05 + 1.2) compared to McPhail (11.81 + 2.0), Multipher (8.1

+ 1.4), or Dome traps (8.9 + 1.4) (P < 0.05, df = 204, n = 52). There were no

significant differences among mean numbers of moths captured in McPhail, Mul-

tipher, and Dome traps baited with molasses solution.

The percentages of L. subjuncta that survived and developed on plants tested

was low (Table 2), but successful development to the adult stage did occur on

dandelion, sow thistle, bindweed, and mallow, indicating that these plants are

potential hosts. Larvae fed on all species of plants tested, as evidenced by the

presence of frass and larval growth.

DISCUSSION

Lacanobia subjuncta is common and widespread in apple growing areas of

central Washington and the contiguous areas of Oregon. Large numbers of adults

were collected in traps with sweet baits from mid May through much of Septem-

ber. One hundred and fifty-six moths were captured in one night with a light trap

in an apple orchard. Larvae of L. subjuncta were collected at six sites in the

Yakima River Valley, as well as near Mattawa, Washington and Milton-Freewater,

Oregon. It is assumed that this species of moth is widely distributed in irrigated

areas of the PNW. According to McCabe (1980) it occurs throughout much of

North America, including Washington.

Lacanobia subjuncta uses apple and possibly other tree fruit species as a pri-

mary host, although it may also utilize other plant species in and around orchards.

All of our records (Table 1) involve larvae collected on apple, pear, or plum trees,

indicating likely feeding and development on those hosts. Collected larvae were

able to finish development on apple, pear, or plum foliage and numerous larvae

(several hundreds) were successfully reared in the laboratory from egg to adult

on apple foliage. One larva was collected while feeding on an apple fruit, but it

is likely that most larvae collected were feeding on foliage.

Previous host records and rearing records indicate broad polyphagy for L. sub-

juncta. This species has been reported on a wide variety of trees and shrubs

including fruit of blueberry, cherry, and sour cherry (Phipps 1930, Prentice 1962,

Rockburne & Lafontaine 1976, Godfrey 1972, Rings et al. 1992). Given its broad

host range on woody plants, it is not surprising that it occurs on apple, pear, and

36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

NO. MOTHS/TRAP/WEEK

15 21 28 4 1118 25 2 9 16 23 30 6 13 20 27 3 10

MAY JUNE JULY AUG SEPT

NO. MOTHS/TRAP/WEEK

15 21 28 4 1118 25 2 9 16 23 30 6 13 20 27 3 10

MAY JUNE JULY AUG SEPT

Figure 1. Seasonal pattern, as weekly trap captures, of Lacanobia subjuncta moths in Golden

Delicious apple orchards near Parker, Yakima Co., Washington, May—September 1996. Numbers are

calendar dates for beginnings of weeks. Data are pooled from two trapping experiments; molasses and

jaggery solutions in McPhail traps (above) and molasses solution in bowl, Dome, McPhail, and Mul-

tipher traps (below). c

1998 LANDOLT: LACANOBIA SUBJUNCTA BIOLOGY 37

Table 2. Development of Lacanobia subjuncta on foliage of weed species, Yakima County, Wash-

ington, 1996. n = 25.

Common name % Pupation % Adult emergence Pupal weight (mg)

Dandelion 24 20 293.3 + 8.4

Black Medic 16 0

Sow Thistle ae 28 285.9 + 17.1

Bindweed 36 28 319.1 + 18.3

Pigweed 0 0

Mallow 24 20 292.2 + 27.6

Lambsquarters 4 0

plum, although these plants have not been reported previously as hosts of L.

subjuncta. Godfrey (1972) also reported several herbaceous plants (weeds and

vegetables) as hosts. It also completed development on four common weed species

found in local orchards; dandelion, sow thistle, mallow, and bindweed. Godfrey

(1972) also listed dandelion as a host.

The possible pest status of L. subjuncta, as well as that of O. hibisci, L. an-

tennata, and A. pyramidoides, on PNW tree fruits, will require additional study.

However, recent reports of damage to fruit in organic apple orchards attributed

to cutworms (Wamer 1996) may have been caused by L. subjuncta. Lacanobia

subjuncta adults were abundant in the Yakima River Valley, while no adults of

O. hibisci, L. antennata, or A. pyramidoides were collected in traps baited with

sweet materials or in intermittent light-trapping in apple orchards. Adults of the

univoltine O. hibisci and L. antennata may fly early in the spring (Howell 1993)

before our trapping tests were initiated in mid-May.

Lacanobia subjuncta appears to be bivoltine. Two distinct peaks in the numbers

of adults captured in traps, first in late May into early June and again in August,

match the presence of larvae on apple trees in late June through July and again

in September through October. Two broods were also indicated by McCabe (1980)

for this species, based on collection dates for 1100 museum specimens examined.

Large numbers of both sexes of this moth were captured in McPhail, Multipher,

and Dome traps baited with solutions of molasses and McPhail traps baited with

diluted jaggery. Molasses and the Multipher and Dome traps are generally avail-

able and could be used to monitor or sample L. subjuncta adults.

ACKNOWLEDGMENT

I thank E. Milecsky, J. E. Turner, L. Lampson, T: R. Unruh and N. Simone for

contributing specimens collected on apple, pear, and plum trees. Tim McCabe

kindly verified the identification of moths obtained as larvae on apple as Laca-

nobia subjuncta. Technical assistance was provided by D. Broer, C. Fein, R. W.

Hofstetter, D. L. Larson, C. L. Smithhisler, and M. Weiss, and R. S. Zack and H.

C. Reed provided helpful critiques of the manuscript.

LITERATURE CITED

Covell, C. V. Jr. 1984. Field Guide to the Moths of Eastern North America. Peterson Field Guide.

Houghton-Mifflin Co.

Crumb, S. E. 1956. The larvae of the Phalaenidae. USDA Tech. Bull. No. 1135. Washington, D.C.

38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Godfrey, G. L. 1972. A review and reclassification of larvae of the subfamily Hadeninae (Lepidoptera:

Noctuidae) of America north of Mexico. USDA, ARS, Tech. Bull., 1450.

Howell, J. E 1993. Green fruitworms and cutworms. Jn Beers, E. H., J. E Brunner, M. J. Willett, &

G. M. Warner (eds.). Orchard Pest Management. A Resource Book for the Pacific Northwest.

Good Fruit Grower, Yakima, Washington.

Landolt, P. J. 1995. Attraction of Mocis latipes (Lepidoptera: Noctuidae) to sweet baits in traps. Fla.

Entomol., 78: 523-530.

McCabe, T. L. 1980. A reclassification of the Polia complex for North America (Lepidoptera: Noc-

tuidae). N. Y. St. Mus. Bull. No. 432, Albany.

Newell, W. 1936. Progress report on the Key West (Florida) fruit fly eradication project. J. Econ.

Entomol., 29: 116-120.

Phipps, C. R. 1930. Blueberry and huckleberry insects. Maine Agric. Exp. Stat. Bull., 356: 107-221.

Prentice, R.M. 1962. Forest Lepidoptera of Canada. Nycteolidae, Noctuidae, Notodontidae, Liparidae.

Can. Dept. For. Biol., Ottawa, Ontario., 128: 77-281.

Rings, R. W., E. H. Metzler, E J. Amold & D.H. Harris. 1992. The owlet moths of Ohio. Order

Lepidoptera, Family Noctuidae. Ohio Biol. Survey Bull. (NS), 9: 1-184.

Rockburne E. W. & J. D. Lafontaine. 1976. The cutworm moths of Ontario and Quebec. Canada

Dept. Agric. Res. Publ. 1593.

SAS Institute. 1985. SAS user’s guide: statistics, version 5 ed. SAS Institute, Cary, NC.

Warner, G. 1996. Goodbye codling moth, but hello cutworms. Good Fruit Grower, 47: 17-18.

Received 31 Mar 1997; Accepted 10 Jun 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 39-46, (1998)

NEW SPECIES OF PHYLLOPHAGA HARRIS FROM

MESOAMERICA (COLEOPTERA:

MELOLONTHIDAE; MELOLONTHINAE)

MIGUEL-ANGEL MORON

Departamento de BiosistemAatica de Insectos

Instituto de Ecologia, A.C.

Apartado Postal 63, Xalapa, Veracruz 91000, México

Abstract.—Phyllophaga ilhuicaminai Mor6n NEW SPECIES is described from western and cen-

tral localities of Mexico, and Phyllophaga caanchaki Mor6n NEW SPECIES is described from

southeastern Mexico and northern Guatemala. Both species have wide geographical distributions,

and are not included in any species group of the genus. Drawings of sclerotized male and female

genital structures are included.

Key Words.—Insecta, May beetles, Phyllophaga, Taxonomy, Mexico, Guatemala.

