The

PAN-PACIFIC

ENTOMOLOGIST

Volume 75 | January 1999 Number |

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

EDITORIAL BOARD

R. V. Dowell, Editor R. M. Bohart

Jeffery Y. Honda, Associate Editor J. T; Doyen

R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.

Julieta F Parinas, Treasurer Warren E. Savary

Published quarterly in January, April, July, and October with Society Proceed-

ings usually appearing in the October issue. All communications regarding non-

receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary;

and financial communications should be addressed to: Julieta F Parinas, Treasurer;

at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-

emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addressed to: William Hamersky, Membership Committee chair, Pacific Coast

Entomological Society, Dept. of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118-4599.

Manuscripts, proofs, and all correspondence concerning editorial matters (but

not aspects of publication charges or costs) should be sent to: Dr. Robert V.

Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,

1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-

tributors, and volume 73(4): 248—255, October 1997, for more detailed informa-

tion. Information on format for taxonomic manuscripts can be found in volume

69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer.

The annual dues, paid in advance, are $25.00 for regular members of the So-

ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for

institutional subscriptions or sponsoring members. Members of the Society receive

The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes

are available; see 72(4): 247 for current prices. Make checks payable to the Pacific

Coast Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 1999

Laura A. Irons, President Vincent E Lee, Managing Secretary

Julieta E Parinas, Treasurer Stanley E. Vaughn, Recording Secretary

THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per

year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate

Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional

mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society,

% California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

This issue mailed 11 June 1999

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

PAN-PACIFIC ENTOMOLOGIST

75(1): 1-7, (1999)

TRANSFER OF THE TAIWANESE PSEUDOPYROCHROA

UMENOI AND THE JAPANESE P. AMAMIANA TO

PSEUDODENDROIDES (COLEOPTERA: PYROCHROIDAE:

PYROCHROINAE)

DANIEL K. YOUNG

Department of Entomology, University of Wisconsin,

Madison, Wisconsin 53706

Abstract.—On the basis of salient characters, particularly those associated with the head, eighth

abdominal sternite and genitalia of the male, both Pseudopyrochroa umenoi K6no of Taiwan,

and Pseudopyrochroa amamiana Nakane of Amami-Oshima Island, Japan are transferred from

Pseudopyrochroa to Pseudodendroides where they are hypothesized to represent species closely

related to the Japanese Pseudodendroides niponensis (Lewis).

Key Words.—Insecta, Coleoptera, Pyrochroidae, Pseudopyrochroa, Pseudodendroides, generic

transfers, phylogeny, confocal microscopy, Japan, Taiwan.

In an effort to redefine Pseudopyrochroa Pic (1906) as a monophyletic taxon,

part of a larger project on the taxonomy of the genus, several taxonomic changes

have become necessary. The body of this paper proposes two changes involving

Pseudopyrochroa and another pyrochroine genus, Pseudodendroides Blair (1914).

Pseudopyrochroa was proposed as a subgenus of Pyrochroa Geoffry for several

Southeast Asian pyrochroids previously assigned to the European Pyrochroa.

Pseudopyrochroa was said to differ by having a smaller head, compound eyes,

and reduced genae. But even Pic applied the name inconsistently, and the generic

use of Pseudopyrochroa did not begin to stabilize until Pic (1913) briefly elab-

orated on his misplacement of several Pseudopyrochroa species that he had orig-

inally attributed to the circumboreal genus Schizotus Newman.

Although the largest genus in the family with approximately 70 species names

attributed to it, Pseudopyrochroa is among the least known of all pyrochroine

genera from both taxonomic and ecological perspectives. Species richness is great-

est along the Pacific coast of the Asian continental plate and forested inland

montane regions. The only previous attempt to synthesize information on Pseu-

dopyrochroa came in the form of Blair’s (1914) provisional comments and key.

This effort was, according to Blair (1914:318), “‘intended merely as a temporary

measure, in the hope of stimulating further study of the genus... .”

Pseudodendroides was proposed for two Indian and two Japanese species pre-

viously assigned to Dendroides Latreille. Two additional species were added by

Pic from China (1938) and the Himalayan regions of Sikkim and Tibet (1955).

At the time of its description, Pseudodendroides was said by Blair (1914: 314)

to differ from Pseudopyrochroa, “by the large eyes, approximate above in the -

male.’ This is certainly the case in males of the Japanese P. niponensis (Lewis),

the type species of Pseudodendroides. However, this derived character state ap-

pears to be homoplasious, having arisen independently several times in both py-

rochroine and pediline pyrochroids (Young, unpublished observations), and Pseu-

dodendroides as characterized by Blair is, at best, paraphyletic.

Depositories, Procedures, and Abbreviations.—Taxonomic material for this

z THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

study came from my personal collection (DYCC), and material borrowed from

the collection of Darren Pollock, Winnipeg (DAPC), the Florida State Collection

of Arthropods, Gainesville (FSCA), the Naturhistorisches Museum Wien, Wein

(NHMV), the Muséum National d’ Histoire Naturelle, Paris (PMNH), the Taiwan

Agricultural Research Institute (TARD, and the Museum fiir Naturkunde der

Humboldt-Universitaét, Berlin (ZMHB). The holotype male of Pseudopyrochroa

umenoi K6éno was also borrowed from the Hokkaido University, Sapporo, Japan

(HUSC), as was the holotype female of Pseudodendroides uraiana K6no. Type

and other pyrochroid material in The Natural History Museum, London (BMNH)

was studied on site.

MATERIALS AND METHODS

As I discuss below, the presence, number and configuration of cranial pits

(Young 1975) in several genera of male pyrochroines offer important characters

for hypothesizing relationships among genera. Unfortunately, these three-dimen-

sionally complex structures are virtually impossible to draw and for many species,

only the type or small type series is known.

Using scanning electron microscopy (SEM) and photomicrography to capture

and illustrate features of insect gross anatomy and exoskeletal ultrastructure has

become a well established research tool. However, SEM techniques have several

unfortunate drawbacks, usually including the necessity of coating the specimen

with a thin layer of a heavy metal to prevent electron charging, exposure of the

specimen to a high vacuum environment, and the possible need to trim or dissect

the specimen to fit the sample holder.

As SEM proved unfeasible for this and related studies, I investigated techniques

of laser-scanning confocal microscopy. Standard confocal techniques such as a

3D z-series reconstruction or projection were unsatisfactory because exoskeletal

opacity inhibited laser penetration and the full vertical range of the cranial com-

plex could not be imaged. Finally, in a nod to well known SEM “‘stereo-pair”’

techniques, images of exoskeletal autofluorescence were collected from a straight

vertical view and a tilted view.

A low-magnification lens for the confocal microscope (3.5X) was selected. Its

wide field of view permitted imaging of the entire dorsoanterior region of the

head, including all of the cranial apparatus, without resorting to making montages

of micrographs. The large depth of field (optical section thickness) of the lens

allowed the protrusions and concavities to be imaged in a single image frame.

Finally, the large working distance of the lens made it possible to image intact

beetles, without resorting to perturbation of any kind. Not only could the speci-

mens remain on their pins, point, or card mounts, but it was not even necessary

to remove any of the specimen labels.

The 488 nm line of the Kr/Ar laser was utilized to excite the autofluorescence

of the exoskeletal material. The standard fluorescein imaging cube was used for

detection of the emitted signal.

Preparation of the specimen for imaging was quickly accomplished by inserting

the mounting pin into a 7.6 X 7.6 X 1.9 cm (3” X 3” X 3/4") foam block. This

entire assembly was able to be placed under the microscope’s objective lens for

imaging. After a straight vertical, ‘head-on’ image was taken, fiducial marks

were made on the microscope’s imaging screen using a felt-tipped marker. Three

1999 YOUNG: PSEUDOPYROCHROA TRANSFERRED 3

standard microscope slides were then placed under one corner of the foam block,

thereby effectively tilting the block (and specimen) by approximately 5—7°.

After re-aligning the “‘live’’ image with the fiducial marks on the screen, a

second image was obtained. When printed side-by-side, these two images form a

stereo pair (Figs. 7 and 8). It should be noted that to create the stereo effect, it

is generally necessary to view the figures with a stereo viewer.

DISCUSSION

The presence of complex, cranial pits in adult males is an important synapo-

morphy establishing the monophyly of several pyrochroine genera. Pyrochroine

genera exhibiting cranial pits, have them located behind the eyes, as in males of

Schizotus, or between the eyes. In the latter case, the pit may consist of a single,

shallow impression, as in the European Pyrochroa and Asian Eupyrochroa Blair,

or pits may be well developed and paired: Pseudodendroides, Phyllocladus Blair,

Neopyrochroa Blair and Pseudopyrochroa. Using this clade of seven genera for

outgroup comparisons, the monophyly of Pseudopyrochroa species may be hy-

pothesized by a synapomorphy associated with the external male genitalia. In

Pseudopyrochroa, the dorsolateral apices of the parameres are bilaterally toothed,

with each tooth projecting basally (Fig. 2). In the plesiomorphic character state,

the apices of the parameres are rounded. Additionally, in Pseudopyrochroa the

apex of the penis (Fig. 3) is provided with a dorsomesal, basally recurved hook

(= apomorphic). In other pyrochroine genera (e.g., Fig. 6) the apex of the penis

is generally rounded and lacking a hook (= plesiomorphic). A similar, apically

hooked penis is present in males of N. flabellata (Fabricius), from the eastern

United States and Canada. However, males of N. femoralis (LeConte) and N.

sierraensis Young lack the modification and its autapomorphic presence in N.

flabellata is hypothesized to be homoplasious with respect to Pseudopyrochroa.

Although exhibiting considerable interspecific variation, the cranial pits of male

Neopyrochroa, Phyllocladus and Pseudopyrochroa are paired and typically well

excavated. Those of Pseudodendroides, as illustrated by P. niponensis (Fig. 7)

have an additional transverse ridge, making them nearly four-chambered (= apo-

morphic); very similar to those of both P. umenoi (Fig. 8) and P. amamiana.

In Pseudodendroides the parameres of the male genitalia are short and widely

separated for approximately half their length (Fig. 5). This character represents a

probable synapomorphy also exhibited by the Asian Phyllocladus, Neopyrochroa,

a genus endemic to North America with two eastern and two western species,

and an undescribed genus and species from the Darjeeling District of India. Males

of Pseudodendroides, Phyllocladus, and the undescribed genus and species from

Darjeeling also share an apomorphy associated with the eighth sternite: the apical

margin is widely emarginate and conspicuously concave (Fig. 4). In males of

other pyrochroine genera, including Pseudopyrochroa, the eighth sternite (Fig. 1)

is tapered distally and narrowly emarginate mesally (= plesiomorphic).

Although there has never been a comprehensive assessment of pyrochroine

antennae, the gross anatomy is rich in characters; this is particularly true in the

case of males (Young, unpublished observations). In males of Pseudodendroides,

the scape is long and parallel-sided (= apomorphic); this condition is not seen in

Pseudopyrochroa or any other pyrochroine genera. Males of both P. umenoi and

P. amamiana have an elongate, parallel-sided antennal scape.

Vol. 75(1)

THE PAN-PACIFIC ENTOMOLOGIST

pre

~ SO ty merenen

1999 YOUNG: PSEUDOPYROCHROA TRANSFERRED 5

Employing these criteria, the Taiwanese Pseudopyrochroa umenoi K6éno and

the Japanese P. amamiana Nakane must be transferred to Pseudodendroides:

Pseudodendroides umenoi (K6no), 1936, NEW COMBINATION

Pseudodendroides amamiana (Nakane), 1988, NEW COMBINATION

Pseudodendroides umenoi was originally described under the generic name

Pseudopyrochroa on the basis of both males and females collected at Numanohira,

Taiwan. In 1960, Nakane outlined and briefly discussed the Pyrochroidae of Japan,

recording what he understood to be P. umenoi from Amami-Oshima Island. In

that paper, Nakane also expressed some uncertainty regarding the relationships

between Pseudopyrochroa and Pseudodendroides, noting that P. umenoi appeared

to lack the retrorsely acuminate processes at the apex of the parameres charac-

teristic of most Pseudopyrochroa. This topic was briefly revisited by Ohbayashi

(1968). He illustrated the genitalia of P. umenoi, noting the “‘small teeth’’ at “‘the

apex of male paramera of the species.’ However, in P. umenoi as in P. niponensis

(Fig. 5), the “‘teeth’’ are formed from the excavate inner margins of the distal

parameres. The tooth-like paramereal processes characteristic of most Pseudopy-

rochroa—as seen, for example, in P. harmondi (Pic) (Fig. 2)—are formed by the

splaying and sclerotization of the outer and dorsal distal surfaces of the parameres,

which are narrowly and shallowly separated distally.

After years of reflecting on the material from Amami-Oshima Island, Nakane

(1988) described Pseudoedendroides amamiana, under the generic name Pseu-

dopyrochroa, stating that it differed consistently from P. umenoi in several char-

acters associated with the head, including secondary sexual characteristics of the

cranium, prothorax, and male genitalia. No further comments relative to generic

relationships were made.

On the basis of the characters discussed above, it is clear that both Pseudo-

pyrochroa umenoi K6no and Pseudopyrochroa amamiana Nakane are incorrectly

placed in Pseudopyrochroa. The evidence suggests that both species belong to

Pseudodendroides.

Material Examined.—Pseudodendroides amamiana: JAPAN. AMAMI OSHIMA: Hatsumo, 1 Apr

1968, ex Tamanuki, 1¢ (DYCC); 28 Mar 1964, Y. Miyake, 12 (DYCC). Pseudodendroides nippo-

nensis: JAPAN. [country only] G. Lewis, 2d ¢, 12 (BMNH); Kiou-Siou (Kiushiu), Bassin Supérieur

de la Sendaigawa, 1906, E. Gallois, 16, 42 2 (PMNH)] KYUSHU: Higo, G. Lewis, 1¢ (BMNH);

Higo, 1881, G. Lewis, 12 (BMNH); Higo, G. Lewis, 12 (BMNH); Kumamoto Pref., Momiki, Izumi

v., 6 Jul 1991, T. Ueno, 16 (DAPC); HONSHU: Akita [underside of mounting card], Nikko, G. Lewis,

1d (BMNH); Akita, G. Lewis, 12 (BMNH); Nikko, 29-31 Oct 1880, G. Lewis, 12 (BMNH); Mi-

yanoshita, G. Lewis, 12 (BMNH); Chiuzenji, 19-24 Aug 1881, G. Lewis, 12 (BMNH); Tokio, [G.

Lewis material], 12 (BMNH); Nagano Pref., Tobira Spa, 31 Jul 1973, S. Hisamatsu, 1¢ (DYCC);

Env. de Tokio et Alpes de Nikko, 1901, J. Harmond, 42 2 (PMNH); Env. de Tokio, 1906, J. Harmond,

222 (PMNH); Kofou, 1906, L. Drouard de Lezey, 12 (PMNH). Pseudodendroides umenoi: TAI-

WAN. NUMANOHIRA: 19 Jun 1932, Umeno & Taira, No. 21, Pseudopyrochroa umenoi Kono, 6

[Holo]Type (HUSC); NANTOU HSIEN: Meifeng, 2150 m, 20—22 Jun 1979, K. S. Lin & H. Chen,

—

Figures 1-3. Pseudopyrochroa harmondi Pic, adult male. Figure 1: Abdominal sternite 8, ventral

view. Figure 2: Tegmen (= basal piece + parameres), dorsal view. Figure 3: Penis, dorsal view. Figures

4-6. Pseudodendroides niponensis (Lewis), adult male. Figure 4: Abdominal sternite 8, ventral view.

Figure 5: Tegmen (= basal piece + parameres), dorsal view. Figure 6: Penis, dorsal view.

6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

= **

2 were tre

anaes

a a Sorat

e

¢

Figures 7-8. Figure 7: Pseudodendroides niponensis (Lewis), adult male, cranium, including cra-

nial pits, stereo-pair, dorsal view. Figure 8: Pseudodendroides umenoi (K6no), adult male, cranium,

including cranial pits, stereo-pair, dorsal view.

222 (TARD; 15 Jul 1982, S.C. Lin & C.N. Lin, 12 (TARI); Tsuifeng, 2300 m, 23-25 Jun 1983,

K. S. Lin & S. C. Lin, 22 2 (TARI); TAICHUNG HSIEN: 8 Mar 1977, HOMEOTYPE: Pseudopy-

rochroa umenoi K6no, Daniel K. Young, Elytra of type a bit lighter, 1d (DYCC); Jul 1977, 12

(DYCC); Anmashan, 2275 m, 6-9 Jul 1979, L. Y. Chou, 12 (TARD); HSINCHU: Kwangou, 2000 m,

24 Jun 1985, J. B. Heppner, 1¢ (FSCA). Pseudodendroides uraiana: TAIWAN. [Formosa], Urai

[underside of label], 24 Apr 1925 [underside of label], T. Kano, Pseudodendroides uraiana Kéno, @

[Holo]Type, “description based on 1, this must be the HOLOTYPE, Daniel K. Young, 1992”

(HUSC); HOOZAN: [Formosa], Sauter, 22 2 (DYCC); [Formosa], Sauter, 22 2 (NHMYV); Apr 1910,

H. Sauter S. G., Zool. Mus. Berlin, 26 ¢ (ZMHB); Apr 1910, H. Sauter S. G., Zool. Mus. Berlin,

HOMEOTYPE: Pseudodendroides uraiana K6no, Daniel K. Young, Type is slightly more teneral -

lighter pn. etc., 12 (ZMHB); [Formosa, (Hoozan) Hosan], Mar 1910, Sauter S., Zool. Mus. Berlin,

13d, 12 (ZMHB); POLISHA: [Formosa], Apr 1910, Sauter S., Zool. Mus. Berlin, 12 (ZMHB).