Restiimen.—Se describe Phyllophaga ilhuicaminai Morén, NUEVA ESPECIE con base en 70

ejemplares recolectados en bosques tropicales caducifolios, encinares, matorrales espinosos y

terrenos agricolas de los estados de Michoacan, Jalisco, México, Guerrero, Morelos y Puebla,

México. También se describe Phyllophaga caanchaki Mor6n, NUEVA ESPECIE con 36 ejem-

plares obtenidos en bosques tropicales caducifolios y perennifolios, y terrenos agricolas de los

estados de Chiapas, Quintana Roo y Campeche, México, asi como en la provincia de Petén,

Guatemala. Debido a la peculiar combinacién de caracteres morfoldégicos, no es posible incluir

a ninguna de estas dos especies en alguno de los grupos de especies del género hasta ahora

definidos. Se incluyen dibujos de las estructuras genitales masculinas y femeninas de las dos

especies y se comentan sus relaciones con otras especies mesoamericanas.

The structure of tarsal claws has been used as one of the primary characters in

the taxonomy of Rhizotrogina and Phyllophaga species. Bates (1887-1889), Say-

lor (1942), Sanderson (1958), Mor6én (1986) and other authors, proposed groups

of species, subgenera and genera mainly based on the shape, proportions and

symmetry of claws. Nearly 500 American species of Phyllophaga (sensu lato)

show toothed, bifid or notched tarsal claws, but less than 20 possess, finely serrate

or micro-dentate claws. These species are not easy to place in subgenera or species

groups and are usually considered as “incerta sedis’’.

During collecting trips to many localities in western, central and southeastern

Mexico, and while examining museum collections from Mexico, the U.S.A. and

Canada, I found nearly one hundred specimens of two species of Phyllophaga

with different, “‘simple’’ tarsal claw structure, not related to any of the species

group defined by Mor6én (1986). This paper describes males, females, and their

variation and gives the distribution of two new species. The characters and terms

used in the descriptions are those of Mordén (1986). Drawings were made with

the aid of a camera lucida and Carl Zeiss stereomicroscope;.measurements were

obtained with ocular micrometer or caliper.

Depository Abbreviations.—California Academy of Sciences, San Francisco

(CASC); the Natural History Museum, London (BMNH); Canadian National Col-

lection, Ottawa (CNC); Instituto de Biologia, UNAM, México City (IBUNAM);

Instituto de Ecologia, Xalapa, Veracruz (IEXA); University of Nebraska State

Museum, Lincoln, Nebraska (UNSM); Field Museum Natural History, Chicago

40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

(FMNH); Universidad del Valle de Guatemala, Guatemala City, (UVG); Instituto

Nacional de Investigaciones Forestales, Agricolas y Pecuarias, SAGAR, Celaya,

Guanajuato (INIFAP); Benemérita Universidad Aut6noma de Puebla (DICA); A.

& H. Howden, Ottawa (AHHC); G. Nogueira, Guadalajara, Jalisco, México

(GNPC); S. Pokorny, Praha, Czech Rep. (SPPC); J. P. Beraud, Cuernavaca, Mo-

relos, México (JBPC); C. Deloya (ACDC) and M. A. Moron, Xalapa, Veracruz

(MXAL) México.

Phyllophaga ilhuicaminai Morén, NEW SPECIES

(Figs. 1-7)

Types.—(described from 46 males and 24 females). Holotype, male; MEXICO.

MICHOACAN: Jungapeo, 29 Jun 1967, G. Halffter; deposited: Canadian National

Collection, Ottawa. Allotype, female; MEXICO. GUERRERO: Chilpancingo, Palo

Blanco, 20 Jun 1987, bosque pino-encino, 1140 m, Morén, Deloya, Delgado;

deposited: M. A. Moré6n collection, Xalapa. Paratypes: same data as holotype (14

males, 3 females); same data as allotype (5 males, 3 females); same data as

allotype except: Apr—May, 1987 (3 males, 1 female). GUERRERO: Acahuizotla,

11 Jul 1986, bosq. trop. Mediano subperennifolio, luz 2000—2300 hrs, 650 m, L.

Delgado (1 female); same data except: 4 Aug 1986, 2000-2200 hrs (1 female);

19 Jun 1987 (1 female); 18 Jun 1985, 750 m (3 males, 1 female); 4 Sep 1985 (1

female); 6 Jun 1986, 2100—2200 hrs, Ruiz and Martinez (1 male). JALISCO:

Chamela, 2 Aug 1985, luz, R. Ayala (2 males); Est. Biol. Chamela, 23 Aug 1985,

luz, R. A. Usela (1 male); Casimiro Castillo, 17 Jun 1994, 390 m, G. Nogueira

(1 male). MICHOACAN: Coalcomaén, Los Laureles, 7 Aug 1983, T. W. Taylor (2

males). MEXICO: Santo Tomas de los Platanos, Jun 1993, S. Pokorny (8 males,

11 females). MORELOS: Tepoztlan, 21 May 1990, J. P. Beraud (1 male). PUEB-

LA: Atlixco, La Joya, 14 Mar 1996, suelo cultivo estatis, 1770 m, A. Aragon (1

male); same data except: 4 Jun 1996, luz 2030—2100 hrs, A. M. Tapia, A. Aragon,

R. Rojas (2 males); Tzicatlacoyan, 17 May 1996, 2000 m, rafces de encino, A.

Aragon (2 males); same data except: 15 Jul 1996, luz 2000—2100 hrs, S. Caselin,

A. Aragon (2 males). Paratypes are deposited at CASC, BMNH, CNC, UNSM,

IBUNAM, IEXA, DICA and in the private collections AHHC, GNPC, SPPC,

JBPC, ACDC and MXAL.

Holotype.—Male. Head and pronotum shining testaceous, elytra similarly colored but slightly

Opaque; mouth parts, sterna, pygidium and legs shining testaceus. Clypeus semicircular, wider than

long (2.5:1), anterior border curved with margins slightly elevated, surface nearly flattened, rugo-

punctate with some scattered microscopic setae. Fronto-clypeal suture slightly sinuated, deeply im-

pressed at sides, obsolete at midline. Frons wider than long (1.3:1) convex, coarsely rugoso-punctate,

with erect, slender setae. Antenna 9 segmented, with 3 segmented club; lamellae 1.5 < longer than

the length of the basal segments combined, segment 5 longer than 4, with a short, rounded tubercle;

segment 6 very short. Frons 2.5 X wider than each eye dorsal diameter. Canthus short and narrowed,

with 13-14 setae. Labrum bilobed, widely sinuated, with scattered slender setae. Mentum slightly

concave, rugo-punctate, with lateral setae slender, anterior border widely sinuated. Pronotum wider

than long (1.5:1) and 2.8 X wider than frons. Pronotal disk with round punctures separated by more

than width of one or two punctures near the midline, becoming confluent on sides; lateral borders

widely angulated, marginal bead crenulate, with long slender setae; anterior angles obtuse, not prom-

inent; posterior angles widely obtuse, rounded. Scutellum 1.3 X wider than long, with 18-20 small

punctures. Elytron 2.8 X longer than wide, densely punctate, glabrous; epipleural border narrow,

extended along the complete margin, provided with dense fringe of short slender setae; humeral calla

rounded, prominent; apical calla rounded. Metathoracic wings completely developed. Propygidium

1998 MORON: NEW MESOAMERICAN PHYLLOPHAGA 41

ny NS ah guasy sug

erly?

Figures 1-7. Phyllophaga ilhuicaminai NEW SPECIES. Figure 1. Male protarsal claw. Figure 2.

Lateral view of paramera and partially exposed aedeagus of holotype. Figure 3. Distal view of the

paramera and aedeagus of the holotype. Figure 4. Detail of the distal angle of the paramerus of paratype

from Atlixco, Puebla, showing the preapical flattened tooth. Figure 5. Lateral view of complete genital

capsule of the same paratype. Figure 6. Female protarsal claw. Figure 7. Female ventral genital plates

of allotype. Scale lines = 1 mm.

42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

shining, fine and dense punctate-setose. Pygidium convex, shining, with dense, round, shallow regu-

larly distributed setiferous punctures; apical margin with 15 long, slender setae; basal margin effaced

at the middle. Pterosternon with long, dense, yellow vestiture. Visible abdominal stermnites 3 and 4

slightly flattened and polished near the midline; sternite 5 with sparce punctures and slender setae at

the middle; anal plate slightly prominent, convex, rugo-punctate, with 70—80 slender, short and long

mixed setae. Protibiae shorter than protarsi (1:1.7), with external border tridentate, preapical spur long,

Straight, acute, shorter than protarsomerus 2. Mesotibiae with one oblique, strong, setiferous, transverse

carina; upper apical spur straight, narrow, 1.3 X longer than the lower spur. Metatibiae shorter than

metatarsi (1:1.4), with one oblique, strong setiferous transverse carina; apical spurs articulated with

the border, upper spur curved, shorter than metatarsomerus 2, and 1.4 X longer than the lower spur.

Tarsomeres semicylindrical, elongated, with enlarged apex, some setae around the apex and two lines

of setae ventrally. Tarsal claws symmetrical, similar on all legs, tooth very short, postmedian, acute

and followed by serrate basal edge (Fig. 1). Genital capsule with shortened paramera, fused in a ring

shaped structure, apex slightly sinuated, bearing a large ventral bidentate projection which is angulated

posteriorly (Figs. 2-3). Paramera with flattened teeth near apex of distal side (Fig. 3). Aedeagus long,

with semitubular, sclerotized support surrounded by patches of spinules, and one dorsal sclerotized

rod with apical setae (Figs. 2, 5). Tectum uniformly convex. Length of genital capsule from the apex

of parameres to the border of basal piece: 4.5 mm. Total body length: 17 mm. Humeral width: 6.5

mm.

Alloty pe-—Female. Similar to the male except as follows: clypeus slightly trapezoidal; clypeus and

frons more rugo-punctate; antennal club 0.6 the length of the basal segments combined; pygidium

semitriangular, more convex toward the apex. Visible abdominal sternites 3 to 5 convex, with scattered

setiferous punctures; anal plate convex, punctate, with 65—70 slender, short and long setae. Apical

spurs of metatibiae curvate. Tarsal claws dentate, with large, median, acute tooth followed by serrate

basal edge (Fig. 6). Ventral genital plates slightly sclerotized, symmetrical, nearly ovate, with setae

on the distal curvate projection (Fig. 7). Total body length: 16.2 mm. Humeral width: 6.1 mm.

Type Locality.—Jungapeo de Juarez, near Zitacuaro, state of Michoacan, Mex-

ico (aprox. 19°28’ N; 100°30’ W).

Variation.—Paratype males are similar to the holotype except as follows: some

Specimens have the antennal club slightly shortened; frontoclypeal carina granu-

lose or rugose; anterior half of the pronotal lateral borders slightly sinuated; py-

gidium with fewer setae; tarsal claws with tooth effaced; flattened teeth on the

distal side of paramera more elongate and more acute and smaller (Fig. 4); ventral

bidentate process of the paramera shortened (Fig. 5); sclerotized dorsal rod of the

aedeagus with ventral preapical tubercle (Fig. 5); total body length: 12.0—16.8

mm, humeral width: 4.0—6.5 mm. Paratype females are similar to the allotype

except as follows: pronotum and pygidium more densely punctate; total body

length: 13.5—16.5 mm; humeral width: 5.5—6.5 mm.

Biological Data—Specimens of P. ilhuicaminai were collected at lights in

tropical deciduous and subdeciduous forests and oak forests located from 150 and

2000 m of altitude, at night between 2000—2300 h. In March an adult was col-

lected from its pupal cell 30 cm deep in soil underneath flower crops of Limonium

sinuatum (Asteraceae); during May adults were collected from organic soil under

the oak trees on which they feed at night. This species is found primarily on the

warmest slopes of Transverse Neovolcanic Axis (Fig. 8). Phenology: May (6),

June (55), July (2), August (6), September (1). Other species of Phyllophaga

flying at the same time were P. (Phytalus) obsoleta (Blanchard), Phyllophaga

(s.str.) ravida (Blanchard) and Phyllophaga (s.str.) vetula (Horn).

Remarks.—Phyllophaga ilhuicaminai is not closely related to any other Mex-

ican species. Specimens from localities in the state of Puebla (eastern population)

have paramera with a shortened ventral bidentate process; specimens from Guer-

1998 MORON: NEW MESOAMERICAN PHYLLOPHAGA 43

@ = Phyllophaga ilhuicaminai

ma =s wPhyllophaga caanchaki

Figure 8. Distribution of Phyllophaga ilhuicaminai NEW SPECIES and Phyllophaga caanchaki

NEW SPECIES.

rero (southern population) have the ventral bidentate process long but less an-

gulate; typical specimens are from Jalisco, Michoacan, México and Morelos (Fig.

8). The elongate body shape, nine-segmented antenna, shape of the tarsal claws

and male genital capsule will aid in the recognition of this species.

Etymology.—Derived from Ilhuicamina, (“‘sky bowman’’) second name of the

fifth Aztec king Motehuzoma I (Sime6én, 1988), who extended and consolidated

the Aztec empire between 1440-1468 A.D. (Leon-Portilla, 1978).

Material Examined—See Types.

Phyllophaga caanchaki Moron, NEW SPECIES

(Figs. 9-14)

Types.—(described from 15 males and 21 females). Holotype, male; MEXICO.

CHIAPAS: Yaxchilan, 7 May. 1981, 150 m, Selva Brosimum, E. Fuentes; depos-

ited: M. A. Mor6én collection, Xalapa. Allotype, female; MEXICO. QUINTANA

ROO: 4 km S Tres Garantias, 25 Jun 1988, selva mediana subperennifolia, trampa

luz 1900-2200 hrs, D. Garrido y V. A. Poot; deposited: M. A. Morén collection,

Xalapa. Paratypes: same data as holotype except 31 May 1981 (1 male); 14 Jun

1981 (3 males); same data as allotype (1 male, 3 females); MEXICO. QUINTANA

ROO: Xul Ha, 3 Jun 1974, Y. Dominguez (1 male); km 146 carr. Chetumal-Puerto

Juarez, 17 Mar 1982, A. Ibarra (1 male); Felipe Carrillo Puerto, 28 May 1984,

10 m, luz neén 2100 hrs, J. E Camal (1 male, 2 females); CAMPECHE: Escar-

cega, 23 Apr 1962, trap light, EK Islas (2 males); GUATEMALA. PETEN: cart.

44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Flores-Tikal, Jun 1993, 150 m, bosq. tropical, M. A. Mor6én (4 males, 9 females);

same data except C. Deloya (1 male, 6 females). Paratypes are deposited at CASC,

BMNH, FMNH, UVG, IBUNAM, INIFAP, IEXA, and in the collections ACDC,

GNPC and MXAL.

Holotype.—Male. Frons shining dark brown, pronotum and elytra testaceous, with dense vestiture

of short, erect setae; clypeus, mouth parts, sterna, pygidium and legs shining testaceous. Clypeus wider

than long (3.6:1), anterior border slightly sinuated with elevated margins, surface concave, with sparse,

shallow punctures and scattered short setae. Fronto-clypeal suture sinuated, impressed at sides, obsolete

at midline. Frons wider than long (2:1) scarcely convex, rugo-punctate, with erect slender setae

throughout. Antennae 9 segmented, with 3 segmented club, lamellae 1.3 X longer than the length of

the basal segments combined; segment 5 without an anterior process; segment 6 very short, ring-like.

Frons 1.5 X wider than dorsal diameter of eye. Canthus narrowed, with 15—16 setae. Labrum bilobed,

widely sinuated, with scattered slender setae. Mentum slightly concave, with sparse punctures and

slender setae, anterior border widely sinuated. Pronotum wider than long (1.5:1) 2.8 X wider than

frons. Pronotal disk setiferous, with shallow, round punctures separated by the width of one puncture

near the midline, becoming slightly confluent on sides; lateral borders widely rounded, marginal bead

crenulate with long slender setae; anterior and posterior angles obtuse, rounded. Scutellum 1.7 X wider

than long with 7 shallow, small punctures. Elytron 3 X longer than wide, setiferous, densely rugo-

punctate; epipleural border narrowly extended along the entire margin, provided with short slender

setae; humeral calla rounded, prominent; apical calla rounded. Metathoracic wings fully developed.

Propygidium shining, with shallow sparse punctures and scattered short setae. Pygidium convex, se-

tiferous, shining, with round, shallow irregularly distributed punctures; apical margin with 10 long,

slender setae; basal margin effaced at the middle. Pterosternon with shortened, yellow setae. Visible

abdominal sternites 2 to 5 slightly depressed, with scattered short setae near the midline; anal plate

slightly prominent, nearly convex, punctate, with 40-50 long, slender setae. Protibiae slightly shorter

than protarsi (1:1.6) with external border tridentate; preapical spur long, straight, acute, as long as

protarsomerus 2. Mesotibiae with one oblique, strong, setiferous transverse carina; upper apical spur

straight, narrow, and 1.2 X longer that the lower spur. Metatibiae slightly shorter than metatarsi (1:

1.3), with one oblique, strong, setiferous transverse carina; apical spurs articulated with the border,

upper spur lanceolate, shorter than metatarsomerus 2, and 1.2 X longer than the lower spur. Tarsomeres

semicylindrical, elongated, with enlarged apex, some setae around the apex and 2 dense lines of setae

ventrally. Tarsal claws symmetrical, similar on all legs, without teeth on the slightly serrate edge (Fig.