ACKNOWLEDGMENT

This study represents part of a larger project relating to the systematics and

phylogeny of Pseudopyrochroa. 1 thank Malcolm Kerley, Christina von Hayek

and Sharon Schute (BMNH) for their kind assistance during my visits, Darren

Pollock (DAPC), Mike Thomas (FSCA), Heinrich Sch6nmann (NHMV), Claude

Girard (PMNH), Liang-yih Chou (TARD), and Fritz Hieke and Manfred Uhlig

1999 YOUNG: PSEUDOPYROCHROA TRANSFERRED 7

(ZMHB) for loans of material, Sadao Takagi, Entomological Institute, Hokkaido

University for loan of type material for Pseudodendroides umenoi and P. uraiana,

and Takehiko Nakane for exchanges of pyrochroid material that enabled me to

make the necessary comparisons. Mr. Charles Thomas, University of Wisconsin

Integrated Microscopy Resource, was instrumental in helping me resolve the prob-

lem of illustrating the cranial pit apparatus. To our knowledge, this represents the

first application of stereo-pair techniques to confocal microscopy. This research

was supported in part by grants from the National Science Foundation (BSR-

9006342), the University of Wisconsin Graduate School (900159), and the Uni-

versity of Wisconsin’s Natural History Museums Council Small Grants Program.

LITERATURE CITED

Blair, K. G. 1914. A revision of the family Pyrochroidae (Coleoptera). Ann. & Mag. Nat. Hist., 13:

310-326.

Kéno, H. 1936. Uber die Kafersammlung des Museums “‘Umeno Konchii Kenkiujo” I. Pyrochroidae,

Meloidae, Rhipiphoridae. Bull. Umeno Entomol. Lab. Kurume, 3: 1-6.

Nakane, T. 1960. On the Pyrochroidae of Japan (Coleoptera). Entomol. Rev. Japan, 11: 59-66.

Nakane, T. 1988. Notes on some little known beetles (Coleoptera) in Japan, 2. Kita-Kyushu no Konchu,

35: 1-6.

Ohbayashi, N. 1968. On two species of Pyrochroidae from Amami-Oshima Island. Entomol. Rev.

Japan, 20: 35-36.

Pic, M. 1906. Contribution a l’étude des Pyrochroides. Echange, 22: 28-30.

Pic, M. 1913. Introduction. Mélanges exotico-ent., 8: 1-2.

Pic, M. 1938. Mutations et Coléoptéres nouveaux. Echange, 54: 14-16.

Pic, M. 1955. Descriptions et notes. Diversités Entomologiques, XV: 9-16.

Young, D. K. 1975. A revision of the family Pyrochroidae (Coleoptera: Heteromera) for North America

based on the larvae, pupae, and adults. Contrib. Amer. Entomol. Inst., 11: 1-39.

Received Jun 1995; Accepted 27 Aug 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 8-12, (1999)

STUDIES ON THE CHRYSOMELIDAE (COLEOPTERA) OF

THE BAJA CALIFORNIA PENINSULA: A NEW SPECIES

OF SCELOLYPERUS (GALERUCINAE), WITH NOTES ON

THE GENUS IN BAJA CALIFORNIA

ARTHUR J. GILBERT! AND FRED G. ANDREWS2

‘Department of Food & Agriculture, Division of Plant Industry, Pest Detection,

2889 N. Larkin #106, Fresno, California 93727

Department of Food & Agriculture, Division of Plant Industry, Analysis &

Identification, 1220 N. Street, Sacramento, California 95814

Abstract——Scelolyperus clarki NEW SPECIES is described from Baja California, Mexico. Notes

on the hosts and distribution for Scelolyperus Crotch species in Baja California are presented.

Key Words.—Insecta, Scelolyperus, clarki, phoxus, torquatus, varipes, Baja California Peninsula,

Baja California, Mexico, Coleoptera, Chrysomelidae, Galerucinae.

The North American species of the genus Scelolyperus Crotch have very re-

cently been reviewed by Clark (1996). During a visit to the University of Cali-

fornia Essig Museum to obtain specimens for our continuing work on the Chry-

somelidae of the Baja California Peninsula, a series of specimens of Scelolyperus

was found that at first was thought to be a very common species, S. torquatus

(LeConte). A precautionary check of the aedeagus revealed a new species. All

species of Scelolyperus found on the Baja California Peninsula occur in the north-

ern part of the State of Baja California (Fig. 7).

Specimen Depositories.—The following abbreviations refer to: CAS—Califor-

nia Academy of Sciences, CDFA—California Department of Food & Agriculture,

UNAM— Universidad Nacional Autonoma de Mexico, UCBC—University of

California, Berkeley Collection.

SCELOLYPERUS CLARKI Gilbert & Andrews, NEW SPECIES

Types.—Holotype (male) and Allotype (female): MEXICO. BAJA CALIFOR-

NIA: 11.3 km (7 mi.) SE Maneadero, 25 Mar 1973, 100’ el., J. Doyen, on Cea-

nothus: Type and Allotype deposited in the University of California, Berkeley

Collection. PARATYPES (20)-(9) Same data as holotype; (11) same data as ho-

lotype, except no host data given (2) [CAS]; (2) [CDFA]; (2) [UNAM]; (14)

[UCBC].

Description—Male (holotype). Length 3.9 mm; width 1.5 mm. Form elongate; prothorax testaceous,

narrower than elytra. Body color black, elytra metallic green or blue. Head dark brown to black, vertex

alutaceous, basically impunctate with metallic luster, a few inconspicuous setae near eyes and along

margin with interocular sulcus; interocular sulcus distinct; interocular width approximately 1.5 times

width of eyes (on a line drawn through center of eyes when viewed head on); eyes entire; frontal

tubercles distinct, smooth, flat, separated from each other by a distinct sulcus; tubercles separated from

interantennal carina by shallow sulci; antennal fossae separated by a distance subequal to length of

antennomere II; interantennal carina well developed, forming a longitudinal, angulate ridge; genal

length subequal to maximum width of antennomere I; antennae extending beyond humeri; antennom-

eres 1—4 totally or partially testaceous; 5-11 dark brown. Pronotum 1.2 times wider than long (width

measured at the widest portion—apical one-third); virtually glabrous (setose punctures visible under

high magnification), alutaceous with shallow punctures that become more coarse and dense posteriorly.

1999 GILBERT & ANDREWS: NEW SCELOLYPERUS SPECIES 9

Scutellum dark brown, polished, impunctate. Elytra 1.9 times longer than wide; slightly rugose (viewed

at an angle), alutaceous, investitus (very scattered inconspicuous setae visible under high magnifica-

tion) with dense, coarse, broad, irregularly placed punctures; most punctures separated by the width

of the punctures or less, occasionally coalescing. Venter black, pubescent; procoxal cavity open; pro-

coxae conical, narrowly separated; last ventrite with a short, broad, truncate lobe. Legs all of approx-

imately equal size, shape; femora black, tibia and tarsi brown (except base of protibia which is

testaceous). Genitalia as in Figs. 1 and 2.

Female (Allotype).—Similar to holotype, differing in the following characters: size slightly larger

(length 3.8 mm; width 1.4 mm); last abdominal sternum not lobed; dorsum shining, only faintly

alutaceous.

Variation.—Male: length 3.3—3.9 mm; width at elytral humeri 1.1—1.5 mm.

Female: length 3.1-3.9 mm; width 1.1—1.5 mm.

Diagnosis.—Scelolyperus clarki NEW SPECIES would key to couplet 11 in

the key presented by Clark (1996). However, it can be readily distinguished from

the two species in this couplet, S. torquatus (LeConte) and S. phoxus Wilcox, by

the aedeagus (Figs. 1—6) and the coarser elytral punctation. Examination of the

aedeagus will provide positive identification. The only other Scelolyperus species

recorded from Baja California, S. varipes, is larger, has a dark pronotum and a

very different aedeagus.

Host.—E leven of the twenty-two specimens in the type series of Scelolyperus

clarki were collected from Ceanothus (Rhamnaceae). No plant association was

given for the other 11 specimens in this series. Adults of S. torquatus are also

associated with Ceanothus and occupy the same habitat in Baja California.

Etymology—Named for Shawn Clark.

Material Examined.—See types.

SCELOLYPERUS PHOXUS Wilcox

Clark (1996) reports this species from Los Angeles and Riverside Counties in

California. A single female specimen collected 3.2 km (2 mi) SE of El Topo (Fig.

7), without host data, appears to be S. phoxus, but without a male specimen this

determination cannot be certain. This would extend the range into northern Baja

California. Most likely this species occupies similar habitat as in that portion of

California between Los Angeles County and Baja California. The senior author

has collected a large series of this species in association with Adenostoma fasci-

culatum Hooker & Arnott (Rosaceae) in the Mt. Baldy area of Los Angeles Coun-

ty. The Adenostoma must be in bloom for the adults to be found. It may be that

the adults are pollen feeding. However, beetles were very concentrated on indi-

vidual plants possibly indicating that they may be congregating for mating or

have just emerged from pupae in the soil beneath these plants. Other species of

perennial plants that were in bloom did not have beetles associated with them. A.

fasciculatum extends into Baja California (Wiggens 1980, Roberts 1989) and may

also have the same association with S. phoxus.

SCELOLYPERUS TORQUATUS (LeConte)

In California S. torquatus is a very common and widely distributed species.

Clark (1996) reports this species collected on a variety of plant species. However,

it can be found most commonly and abundantly on Ceanothus and Adenostoma

when these plants are in bloom. Clark (1996) and Wilcox (1965) both report S.

torquatus from Baja California. We have examined numerous specimens also

THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

1999 GILBERT & ANDREWS: NEW SCELOLYPERUS SPECIES

11

5 ALIFORNIA

tie Se

Tijuana 7

*Ht, f zon

! 4

o H

oO

A.

» 8, Ss

a

ot

(A) S.clarki Gilbert & Andrews

(*) S. phoxus Wilcox

di (@ ) S. torquatus (LeConte)

(@ ) S. varipes (LeConte)

Re Me

BAJA CALIFORNIA ol

§ in

ote és

» BS

es Guerrero

Sos

v i

: d

BAJA CALIFORNIA SUR | oe:

La Paz

1 inch = 72 miles

Figure 7. Known geographical distribution of Scelolyperus species in the Baja California Peninsula.

e

Figures 1-6. Male aedeagus. Figure 1. Scelolyperus clarki, lateral view. Figure 2. Scelolyperus

clarki, ventral view. Figure 3. Scelolyperus torquatus, lateral view. Figure 4. Scelolyperus torquatus,

ventral view. Figure 5. Scelolyperus phoxus, \ateral view. Figure 6. Scelolyperus phoxus, ventral view.

12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

collected in the very northern portion of Baja California; 0.62 km (1 m1) S El

Condor, 3.2 km (2 mi) (no direction given) Santo Tomas Arroyo, upper Canyon

del Cantil, Sierra Juarez, Ejido Uruapan, 23 km E. Ensenada, 47 km E. Ensenada,

77 km SE. Ensenada and 98 km SE. Ensenada (Fig. 7).

SCELOLYPERUS VARIPES (LeConte)

Scelolyperus varipes is distributed from British Columbia to Montana to New

Mexico and California (Clark 1996). We have examined five specimens from the

following five localities in Baja California: El Topo; 4.8 km (3 mi) S Laguna

Hansen; Las Encinas, Sierra San Pedro Martir; 9.7 km (6 mi) N Laguna Hansen,

Sierra Juarez and 3.5 km (2.2 mi) S El Topo, Sierra Juarez (Fig. 7).

ACKNOWLEDGMENT

Specimens were made available for this study by Cheryl Barr, University of

California, Berkeley—California Insect Survey, E. L. Sleeper, California State

University, Long Beach and Dave Faulkner, San Diego County Natural History

Museum.

LITERATURE CITED

Clark, Shawn M. 1996. The Genus Scelolyperus Crotch in North America (Coleoptera: Chrysomelidae:

Galerucinae). Insecta Mundi, 10: 261—280.

Roberts, Norman C. 1989. Baja California plant field guide. Natural History Publishing Company, La

Jolla, California.

Wilcox, J. A. 1965. A synopsis of the North American Galerucinae (Coleoptera: Chrysomelidae). Bull.

N.Y. St. Mus. Sci. Serv., no. 400: 1-226.

Wiggens, I. L. 1980. Flora of Baja California. Stanford University Press, Palo Alto, California.

Received 18 May 1998; Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 13-17, (1999)

A NEW SPECIES OF METAPHYCUS (HYMENOPTERA:

ENCYRTIDAE) PARASITIC ON SAISSETIA OLEAE

(OLIVIER) (HOMOPTERA: COCCIDAE)

KENT M. DAANE AND LEOPOLDO E. CALTAGIRONE

Center for Biological Control, Division of Insect Biology,

Department of Environmental Science, Policy & Management,

University of California, Berkeley, California 94720

Abstract.—A new encyrtid species of the zebratus-group of Metaphycus is described: Metaphy-

cus hageni NEW SPECIES. This parasitoid was reared from black scale, Saissetia oleae

(Olivier), collected on olives near Almufiécar, Spain. This species is similar to M. lounsburyi

(Howard)!, but can be distinguished by the relative length of the ovipositor, the shape of the

male genitalia, and the shape of the antennal club in both females and males. Characters that

differentiate M. hageni from closely related species are given.

Key Words.—Insecta, Hymenoptera, Encyrtidae, Metaphycus, Saissetia oleae.

Saissetia oleae (Olivier) were collected on olives, Olea europaea L., near Al-

mufiécar in southern Spain, in 1985, and shipped to the quarantine facility at the

(former) Division of Biological Control, University of California, Berkeley. Nu-

merous specimens of an encyrtid, identified by K. S. Hagen as Metaphycus sp.

nr. lounsburyi (Howard), using the Annecke & Mynhardt (1971) key to the ze-

bratus-group of Metaphycus species, emerged from the scales. We consider this

an unnamed and undescribed species, which we name and describe here. Our

description is based on specimens reared from S. oleae collected in Almufiécar,

individuals from their progeny reared in the insectary using S. oleae on oleander

(Nerium oleander L.), and specimens recovered from various sites in California

where the parasitoid was released in olive orchards infested with S. oleae (Daane

et al. 1991).

METHODS AND MATERIALS

Described specimens were preserved by different methods. Some specimens

were mounted dry, without previous treatment, on paper cards or points using

book-binders’ glue (Yes®) as adhesive. Others were mounted on glass slides, some

whole, some dissected as needed. Some specimens were cleared in chloral-phenol

(10 g phenol crystals, 10 g chloral hydrate, 3 ml distilled water) and mounted in

Faure’s medium (60 g lump gum arabic, 100 g chloral hydrate, 25 ml 50% glu-

cose, 25 ml glacial acetic acid, 120 ml distilled water). Measurements of various

structures were taken from slide-mounted specimens. The holotype is card mount-

ed, paratypes are both card mounted and slide mounted in Canada balsam follow-

ing Noyes (1982). Specimens are deposited in the Essig Museum (UCB), Uni-

versity of California, Berkeley; United States National Museum (USNM), Wash-

ington D.C.; and the National History Museum (BMNH), London.

' According to E. Guerrieri and J. S. Noyes (personal communication) the name M. lounsburyi

(sensu Compere [1940] and Annecke & Mynhardt [1971]) is based on a misidentification of the type

material (Howard 1898), which is redescribed in their manuscript (in preparation) that deals with the

European species of Metaphycus.

14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

METAPHYCUS HAGENI Daane and Caltagirone, NEW SPECIES

Types.—Holotype: female; data: SPAIN, ANDALUCIA: ~5 km west of Al-

mufiécar near “‘La Punta de la Mona’ tunnel, 6 Jun 1985, L. E. Caltagirone,

reared from Saissetia oleae (Olivier) collected on olive (Olea europaea L.), de-

posited: UCB. Paratypes: same data as holotype, 6 females, 4 males, deposited:

UCB. Albany, CALIFORNIA, ALAMEDA Co.: Insectary colony, University of

California, 11 Jan 1986,.K. M. Daane, reared from S. oleae on oleander (Nerium

oleander), 6 females, 4 males, deposited: USNM; 6 females, 4 males, deposited:

BMNH.

Female.—Length of air-dried specimens 1.3 mm (from frons to tip of ovipositor sheath, range

1.025-1.5, n = 11). Color variable: white to pale yellow to golden brown. -Frons, upper half of scrobes,

mesoscutum, axillae, and scutellum orange yellow, sometimes with a brownish hue; spot on middle

of scape extending longitudinally ventrally and dorsally, basal: half of pedicel, basal 2-or 3 funicular

segments, club, occiput except ridges, a spot on each side and middle of pronotum, anterior of me-

soscutum (seen in dissected, slide-mounted specimens), metanotum, propodeum, metasoma dorsally

(except for a narrow outer margin) dark brown. Tibiae basally, two oblique rings medially and apically

brown, sometimes with faded sections. Mandibles reddish brown fading to pale yellow at base. Ocelli

reddish brown. Eyes gray with greenish hue. Wings hyaline. Ocelli forming an acute triangle (~45°),

lateral ocelli about one-half their longest diameter from eye margin, distance from each other about

their longest diameter; middle ocellus separate from lateral ocelli by a distance about twice its longest

diameter. Scape width 0.36X length (range 0.32-0.40, n = 32); pedicel width 0.53 length (range

0.50-0.57, n = 20); pedicel 1.08X length of basal 3 funicular segments combined (range 1.05-1.27,

n = 13); basal 3 funicular segments subequal in width, the apical 3 gradually widening so that the

apical segment width 1.94 basal segment width (range 1.69-—2.12, n = 14); club.3-segmented, ovate,

its width 0.66X its length (range 0.55—-0.74, n = 31), its length 0.81X length of funicle (range 0.69-

0.93, n = 31) (Fig. 1). Fore wing width 0.44% its length (range 0.41—0.46, n = 30); length of stigmal

vein 0.22X length of submarginal vein (range 0:17—0.29, n = 32). Length of middle tibia 0.98 X length

of middle femur (range 0.94-1.04, n = 18); length of middle tibial spur 0.81 length of middle

basitarsus (range 0.71-0.85, n = 30). Length of hind tibia 1.08 length of hind femur (range 1.08—

1.22, n = 17); length of hind tibial spur 0.45X length of hind basitarsus (range 0.41-0.50, n = 18).

Length of ovipositor (measured as length of 2nd valvulae) 0.80 length of metasoma (range 0.71—

0.85, n = 13), and 1.06X length of hind tibia (range 1.0-1.15, n = 14); length of ovipositor sheaths

(3rd valvulae) 0.19X length of 2nd valvulae (range 0.17—0.21, n = 14) (Fig. 5).

Male.—Similar to female. Club entire, its length 0.74 (range 0.66-0.78, n = 12) length of funicle

(Fig. 3). Genitalia long-elliptical (Fig. 6).

Diagnosis.—Females of M. hageni can be separated from those of M. louns-

buryi (Howard) and Metaphycus bartletti Annecke & Mynhardt, the two mor-

phologically similar species found in California, by the following characters. M.

hageni ovipositor is as long or slightly longer than hind tibia, M. lounsburyi and

M. bartletti ovipositors are shorter than respective hind tibia. The antennal club

of M. hageni (Fig. 1) and M. bartletti is ovate, the apex gradually narrowing, the

antennal club of M. lounsburyi is truncate (barrel-shaped) (Fig. 2). Torular sen-

sillae present in M. hageni, absent in M. lounsburyi and M. bartletti. The genitalia

of male M. hageni is long-elliptical (Fig. 6), the antennal club rounded at apex;

the genitalia of M. lounsburyi is wedge-shape (Fig. 8), the antennal club truncate

at apex (Fig. 4).

DISCUSSION

Species of Metaphycus Mercet are important natural enemies of soft scale (Ho-

moptera: Coccidae). Metaphycus species in the zebratus-group are parasitic on

lecaniine scale (Annecke & Mynhardt 1971). The most commonly known of these

1999 DAANE & CALTAGIRONE: METAPHYCUS HAGENI NEW SPECIES 15

Figures 1-4. Figure 1. Metaphycus hageni NEW SPECIES, female antenna. Figure 2. Metaphycus

lounsburyi (Howard), female antennal club in outline. Figure 3. Metaphycus hageni NEW SPECIES,

male antennae. Figure 4. Metaphycus lounsburyi (Howard), male antennal club in outline. Scale =

0.1 mm.

is M. lounsburyi, a parasitoid of black scale, S. oleae. Metaphycus lounsburyi has

a wide geographic distribution, a result of South African material being imported

to many areas for improved control of S. oleae (Bartlett 1978). During initial

taxonomic and behavioral studies with the material imported from Spain, one of

us (LEC) and K. S. Hagen noted inconsistencies between specimens of M. louns-

buryi from California and the imported Metaphycus specimens. These observa-

tions led to taxonomic, cross-mating, and behavioral studies to determine if im-

ported material was a biotype of M. lounsburyi, as has been reported to occur by

Panis & Marro (1978), or a different species. Females were never produced in

cross-mating experiments (Barzman et al. in press). Observations of oviposition

behavior and host-feeding revealed M. lounsburyi females deposit eggs through

the ventral side of the scale and were not observed host-feeding. Metaphycus

hageni females deposit eggs through the dorsum of the scale and frequently feed

on host body fluids exuding from a puncture through which an egg is never

deposited. The collective evidence indicates that M. lounsburyi and M. hageni are

separate species.

Etymology.—This species is named in honor of our esteemed colleague, the

late Kenneth S. Hagen, who was an invaluable adviser during the early stages of

our research with this parasitoid and a friend throughout our careers.

Material Examined.—See types. CALIFORNIA, ALAMEDA Co.: Insectary of the (former) Division

of Biological Control, University of California, Albany, California, 11 Jan 1986, K. M. Daane, reared

from S. oleae on oleander (N. oleander), 20 females, 9 males; slide mounted in Faure’s; deposited:

16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Figures 5-8. Figure 5. Metaphycus hageni NEW SPECIES, female ovipositor, ventral view, show-

ing position on metasoma. Scale = 0.1 mm. Figure 6. Metaphycus hageni NEW SPECIES, male

genitalia, ventral view. Scale = 0.05 mm. Figure 7. Metaphycus lounsburyi (Howard), female ovi-

positor, ventral view, showing position on metasoma. Scale = 0.1 mm. Figure 8. Metaphycus louns-

buryi (Howard), male genitalia, ventral view. Scale = 0.05 mm.

Kearney Agricultural Center (KAC), University of California, Parlier. CALIFORNIA, TEHAMA Co.:

1 km east of Corning, 200 m, 13 Nov 1985, K. M. Daane, reared from S. oleae on olive (O. europaea),

3 females; slide mounted in Faure’s; deposited: KAC.

ACKNOWLEDGMENT

We thank John S. Noyes (The National History Museum, London) for review-

ing this manuscript and help with diagnostic characters of species in the zebratus-

group of Metaphycus and Gregory Zolnerowich (Texas A&M University) for

discussions on Metaphycus species taxonomy.

LITERATURE CITED

Annecke, D. P. & M. J. Mynhardt. 1971. The species of the zebratus-group of Metaphycus Mercet

(Hymenoptera: Encyrtidae) from South Africa with notes on some extralimital species. Rev.

Zool. Bot. Afr., 83: 322-360.

1999 DAANE & CALTAGIRONE: METAPHYCUS HAGENI NEW SPECIES 17

Bartlett, B. R. 1978. Coccidae. In Introduced parasites and predators of arthropod pests and weeds: a

world review. Agriculture Handbook no. 480, Agricultural Research Service, USDA.

Barzman, M. S., K. M. Daane, L. E. Caltagirone & K. S. Hagen. Metaphycus hageniand M. lounsburyi

(Hym.: Encyrtidae): two discrete species parasitic on the black scale, Saissetia oleae (Hom.:

Coccidae). BioControl, (In Press).

Compere, H. 1940. The African species of Metaphycus, Mercet. Bull. Entomol. Res., 31: 7-33.

Daane, K. M., M. S. Barzman, C. E. Kennett & L. E. Caltagirone. 1991. Parasitoids of black scale in

California: establishment of Prococcophagus probus Annecke & Mynhardt and Coccophagus

rusti Compere (Hymenoptera: Aphelinidae) in olive orchards. Pan-Pac. Entomol., 67: 99-106.

Howard, L. O. 1898. On some new parasitic insects of the subfamily Encyrtinae. Proc. U.S. National

Museum. 21: 231-248.

Noyes, J. S. 1982. Collecting and preserving chalcid wasps (Hymenoptera: Chalcidoidea). J. Natur.

Hist., 16: 315-334.

Panis, A. & J. PR. Marro. 1978. Variation du comportement chez Metaphycus lounsburyi [Hym.: En-

cyrtidae]. Entomophaga, 23: 9-18.

Received 14 Jul 1998; Accepted 29 Sep 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 18-22, (1999)

SYNONYMY OF DASYMUTILLA NOCTURNA MICKEL

(HYMENOPTERA: MUTILLIDAE)!'

DONALD G. MANLEY

Department of Entomology, Clemson University,

Pee Dee Research and Education Center,

2200 Pocket Road, Florence, South Carolina 29506-9706

Abstract—Dasymutilla nocturna Mickel and D. subhyalina Mickel were thought to be female

and male, respectively, of the same species since their description in 1928. One female is known

from Blythe, Riverside County, California. Another female bears a collection label from Preston,

Nevada. All other known females and males are from the Colorado Desert of Imperial County,

California. In 1947, D. paranocturna Barr & Hurd was described. The known specimens of D.

paranocturna probably represent two different species, D. nocturna and D. arenivaga Mickel.

Subsequent collections, the use of caged females, and comparison with type specimens have led

to the conclusion that D. nocturna, D. subhyalina, and at least part of the D. paranocturna

specimens are the same species. A complete synonymy is included.

Key Words.—Insecta, Hymenoptera, Mutillidae, Dasymutilla nocturna, Dasymutilla subhyalina,

Dasymutilla paranocturna, Dasymutilla arenivaga, California.

Dasymutilla nocturna was first described by Mickel (1928) on the basis of two

females. At the same time, Mickel described D. subhyalina from two males,

acknowledging that these specimens most probably were female and male of the

same species. The holotype female and both the holotype and paratype males

were collected at light at about 23:00 h on 10 Aug 1917 near Andrade, California,

Colorado Sand Desert (Imperial County), by J. Bequaert. The paratype female

was collected on 9 Aug 1914 near Brawley, Imperial County, California, by J. C.

Bradley. The male paratype was in the collection of J. Bequaert. It is now in the

Museum of Comparative Zoology, labeled as D. nocturna. All of these specimens

have been examined by this author.

Dasymutilla paranocturna was described by Barr & Hurd (1947) on the basis

of two female specimens. The holotype was collected from Blythe, Riverside

County, California, on 6 Jul 1946, by W. E Barr. The paratype was collected from

San Felipe Creek, Imperial County, California on 17 Jun 1940, by R. G. Dahl. It

is in the collection of the University of California, Berkeley. No mention was

made of the time of day when these two specimens were collected although they

were most likely collected at night (Barr, personal communication). Both of these

specimens have also been examined by this author.

Nearly all of the approximately 150 known species of Dasymutilla are diurnal.

There are only five known exceptions. These include D. nocturna and some of

the D. paranocturna, both of which are known from females only, and D. sub-

hyalina, known from males only. All three of these occupy the same geographic

range. They also include D. arenivaga Mickel and the remaining D. paranocturna,

known from females only, and D. megalophthalma Mickel, known from males

only. These occupy the same geographic range.

' Technical contribution no. 4320 of the South Carolina Agricultural Experiment Station, Clemson

University.

1999 MANLEY: SYNONYMY OF DASYMUTILLA 19

Material Examined.—In addition to the type specimens mentioned above, the following material

has been examined (all are females of D. nocturna and males of D. subhyalina, except as noted):

USA. CALIFORNIA. IMPERIAL Co.: Westmoreland, 20 Jul 1928, 1 female; Holtville, 22 Oct 1936,

A. T. McClay, 1 female; Laguna Lake, 9-11 Jun 1950, 1 female (identified as D. paranocturna,

probably D. arenivaga); Grays Well, 6 Jun 1951, D. J. and J. N. Knull, 1 male; Fort Yuma, near

Colorado River, 1 Jul 1951, 1 female (identified as D. paranocturna, probably D. arenivaga); Algo-

dones Sand Dunes, 9.6 km W of Glamis, 18 Nov 1963, M. E. Irwin, 2 females; 9.6 km W of Glamis,

18 Nov 1963, E. I. Schlinger, 1 male; Glamis, 8 February 1964, M. E. Irwin, E. I. Schlinger, 1 female;

32 km E of Brawley, 13 Jun 1965, G. R. Balmer, 1 male; Glamis, 11 Oct 1972, C. Goodpasture, 1

male; 4.8 km SW of Glamis, 10 Jul 1974, J. Doyen, 2 males; 1.6 km NW of Glamis, 11 Jul 1974,

D. G. Manley, 4 males; 4.8 km SW of Glamis, 12 Jul 1974, J. Doyen, 5 males; 1.6 km NW of Glamis,

7 Aug 1974, D. G. Manley, 13 males; 3.2 km NW of Glamis, 4 Nov 1974, J. A. Powell, 4 females;

6.4 km NW of Glamis, 1 May 1975, R. Aalbu, 7 females; 3.2 km NW of Glamis, 22 May 1975, D.

G. Manley, 4 females; 3.2 km NW of Glamis, 4 Jun 1975, T. Allen, 1 female; 1.6 km S of Glamis,

29 Mar 1978, R. Dietz and J. Powell, 2 females; 1.6 km S of Glamis, 31 Mar 1978, R. Dietz, 1

female; Glamis, 19 Sep 1980, K. A. Smith, 1 male; 1.6 km NW of Glamis, 12 May 1992, D. G.

Manley, 8 females; 1.6 km NW of Glamis, 13 May 1992, D. G. Manley, 9 females; 1.6 km NW of

Glamis, 11 Jul 1992, D. G. Manley, 6 females; 1.6 km NW of Glamis, 11 Jul 1992, D. G. Manley,

1 female (D. paranocturna); 1.6 km NW of Glamis, 28 Aug 1992, D. G. Manley, 7 females; 1.6 km

NW of Glamis, 28 Aug 1992, D. G. Manley, 2 females (D. paranocturna); 1.9 km W of Glamis, 24

Apr 1993, J. D. McCarty, 2 females; Glamis, 24 Jul 1995, D. G. Manley, 5 females and 22 males.

RIVERSIDE Co.: Salton Sea, 22 Jul 1952, H. L. Mathis, 1 female (identified as D. nocturna, clearly

D. arenivaga). SAN BERNARDINO Co.: 8 km NE of Yermo, 26 Jun 1939, W. M. Pearce, 1 female

(identified as D. paranocturna, probably D. arenivaga),; Kelso Dunes, 12.8 km SW of Kelso, 14-15

Jul 1974, J. Doyen, 2 females (identified as D. nocturna, clearly D. arenivaga). ARIZONA. YUMA

Co.: Yuma, 6 May 1939, R. M. Bohart, 1 female (identified as D. nocturna, probably D. arenivaga).

NEVADA. WHITE PINE Co.: Preston, Oct 1941, U. N. Lanham, 1 female. NO DATA. 1 male.

DISCUSSION

From the time of their original descriptions, it seemed likely that D. nocturna

and D. subhyalina were female and male of the same species. However, Mickel

(1928) did not want to make such a definitive statement solely on the basis of

the fact that one female and two males were collected at the same time and

location.

Females and males of many species of Dasymutilla are very similar in color

and pattern. However, in many other species, male and female color patterns are

very different from each other. This has made sex correlation of many Dasymutilla

species quite difficult.

Nearly all species in the genus Dasymutilla are diurnal. The only five excep-

tions have already been mentioned. Of those, D. nocturna, some of the D. par-

anocturna specimens, and D. subhyalina all share the same colors, pattern, and

geographic range. Dasymutilla arenivaga, some D. paranocturna specimens, and

D. megalophthalma have different colors, pattern, and geographic range.

Two other specimens of D. nocturna were collected prior to the description of

D. paranocturna (Hurd, 1951), both from Imperial Co., California. On the basis

of the type specimens, and the slight variations between those examined by Mickel

and those examined by Barr and Hurd, the latter saw fit to describe their speci-

mens as a new species.

The holotype of D. paranocturna is undoubtedly a specimen of D. nocturna,

with only slight variation in color of the pubescence and integument. Likewise,

the paratype of D. paranocturna is undoubtedly a specimen of D. arenivaga, with

only slight variation in color of the pubescence and integument. The specimens

20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

identified as D. paranocturna that more closely resemble D. nocturna tend to be

from the Algodones Sand Dunes or very nearby. The specimens identified as D.

paranocturna that more closely resemble D. arenivaga are more widely distrib-

uted.