9). Genital capsule with elongate paramera, fused at base and apex, the apex is slightly acute and

without a ventral process (Figs. 10-12). Aedeagus long, with sclerotized support rod and dorsal needle-

like process (Fig. 12). Tectum uniformly convex. Length of genital capsule from the apex of paramera

to the border of basal piece: 4 mm. Total body length: 12 mm. Humeral width: 4.5 mm.

Alloty pe-—Female. Similar to the male except as follows: antennal club as long as the basal seg-

ments combined; pygidium nearly flattened, with the apex slightly narrowed; visible abdominal ster-

nites 3 to 5 convex, with scattered setiferous punctures. Protibiae as long as the protarsi; apical spurs

of metatibiae with rounded apex; tarsal claws dentate, acute, large, antero-median tooth with the edge

feebly serrate (Fig. 13). Ventral genital plates poorly sclerotized, symmetrical, nearly ovate, with setae

on the distal curved process (Fig. 14). Total body length: 12 mm. Humeral width: 5 mm.

Type Locality.—Yaxchilan, Ocosingo, state of Chiapas, México (aprox. 17°10’

N; 91°18’ W).

Variation.—Paratype males are similar to the holotype except as follows: some

specimens have antennal segments 3 and 4 nearly fused; the dorsal and pygidial

vestiture is more or less dense; total body length: 11.0—12.5 mm; humeral width:

4—5 mm. Female paratypes are similar to the allotype except as follows: the

pronotum is more densely punctate and the teeth on the external border of the

protibiae are more rounded; total body length: 11.0-12.5 mm; humeral width:

4.5—5.0 mm.

Biological Data—Specimens of P. caanchaki were collected at lights in the

tropical subdeciduous forests and tropical rain forests located between 10 and 150

1998 MORON: NEW MESOAMERICAN PHYLLOPHAGA 45

Figures 9-14. Phyllophaga caanchaki NEW SPECIES. Figure 9. Male protarsal claw. Figure 10.

Ventral-apical view of paramera of holotype. Figure 11. Distal view of same paramera. Figure 12.

Lateral view of complete genital capsule of holotype. Figure 13. Female protarsal claw. Figure 14.

Female ventral genital plates of allotype. Scale lines = 1 mm.

m of altitude, at night from 1900—2200 h. This species is known from tropical

lowlands of the Petén, Guatemala and the southern Yucatan Peninsula (Fig. 8).

Phenology: March (1), April (2), May (5), June (29). Other species of Phyllo-

phaga flying at the same time were P. (Chlaenobia) vexata (Horn) and P. (s.str.)

menetriesi (Blanchard).

Remarks.—Phyllo phaga caanchaki is not closely related to any other Mexican,

Central American or Antillean species. The small size, elongate body shape, nine

segmented antennae, concave clypeus, dorsal vestiture, shape of tarsal claws and

male genital capsule will aid in the recognition of this species. Males of P. pan-

orpa Sanderson from Florida, U.S.A. also possess tarsal claws without teeth and

dorsal vestiture composed of short dense setae (Woodruff & Beck 1989), but the

number of antennal segments, body size and genital structure are different.

Etymology.—Derived from the second part of the name Ah cacao-caan chak,

celebrated Mayan king who was born in Campeche and promoted important de-

46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

velopment of the city of Tikal around 700 A.D. (data from exhibitions in Ar-

chaeological Museum at Tikal, Guatemala).

Material Examined.—See Type.

ACKNOWLEDGMENT

I am indebted to the following individuals and institutions for the loan or

donation of specimens from their collections: J. MacNamara, Canadian National

Collection; H. E Howden, Ottawa; B. C. Ratcliffe, University of Nebraska, (Lin-

coln, Nebraska); E. Smith, Field Museum Natural History, Chicago; S. Pokorny,

Praha, Czech Republic; EK Noguera, Est. Biol. Chamela, Universidad Nacional

Auténoma de México; A. Aragon, Benemérita Universidad Aut6énoma de Puebla,

Puebla; A. Marin, Instituto Nacional de Investigaciones Forestales Agricolas y

Pecuarias-Secretaria de Agricultura Ganaderia y Desarrollo Rural, Celaya, Gua-

najuato; J. P. Beraud, Cuernavaca, Morelos; G. Nogueira, Guadalajara, Jalisco; E.

Fuentes, Mexico City; D. Garrido, Mérida, Yucatan; J. EK Camal, Carrillo Puerto,

Quintana Roo; and C. Deloya, Xalapa, Veracruz.

This study was undertaken while the author was on sabbatical (1996) at the

Departamento de Investigaciones en Ciencias Agricolas (DICA), Benemérita

Universidad Aut6noma de Puebla (BUAP), México. I acknowledge the aid of

Ana Maria Tapia, Sandra Caselin, Agustin Aragon and Ratl Rojas (DICA) during

the capture and preparation of some specimens described in this paper. Field trips

in the state of Puebla were conducted with the support of project FB286/H125/

96, CONABIO, México-BUAP. The present paper was published with the funds

of Departamento de Biosistematica de Insectos, Instituto de Ecologia, Xalapa

(902-02).

LITERATURE CITED

Bates, H. W. 1887-1889. Biologia Centrali Americana, Insecta, Coleoptera. Vol. 2, Part II: 161-416.

Leén-Portilla, M. 1978. ‘‘Casi cien afios de grandeza del pueblo del sol’. pp. 787-804. Jn Historia

de México, tomo IV. Salvat Mexicana de Ediciones, México.

Moron, M. A. 1986. El género Phyllophaga en México. Morfologia, distribucién y sistematica su-

praespecifica (Insecta: Coleoptera). Publ. No. 20, Instituto de Ecologia, México.

Sanderson, M. W. 1958. Faunal affinities of Arizona Phyllophaga, with notes and descriptions of new

species. J. Kansas Entomol. Soc., 31: 158-173.

Saylor, L. W. 1942. Notes on beetles related to Phyllophaga Harris with descriptions of new genera

and subgenera. Proc. U.S. Nat. Mus. 92 (3145): 157-165.

Simeon, R. 1988. Diccionario de la lengua nahuatl o mexicana. Siglo XXI Editores, México.

Woodruff, R. E. & B. M. Beck. 1989. The scarab beetles of Florida (Coleoptera: Scarabaeidae). Part

II. The May or June beetles (Genus Phyllophaga). Arthropods of Florida and Neighboring Land

Areas, Vol. 132. Florida Department of Agriculture and Consumer Services, Gainesville.

Received 28 Oct 1996; Accepted 31 July 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 47-54, (1998)

COLLEMBOLA FROM THE CANOPY OF A MEXICAN

TROPICAL DECIDUOUS FOREST'

JOSE G. PALACIOS-VARGAS, GABRIELA CASTANO MENESES?, &

JosE A. GOMEZ-ANAYA

Laboratorio de Ecologia y Sistematica de Microartrépodos

Depto. Biologia, Fac. Ciencias, UNAM, 04510 México, D.F

Abstract.—Sampling of the Arthropoda in a tropical deciduous forest was carried out at Chamela,

Mexico. The Collembola community was emphasized because it was the most abundant arthro-

pod group in the samples. Salina banksi (Paronellidae) was the most abundant and constant

Collembola species. Seasonal variation of the abundance was positively correlated with precipi-

tation and temperature. Canopy and shrub layer from Chamela shared only 12 Collembola species

with the leaf litter and soil from the same study area. From reports in the literature, it was

determined than Chamelan Collembola are more abundant than elsewhere.

Key Words.—Insecta, Collembola, Chamela, Canopy.

Collembola are usually associated with edaphic environments, where they con-

stitute an important group because of their abundance and their role in nutrient

cycling in the soil. Collembola can be abundant in other environments such as in

the soil and litter accumulated in epiphytic plants as demonstrated by Palacios-

Vargas (1981) e.g., so called suspended soils of Delamare-Debouteville (1948).

The role of Collembola in the canopy can be as important as that in the soil,

especially in environments where there is an accumulation of organic matter in

the canopy, as occurs in the tropical deciduous forest of Chamela.

Current studies of South American epiphytic environments provided descrip-

tions of new collembolan taxa (Bretfeld 1994), and studies using insecticidal

fogging have shown that Collembola are one of the dominant groups in the canopy

(Guilbert et al. 1995).

MATERIALS AND METHODS

The study was carried out in the Tropical deciduous forest at the Chamela

Biological Station in the state of Jalisco, México (19°30’ N, 105°03’ W). Bullock

(1988) showed that rainy season is from July to October and dry season from

September to June. The vegetation was described in detail by Lott (1985).