It is possible that all five of the ‘nocturnal’? Dasymutilla represent a single

species. However, numerous specimens of D. arenivaga and D. megalo phthalma

have been examined, including both holotypes. Although they are obviously

closely related to D. nocturna and D. subhyalina, respectively, they appear to be

distinct. These undoubtedly represent female and male, respectively, of the same

species.

The specimen of D. nocturna that bears the collection locality of Preston, Ne-

vada presents another problem. There is no question that the specimen is D.

nocturna. However, no other specimen of this species has been found within 1100

km of Preston, Nevada. It seems much more likely that the specimen was collected

in or near Imperial Co., California, and that it was mislabeled.

In July and August of 1974, 17 additional specimens of D. subhyalina were

collected by the author from the Algodones sand dunes in Imperial County, Cal-

ifornia. No females were collected during that time. In May and June of 1975,

the author collected 12 additional specimens of D. nocturna from the same lo-

cation. However, no males were collected at that time. All specimens were col-

lected at night. These specimens were subsequently compared to the respective

holotypes and found to be identical. Although these additional specimens were

all collected at night from the same location, no definitive statement could be

made on conspecificity as specimens were not collected at the same time.

Additional specimens of D. nocturna were collected by the author in 1992, all

from the same location. All were either crepuscular, matinal, or nocturnal in their

habits. Eight specimens were collected on 12 May, and nine others on 13 May.

Seven additional females were collected on 11 July. One of these specimens

differed slightly with respect to color of pubescence and integument. It was sub-

sequently compared to the holotype of D. paranocturna and found to be mor-

phologically identical. On 28 August, an additional nine specimens were collected

by the author from the same location. Seven had the pubescence and integumental

coloration characteristic of D. nocturna; two had the slightly lighter-colored pu-

bescence and reddish integument characteristic of D. paranocturna. Again, no

males were taken during any of these collection dates. Diurnal collecting was

done during all of the preceding collection dates. However, no mutillid specimens

were ever taken except at dusk, during darkness, or right at dawn.

A female of D. nocturna was collected on 24 Jul 1995 on the Algodones sand

dunes shortly after dark, and placed in a small, plastic cage from which the ends

had been cut out and replaced with wire screen. A short time later, a male of D.

subhyalina was attracted to the caged female, and attempted to mate with her

through the wire screen. Later in the evening, another male was attracted to an

uncaged female. Both were collected. In all, four females and 22 males were

collected between 19:45 h (PDT) and 22:00 h (PDT).

Caged females have been used previously to attract males, both in cases where

the male was already known and where the male was unknown. Males are not

always attracted to the caged females. However, in all cases in which males have

been attracted to the caged females, they have been subsequently shown to be

1999 MANLEY: SYNONYMY OF DASYMUTILLA 21

conspecific. And in cases where the males were previously known, they have

always been of the same species as the caged female. Thus, the use of caged

females is a reliable method for use in sex correlations among Dasymutilla.

CONCLUSIONS

Even before the addition of the present evidence, it seemed likely that D.

nocturna and D. subhyalina represented female and male, respectively, of the

same species. Furthermore, it seemed likely that D. paranocturna was also just a

slight variant of the same species. The following facts lead me to believe that D.

nocturna, many of the specimens of D. paranocturna, and D. subhyalina are

members of a single species: 1) all are nocturnal, 2) all share the same geographic

range, the Colorado Desert, 3) new evidence that numerous individuals have been

found in the same place at the same time, 4) males have been observed attracted

to and trying to mate with caged females. Because the name D. nocturna has

precedence over the other two, that name shall stand. A synonymy for the species

follows.

DASYMUTILLA NOCTURNA MICKEL

Dasymutilla nocturna Mickel, 1928: 279. Type locality: CALIFORNIA. IMPE-

RIAL Co.: Colorado Sand Desert, near Andrade. Holotype deposited University

of Minnesota. @

Dasymutilla subhyalina Mickel, 1928: 281. Type locality: CALIFORNIA. /M-

PERIAL Co.: Colorado Sand Desert, near Andrade. Holotype deposited Uni-

versity of Minnesota. NEW SYNONYM. 3

Dasymutilla paranocturna Barr & Hurd, 1947: 88. Type locality: CALIFORNIA.

RIVERSIDE Co.: Blythe. Holotype deposited California Academy of Sciences,

Entomology (No. 5619). NEW SYNONYM. &

ACKNOWLEDGMENT

I thank Rolf Aalbu for assistance in the collection of specimens from the Al-

godones sand dunes in 1974 and 1975; Jeff Bolton, Stephanie Manley and Char-

lotte Singleton for assistance in the collection of specimens, and Ben Kofki and

Nancy Nicoli of the El Centro Resource Area, and the Bureau of Land Manage-

ment for permission to collect and study specimens from the Algodones sand

dunes in 1995; Phil Clausen, University of Minnesota, for allowing me to examine

the holotypes of Dasymutilla nocturna and D. subhyalina; Wojciech Pulawski,

California Academy of Sciences, for allowing me to examine the holotype of D.

paranocturna; Philip Perkins, Museum of Comparative Zoology, for allowing me

to examine the paratype of D. subhyalina; Cheryl Barr, University of California,

Berkeley, for allowing me to examine the paratype of D. paranocturna as well

as other specimens of all of these species; Steve Heydon, University of California,

Davis, for allowing me to examine specimens of these species; and Serguei Triap-

itsyn, University of California, Riverside, for allowing me to examine specimens

of these species. Fieldwork for collection and study of the specimens in 1992 was

supported, in part, by a grant from the National Science Foundation (BSR-

9106369).

22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

LITERATURE CITED

Barr, W. E & P. D. Hurd, Jr. 1947. Notes on the Dasymutilla of the Palo Verde Valley, California with

the description of a new species (Hymenoptera: Mutillidae). Pan-Pac. Entomol., 23: 85-90.

Hurd, P. D., Jr. 1951. The California velvet ants of the genus Dasymutilla Ashmead (Hymenoptera:

Mutillidae). Bull. Calif. Insect Surv., 1: 89-118.

Mickel, C. E. 1928. Biological and taxonomic investigations of the mutillid wasps. U.S. Nat. Mus.,

Bull., 143: 1-351.

Received 4 Aug 1997; Accepted 30 Sep 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 23-31, (1999)

HOST RECORDS OF BRACONIDAE (HYMENOPTERA)

OCCURRING IN MIRIDAE (HEMIPTERA:

HETEROPTERA) FOUND ON LODGEPOLE PINE

(PINUS CONTORTA) AND ASSOCIATED CONIFERS

J. D. LATTIN! AND N. L. STANTON?

‘Systematic Entomology Laboratory,

Department of Entomology,

Oregon State University,

Corvallis, Oregon 97331

*Department of Zoology and Parasitology,

University of Wyoming,

Laramie, Wyoming 82071-3166

Abstract—The plant bug (Hemiptera: Heteroptera: Miridae) fauna of lodgepole pine (Pinus

contorta Douglas ex Loud) and associated conifers was examined in Oregon and Wyoming,

United States of America. Parasitoid larvae of Braconidae were recovered from 20 species of

Miridae previously unrecorded as braconid hosts, representing 10 genera of bugs, four of which

have not been previously recorded as hosts. The parasitoid larvae were found only in the im-

mature stages of the bugs. A sequence of species of Miridae occurs on the host tree through

time but only the earlier species are parasitized. Pinus contorta is the most widespread conifer

in North America, its four subspecies extending from Baja California Norte to the Yukon Ter-

ritory, Canada. Over 50 species of Miridae have been found on this tree species. Other parasitized

species will certainly be found.

Key Words.—Insecta, Braconidae, conifers, Miridae, parasitoids, plant bugs, Pinus.

The only known Hymenoptera parasitoids of immature and adult Miridae are

the Braconidae, Leiophron Nees and Peristenus Foerster (Brindley 1939; Leston

1959, 1961; Loan 1974a, b, 1980, 1983; Glen 1977; Marsh 1979; Wheeler &

Loan 1984). Some of the Miridae genera contain economically important species

(e.g., Adelphocoris Reuter, Leptopterna Fieber, and Lygus Hahn) and work is

being carried out to utilize these parasitoids for biological control. Considerable

effort has been made to collect these two genera of Braconidae from various parts

of the world and bring them into North America for ultimate release. Leston

(1961) reported 31 genera and 51 species of Miridae parasitized by Braconidae

in Great Britain and Lattin & Ozanne (1993) added additional species there. Marsh

(1979) listed 19 genera and 28 species parasitized by species of Leophron and

Peristenus in North America. Loan (1980) reported additional bug genera and

species, bringing the total to 22 genera and 34 species. Here, we add 20 additional

bug species found in 10 genera that were parasitized by braconids, including four

additional genera.

We collected Miridae at six sites in Oregon and Wyoming during the 1986

season. Seven sites in Oregon, including the three in this study, were sampled

every two weeks in 1985 and in 1988. Most of the effort was made on three

subspecies of Pinus contorta Douglas ex Loud and, to a limited extent, the other

species of conifers found in association with these pines. Pinus contorta is the

most widespread species of pine in North America and is considered to have four

subspecies (Critchfield 1957, 1980, 1984, 1985; MacDonald & Cwynar 1985, but

24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

see Forrest 1980a, b). Three subspecies were sampled: Pinus contorta latifolia

Engelmann in Wyoming and Oregon, P. contorta contorta Douglas ex Loud and

P. contorta murrayana (Greville and Balfour) Engelmann in Oregon. Approxi-

mately 20 genera and 50 species of Miridae are found on Pinus contorta based

on the available literature and our studies, and material supplied to us by several

mirid specialists. Because parasitism was known to occur chiefly in the nymphal

stages, samples of late instar nymphs were dissected, but samples of adults also

were examined. We found 10 genera and 18 species of Anthocoridae (Hemiptera:

Heteroptera) on the same sites and “‘host”’ trees (Lattin & Stanton 1992).

Leston (1959, 1961) described a group of mirids he referred to as ‘“‘arboreal

and early’? whose parasitoids emerged from the adult stage. At least four other

species he listed had the larva emerging from mature bug nymphs during the

same period. Brindley (1939) stated that the mirid is usually parasitized as a

second- or third-instar nymph. That would place the time of parasitoid oviposition

about three or four weeks ahead of emergence of the mature parasitoid larva.

Parasitism prolongs the time required for the bug to reach maturity, if it ever does

(Leston 1959, Loan 1983). The stylized life history of a mirid parasitoid is shown

in Fig. 1 (after Loan 1983). We found mature parasitoid larvae only in nymphs

and none in adults. Observation numbers were low for some taxa because of the

scarcity of parasitized nymphs but species of other genera (e.g., Phoenicocoris

Reuter and Microphylellus Reuter) were well represented.

MATERIALS AND METHODS

We collected Hemiptera: Heteroptera from six sites in Oregon and Wyoming

during the 1986 season. Seven sites in Oregon, including the three in this study,

also were sampled regularly during 1985. Sampling was done by a beating sheet

held under a branch. Ten beats with an ax handle were made and all bugs col-

lected. If any bugs were found, another ten beats were made. This effort was

continued until no further bugs were collected in a ten beat effort. Ten trees were

sampled at each site every two weeks throughout the season. At times, such efforts

would produce over 100 specimens of Miridae on a single branch. Such regular

sampling collected all instars of most species. Sampling dates began before most

bugs appeared and continued into late summer until no further specimens were

recovered.

Site Descriptions

Oregon

Site 5—South Beach State Park (SBSP) is 1.6 km south of Newport, Lincoln

County, Oregon at an elevation of 7 m (SW % of Sec. 20, T11S, R11W) and is

adjacent to the Pacific Ocean. A vigorous young stand of shore pine (P. contorta

contorta) was sampled on a partially stabilized low dune behind the foredune. A

few, very small, Sitka spruce (Picae sitchensis) (Bongard) Carriére) were scattered

through the stand with low willow (Salix sp.) adjacent. Older, stabilized dunes to

the east contained larger shore pine and some large Sitka spruce.

Site 7—Ochoco Mountains (OM) is 40 km east of Prineville, Crook County,

Oregon at an elevation of 1476 m; specifically 0.3 km south of Hwy 25 on FS

road 2620 (Sw % of Sec. 2, T13S, R19E). Most sampling was done on the west

side of FS 2620, on an easterly facing slope. The site contained an open stand of

1999 LATTIN & STANTON: MIRIDS PARASITIZED BY BRACONIDS 25

WINTER

Parasitoid Wasp

adult diapauses

overwinters in cocoon in soil

Host Bug

usually overwinters as egg

in conifer branches

Wasp

adult emerges April-May

Wasp Bug N

cocoon in soil egg hatches PS

a nymph develops ye)

= Bu _—

<_ pug . 2

eggs in conifer Wasp

Fy brenches adult oviposits in first -)

or second instar nymph

wasp larva emerges

from fifth instar nymph

or (rarely) adult May-June

Wasp),

larva pupates in soil

Bug

adult lays eggs

in conifer branches

SUMMER

Figure 1. Generalized life cycle of braconid parasitoid and host mirid (after Loan 1983).

mature ponderosa pine (P. ponderosa Douglas ex Lawson) with some regenera-

tion, a few larch (Larix occidentalis Nuttall), and mostly young lodgepole pine

(P. contorta latifolia). A few mature trees also were present.

Site S—Three Creeks Meadow (TCM) site is 26 km south of Sisters, Deschutes

County, Oregon at an elevation of 2069 m (Sw % of Sec. 13, T17S, RYE). It is

a moist, subalpine meadow dissected by several small streams. Sierra lodgepole

pine (P. contorta myrrayana) occurs around the edge of the meadow and up onto

the drier slopes surrounding it. Engelmann spruce (Picea engelmanni Parry ex

Engelmann) was scattered within the pine stand in the meadow. Subalpine fir

(Abies lasiocarpa (Hooker) Nuttall occurred among the pine on the drier slopes

surrounding the meadow. Sampling was done on trees in the southeastern corner

of the meadow, which consisted of both mature and young trees. A few of the

older trees were being attacked by the mountain pine beetle, Dendroctonus pon-

26 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

derosa Hopkins. The dwarf mistletoe, Arcenthobium americanum Nuttall ex En-

gelmann, was common on the pines, especially on the older trees.)

Wyoming

Site 2—Happy Jack road (HJR) sites are in the Laramie Range, Medicine Bow

National Forest, 19 km east of Laramie, Albany County, Wyoming at an elevation

of 2500 m. Site 2.1 (Yellow Pole Campground). They had open stands of mature

timber (P. flexilis James) and ponderosa pine (P. ponderosa) on a south-facing

slope (Sec. 25, TI5N, R72W). Stands of mature lodgepole pine (P. contorta

latifolia) were sampled a few km east of this spot at site 2.2 (Sec. 30, T15N,

R71W). Lodgepole pines were sampled in both open stands on a south-facing

slope, and closed stands on a north facing slope.

Site 3—Sand Lake Road (SLR) site is on the east side of the Snowy Range

Mountains about 60 km west of Laramie, Albany County, Wyoming (Sec. 17,

T16N, R78W). The specific site lies immediately east of the North Fork camp-

ground at an altitude of 2800 m. The stand had been thinned and consisted of

second growth lodgepole pine (P. contorta latifolia) interspersed with subalpine

fir (A. lasiocarpa) and some Engelmann spruce (P. engelmannii).

Site 4—French Creek Road (FCR) sites are about 90 km west of Laramie

(Carbon County) on the west side of the Snowy Range and lie along a back road

cutting south from Hwy 130 to French Creek. Site 4.1, at an altitude of 2750 m,

was a Clear cut east of and adjacent to the highway. (Sec. 1 and 17, TI5N, R81W).

Regeneration consisted primarily of lodgepole pine (P. contorta latifolia) with

occasional fir and spruce. Site 4.2 (Sec. 14, TI5N, R81W) was at 2650 m. Site

4.3 (Sec. 19, TISN, R8OW) was at 2600 m.

PARASITIZED TAXA

Taxonomic problems in several of these plant bug genera prevent precise spe-

cies assignment and, in at least one instance, involves an undescribed species (to

be described later). The presentation of species follows the catalog of Henry &

Wheeler (1988) and several later publications that appeared after that date (Stone-

dahl 1988, 1990; Schuh 1995; Stonedahl & Schwartz 1996).

Deraecorinae: Clivinematini

Largidea Van Duzee is a North American genus containing 10 species (Henry

& Wheeler 1988) all confined to conifers and most on Pinus spp. Largidea sho-

shonea Knight was the only species found with braconid larvae: Collection was

made in Wyoming, site 2.2 (HJR) ex. Pinus contorta latifolia, 9 and 25 Jul 1986

from V instar nymphs.

Deraeocorinae: Deraeocorini

Deraeocoris Kirshbaum is a large, cosmopolitan genus containing around 200

species world-wide (Carvahlo 1957, Razafimahatratra 1980, Razafimahatratra &

Lattin 1982). Most known species of Deraeocoris are predaceous. Deraeocoris

brevis (Uhler) is a predator of the pear psylla (Westigard 1973) but Fichter (1984)

found it to feed on the douglas-fir tussock moth. Parasitized nymphs of Dereacoris

diveni Knight were taken in Oregon: site 8 ex Picea engelmanni, 23 Jul 1986

from IV and V instar nymphs. Deraeocoris kennicotti Knight yielded parasitized

1999 LATTIN & STANTON: MIRIDS PARASITIZED BY BRACONIDS 27

V instar nymphs in Wyoming: site 2.2 ex P. ponderosa and P. flexilus on 25 Jul

1986. A Deraeocoris rubroclarus Knight V instar nymph was collected in

Oregon, Benton County, Lobster Valley ex Pseudotsuga menzessii on 3 Aug 1986.

Mirinae: Mirini

Dichrooscytus Fieber is Holarctic with 48 described species in America north

of Mexico (Henry & Wheeler 1988) where they occur on conifers and usually

are regarded as phytophagous (but see Fichter 1984). Loan (1974 b) described

the Braconidae, Peristenus juniperinus, from Canada, based on species from Di-

chrooscytus tinctipennis Knight. Fifth instar nymphs of Dichrooscytus sp. A from

Oregon: site 7, ex Pinus contorta murrayana were collected 25 Jun—9 Jul 1986

while V instar nymphs of Dichrooscytus sp. C were collected from Oregon: site

8 ex Picea engelmanni on 6 Aug 1986. Species of Dichrooscytus were found at

five of the six sites sampled but parasitized nymphs were found only at the sites

listed above.

Phytocoris Fallén has world-wide distribution and may be the largest genus in

the family. Stonedahl (1988) stated that over 200 species occur in North America

alone. Most species are thought to be predaceous (Fichter 1984) but many show

remarkable fidelity to “host trees’’. Leston (1961) reported 2 species that were

parasitized. Marsh (1979) reported Peristenus dumestris Loan as a parasitoid of

Phytocoris sp. and Loan (1980) reported the same braconid as a parasitoid of

Phytocoris lasiomerus Reuter and P. pallidocornis Reuter. He reported euphorine

larvae from Phytocoris tibialis Reuter and Phytocoris sp. Four additional species

of Phytocoris were found parasitized in this present study.

Phytocoris comulus Knight was collected at Wyoming: site 2 ex Pinus contorta

latifolia and IV instar nymphs were parasitized as was a parasitized V instar

nymph ex P. ponderosa on 9 Jul 1986. Phytocoris fraterculus Van Duzee was

collected in Oregon, Deschutes County, 4 km S. of Sisters ex Pinus ponderosa

on 29 Aug 1989 with V instar nymphs parasitized. Phytocoris heidemanni Reuter

was collected at Wyoming: site 2 from Pinus ponderosa on 9 Jul 1986 where

seven IV instar nymphs contained parasite larvae. Phytocoris stellatus Van Duzee

was recovered from Wyoming: site 2.2 ex Pinus contorta latifolia on 9 Jul 1986

where one IV instar nymph was parasitized while a IV instar nymph from site 3

ex Pinus contorta latifolia was collected 11 Jul 1986 and a V instar parasitized

nymph from the same host tree was collected from site 4 on 17 Jul 1986. This

was a common species at Oregon: sites 5 and 8 but no parasites were recovered.

Platylygus Van Duzee is a North American genus containing 31 species (Kelton

& Knight (1970). Although four species have been reported on various subspecies

of Pinus contorta (Kelton and Knight 1970), only two of these were found par-

asitized. Species occur early in the season and appear to feed on both male and

female cones. Rauf et al. (1984a) and Rauf et al. (1984b) reported that Platylygus

luridus (Reuter) caused conelet abortion on Pinus banksiana resulting in 74%

damage. Platylygus luridus (Reuter) was taken at Wyoming: site 2.2 ex Pinus

ponderosa on 9 Jul 1986 where four of the seven V instar nymphs contained

larvae. Platylygus rubripes Knight was recovered at Oregon: site 7 ex Pinus

contorta latifolia, 10 Jun 1986 where a IV instar nymph contained a parasitic

larva.

28 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Phylinae: Phyline

Knightomiroides Stonedahl and Schwartz contains a small group of species

found on conifers, largely Pinus (Stonedahl & Schwartz 1996). Knightomiroides

contortae Stonedahl & Schwartz was collected at Wyoming: site 2 ex Pinus con-

torta latifolia where a V instar nymph, collected on 25 Jul 1986, and a V instar

nymph from Wyoming: site 4 from the same host tree contained parasitoid larvae.

Microphylellus Reuter, a North American genus, contains 21 species including

one undescribed species found during this study. Knight (1923) reported M. mo-

destus Reuter as a predator. Species occur on both broad-leaf and coniferous trees.

Condit & Cate (1982) stated that Peristenus stygicus Loan, a Braconidae, attacked

Microphylellus maculipennis (Knight) in the laboratory. Microphylellus alpinus

Van Duzee was collected at Oregon: site 8 ex Pinus contorta murrayana on 25

Jun 1986 and parasite larvae were recovered from IV and V instar nymphs. M7-

crophylellus sp. A. occurred at Oregon: site 5 ex Pinus contorta contorta where

parasitized V instar nymphs were collected on 26 May and 2 Jun 1986.

Phoenicocoris Reuter is a Holarctic genus with nine species found in North

America (Schuh 1995). Stonedahl (1990) moved a number of species into this

genus, chiefly from Lepidopsallus Knight. Phoenicocoris hes perus (Knight) was

taken at Oregon: site 5 ex Pinus contorta contorta where a parasitized V instar

nymph was collected on 18 Jun 1986. Phoenicocoris longirostris (Knight) oc-

curred at Oregon: site 8 ex Pinus contorta murrayana where V instar nymphs

collected on 25 Jun and 9 Jul 1986 were parasitized. This bug was also found at

Wyoming: site 2 ex Pinus contorta latifolia where a V instar nymph collected on

9 Jul 1986 was parasitized.

Sthenarus Fieber is another Holarctic genus with four species described from

North America. There is confusion over the exact placement of the species re-

ported here that will be clarified when a new generic placement is established.

Stenaris sp. A. was collected at Wyoming: site 2 ex Pinus contorta latifolia from

V instar nymphs collected on 9 Jul 1986 and a V instar nymph from Pinus

ponderosa collected 26 Jul 1986, which contained parasitoids.

Phylinae: Pilophorini

Pilophorus is a Holarctic genus with 44 species in the New World (Schuh &

Schwartz 1988). These anti-mimics are found chiefly on trees and shrubs with

many on conifers where some species feed on aphids. Pilophorus americanus

Poppius was the only species found parasitized in this study and parasitoid. V

instar nymphs were taken at Wyoming: site 2.2 on Pinus flexilus on 25 Jul 1986

and at site 3 on Pinus contorta latifolia on 11 Jul 1986. Figure 2 shows a fifth

instar nymph of Pilophorus americanus with a mature braconid larva inside.

DISCUSSION

The occurrence of braconid larvae in the nymphs of Miridae is not surprising

(Leston 1959, 1961; Loan 1980). The absence of such parasitoids in some species

of Miridae we collected should not be considered non-occurrence because some

mirid species were collected in small numbers. As we learn more about the spe-

cific habits of both parasitoids and bugs, some of these points will be clarified.

It does appear that parasitism is chiefly an early season phenomenon, at least of

tree-inhabiting Miridae, as Leston (1959, 1961) suggested. We collected no par-

1999 LATTIN & STANTON: MIRIDS PARASITIZED BY BRACONIDS 29

Figure 2. Fifth instar nymph of Pilophorus americanus from below, showing parasitoid larva

occupying most of the body cavity.

asitized individuals after July so it is possible that the species of bugs occurring

later may escape the parasitoids. For example, Ceratocapsus apicatus Van Duzee

was one of the last species to appear in the season from early July to late Sep-

tember, was quite abundant at several sites and yet no parasitoids were recovered

from it.

Just why parasitism is more common in the earlier part of the season is not

clear. Perhaps only a few polyphagous parasitoid species might be involved at

any given site. For example, Peristenus pallipés (Curtis) uses at least 9 species

of mirid hosts, while P. juniperinus appears to attack only a single bug species.

Species of braconids attacking certain grass-feeding mirids (i.e., Acetropis Fieber,

Irbisia Reuter, Leptopterna Fieber, and Stenotus Jakovlev) also must mature early

in the season. The tree- and grass-inhabiting mirid species are almost all univol-

tine. The explanation may be quite simple—if the bug occurs early and is uni-

voltine, the parasitoid must occur at the same time. An examination of the par-

asitoid system involving the bivoltine, grass-feeding mirid genus Notostira Fieber

in Europe would be of considerable interest. Are both generations attacked? The

same would be true of Phytocoris stellatus, normally univoltine in Oregon, Wash-

30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

ington, and on north and east, but multivoltine in southern, coastal California

(Stonedahl 1988). Alternatively, parasitoids attacking Lygus may occur throughout

the season because many species of Lygus are multivoltine (Schwartz & Foottit

1998).

The emphasis in field work must shift to the parasitoids themselves to solve

these problems. It was our purpose to expand the records of the occurrence of

parasitoids in conifer-inhabiting mirid communities and call attention to their ex-

istence in many species of our Miridae.

ACKNOWLEDGMENT

We thank L. M. Shults (University of Wyoming), A. R. Moldenke, J. A.

DiGiulio, P. E. Hanson (Oregon State University) and G. Cassis (Australian Mu-

seum, Sidney, Australia) for assistance in collecting specimens; B. B. Hall

(Oregon State University) for the figures; A. Asquith (U.S. Department of Inte-

rior), R. T. Schuh (American Museum of Natural History, New York), M. D.

Schwartz and C. Loan (Canadian National Collection, Ottawa), and G. M. Stone-

dahl (Commonwealth Institute of Entomology, London) for discussion and tech-

nical assistance; P. Amaud (California Academy of Sciences, San Francisco) for

the loan of type materials; and L. Parks and D. Watkins for manuscript prepara-

tion.

LITERATURE CITED

Brindley, M. D. 1939. Observations on the life-history of Euphorus pallipes (Curtis) (Hym.: Bracon-

idae, a parasite of Hemiptera-Heteroptera. Proc. R. Entomol. Soc. London (A), 14: 51-56.

Carvalho, J. C. M. 1957. Catalogue of the Miridae of the world. Arq. Mus. Nac., Rio de J. Part I.

Cylapinae, Deraeocorinae, Bryocorinae, 44: 1-158.

Condit, B. P & J. R. Cate. 1982. Determination of host range in relation to systematics for Peristenus

stygicus (Hym.: Braconidae), a parasitoid of Miridae. Entomophaga, 27: 203-210.

Critchfield, W. B. 1957. Geographic variation in Pinus contorta. Maria Moors Cabot Foundation

Publication No. 3. Harvard University, Cambridge, Massachusetts.

Critchfield, W. B. 1980. The genetics of lodgepole pine. U.S. Dept. Agric., For. Serv., Res. Pap.,

WO-37.

Critchfield, W. B. 1984. Impact of the Pleistocene on the genetic structure of North America conifers.

pp. 70-118. In R. M. Lanner (ed.). Proc. Eighth N. Amer. For. Biol. Workshop, Logan, Utah.

Critchfield, W. B. 1985. The late quaternary history of lodgepole and jack pines. Can. J. For. Res.,

15: 749-772.

Fichter, B. L. 1984. Arboreal arthropod predation on early instar Douglas-fir tussock Moth. Ph.D.

Thesis, Oregon State University, Corvallis.

Forrest, G. I. 1980a. Variation in monoterpene composition of the shoot corical oleoresin within and

between trees of Pinus contorta. Biochem. Syst. Ecol., 8: 337-341.

Forrest, G.I. 1980b. Geographical variation in the monoterpenes of Pinus contorta oleoresin. Biochem.

Syst. Ecol., 8: 343-359.

Glen, D. M. 1977. Ecology of the parasites of predatory bug, Blepharidopterus angulatus (Fall.). Ecol.

Entomol., 2: 47-55.

Henry, T. J. & A. G. Wheeler, Jr. 1988. Family Miridae. pp. 251-507, In T. J. Henry and R. C.

Froeschner (eds.). Catalog of the Heteroptera, or true bugs, of Canada and the continental United

States. E. J. Brill, Leiden.

Kelton, L. A. & H. H. Knight. 1970. Revision of the genus Platylygus, with descriptions of 26 new

species (Hemiptera: Miridae). Can. Entomol., 102: 1429-1460.

Knight, H. H. 1923. Family Miridae (Capsidae). pp. 422-658. Jn Britton, W. E. (ed.). The Hemiptera

or sucking insects of Connecticut. Conn. Geol. Nat. Hist. Surv. Bull., 34: 1-807.

Lattin, J. D. & M. L. Stanton. 1992. A review of the species of Anthocoridae (Hemiptera: Heteroptera)

found on Pinus contorta J. N.Y. Entomol. Soc., 100: 424—479.

1999 LATTIN & STANTON: MIRIDS PARASITIZED BY BRACONIDS 31

Lattin, J.D. & C. M. Ozanne. 1993. New Miridae hosts of Braconidae (Hymenoptera). Entomol. Mon.

Mag., 129: 254.

Leston, D. 1959. The mirid (Hem.) hosts of Braconidae (Hym.) in Britain. Entomol. Mon. Mag., 95:

97-100.

Leston, D. 1961. Observations on the mirid (Hem.) hosts of Braconidae (Hym.) in Britain. Entomol.

Mon. Mag., 97: 65-71.

Loan, C. C. 1974a. The European species of Leiophron Nees and Peristenus Foerster (Hymenoptera:

Braconidae, Euphorinae). Trans. R. Entomol. Soc. London, 126: 207-238.

Loan, C. C. 1974b. The North American species of Leiophron Nees, 1818 and Peristenus Foerster,

1862 (Hymenoptera: Braconidae, Euphorinae) including the description of 31 new species. Nat.

Can., 101: 821-860.

Loan, C. C. 1980. Plant bug hosts (Heteroptera: Miridae) of some euphorine parasites (Hymenoptera:

Braconidae) near Belleville, Ontario, Canada. Nat. Can., 107: 87-93.

Loan, C. C. 1983. Host and generic relations of the Euphorini (Hymenoptera: Braconidae). Contrib.

Amer. Entomol. Inst., 20: 388-397.

MacDonald, G. M. & L. C. Cwynar. 1985. A fossil pollen based reconstruction of the late Quaternary

history of lodgepole pine (Pinus contorta ssp. latifolia) in the western interior of Canada. Can.

J. For. Res., 15: 1039-1044.

Marsh, P. M. 1979. Braconidae. pp. 144—295. Jn Krombeim, K. V., P. D. Hurd, Jr., D. R. Smith & B.

D. Burks (eds.). Catalog of Hymenoptera in America North of Mexico. Volume 1. Smithsonian

Institution Press, Washington, D. C.

Rauf, A., D. M. Benjamin & R. A. Cecich. 1984. Bionomics of Platylygus luridus (Hemiptera: Mir-

idae) in Wisconsin jack pine seed orchards. Can. Entomol., 116: 1219-1225.

Rauf, A., R. A. Cecich & D. M. Benjamin. 1984. Conelet abortion in jack pine caused by Platylygus

luridus (Hemiptera: Miridae). Can. Entomol., 116: 1213-1218.

Razafimahatratra, V. D. 1980. A revision of the genus Deraeocoris Kirschbaum (Heteroptera: Miridae)

from western America north of Mexico. Ph.D. Thesis, Oregon State University, Corvallis.

Razafimahatratra, V. D. And J. D. Lattin. 1982. Five new species and new synonomies for the genus

Deraeocoris (Heteroptera: Miridae) from western North America. Pan-Pac. Entomol., 58: 352—

364.

Schuh, R. T. 1995. Plant bugs of the world (Insecta: Heteroptera: Miridae): Systematic catalog, dis-

tributions, host list, and bibliography. New York Entomological Society, New York.

Schuh, R. T. & M. D. Schwartz. 1988. Revision of the New World Pilophorini (Heteroptera: Miridae:

Phylinae). Bull. Amer. Mus. Nat. Hist., 187: 101-120.

Schwartz, M. D. & R. G. Foottit. 1998. Revision of the Nearctic species of the genus Lygus Hahn,

with a review of the Palaearctic species (Heteroptera: Miridae). Associated Publishers, Gaines-

ville, Florida.

Stonedahl, G. M. 1988. Revision of the mirine genus Phytocoris Fallén (Heteroptera: Miridae) for

western North America. Bull. Amer. Mus. Nat. Hist., 188: 1-257.

Stonedahl, G. M. 1990. Revision and cladistic analysis of the Holarctic genus Atractotomus Fieber

(Heteroptera: Miridae: Phylinae). Bull. Amer. Mus. Nat. Hist., No. 198.

Stonedahl, G. M. & Schwartz, M. D. 1996. Two new genera for pine-inhabiting species of Phylini in

North America (Heteroptera: Miridae: Phylinae). Amer. Mus., Novitates, No. 3166.