For sampling, three watersheds, designated 1, 4 and 4A by Cervantes et al.

(1988), were selected because their primary production is similar (Saharukaén &

Maass 1990). The tree layer sampled was 25 m tall.

Seven fumigations were carried out to include both rainy (August and Septem-

ber 1992, July 1993) and dry seasons (May and November 1993, and February

and May 1994). We used a fogging machine (Dyna fog) and a natural pyrethum

as the insecticide (Resmethrin 3% in kerosene solution) applied before sun rise

(0400 to 0600 h), following the method of Erwin (1983). For every fumigation,

a 100 m’ area was delimited and 50 funnels (0.5 m of diameter) were hung

randomly in the shrub layer about 50 cm above the floor forest. Specimens that

' Project DGAPA IN2078/91, UNAM.

2 Project PADEP, UNAM 003004.

48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

fell on the funnels were collected by washing with alcohol (80%) some 5 h latter.

In 5 fumigations, the fauna of ten funnels were quantified separately to estimate

group distribution on the sampling surface.

Sampling of the shrub layer was done every two weeks from August 1991 to

July 1993 using five malaise traps per watershed. In the first year, sampling was

done in watersheds 1 and 4 and during the second year it was done in watersheds

1 and 4A.

The specimens were sorted and counted into orders, except the Collembola

which were mounted and identified to species. Differences in the abundance of

Collembola among watersheds and months were tested with ANOVA. The dis-

tribution of the Collembola on the sample surface was analyzed by standard de-

viation among some funnels samples in the same region and time. Simple re-

gression analysis (Zar 1984) was performed between abiotic (temperature and

precipitation) and biotic parameters, such as number of Collembola and abundance

of other arthropods at all sites.

Rainy and dry seasons were compared in terms of species richness and com-

position, Hill Diversity Indices, Shannon-Weaver Diversity Index and Pielou’s

Evenness Index (Ludwig & Reynolds 1988) for the canopy and shrub layer.

RESULTS

A total of 25 species of Collembola were found in both habitats (Appendix I).

Twenty-five groups of Arthropoda were represented in the canopy samples (Table

1). A total of 1,098,248 specimens were collected with a mean density of 15,986

specimens/m?’; 95% of which were Collembola (1,044,032) with nineteen species

found (Table 2). Salina banksi MacGillivray (Paronellidae) was the most abundant

species of Collembola, reaching 89.59% of the total Collembola. A remarkable

seasonal behavior was noted. This species was most abundant in the rainy season

(July to October) when it represented 89.9% of all Collembola trapped but in the

dry season this declined to 65% of all Collembola. Seira spp. and Deuterosmin-

thurus maassius Palacios-Vargas & Gonzalez were the next most abundant species

of Collembola collected during both the rainy season (8.4% of the total abun-

dance) and the dry season (19% of the total abundance). Mean density of Col-

lembola from the canopy was estimated to be 15,197 specimens/m?.

Both seasons had 19 species and similar composition, the dominant species was

Salina banksi in rainy season according to N2 of Hill diversity, whereas the same

species and Seira dubia were dominant in dry season (Table 3). Shannon and

evenness indices were higher in dry season because of the decreased number of

S. banksi (Table 3).

The Collembola abundance showed heterogenety among foggings (Table 4)

with distribution being more homogenous in the rainy months than in dry months.

There was significant positive correlation between abundance of Collembola

and monthly average of precipitation (r = 0.83, df = 5, P = 0.02). Similar results

were obtained when this parameter was correlated with more abundant species,

such as Salina banksi (r = 0.83, df = 5, P = 0.02), Seira dubia (r = 0.82, df =

5, P = 0.02), S. knowltoni (r = 0.83, df = 5, P = 0.02) and S. bipunctata (r =

0.83, df = 5, P = 0.02). However, there was no significant correlation between

temperature and abundance of Collembola (yr = 0.31, df = 5, P = 0.49). There

was a Significant negative correlation between temperature and abundance of Co-

1998 PALACIOS-VARGAS ET AL.: CANOPY COLLEMBOLA

Table 1. Mean density and relative abundance of Arthropoda from canopy in Chamela.

Taxa

Collembola

Acari

Hymenoptera

Aranea

Diptera

Coleoptera

Homoptera

Psocoptera

Thysanoptera

Larvae

Orthoptera

Hemiptera

Isopoda

Dyctioptera

Lepidoptera

Pseudoscorpionidae

Isoptera

Chilopoda

Thysanura

Neuroptera

Embioptera

Scorpionida

Mecoptera

Solifuga

Odonata

Total

Abundance

1,044,032

13,691

9850

7333

4920

3872

3598

2314

2310

2114

919

838

688

1,098,248

Percentage

95.06

1.25

0.90

0.67

0.45

0.35

0.33

0.21

0.19

0.08

0.06

0.04

0.03

0.01

<0.01

100

Table 2. Seasonal abundance of canopy Collembola from Chamela.

Taxa

Ceratophysella gibbosa

Xenylla humicola

Brachystomella minimucronata

Pseudachorutes subcrassoides

Aethiopella sp.

Neotropiella quinqueoculata

Entomobrya ca. californica

Seira bipunctata

S. dubia

S. knowtoni

Lepidocyrtus finus

L. gr. lanuginosus

Salina banksi

Sphaeridia pumilis

Sminthurinus ca. conchyliatus

S. ca. latimaculosus

S. (Polykatianna) ca. radiculus

Sphyroteca ca. mucroserrata

Deuterosminthurus maassius

Total

Rainy season

1009

2236

TAF

319

953

907

899

22,073

22,700

32,940

1830

3385

925,388

798

595

1028

934

1040

8940

1,028,751

Dry season

Total

1119

2484

900

410

1059

1057

1120

22,789

23,562

33,656

1955

3620

935,321

1022

810

1232

1213

1154

9549

1,044,032

Density specimens/m?

15,196.97

199.29

143.38

106.74

71.62

56.36

52.37

33.68

33.62

30.77

13.38

12.20

10.01

6.94

5.94

5.23

Sal?

1.00

0.51

0.42

0.38

0.12

0.01

15,986.14

16.29

36.16

13.10

Dee

15.41

15.39

16.30

331,72

342.97

489.90

28.48

52.69

13,614.57

14.88

11.79

17.93

17.66

16.80

139.00

15,196.97

Density specimens/m?

50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Table 3. Parameters of Collembola communities (see text for explanation).

Canopy Shrub layer

Parameter “Rainy season ~Dryseason. Rainy season Dry season

S) 19 19 12 14

N1 2 5 5 5

N2 1 2 4 3

HH? 0.52 1.56 1.68 1.52

Atk 0.18 0.53 0.68 0.58

S = Richness species; N1] = Abundant species; N2 = Very abundant species; H’ = Shanon-Weaver

diversity index; J’ = Pielou evenness index.

leoptera (r = —0.91, df = 5, P = 0.04), Pseudoscorpionida (r = —0.97, df = 5,

P = 0.02) and Hymenoptera (r = —0.72, df = 5, P = 0.03).

Malaise traps are not suitable for sampling Collembola, nevertheless, the spec-

imens obtained in them allowed an estimate of the general richness and compo-

sition of Collembola of the shrub layer. A total of 41,443 specimens, belonging

to 24 arthropod groups were collected (Table 5) in malaise traps. Collembola were

the fourth group in abundance (6.3%) after Diptera (35%), Coleoptera (13.4%)

and Hymenoptera (13%).

A total of 14 species of Collembola were collected in the shrub layer; the most

abundant species were Lepidocyrtus finus Christiansen & Bellinger, Calvatomina

rufescens Reuter and S. banksi, and four are new records for the Jalisco State,

Sminthurus butcheri Snider, Temeritas macrocerus Denis, Isotoma (Desoria) hie-

malis Schott and Calvatomina rufescens (Table 6). Twelve species were found

during the rainy season whereas 14 were collected during the dry season; however,

diversity and evenness were higher in the rainy season (Table 3).

Collembola abundance in the shrub layer was significantly correlated with pre-

cipitation (r = 0.43, df = 438, P = 0.03), but not with temperature (r = 0.09,

df = 438, P = 0.27). There were significant correlations only between Collembola

and Acari (r = 0.81, df = 459, P = 0.01), Aranea (r = 0.49, df = 459, P =

0.02), Chilopoda (r = 0.59, df = 459, P = 0.04) and Isopoda (r = 0.49, df =

460, P = 0.03).

ANOVA tests showed significant differences between Collembolan abundance

of the two watersheds (F = 6.31, df = 1,459; P < 0.001) as well as among

months (F = 5.07, df = 11,459, P < 0.001) and the interaction between months

and watersheds was significant (F = 3.08, df = 11,436 P < 0.0005) indicating

spatial and temporal heterogenity of Collembolan abundance in the shrub layer.