Westigard, P. H. 1973. The biology of and effect of pesticides on Deraeocoris brevis piceatus (Het-

eroptera: Miridae). Can. Entomol., 105: 1105-1111.

Wheeler, A. G., Jr, & C.C. Loan. 1984. Peristenus henryi (Hymenoptera: Braconidae, Euphorinae),

a new species parasitic on the honeylocust plant bug, Diaphnocoris chlorionis (Hemiptera:

Miridae). Proc. Entomol. Soc. Wash., 86: 669-672.

Received 13 Jul 1998; Accepted 28 Dec 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 32-34, (1999)

A SYNONYMY FOR PSEUDOMETHOCA DONAEANAE

(COCKERELL & FOX) (HYMENOPTERA: MUTILLIDAE)!

DONALD G. MANLEY

Department of Entomology, Clemson University,

Pee Dee Research and Education Center,

2200 Pocket Road, Florence, South Carolina 29506-9706

Abstract.—Pseudomethoca donaeanae (Cockerell & Fox) was described in 1897 based on fe-

males only. Its known range is recorded as Arizona, California, New Mexico and Texas. Pseu-

domethoca russeola Mickel was described in 1924 based on a single male collected in Texas.

Additional specimens examined here expand that range to include Arizona, Kansas and New

Mexico. Numerous females of P. donaeanae were collected in Dofia Ana County, New Mexico

in 1992, as well as two males of P. russeola that were attracted to a caged female. This, and

the similar geographic range, have led to the conclusion that these two are the same species.

The name P. donaeanae has precedence over P. russeola. A complete synonymy is included.

Key Words.—Insecta, Hymenoptera, Mutillidae, Pseudomethoca donaeanae, Pseudomethoca

russeola, taxonomy, synonymy.

Pseudomethoca donaeanae was first described by Cockerell & Fox (1897) as

Sphaerophthalma donae-anae. The description was apparently made on the basis

of four females collected in 1896 in the Mesilla Valley in Donia Ana County,

New Mexico. The holotype is in the collection of the American Entomological

Society in Philadelphia, and has been examined by this author. Mickel examined

two additional female specimens from Arizona and California (1924), and seven

female specimens from Arizona, New Mexico and Texas (1935). Until now, this

species has been known only from these few female specimens.

Pseudomethoca russeola was described by Mickel (1924) on the basis of a

single male collected on 4 May 1901 in San Diego, Texas. This holotype is in

the collection of the United States National Museum in Washington, D.C., and

has been examined by this author. No other specimen of this species has been

recorded in the literature.

PSEUDOMETHOCA DONAEANAE (COCKERELL AND FOx)

Sphaerophthalma donae-anae Cockerell and Fox, 1897: 136. 2

Mutilla donae-anae Fox, 1899: 224. 2

Pseudomethoca Donae-Anae André, 1903: 28 &

Pseudomethoca russeola Mickel, 1924: 44. NEW SYNONYMY. 6

Pseudomethoca donaeanae Krombein, 1979: 1302. 2

Material Examined.—In addition to the type specimens mentioned above, the following material

has been examined (all are females of P. donaeanae or males of P. russeola): U.S.A. ARIZONA.

COCHISE Co.: Portal, 2 Sep 1959, H. E. Evans, 1 male. KANSAS. DOUGLAS Co.: 8 km E of

Lawrence, 15 Aug—15 Sep 1988, W. T. Wcislo and R. L. Minckley, 1 male. NEW MEXICO. DONA

ANA Co.: 3.2 km E of Radium Springs, 1 Sep 1992, D. G. Manley, 12 females; 3.2 km E of Radium

Springs, 2 Sep 1992, D. G. Manley, 16 females and 2 males; 1.6 km E of Vado, 2 Sep 1992, D. G.

Manley, 18 females; 3.2 km E of Radium Springs, 3 Sep 1992, D. G. Manley, 12 females. HIDALGO

' Technical contribution no. 4377 of the South Carolina Agricultural Experiment Station, Clemson

University.

1999 MANLEY: PSEUDOMETHOCA DONAEANAE SYNONYM 33

Co.: Rodeo, 18 Aug 1959, H. E. Evans, 1 female; Rodeo, 28 Aug 1959, H. E. Evans, 1 female and

1 male. TEXAS. BEXAR Co.: U-Bar Ranch, 9.6 km E of Castroville, 29 May 1992, D. G. Manley, 1

Male. DIMMIT Co.: Chaparral W. M. A., 20-30 May 1991, A. W. Hook, 1 male. FRIJO Co.: 9.6 km

SE of Pearsall, 7 Jul 1972, E. E. Grissell and J. Smith, 1 male. HIDALGO Co.: Bentsen Rio Grande

Valley State Park, 13 Jun 1978, C. C. Porter, 1 male; Bentsen Rio Grande Valley State Park, 30 Nov—

2 Dec 1978, E. E. Grissell and A. S. Menke, 1 male. KLEBERG Co.: Site 55, 10 Oct 1978, G. E.

Gillaspy, 1 male.

DISCUSSION

Both the females of P. donaeanae and the males of P. russeola have been

known from only a few specimens since their original descriptions. The known

geographic ranges of southeastern Arizona, New Mexico and Texas are similar

for the two. I have not examined the specimen from Calexico, California that was

noted by Mickel (1924). Even if correctly identified, I find the collection data on

that specimen to be suspect. I have collected extensively throughout Arizona and

southern California, including the Calexico area, and have not found (or seen)

specimens of either of these species from those areas. The female specimens of

P. donaeanae and the male specimens of P. russeola collected by Evans in the

Portal, Arizona/Rodeo, New Mexico area (only about 16 km apart) on or about

the same collection dates add further evidence for the synonymy of these two

species.

A female of P. donaeanae was collected on 2 Sep 1992 about 1.6 km E of

Radium Springs, New Mexico. It was placed in a small, plastic cage from which

the ends had been cut out and replaced with wire screen. Two males were attracted

to the caged female and attempted to mate with her through the screen. They

were subsequently captured a short time later and determined to be P. russeola.

No other males were attracted to the caged female, although there were numerous

males and females of other mutillid species in the area at the time. The use of

caged females has been used previously to attract males, both in cases where the

male was already known and where the male was unknown. Males are not always

attracted to the caged females. However, in all cases in which males have been

attracted to the caged females, they have been subsequently shown to be of a

single species. And in cases where the males were previously known, they have

always been of the same species as the caged female. Thus, the use of caged

females is a reliable method for obtaining sex correlations among mutillid wasps.

CONCLUSIONS

The known geographic ranges for P. donaeanae and P. russeola, the present

collection data, and the use of caged females for the attraction of conspecific

males, all lead to the conclusion that these two are a single species. Because P.

donaeanae has precedence over P. russeola, that name shall stand.

ACKNOWLEDGMENT

I thank Donald Azuma, Philadelphia Academy of Sciences, for allowing me to

examine the holotype of Pseudomethoca donaeanae, and Arnold Menke, United

States National Museum, for allowing me to examine the holotype of Pseudome-

thoca russeola. Fieldwork for collection and study of the specimens in 1992 was

supported, in part, by a grant from the National Science Foundation (BSR-

9106369).

34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

LITERATURE CITED

André, E. 1903. Mutillidae. Gen. Ins. Volume 1, fasc. 11, 1-77.

Cockerell, T. D. A. & W. J. Fox. 1897. Descriptions of new Hymenoptera from New Mexico. Acad.

Nat. Sci. Phila., Proc., 49: 135-141.

Fox, W. J. 1899. The North American Mutillidae. Amer. Entomol. Soc., Trans., 25: 219-292.

Krombein, K. V., P. D. Hurd, Jr, D. R. Smith & B. D. Burks. 1979. Family Mutillidae. Catalog of

Hymenoptera in America North of Mexico. Volume 2: 1276-1314.

Mickel, C. E. 1924. A revision of the mutillid wasps of the genera Myrmilloides and Pseudomethoca

occurring in America North of Mexico. U.S. Nat. Mus., Proc., 64: 1-52.

Mickel, C. E. 1935. Descriptions and records of nearctic mutillid wasps of the genera Myrmilloides

and Pseudomethoca (Hymenoptera: Mutillidae). Amer. Entomol. Soc., Trans., 61: 383-98.

Received 13 Feb 1998; Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 35-47, (1999)

THE LEIODIDAE (COLEOPTERA) OF THE CARNEGIE

MUSEUM OF NATURAL HISTORY. NEW DATA AND

DESCRIPTION OF TWO NEW SPECIES

J. M. SALGADO

Department of Animal Biology, University of Leén,

24071 Leén, Spain

Abstract—The collection of the family Leiodidae at the Carnegie Museum of Natural History

(CMNH) was studied. Two new species, Catops davidsoni, NEW SPECIES (USA) and Adelo psis

chapadaensis, NEW SPECIES (Brazil) are described. Also, new biogeographic data on several

species from America and Europe are given.

Key Words.—Insecta, Coleoptera, Leiodidae, systematic, Catops davidsoni n.sp., Adelopsis cha-

padaensis n.sp., new data.

Resumen.—Se ha examinado la coleccién de la familia Leiodidae del Carnegie Museum of

Natural History (CMNH). A partir de tan interesante material entomoldégico se han descrito dos

especies nuevas, Catops davidsoni n. sp. y Adelopsis chapadaensis n. sp., y ademas se propor-

cionan nuevos datos biogeograficos de varias especies de la familia Leiodidae de América y de

Europa.

This paper widens our knowledge of the beetle family Leiodidae by studying

745 specimens belonging to the Carnegie Museum of Natural History (CMNH)

collection, Pittsburg, Pennsylvania, U.S.A.

The material examined has enabled me to differentiate 45 American species

and 15 European ones. The American species include two new ones, Catops

davidsoni, NEW SPECIES and Adelopsis chapadaensis, NEW SPECIES; of the

remaining 43 species, the number of specimens studied and their distribution is

reported and attention is brought to those species which are new state records in

the U.S.A. An appendix of the 15 European species is included at the end.

MATERIAL STUDIED

I follow the suprageneric classification proposed by Newton & Thayer (1992).

Family Leiodidae Fleming, 1821

Subfamily Coloninae Horn, 1880

Colon (Colon) horni Szymczakowski, 1981

Widespread species, known from Nova Scotia and Massachusetts, west to On-

tario and Minnesota (Peck & Stephan 1996).

Records: Pittsburgh (PA), 1m; 1f. This record is new for Pennsylvania.

Colon (Colon) bidentatum Sahlberg, 1834

This species is known from central and northern Europe (Szymczakowski

1969a, von Peez 1971) and from North America, northern transcontinental (Peck

& Stephan 1996).

Records: DC, Im, NY, 1f. PA, 2m; 6f. VA, 1m; 1f. New record for the District

of Columbia.

36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Colon (Colon) tibiale Hatch, 1957

Known from North America, northern transcontinental distribution.

Records: CA, lf.

Colon (Colon) asperatum Horn, 1880

Very similar distribution to the previous species, northern transcontinental.

Records: DC, Im. New record for the District of Columbia.

Colon (Euricolon) magnicolle Mannerheim, 1853

Widespread species, northern transcontinental.

Records: LA, 1f. PA, 1m. Both are new records for the states of Louisiana and

Pennsylvania.

Colon (Striatocolon) thoracicum Horn, 1880

Widespread in eastern North America (Peck & Stephan 1996).

Records: DC, 1m; 1f. PA, 1m.

Colon (Myloechus) hubbardi Horn, 1880

Across all of North America. Fairly commom.

Records: DC, Im. IA, 1f. MI, 1m; 1f. PA, Im. TN, lm. The records from the

states of Pennsylvania and Tennessee are new.

Colon (Myloechus) megasetosum Peck & Stephan, 1996

Widespread in eastern North America, Ontario and Quebec to Alabama and

Georgia (Peck & Stephan 1996).

Records: DC 1f. PA, 4m; 6f.

Colon (Myloechus) celatum Horn, 1880

This species is known from southern British Columbia and Alberta to Cali-

fornia.

Records: NV, If.

Colon (Myloechus) longitorsum Peck & Stephan, 1996

Distribution from North Carolina to Oregon (Peck & Stephan 1996)

Records: CA, If.

Colon (Myloechus) serratum Hatch, 1957

Western North America from southern Alaska to northern California (Peck &

Stephan 1996)

Records: CA, 1m.

Colon (Myloechus) dentatum LeConte, 1853

This species is known in most of eastern North America.

Records: DC, 8m; 13f. IL, Im; 3f. MD, Im; I1f. MO., Im; 2f. PA, 7m; 10f.

TN, Im. VA, 1m. The records for Montana and Tennessee are new.

1999 CARNEGIE MUSEUM LEIODIDAE 37

Subfamily Cholevinae Kirkby, 1837

Tribe Anemadini Hatch, 1928

Nemadus (Laferius) brachycerus (LeConte, 1863)

Eastern half of North America and more abundant in the south.

Records: SD, 6m; 10f.

Nemadus (Nemadus) hornii Hatch, 1933

Similar distribution to that of previous species (Hatch 1933, 1957).

Records: IL, 1m; 2f. MI, Im; 1f. PA, 7m; 9f. OH, 2m; 1f. SD, 5m; 7f. The

records for Illinois and South Dakota are new.

Nemadus (Nemadus) tenuitarsis Jeannel, 1936

Known only from Ohio and Pennsylvania (Jeannel 1936).

Records: PA, If.

Nemadus (Nemadus) parasitus (LeConte, 1853)

Widespread in northern, central and western states of North America (Hatch

1933).

Records: DC, 10m; 13f. IL, Im. NJ, lm. NY, 1m; 1f. PA, 12m; 17f. VA, 1f.

New locations for the states of Illinois and Virginia.

Nemadus (Nemadus) pusio (LeConte, 1853)

Known from eastern North America from British Columbia to California.

Records: CA, 1m.

Dissochaetus hetschkoi Reitter, 1884

This species is known from Brazil (Jeannel 1936, Szymczakowski 1963, Gnas-

pini 1991, Salgado 1991), Venezuela (Szymczakowski 1969b) and Mexico (Peck

1977a).

Records: This species was located for the first time in Rio de Janeiro, 1f; San-

tarem, 1f; Minas Gerais, Im, and Chapada dos Guimaraes (Mato Grosso), 1m; 2f.

Dissochaetus murrayi murrayi Reitter, 1884

Widespread from Argentina—state of Corrientes—(Salgado 1991) to the entire

central and eastern area of Brazil (Jeannel 1936, Gnaspini 1991).

Records: Aguas Vermelhas (Minas Gerais), 1m; 1f. and Alagoas (Serra Branca),

1f. The Alagoas location is new for the state of Serra Branca.

Dissochaetus oblitus (LeConte, 1853)

Present in all central and eastern states in the U.S.A.

Records: DC; 3m; 3f. PA, 3m; 3f.

Dissochaetus mexicanus Jeannel, 1936

Known from the state of Mexico (Jeannel 1936, Szymczakowski 1968, Peck

1977a).

Records: Guadalajara (Mexico), If.

Prionochaeta opaca (Say, 1825)

38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Widespread in the centre and east of North America (Peck 1977b).

Records: DC, 2m; 1f. MD, 1m; 1f. PA, 26m; 25f. WV, 2m; 3f.

Tribe Cholevini Kirby, 1837

Catops davidsoni Salgado, NEW SPECIES

Figs. 1-6

Type series PENNSYLVANIA. Allegheny: Holotype male deposited in

CMNH. Paratypes: PENNSYLVANIA. Allegheny. 1m, Pittsburg. 1m, St. Vicent.

lf. District of Columbia, 3m, 1f. Deposited: CMNH (6) and Col. Salgado (Uni-

versity of Leén, Spain) (1).

Description—Large size; length 3.10 to 3.70 mm; width 1.50 to 1.80 mm. (Fig. 1).

Elongate elyptical body. Head and protonum dark brown, elytra more light brown. Head coarsely

punctured. Antenna robust, 1.6 times as long as protonum, passing base of protonum when laid back;

proportions of length of each segment and that of the 9th from Ist to 11th (Fig. 2); 1.42, 1.03, 1.00,

0.50, 0.50, 0.45, 1.00, 0.20, 1.00, 0.96, 1.45, the equal length of 3rd, 7th and 9th segments stands

out; proportions of length and width of each segment from Ist to 11th, 2.43, 1.77, 1.50, 0.75, 0.63,

0.53, 0.75, 0.19, 0.75, 0.72, 1.23, the strongly transverse sixth antennal segment stands out, almost

twice as wide as long in frontal view. Pronotum densely haired and finely granulate with setal bases;

base narrower than the elytra; sides very curved in both sexes; widest at middle; 1.7 times as wide

as long; hind angles almost rounded. Elytra together 1.2 times as long as wide; widest at basal third;

densely granulate from setal bases. Sutural striae well marked and sutural angle feebly pointed. Flight

wings fully formed. First segment of male protarsus 0.9 times as wide as the maximum width of tibia.