Table 4. Simple statistics within samples of Collembola in five fumigations by funnels.

Fumigation Mean SD

August 1992 301.6 98.46

May 1993 1.28 1.31

November 1993 201.5 20.14

February 1994 7.7 6.03

May 1994 5.23 bt Be)

1998

PALACIOS-VARGAS ET AL.: CANOPY COLLEMBOLA

Table 5. Abundance and percentage of Arthropoda collected by malaise trap in Chamela.

Table 6. Seasonal abundance of malaise trap Collembola from Chamela.

Taxa

Diptera

Coleoptera

Hymenoptera

Collembola

Lepidoptera

Acari

Homoptera

Aranea

Psocoptera

Orthoptera

Larvae

Hemiptera

Thysanoptera

Isoptera

Undetermined

Neuroptera

Dictyoptera

Thysanura

Others

Embioptera

Opilionida

Chilopoda

Solifuga

Isopoda

Mecoptera

Scorpionida

Pseudoscorpionida

Strepsiptera

Total

Taxa

Ceratophysella sp.

Rapoportella sigwalti

Seira bipunctata

S. dubia

S. knowltoni

Lepidocyrtus finus

L. gr. lanuginosus

Salina banksi

Isotoma (Desoria) hiemalis

Sminthurus ca. butcheri

Sminthurus sp.

Calvatomina rmfescens

Temeritas macroceros

Deuterosminthurus maassius

Total

Abundance

14,600

5566

5401

2633

3863

3411

2144

1162

795

518

Rainy

season

Dry season

Percentage

35.23

13.43

13.03

6.35

9.32

8.23

5.17

2.80

192

1.25

0.95

0.73

0.34

0.31

0.22

0.21

0.08

0.06

0.05

0.03

0.02

0.01

<0.01

100.00

a THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

DISCUSSION AND CONCLUSIONS

We found more Collembola species in canopy fogging than in malaise traps

(19 and 14 species respectively). Guilbert et al. (1995) reported 32 Collembola

species by fogging in New Caledonia. This result is higher than our observations

in Chamela.

Palacios-Vargas (1981) reported 22 genera (31 species) of Collembola living

in Tillandsia (Bromeliaceae) on the basaltic flows of Chichinautzin (Morelos,

Mexico). Of these genera, 11 are also present in Chamela canopy and, it is pos-

sible they are also living in Tillandsia which are abundant at the Station (Lott et

al. 1987).

The greatest abundance of Collembola reported by Guilbert et al. (1995)

reached 18.4% of total arthropods, but in Chamela it was 98% and with one

species, S. banksi accounting for 89.6% of all Collembola. This means that one

species of Collembola is dominant in the Arthropoda of the canopy. Density of

Collembola in the canopy was estimated to be 30,438 specimens/m~? and this figure

is higher than that of: Guilbert et al. (1995) for a primary forest in New Caledonia

(107 specimens/m?); Watanabe & Ruaysoongnern (1989) for a dry evergreen for-

est in Northeastern Thailand (195 specimens/m7), and Hijii (1986) for a Crypto-

meria japonica (Linnaeus f.) D. Don forest in Japan (3754 specimens/m?). Dif-

ferences in densities might be explained by differences in height and composition

plant cover, the amount of suspended soil acumulated in Tillandsia (and on

trunks), or amount of dead plant matter in the canopy, which is 75% of the plant

matter (J. M. Maass, personal communication).

Number of Collembola species in true soil and leaf litter from Chamela are 49

and 33 respectively (Palacios-Vargas & Gdémez-Anaya 1993). There is greater

similiarity between leaf litter and shrub layer then between shrub layer and canopy

layer. Salina banksi is most abundant in the canopy and decreases significantly

in the shrub layer and is rare in leaf litter and soil (two specimens in a year of

montly sampling). Three species occur solely in shrub layer, Jsotoma (Desoria)

hiemalis, Calvatomina rufescens and Sminthurus butcheri.

According to Guilbert et al. (1995) most Collembolans from shrub layer and

canopy are epiedaphic forms such as Entomobryomorpha and Symphypleona. We

found that 85% and 68% of the shrub layer and canopy species belong to these

groups. The other species may occur in the Tillandsia and bark.

Precipitation affects the abundance of Collembola in a determinant way both

in the soil (G6mez-Anaya 1998) and in canopy. Seasonal variation of the Collem-

bola from the canopy and the shrub layer are associated with humidity, but not

temperature. There is a greater correlation with humidity in the canopy due to

insolation and the exposure possibly resulting in dehydratation.

Humidity is correlated with the availability of food for the Collembola in each

layer. The most abundant genera from canopy are mainly detritophagous, myce-

tophagous and polyphagous, but in the shrub layer the most common groups are

saprophagous, mycetophagous and detriphagous (MacNamara 1924, De Bernardi

& Parisi 1968, Ellis 1978). High accumulation of organic matter exists in the

canopy of Chamela and this is correlated with the different feeding habits of

Collembola been found there.

The abundance of juveniles of Hypogastruridae, Entomobryidae and Symphy-

1998 PALACIOS-VARGAS ET AL.: CANOPY COLLEMBOLA oe

pleona founded in shrub layer suggest the possibility of vertical migrations be-

tween this layer and canopy.

Few species of Collembola live only in the canopy, Neotropiella quinqueocu-

lata Denis and the three species of Sminthurinus, the remaninder can be found in

shrub layer, soil or leaf litter. Jsotoma (Desoria) hiemalis, Sminthurus ca. butcheri,

Sminthurus sp., Temeritas macrocerus and Calvatomina rufescens, however, are

found only in shrub layer.

Our results show that the fauna living in the canopy, particularly the Collem-

bola, should have an important role in the cycling nutrients and energy in the

deciduous forest.

ACKNOWLEDGMENT

We thank Dr. Eric Guilbert (Laboratoire d’Entomologie, Muséum National

d’ Histoire Naturelle) and Dr. Zenén Cano Santana (Laboratorio de Ecologia, Fac.

de Ciencias, UNAM) for critical review to the manuscript and Ms. Blanca E.

Mejia Recamier, Ms. Alicia Palafox Rodriguez and Dr. Alfonso Pescador Rubio

(Depto. de Biologia, Fac. Ciencias, UNAM) for their assistence during this pro-

ject.

LITERATURE CITED

Bretfeld, G. 1994. Sturmius epiphytus n. gen. n. spec. from Colombia, a taxon of the Symphypleona

(Insecta: Collembola) with an unexpected character combination. J. Zool. Syst. Evol. Res., 32:

264-281.

Bullock, S. H. 1988. Rasgos del Ambiente Fisico y Biol6gico de Chamela, Jalisco, México. Folia

Entomol. Mex., 77: 5—17.

Cervantes, L., R. Dominguez & M. Maas. 1988. Relacién lluvia-escurrimiento en un sistema pequefio

de cuencas de selva baja caducifolia. Ingenierfa Hidrdulica en México, III: 30-41.

De Bernardi, FE & V. Parisi. 1968. Osservazioni sul regime alimentare di alane especie di Orchesella

e Tomocerus (Collembola) in una valle alpina (Val Malenco). Licei. Rend. Sc. fis. mat. e nat.,

45: 98-106.

Delamare-Debouteville, C. 1948. Etude quantitative du peuplemente animal des sols suspendu et des

ephiphytes en forét tropicale. Comptes Rendus de I Academie du Science, 226: 1544-1546.

Ellis, W. N. 1978. Pollen feeding of Deuterosminthurus repondus (Agren) in the High Pyrenees and

some systematic notes (Collembola. Sminthuridae). Entomol. Berichtzn, Deel, 38: 61-63.

Erwin, T. L. 1983. Beetles and other insects of tropical forest canopies at Manaus; Brazil, sampled

by insecticidal fogging. pp. 59-79. Jn Sutton S. L. et al. (eds.). Tropical rain forest ecology

and manegament. Blackwell Scientific Publications, London.

Goémez-Anaya, J. A. 1998. Ecologia de Collembola (Hexapoda: Apterygota) de Chamela, Jalisco,

México. M. Sc. Thesis. Facultad de Ciencias, UNAM, México.

Guilbert, E, M. Baylac & J. Najt. 1995. Canopy arthropod diversity in a New Caledonian primary

forest sampled by fogging. Pan-Pac. Entomol., 71: 3-12.

Hijii, N. 1986. Density, biomass and guild structure of arboreal arthropods as related to their inhabited

tree size in Cryptomeria japonica plantation. Ecol. Res., 1: 97-118.

Lott, E. J. 1985. Listados floristicos de México. III. La Estaci6n de Biologia Chamela, Jalisco. Instituto

de Biologia, Univ. Nac. Autén., Méx. México.

Lott, E. J., S. H. Bullock & J. A. Solis-Magallanes. 1987. Floristic diversity and structure of upland

and arroyo forests in coastal Jalisco. Biotropica, 19: 228-235.

Ludwig, J. A. & J. E Reynolds. 1988. Statistical ecology. A primer methods and computing. Wiley

Intersciences Publiction, New York.

MacNamara, C. 1924. The food of Collembola. Can. Ent., 56: 99-105.

Palacios-Vargas, J. G. 1981. Collembola asociados a Tillandsia (Bromeliacea) en el derrame lavico

del Chichinautzin, Morelos, México. Southwestern Ent., 6(2): 87-98.

54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

Palacios-Vargas, J. G. & J. A. Gémez-Anaya. 1993. Los collembola (Hexapoda: Apterygota) de

Chamela, Jalisco, México (Distribuci6n ecolégica y claves). Folia Entomol. Mex., 89: 1-34.

Sarukhan J. & M. Maass. 1990. Bases ecol6gicas para un manejo sostenido de los ecositemas: El

sistema de cuencas hidrologicas. pp. 81-114. Jn Leff, E. (ed.). Medio Ambiente y Desarrollo

en México. Vol. I. Centro de Investigaciones Interedisciplinarias en Humanidades, UNAM.

México.

Watanabe, H. & S. Ruaysoongnern. 1989. Estimation of arboreal arthropod density in a dry evergreen

forest in northeastern Thailand. J. Trop. Ecol., 5: 151-158.

Zar, J. H. 1984. Bioestatistical analysis (2nd ed.). Prentice-Hall. Englewood Cliffs, New Jersey.

Received 24 Mar 1997; Accepted 21 May 1997.

Appendix I. Systematic list of Collembola from canopy and shrub layer.

HY POGASTRURIDAE

Ceratophysella gibbosa (Bagnall)

Xenylla humicola (Fabricius)

BRACHYSTOMELLIDAE

Brachystomella minimucronata Palacios-Vargas & Najt

B. sigwalti Palacios-Vargas & Najt

ANURIDIDAE

Neotropiella quinqueoculata (Denis)

Pseudachorutes ca. subcrassoides Mills

NEANURIDAE

Aethiopella sp.

ISOTOMIDAE

Isotoma (Desoria) hiemalis Schott

ENTOMOBRYIDAE

Entomobrya (Drepanura) californica Schott

Lepidocyrtus finus Christiansen & Bellinger

L. gr. lanuginosus (Gmelin)

Seira bipunctata (Packard)

S. dubia Christiansen & Bellinger

S. knowltoni (Wray)

PARONELLIDAE

Salina banksi MacGillivray

SMINTHURIDIDAE

Sphaeridia pumilis (Krausbauer)

KATIANNIDAE

Sminthurinus (Polykatianna) ca. radiculus Maynard

S. ca. conchyliatus Snider

S. ca. latimaculosus Maynard

SMINTHURIDAE

Sphyrotheca ca. mucroserrata Snider

Sminthurus ca. butcheri Snider

Sminthurus sp.

Temeritas macroceros Denis

Calvatomina rmufescens (Reuter)

BOURLETIELLIDAE

Deuterosminthurus maassius Palacios-Vargas & Gonzalez

PAN-PACIFIC ENTOMOLOGIST

74(1): 55-57, (1998)

Scientific Note

DISCOVERY OF THE GALL MITE ACERIA GENISTAE

(NALEPA) (ACARINA: ERIOPHYIDAE) ON GORSE AND

FRENCH BROOM IN THE UNITED STATES

Gorse (Ulex europaeus L.) and French broom (Genista monspessulana [L.] L.

Johnson) are noxious and invasive leguminous shrubs native to western and Med-

iterranean Europe respectively (Hickman, J. C. [ed.] 1993. The Jepson manual:

higher plants of California. Univ. California Press, Berkeley, California). Having

escaped from cultivation, they are weeds of disturbed sites in the coastal Pacific

Northwest region from the San Francisco Bay Area to Washington and British

Columbia, and in higher elevation sites in Maui, Hawaii and Kauai (Haselwood,

E. L., G. G. Motter & R. T. Hirano (eds.). 1983. Handbook of Hawaiian weeds

[2nd ed.]. Univ. Hawaii Press, Honolulu, Hawaii; Markin, G. P, E. R. Yoshioka

& R. E. Brown. 1995. Chap. 78. Jn Biological control in the western United

States. Univ. California DANR Publ. 3361, Oakland, California). Although bio-

logical control programs have purposely reunited gorse with two European natural

enemies (Apion ulicis [Forster] [Coleoptera: Apionidae] and Tetranychus lintear-

ius Dufour [Acarina: Tetranychidae]) in the continental United States (Rees, N.

E., P C. Quimby, Jr., G. L. Piper, E. M. Coombs, C. E. Turner, N. R. Spencer &

L. V. Knutson. 1996. Biological control of weeds in the west. Western Society of

Weed Science, Bozeman, Montana), we report here the detection of an acciden-

tally introduced European gall mite, Aceria genistae (Nalepa) (Acarina: Eriophy-

idae), on both gorse and French broom in the San Francisco Bay Area.

In late October 1994 we transplanted gorse seedlings from a vacant lot in Daly

City to our greenhouse in Albany, California. By mid-Jan 1995 several plants

exhibited abnormal growth: continued production of juvenile leaves and stunted

shoot tips with reduced, thickened scale leaves or even clublike apices. Exami-

nation of the shoot tips revealed colonies of eriophyoid mites living among the

appressed leaves. These mites were subsequently determined by James W. Am-

rine, Jr. of West Virginia University as Aceria genistae (Nalepa), originally de-

scribed from scotch broom (Sarothamnus scoparius Koch. [= Cytisus scoparius

(L.) Link]) in Lorraine, France. We collected shoots of gorse and French broom

from the Daly City field site on 18 Apr 1996 and 14 May 1996. Presence of the

same mite species on seedlings of both legumes confirmed that the mites living

on our gorse in Albany had originated from Daly City. It is unknown how or

when the mites arrived in California. This is the first field record of A. genistae

in the United States.

In Europe, Aceria genistae has been recorded from two species of Cytisus

(including C. scoparius [scotch broom]), four species of Genista (but not G.

mons pessulana) and two species of Ulex (including U. europaeus) (Baker, E. W.,

T. Kono, J. W. Amrine, Jr., M. Delfinado-Baker & T. A. Stasny. 1996. Eriophyoid

mites of the United States. Indira Publishing House, West Bloomfield, Michigan).

Its native range includes Great Britain, Spain, Italy and Central Europe, and it

has been reported on U. europaeus in New Zealand where gorse is also a natu-

56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 74(1)

ralized weed (Manson, D. C. M. 1989. New Zealand J. Zool., 16: 37—49). Females

are 165—225 um long and apparently do not have a deuterogyne stage (Castagnoli,

M. 1978. Redia, 61: 539-550). Recent descriptions and figures may be found in

Castagnoli (1978) and Baker et al. (1996). Castagnoli (1978) reports that in Italy,

A. genistae produces several generations throughout spring and summer on scotch

broom, and causes glomerule-like galls on stem bud tissue.

We observed that the mite prefers young, soft, pubescent foliage on gorse and

French broom. We now see mites on the new growth of mature gorse plants in

the shadehouse into which we moved our original gorse transplants. The colonies

have persisted without killing or further deforming the plants. However, the nat-

urally occurring mite predators present in the shadehouse may be preventing the

eriophyoids from causing appreciable damage. Perhaps feeding by the mites under

protected, warm and humid greenhouse conditions produced the unusual mor-

phology that we initially saw.

The mite’s damage to gorse and French broom in Daly City appears to be

negligible and is often inconspicuous. Based on our observations in the San Fran-

cisco Bay Area, we do not anticipate that A. genistae will play a substantial role

in the biological control of either gorse or French broom. Rather, the eriophyoid

joins a suite of unintentionally introduced European broom and gorse feeders

(Waloff, N. 1966. J. Appl. Ecol. 3: 293-311; Andres, L. & E. Coombs. 1995.