Male profemur with broad raised area on inner margin. Male metatrochanter (Fig. 3) with very pointed

curved tip, a form unique in American species of the genus Catops. Female metatrochanter pointed,

not curved. Aedeagus in lateral view (Fig. 4) in regular curve with widened apex area; in ventral view

(Fig. 5) sides parallel in centre of median lobe, becoming wider and ending in a rounded tip. Internal

sac armed with very characteristic structures, such as a strongly scleroticized mid basal plaque, formed

by 6 symetrically placed robust teeth and two symetrical rows of thick bristles along the middle.

Genital segment complete, longer than broad, wide and indented lobes with some pointed setae,

numerous setae on sternal face (Fig. 6).

Diagnosis.—The species is readily distinguished by the shape of the aedeagus

and the curved pointed tip of the male metatrochanter.

Discussion.—lt is possible that some specimens of Catops davidsoni have been

mistaken for C. americanus, because the latter shows a wide polymorphism

(Hatch 1933), and is also widespread, including the area of the new species with

which it lives.

The differences between C. davidsoni and C. americanus are evident and are

observed in the general form of the aedeagus, in the structures of the internal sac,

the genital segment and the metatrochanter.

KEY

In order to assess the taxonomic status of C. davidsoni, I have followed the

key proposed by Hatch (1933, 1957) for the genus Catops species.

1. Third antennal segment longer than second; eighth antennal segment more

than half as long as ninth and subequal to the sixth in length; male

prefemora and metatrochanters and female abdomen unmodified.

1999 CARNEGIE MUSEUM LEIODIDAE 39

Two species under this heading: Catops alsiosus and C. gratiosus.

1’. Third antennal segment equal to, or a little shorter than second; eighth

antennal segment less than half as long as ninth, strongly transverse .. 2

2. Antennae with sixth segment strongly transverse, nearly twice as wide as

long in frontal view; pronotum slightly arcuate at sides; female with

fifth abdominal sternite emarginate behind.

Three species under this heading: C. basilaris, C. egenus and C. mathersi.

2’. Antennae with sixth segment slightly transverse in frontal view; pronotum

more strongly arcuate at sides; female with abdominal sternite not emar-

SiNAteUDSMINGs le OP ye ees ee A Bn ae fe ee 3

3. Male with prefemora tuberculate below, the edentate metatrochanter, fe-

male with abdominal sternites third to sixth more or less impressed.

Two species under this heading: C. luridipennis and C. simplex.

3’. Male with prefemora not tuberculate below, flattened, the metatrochanter

dentate at about apical fourth; female with abdominal sternites unim-

pressed or only impressed sternites fifth and sixth .................. 4

4. Male metatrochanter with very pointed curved tip; apical zone of median

lobe of aedeagus strongly narrow and elongate, with rounded tip; base

of internal sac with six symmetric robust teeth ................

FO ese Beet a Bae OR carat ee ee RR oe ee Catops davidsoni n. sp.

4'. Male metatrochanter pointed, not curved; apical zone of median lobe of

aedeagus wider or in the shape of a spear head; base of internal sac

without robust teeth, only some bristle formations ... Catops americanus

Etymology.—tThe species is dedicated to Robert L. Davidson, Curator of the

Carnegie Museum, who has generously made it possible for me to study the many

Leiodids of the Museum collection.

Distribution.—The species is known from Georgetown (District of Columbia)

and from Allegheny, Pittsburgh and St. Vincent (Pennsylvania):

Catops alsiosus (Horn, 1885)

Distribution northern transcontinental.

Records: AK, 1m.

Catops mathersi Hatch, 1957

This species is known in the central-northern states of the USA and the central-

south of Canada.

Records: AB, 1m.

Catops gratiosus (Blanchard, 1915)

Widespread in northern and eastern North America (Hatch 1933, 1957).

Records: DC, Im. KY, 2m; 1f. PA, Im. VA, If. It is a new record for the

District of Columbia.

40

Figures 1-3.

view.

THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

1mm,

05mm

Catops davidsoni. i: habitus; 2: antenna, lateral view; 3: male metatrochanter; lateral

1999 CARNEGIE MUSEUM LEIODIDAE 41

0,20mm

0,20 mm

Figures 4-6. Catops davidsoni. 4: aedeagus, lateral view; 5: aedeagus, ventral view; 6: genital

segment.

42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Catops luridipennis luripennis Mannerheim, 1853

Known in Alaska, almost all states in Canada and in central western states of

the USA.

Records: AB, Im. AZ, 1m; If.

Catops simplex Say, 1825

Widespread _ in all North America.

Records: AL, Im; 1f. DC, 5m; 7f. IL, 1m; 1f. PA, 7m; 9f.

Catops egenus (Horn, 1880)

Very frequent in the states of western North America.

Records: AK, 5m; 6f. AR. lm. CA, 1m; 1f. OR, Im. The record is new for

the. state of Arkansas.

Catops basilaris (Say, 1823)

Distributed throughout most states of North America. Already indicated by

Hatch (1933, 1957) in 23 states in North America.

Records: AL, 2m; 1f. CA, Im; 1f. DC, 1f. IL, Im; 1f. LA, 1f. MD, 1f. NC,

13m; 15f. NH, 1f. PA, 1m; 2f. The records are new for the states of Illinois and

Louisiana.

Catops americanus (Hatch, 1928)

Found in most states of North America (Hatch 1933, 1957). This species is

markedly variable in the form of the middle lobe of the aedeagus, which may

cause a certain degree of confusion. Hence the diagrams of the two species, one

of NC (Figs. 7—9) and another of DC (Figs. 10-11).

Records: DC, 4m; 1f. NC, Im. PA, 1m.

Catoptrichus frankenhauseri Mannerheim, 1853

Known only from British Columbia, Alaska, Idaho, Oregon and Washington.

Records: I have seen 33 specimens from AK, ID, and OR.

Sciodre poides fumatus terminans (LeConte, 1850)

Very frequent in most states of North America.

Records: I have seen 52 specimens from DC, IA, IL, NB, NY, OH, and PA.

The record for the state of Illinois is new.

Sciodrepoides watsoni hernianus (Blanchard, 1915)

It lives with the above-mentioned species in several states and is likewise fairly

common.

Records:. I have seen 47 specimens from DC, KS, MD, PA and VT. The record

for Vermont is new.

Tribe Leptodirini Lacordaire, 1854

Platycholeus leptinoides (Crotch, 1874)

Species known from the states of California, Nevada and Oregon.

Records: CA, 1f. NV, Im, If.

43

CARNEGIE MUSEUM LEIODIDAE

1999

=

my

Soe

nyt

AW

, ae

v

RNS YG x

we ANA 3 N a

35 ae de

it Z >

ZA: % fe fi, ay

eC TO

RRR

SUG

CAN

ra

ww OZ’O

0,20 mm

WwW O20

Figures 7-11. Catops americanus. 7: aedeagus, lateral view; 8: aedeagus, ventral view (specimen

from North Carolina); 9: genital segment; 10: aedeagus, ventral view (specimen from D.C.); 11:

internal sac armature (specimen from D.C.).

Platycholeus opacellus Fall, 1909

This species, not so widespread as the previous species, only known from

California.

Records: CA, Im.

44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Tribe Eucatopini Jeannel, 1921

Eucatops glabricollis (Reitter, 1884)

This species is known from Brazil, Santa Catharina and Sao Paulo (Jeannel

1936) and from Blumenau (Szymczakowski 1963). Recently mentioned by Gnas-

pini (1994).

Records: Alagoas (Serra Branca), 1m and Para, 1f. Both records are new for

Serra Branca and Para.

Tribe Ptomaphagini Jeannel, 1911

Adelopsis chapadaensis Salgado, NEW SPECIES

Figs. 12—15

Type series BRAZIL. Mato Grosso. Chapada dos Guimaraes. Holotype male

and paratype female deposited in CMNH.

Diagnosis and description—Length: 1.85-1.95 mm; width: 0.90-0.97 mm (Fig. 12). Pubescence

golden, with many short recumbent setae, setal sockets forming strigae on the head, protonum and

elytra. Color light brown. All antenna color lightening, fairly robust with flattened club, 1.22 times as

long as protonum; proportions of length of each segment and that of the 9th from Ist to 11th: 2.00,

1.85, 0.95, 0.90, 0.80, 0.65, 1.10, 0.45, 1.00, 1.00, 1.75; proportions of length and width of each

segment of the club, from 7th to 11th: 0.9, 0.38, 0.79, 0.74, 1.12. Pronotum 1.54 times as wide as

long, as wide as the elytra, maximum width in hind angles. Hind angles weakly pointed but slightly

protruding backwards. Elongated elytra, not very convex, not pointed at back and weakly arched over

the sides; together 1.40 times as long as wide; with dense oblique strigae. Sutural striae entire and

deep. Mesosternal carina high and cutting, smooth rounded profile view. Summits of tibia the typical

of Ptomaphagini, armed with a comb of many short and equal spines; with the first segment of male

protarsus 0.75 times as broad as the maximum width of tibia. Aedeagus (Figs. 13-14) with apical

plaque transversally produced, bearing a medial cavity, margins slightly protruding and 7 small setae

inserted in groups of 3 and 4 setae. Genital segment (Fig. 15) slightly broader than long, lateral lobes

bearing several long and short setae. Spiculum gastrale short and straight, also somewhat wider at

base.

Etymology.—The name is derived from the area in Brazil where it was cap-

tured, ““Chapada’’ (type locality).

Discussion.—Using the characteristics of the aedeagus as a basis of differen-

tiation, A. chapadaensis is clearly unlike A. bellatrix Szymczakowski, 1968; A.

peruviensis Blas, 1980 and A. confluens Gnaspini & Peck, 1996 in that it does

not present a small right lobe (lateral view), usually with setae surrounding the

apical orifice of the aedeagus.

Moreover, A. chapadaensis could form a group of species with A. brunnea

Jeannel, 1936 and their different shapes (Szymczakowski 1975), A. coronaria

Gnaspini & Peck 1996 and A. galea Gnaspini & Peck 1996, having the same

model of aedeagus and gastral spiculum in the genital segment. However, differ-

ences amongst these three species can be observed in the size, form of antennae

and the protarsus of the male, but particularly in the shape of the aedeagus and

number of setae on the apical plaque, and also in the shape, insertion and number

of setae of the parameres. Finally, it should be pointed out that Gnaspini (1996)

modified the generic concepts in Ptomaphagini and consequently the subgenera

of Adelopsis have been dropped.

45

CARNEGIE MUSEUM LEIODIDAE

0,20 mm.

Figures 12-15. Adelopsis chapadaensis. 12: habitus; 13: aedeagus, lateral view; 14: aedeagus,

dorsal view; 15: genital segment.

Paulipalpina claudicans (Szymczakowski, 1980)

The genus Paulipalpina was established by Gnaspini & Peck (1996) using

Adelopsis claudicans described from Novia Teutonia, Santa Catharina (Brazil) by

Szymczakowski (1980) as type species.

Records: Rio de Janeiro (Brazil), 1f. It is new record from Rio de Janeiro.

46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Ptomaphagus (Adelops) ulkei Horn, 1885

According to Peck (1973), this species is known from central New York, west-

ward to central Indiana and southward to northwestern Georgia.

Records: DC, 1m. The record for the District of Columbia is new.

Ptomaphagus (Adelops) fisus Horn, 1885

Known in several states in western USA (AZ, CA, CO, ID, NV, TX) and one

record from north of Mexico.

Records: CA, 2m; 1f. NB, 1m. The Nebraska record is new.

Ptomaphagus (Adelops) nevadicus Horn, 1880

This species has the widest range of any known Adelops. It is distributed in

southern Canada, most of the western states of the USA and Durango (Mexico).

Records: CA, 6m; 8&f.

Ptomaphagus (Adelops) brevior Jeannel, 1949

Widespread in eastern North America.

Records: DC, 3m; 5f. IL, Im. NC, 1f. PA, 1m; 1f. TN, Im. The record for the

state of Tennessee is new. |

Ptomaphagus (Adelops) consobrinus (LeConte, 1853)

This species is distributed in the south from Texas to Florida, and towards the

north of Colorado to New Jersey (Peck 1973).

Records: PA, 4m; 2f. SC, Im. TX, 1m. The record for Pennsylvania is new.

Ptomaphagus (Adelops) hirtus (Tellkampf, 1844)

This species only is known from the Mammoth Cave area of Kentucky.

Records: I have seen 18 specimens, Mammoth Cave (KY).

European species checklist—Colon (Euricolon) latum Kraatz, 1850. 4 speci-

mens, Sweden. Bathysciola schiodtei schiodtei (Kiesenwetter, 1850). 5 specimens,

France. Nargus (Nargus) anisotomoides (Spence, 1815). 2 specimens, France.

Nargus (Demochrus) wilkini (Spence, 1815). 3 specimens, Austria. Choleva

(Choleva) oblonga oblonga Latreille, 1807. 3 specimens, Austria. Choleva (Chol-

eva) sturmi Brisout, 1863. 1 specimen, Austria. Choleva (Choleva) cisteloides

(Frélich, 1799). 2 specimens, Hungary. Catops morio (Fabricius, 1792). 2 spec-

imens, Austria; 5 specimens, Sweden. Catops borealis Krogerus, 1931. 11 spec-

imens, Sweden. Catops fuscus fuscus (Panzer, 1794). 12 specimens, Sweden. Ca-

tops tristis (Panzer, 1794). 6 specimens, Austria. Catops subfuscus subfuscus Kell-

ner, 1846. 3 specimens, Austria; 3 specimens, Switzerland. Apocatops nigrita

(Erichson, 1837). 2 specimens, Francia; 3 specimens, Sweden. Sciodrepoides wat-

soni watsoni (Spence, 1815). 5 specimens, Austria; 8 specimens, Sweden. Pto-

maphagus sericatus septentrionalis Jeannel, 1934. 9 specimens, Sweden.

ACKNOWLEDGMENT

I would like to express my thanks to Mr. Davidson, Curator of the Camegie

Museum, for allowing me to study the interesting collection at the Carnegie Mu-

1999 CARNEGIE MUSEUM LEIODIDAE 47

seum of Natural History and to Professor Peck, Carleton University, for revising

this paper and for his much appreciated and valuable comments.

LITERATURE CITED

Gnaspini, P. 1991. Brazilian Cholevidae (Coleoptera) with emphasis on cavernicolous species I. Genus

Dissochatus. G. it. Entom., 5: 325-340.

Gnaspini, P. 1994. The genus Eucatops (Coleoptera, Cholevidae, Eucatopinae), description of new

species and considerations on its systematic position. Iheringia (Zoologia), 76: 33-42.

Gnaspini, P. 1996. Phylogenetic analysis of the tribe Ptomaphagini, with description of new Neotrop-

ical genera and species. Papéis Avulsos Zool., S. Paulo, 39: 509-556.

Gnaspini, P. & S. B. Peck. 1996. The Adelopsis of Costa Rica and Panama (Coleoptera, Leiodidae,

Cholevinae, Ptomaphagini). Papéis Avulsos Zool., S. Paulo, 39: 405-441.

Hatch, M.H. 1933. Studies on the Leptodirinae (Catopidae) with description of new species. J. New

York Entomol. Soc., 41: 187-239.

Hatch, M. H. 1957. The beetles of the Pacific northwest, Part II. Staphyliniformia. Univ. Washington,

Publ. Biol., 16: 1-384.

Jeannel, R. 1936. Monographie des Catopidae. Mém. Mus. Natl. Hist. Nat., ns. 1: 1-438.

Newton, A. F, Jr. & M. K. Thayer. 1992. Current classification and family-grupo names in Staphylin-

iformia (Coleoptera). Fieldiana Zoology, n.s., 67: 1-92.

Peck, S. B. 1973. A systematic revision and the evolutionary biology of the Ptomaphagus (Adelops)

beetles of North America (Coleoptera; Leiodidae; Catopinae), with emphasis on cave-inhabiting

species. Bull. Mus. Comp. Zool. 145: 29-162.

Peck, S. B. 1977a. The subteranean and epigean Catopinae of Mexico (Coleoptera: Leiodidae). Assoc.

Mex. Cave Stud. Bull., 6: 185-213.

Peck, S. B. 1977b. A review of the distribution and biology of the small carrion beetle Prionochaeta

opaca of North America (Coleoptera; Leiodidae; Catopinae). Psyche, 83: 299-307.

Peck, S. B. & K. Stephan. 1996. A revision of the genus Colon Herbst (Coleoptera; Leiodiade;

Coloninae) of North America. Can. Entom., 128: 667-741.

Salgado, J. M. 1991. Nota sobre algunos Dissochaetus (Coleoptera, Catopidae) de Brasil y Argentina.

Bull. Annls. Soc. r. belge Ent., 127: 211-215.

Szymezakowski, W. 1963. Catopidae (Coleoptera) récoltés au Brésil par J. Mraz. Act. Ent. Mus. Nat,

Pragae, 35: 667-680.