Chap. 79. In Biological control in the western United States. Univ. California

DANR Publ. 3361, Oakland, California; Markin et al. 1995). To date, these ar-

thropods have not curbed scotch broom, French broom or gorse populations in

the Pacific Northwest. However, a European mite collected from scotch broom

and identified as A. genistae is currently under consideration as a potential bio-

logical control agent of scotch broom in New Zealand and Australia (G. P. Markin,

personal communication). Preliminary host range tests in Europe indicate that this

mite may cause galling only on C. scoparius, but further testing is needed to

determine if gorse or French broom are not suitable hosts also (Q. Paynter, per-

sonal communication). Interestingly, the A. genistae reported on gorse in New

Zealand has not been found on scotch broom there and did not transfer success-

fully from gorse to scotch broom in a laboratory experiment (P. Syrett, personal

communication). Scotch broom is not present at our Daly City field site, so we

cannot comment on the suitability of scotch broom as a host for the A. genistae

we found in California. These incongruent host associations suggest that further

taxonomic work is needed to determine if A. genistae is actually a complex of

sibling species. Castagnoli (1978) has already reaffirmed the validity of A. spartii

(G. Canestrini) as a separate species rather than a synonym of A. genistae, based

on morphological differences and A. spartii’s specificity to Spartium junceum L.

We intend to survey scotch broom and other gorse and French broom populations

in California for eriophyoids. Additional collection records should help solve this

taxonomic puzzle.

Records—CALIFORNIA. ALAMEDA Co.: Albany, 1 Feb 1995, K. L. Chan,

Ulex europaeus, shoot tips. SAN MATEO Co.: Daly City, 14 May 1996, K. L.

Chan, Genista monspessulana, shoot tips.

Acknowled gment.—We thank the following for reviewing an earlier draft of the

manuscript and for their specific contributions to it: J. W. Amrine, Jr. (West Vir-

ginia University, Morgantown) determined our specimens and provided additional

1998 SCIENTIFIC NOTE wy,

information on the mite, R. Somerby (California Department of Food & Agri-

culture, Sacramento) examined our initial specimens, G. P. Markin (United States

Department of Agriculture, Forest Service, Bozeman, Montana), Q. Paynter (In-

ternational Institute of Biological Control, Montpellier, France) and P. Syrett (Ma-

naaki Whenua—Landcare Research, Lincoln, New Zealand) contributed infor-

mation on the mite’s status as a potential biological control agent, J. Xerogeanes

(Mendocino College, Ukiah) informed us of the Daly City gorse site, and J. Herr

and R. Takumi provided technical assistance. KLC dedicates this paper to the

memory of her co-author whose untimely passing in April 1997 cut short a ded-

icated career of insightful contributions to the biological control of weeds.

Kathleen L. Chan & Charles E. Turner, United States Department of Agricul-

ture, Agricultural Research Service, Western Regional Research Center, Albany,

California 94710.

Received 23 Jul 1997; Accepted 10 Nov 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 58, (1998)

Scientific Note

NEOHERMES CALIFORNICUS (WALKER)

(MEGALOPTERA: CORYDALIDAE) IN IDAHO: A NEW

STATE RECORD

The genus Neohermes is represented in North America by five species. Three

are eastern and two, N. californicus (Walker) and N. filicornis (Banks), occur in

the west (Flint, O. S. 1965. Psyche, 72: 255-263). Neohermes filicornis has been

collected in California, Arizona and New Mexico; N. californicus has been col-

lected in California, Oregon, and Nevada.

Recently we have collected in the north-central, montane area of Idaho on

private land in a tributary of Wall Creek on the south fork of the Clearwater River

in Idaho County at an elevation of 731 m. This stream is intermittent. The area

receives up to 1.3 m of snow annually; the snow contributes most of the runoff

for the site (Ronald C. Miller & James M. Goodwin, pers. comm., ranchers living

near the site). The large cobbles on sandy substratum in the riffle areas and the

flow intermittency match up well with the description of the species’ habitat in

California and Oregon (Evans, E. D. 1972. unpubl. dissertation, Oregon St. Univ.).

On 14 Jun 1996, we found a 19 mm corydalid larva clinging to the underside

of a rock at this location. Its small size made generic identification uncertain. On

21 Jun 1996, kick-screen collecting yielded a late instar larva (48 mm) from a

riffle area of the stream. On 13 Jul 1996 at 22:20 h an adult male (55 mm) was

collected at an artificial white light provided by a Coleman gas lantern. We de-

termined the species to be N. californicus (Walker) using keys found in Flint

(1965), and Evans, E. D. & H. H. Neunzig. (1996 pp. 298-308, Jn Merritt, R.

W., & K. W. Cummins (eds.), An Introduction to the Aquatic Insects of North

America, Kendall, Hunt Publ.) and Evans (1972). The determination was con-

firmed by E. D. Evans.

This collection represents a new state record and extends the range of this

Species approximately 519 km north and east into. north-central Idaho (E. D.

Evans, pers. comm.) from its previous limits of Linn County in northwestern

Oregon, and Kern County, California near Kernville (Evans 1972). Specimens are

deposited in the William E Barr Entomological Museum at the University of

Idaho.

Acknowled gment.—We acknowledge Dale’s wife, Brandy L. Kellar, for the col-

lection of the adult male specimen, Elwin D. Evans for confirming the identifi-

cations, Ronald C. Miller and James M. Goodwin for their knowledge of the

study area, and E. D. Evans, M. A. Brusven, Sanford D. Eigenbrode and James

B. Johnson for reviewing this note. This note is published with the approval of

the director of the Idaho Agricultural Experiment Station as paper no. 96746.

Russell C. Biggam, Dept. Plant, Soil and Entomological Sciences (PSES), Uni-

versity of Idaho, Moscow, Idaho 83844-2339, & Dale S. Kellar, 3525 12th St.,

Apt. #6, Lewiston, Idaho 83501.

Received 5 Dec 1996; Accepted 19 Jun 1997.

PAN-PACIFIC ENTOMOLOGIST

74(1): 59, (1998)

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PAN-PACIFIC ENTOMOLOGIST

74(1): 60, (1998)

ERRATUM

In the article by W. D. Wiesenborn entitled Hesperopsis graciliae (MacNeill)

(Lepidoptera: Hesperiidae) flight between hostplants and Prosopis glandulosa (73:

186), the name of skipper should have been Hesperopsis gracielae not H. gra-

ciliae.

ADDENDUM

The following authors received C. P. Alexander funds to partially offset their

page charges:

Barthell, J. EK, T. L. Griswold, G. W. Frankie & R. W. Thorp. Osmia (Hymenop-

tera:Megschilidae) diversity at a site in centreal coastal California. 73: 141-—

151.

Brailovsky, H. A. Sibuyanhygia, a new genus of Colpurini from the Philippine

Republic, with descriptions of three new species (Heteroptera: Coridae) 73: 70—

78.

Burquez, A. Distributional limits of Euglossine and Meliponine bees (Hymenop-

tera: Apidae) in northwestern Mexico. 73: 137-140.

Conway, J. R. Foraging activity, trails, food sources and predators of Formica

obscuripes Forel (Hymenoptera: Formicidae) at high altitude in Colorado. 73:

172-183.

Gulmahamad, H. Gnathamitermes perplexus (Banks) (Isoptera: Termitidae): a nui-

sance structural termite pest in southern California. 72: 37-38.

Hynes, C. D. The immature stages and biology of the craneflies Toxorhina ca-

ledonica and Elephantomyia garrigouana (Diptera: Limoniidae). 73: 93-99.

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Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

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THE PAN-PACIFIC ENTOMOLOGIST

Volume 74 January 1998 Number 1

Contents

BROWN, J. W. & S. PASSOA—Larval foodplants of Euliini (Lepidoptera: Tortricidae) from

Abies to Vitis

CHEMSAK, J. A. & EF A. NOGUERA—Review of the genus Sphaenothecus Dupont (Cole-

optera: Cerambycidae)

LANG, R. F,.G. L. PIPER & E. M. COOMBS—Fstablishment and redistribution of Sphen-

optera jugoslavica Obenberger (Coleoptera: Buprestidae) for biological control of diffuse

knapweed (Centaurea diffusa Lamarck) in the midwestern and western United States --

LANDOLYT, P. J.—Lacanobia subjuncta (Lepidoptera: Noctuidae) on tree fruits in the Pacific

Northwest

MORON, M.-A.—New species of Phyllophaga Harris from Mesoamerica (Coleoptera: Melo-

lonthidae: Melolonthinae)

PALACIOS-VARGAS, J. G., G. C. MENESES & J. A. GOMEZ-ANAYA—Collembola from

the canopy of a Mexican tropical deciduous forest

SCIENTIFIC NOTES

CHAN, K. L. & C. E. TURNER—Discovery of the gall mite Aceria genistae (Nalepa) (Acarina:

Eriophyidae) on gorse and French broom in the United States __

BIGGAM, R. C. & D. S. KELLAR—WNeohermes californicus (Walker) (Megaloptera: Cory-

dalidae) in Idaho: a new state record _

Pacific Coast Entomological Society: Sponsoring Members 1997

Pan-Pacific Entomologist: Erratum, Addendum & Editorial Notice

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PAN-PACIFIC

ENTOMOLOGIST

Volume 74 April 1998 Number 2

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

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