Szymezakowski, W. 1968. Sur quelques Catopidae (Coleoptera) de la région néotropicale. Acta Zool.

Cracov., 13: 13-27.

Szymczakowski, W. 1969a. Die mitteleuropdischon Arten der Gattung Colon Herbst. Entom. Abhand.

(Dresden), 36: 303-339.

Szymczakowski, W. 1969b. Note sur quelques Catopidae (Coleoptera) du Venezuela. Bull. Acad.

Polon. Sci., Sér. Sci. Biol., 17: 407-412.

Szymczakowski, W. 1975. Formes cavernicoles d’Adelopsis brunneus Jeann. Du Venezuela et d’Ile

de Trinidad (Coleoptera: Catopidae). Bol. Soc. Venezolana Espel., 6: 13-24.

Szymczakowski, W. 1980. Deux espéces nouvelles de Ptomaphaginae (Coleoptera, Catopidae). Pol.

Pismo ent. 50: 515-521.

von Peez, A. 1971. Family Colonidae. pp. 237-243. In Freude, H., K. W. Harde & G. A. Lohse (eds.).

Die Kafer Mitteleuropas Band 3. Goecke and Evers, Krefel.

Received 28 Jan 1998; Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 48-51, (1999)

A SECOND SPECIES OF FOSSIL DASYMUTILLA

(HYMENOPTERA: MUTILLIDAE) FROM

DOMINICAN AMBER!

DONALD G. MANLEY? AND GEORGE O. POINAR, JR.?

*Department of Entomology, Clemson University,

Pee Dee Research and Education Center,

Florence, South Carolina 29506

3Department of Entomology, Oregon State University,

Corvallis, Oregon 97331

Abstract—Dasymutilla albifasciatus, a NEW SPECIES of mutillid wasp, is described from a

fossil specimen embedded in a piece of Dominican amber estimated to be anywhere from 15—

45 million years old. Very few fossil mutillids have been recorded, and this represents only the

second description of a fossil mutillid from the New World. This male is thought to belong to

the bioculata species group, and can be distinguished from closely related species by the band

of white integument extending from the top of the propodeum along the sides of the thorax to

the lower half of the pronotum.

Key Words.—Insecta, Dasymutilla albifasciatus, Mutillidae, new species, fossil, Dominican am-

ber.

Fossil velvet ants, or mutillids, are extremely rare and to date all except one

have been limited to Palaearctic forms. Menge (1856) found six specimens of this

family in Baltic amber and Brischke (1886) cites another three from the same

deposits. Larsson (1978) mentioned an additional three in the Copenhagen col-

lection of Baltic amber insects. To our knowledge, none of the above have been

described. Scudder (1891) referred to an undescribed Mutilla from the mid-Oli-

gocene beds (not amber) located in Brunstatt, Alsatia (now Alsace), in north-

eastern France.

Bischoff (1915) described seven species of fossil mutillids from Baltic amber,

placing them in a new genus, Protomutilla. Krombein (1979) suggested that at

least one of those species may not be correctly assigned to the family Mutillidae.

However, that still appears to have been the first description of a fossil mutillid.

Sharov (1957) described another new species of aculeate Hymenoptera from

the Cretaceous of Siberia, which was ultimately assigned to Mutillidae. That fossil

specimen, estimated to be about 80 million years old, was called Cretavus sibir-

icus Sharov, for which the author created a new family, Cretavidae. Rasnitsyn

(1975) later placed it in the family Mutillidae.

Manley & Poinar (1991) described Dasymutilla dominica Manley & Poinar

from Dominican amber estimated to be from 25 to 40 million years old. That

represented the first record and description of a fossil mutillid from the New

World. Here we describe another new species of fossil Dasymutilla, also from

Dominican amber.

' Technical contribution no. 4376 of the South Carolina Agricultural Experiment Station, Clemson

University.

1999 MANLEY & POINAR: NEW FOSSIL DASYMUTILLA 49

Figure 1. Holotype of Dasymutilla albifasciatus Manley & Poinar. Lateral view showing pale felt

line (arrow) and white band on thorax (fleck).

METHODS AND MATERIALS

The specimen is believed to have originated from mines in the Cordillera Sep-

tentrional of the Dominican Republic. These mines are in the El Mamey For-

mation (Upper Eocene), which is a shale-sandstone interspersed with a conglom-

erate of well-rounded pebbles (Eberle et al. 1980). The exact age of the amber is

unknown but estimates based on various microfossils and chemical analyses pro-

vide a range from 15—20 million years (Iturralde-Vincent & MacPhee 1996) to

30—45 million years (Cepek in Schlee 1990). The amber piece containing the

fossil mutillid is possibly between 20 to 40 million years. It is roughly triangular

in shape, with the sides measuring 28 mm by 40 mm by 25 mm. The thickness

of the piece varies from 5—7 mm and the weight is 3.3 g. Other prominent inclu-

sions in the amber piece are a drosophilid fly, bush cricket, ciid beetle, erythraeid

mite, cranefly and bark fragment. The amber was re-shaped and re-polished by

one author (GOP) to obtain the best viewing angle.

DASYMUTILLA ALBIFASCIATUS MANLEY & POINAR, NEW SPECIES (Fig. 1)

Type.—Holotype male, data: DOMINICAN REPUBLIC. Cordillera Septentri-

onal mountains; deposited in the Poinar amber collection, Department of Ento-

mology, Oregon State University, Corvallis, Oregon, U.S.A.

Description—Male. Length 10 mm. Integument predominantly black; pubescence sparse, both pale

and black, erect and semierect. Head black, rounded, with sparse, pale pubescence throughout; head

appears narrower than thorax, although this character is not clear. Eyes and ocelli normal; eyes large,

not bulging; ocelli small; ocello-ocular distance about 3.0X greatest width of ocelli. Punctures rela-

tively large, well-separated where visible. Mandibles bidentate, black throughout. Clypeus bidentate.

Antennae black; flagellomeres subequal in length; scape bicarinate beneath; punctation not apparent,

antennal scrobes appear slightly carinate, although not clearly visible. Thorax black, except broad

white band extending from upper part of propodeum laterally through lower half of propleura (Fig.

1); white band void of pubescence except a few sparse white hairs on the propodeum and posterior

50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

part of the mesopleuron of the band. Thorax appears longer than broad, although this character not

clearly visible. Pronotum with humeral angles rounded; anterior margin apparently not emarginate

(consistent with evenly rounded posterior margin of head). Mesonotum black except white circular

spot in middle; without laterally expanded lobes. Propodeum coarsely reticulate. Tegulae subhemi-

spherical, appear glabrous, smooth and shining; anterior margin with sparse, black pubescence. Pleura

coarsely punctate throughout, with relatively large, separated punctures. Prothorax and propodeum

with sparse, erect pubescence. Mesonotum, scutellum, and pleura with black, recumbent pubescence.

Legs black, with sparse, pale, erect pubescence; calcaria pale; apices of middle and hind femora

rounded; posterior trochanters appear normal, not toothed, although not clearly visible; hind tibiae

cylindrical, not flattened on inner margin. Wings folded, not clearly visible, but appear banded biack

and white. Abdomen black throughout. Abdomen with sparse, shallow punctures, except second tergite

laterally with relatively coarse punctation. First sternite with a longitudinal carina about nine-tenths

length of segment. Tergite I and tergite II anteriorly with sparse, pale, erect pubescence; tergite I

terminating in apical band of pale pubescence. Sternum with sparse, pale, erect pubescence scattered

throughout; sterna II-IV with apical bands of pale pubescence; sterna V-VI with apical bands of black

pubescence. Apical half of tergite II, and remainder of tergum clothed with recumbent and erect, black

pubescence. Sternite II with a round, median pit filled with pale hairs. Felt lines with pale pubescence

(Fig. 1). Pygidium not clearly visible, but apparently with apical fringe of hairs. Hypopygium black,

punctation not readily visible; posterolateral angles of last sternite rounded, not dentate.

Diagnosis.—Dasymutilla albifasciatus, like D. dominica, is placed in the bio-

culata species group. It is distinguished from all other members of the bioculata

Species group by the white band along each side of the thorax and the white spot

on the mesothorax. In no other known Dasymutilla is there any evidence of white

integument.

Etymology.—The specific name is taken from the Latinized albus, meaning

white, and fasciatus, meaning broad transverse stripe, thus reflecting the speci-

men’s diagnostic character of a transverse white band of integument along the

side of the thorax.

Distribution.—Known only from the type specimen.

Material Examined.—Type.

DISCUSSION

Dasymutilla albifasciatus, like D. dominica, exhibits the characteristics attri-

buted to males of the genus Dasymutilla by Mickel (1928). These include eyes

being round, prominent, almost hemispherical in shape, polished, with the facets

usually (but not always) very indistinct, and the first abdominal segment either

distinctly petiolate, subpetiolate, or subsessile, but never completely sessile with

the second. Another important character present only in some Dasymutilla and

some Jraumatomutilla is the median pit filled with hairs on the second abdominal

sternite. That pit is present in the specimen of D. albifasciatus, just as it is in the

specimen of D. dominica.

Mickel (1928) pointed out that many of the characters present in Dasymutilla

are present in more than one species. He used the term “species group,’ with

members of each group being closely related. In Mickel’s (1928) key to males of

Dasymutilla, D. albifasciatus keys to couplet 54 or 55, with species in couplets

54—56 belonging to the bioculata species group. As noted in the description, it

is unclear whether or not the antennal scrobes are carinate. Thus, it is not known

for certain whether this species would key to couplet 54 or 55. Characters shared

by males of this group include the presence of a median pit filled with hairs on

the second abdominal sternite, the presence of an apical fringe of hairs on the

1999 MANLEY & POINAR: NEW FOSSIL DASYMUTILLA 51

last tergite, the absence of a prominent tooth on the posterior trochanters, normal

size ocelli and eyes, cylindrical posterior tibiae, and rounded apices on the middle

and hind femora. As also noted in the description, it is not completely clear

whether or not the apical fringe of hairs on the last tergite is present, although it

certainly appears to be there. And the posterior trochanters are partially obscured

in the specimen, although they do not appear to have the prominent tooth present

in a few species.

This species is very similar to D. dominica, especially in the color of the

integument almost entirely black. However, D. albifasciatus can be distinguished

from the former by the presence of the broad lateral band of white integument

extending from the top of the propodeum to the lower half of the pronotum.

Dasymutilla albifasciatus also has a small, circular spot of white integument on

the mesonotum. It also can be distinguished from D. dominica by the white hairs

in the felt lines (the felt lines are black in D. dominica).

This species, like D. dominica, appears to be a highly derived species, based

on the presence of the pit on the second abdominal sternite. Thus, this provides

further evidence that the genus Dasymutilla likely radiated from a more primitive

Sphaeropthalmina well over 40 million years ago.

LITERATURE CITED

Bischoff, H. 1915. Bernsteinhymnenopteren. Schrift. Phys.-Oekon. Ges., Konigsberg, 56: 139-144.

Brischke, C. G. A. 1886. Die Hymenopteren des Bernsteins. Schr. Nat. Ges. Danzig, N. FE, 6: 278-

219:

Eberle, W, W. Hirdes, R. Muff & M. Pelaez. 1980. The geology of the Cordillera Septentrional. pp.

619-632. In Proceedings of the 9th Caribbean Geological Conference, held in Santo Domingo,

August 1980.

Iturralde-Vincent, M. A. & R. D. E. MacPhee. 1996. Age and paleogeographical origin of Domincan

amber. Science, 273: 1850-1852.

Krombein, K. V. 1979. Biosystematic studies of Celonese wasps, IV. Kudakrumiinae, a new subfamily

of primitive wasps (Hymenoptera: Mutillidae). Trans. Am. Entomol. Soc., 105: 67-83.

Larsson, S. G. 1978. Baltic amber—a palaeobiological study. Entomonograph. Volume I. Scandinavian

Science Press, Klampenborg, Denmark.

Manley, D. G. & G. O. Poinar, Jr. 1991. A new species of fossil Dasymutilla (Hymenoptera: Mutillidae)

from Dominican amber. Pan-Pacific Entomol., 67: 200-205.

Menge, A. 1856. Lebenszeichen vorweltlicher, im Bernstein eingeschlossener Tiere. Programm Pe-

trischule, Danzig, 1-32.

Mickel, C. E. 1928. Biological and taxonomic investigations on the mutillid wasps. U.S. Nat. Mus.

Bull., 143: 1-351.

Rasnitsyn, A. P. 1975. Hymenoptera Apocrita of the Mesozoic. Trudy Paleontol. Inst. Akad. Nauk

SSSR, 147: 1-140.

Schlee, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitrager fur Naturkunde, Ser. C, No. 28: 1-100.

Scudder, S. H. 1891. Index to the known fossil insects of the world including myriapods and arachnids.

Bull. U.S. Geol. Survey, 71.

Sharov, A. P. 1957. First discovery of a Cretaceous aculeate hymenopteron. Dokl. Akad. Nauk SSSR,

112: 943-944.

Received 13 Feb 1998; Accepted 2 Sep 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 52-54, (1999)

Scientific Note

CHILACIS TYPHAE (PERRIN) AND HOLCOCRANUM

SATUREJAE (KOLENATI) (HEMIPTERA: LYGAEOIDEA:

ARTHENEIDAE): FIRST WESTERN NORTH AMERICAN

RECORDS OF TWO PALEARCTIC CATTAIL BUGS

Until the mid- to late 1980s, no members of the lygaeid subfamily Artheneinae

were recorded from the New World. The subfamily’s presence in the Western

Hemisphere was recognized with the transfer of an indigenous neotropical species,

Polychisme ferruginosus (Stal), from Ischnorhynchinae to Artheneinae (Slater, J.

A. & H. Brailovsky. 1986. J. N. Y. Entomol. Soc., 94: 409-415), but this species

has now been returned to the Ischnorhynchinae (Kerzhner, I. M. 1997. Zoosyst.

Ross., 6: 213—222). Two Palearctic cattail-associated artheneines—Chilacis typhae

(Perrin) and Holcocranum saturejae (Kolenati)—have since been detected in east-

ern North America (Wheeler, A. G. & J. E. Fetter. 1987. Proc. Entomol. Soc.

Wash., 89: 244—249; Hoffman, R. L. & J. A. Slater. 1995. Banisteria, 5: 12—15).

In addition, with reassessment of pentatomomorphan phylogeny, the Artheneinae

have been proposed as a separate lygaeoid family, the Artheneidae (Henry, T. J.

1997. Ann. Entomol. Soc. Am., 90: 275-301).

Chilacis typhae, belonging to a monotypic genus, was first reported in North

America from Delaware, Maryland, New York, and Pennsylvania (Wheeler &

Fetter 1987). It has since been found in Tennessee and Virginia (Hoffman, R. L.

1996. The insects of Virginia No. 14: seed bugs of Virginia (Heteroptera: Ly-

gaeidae). Virginia Museum of Natural History, Martinsville, Virginia). This seed-

feeding bug develops on and in pistillate heads (“‘spikes’’) of cattails (Typha spp.;

Typhaceae), typically those that have been tunneled by larvae of the Holarctic

cosmopterygid Limnaecia phragmitella Stainton (Wheeler & Fetter 1987). In Eu-

rope, it sometimes can be found in cattail heads uninfested by this microlepidop-

teran (Stehlik, J. L. & I. Vavirinova. 1996. Acta Mus. Moraviae Sci. Nat., 80:

163-233). This bug is restricted to feeding on Typha spp. throughout its native

West Palearctic range (Hoffman & Slater 1995; Stehlik & Vavrinova 1996). Pop-

ulations of this apparently bivoltine species are most abundant in cattail heads

from May to October (Stehlik & Vaviinova 1996).

The first North American records of H. saturejae were from Florida, New

Jersey, North Carolina, South Carolina, and Virginia (Hoffman & Slater 1995).

Hoffman (1996) noted that Wheeler & Fetter’s (1987) records of C. typhae from

New Castle Co., Delaware, and from Chester Co., Pennsylvania, were based in

part on material of H. saturejae, thus adding those states to its known North

American distribution. Holcocranum saturejae has been found mainly at low el-

evations from southeastern Pennsylvania to northern Florida, with populations

also recorded west of the Blue Ridge Mountains of Virginia up to approximately

760 m above sea level (Hoffman 1996). Although all eastern U.S. collections

have been made from cattail heads, this species sometimes develops in willow

(Salix spp.; Salicaceae) catkins in Europe (Hoffman & Slater 1995). Adults over-

winter in cattail heads, and in some parts of Europe, adults are found on Typha

1999 SCIENTIFIC NOTE 53

spp. only from October to January. Some Old World populations, however, use

cattail seeds for nymphal development (Stehlik & Vavrinova 1996).

On the basis of recent collecting, we add to the western North American fauna

both European cattail bugs, whose North American distributions have been as-

sumed to be strictly eastern: C. typhae from Oregon and Washington and H.

saturejae from California to Texas. Voucher specimens are deposited in the Na-

tional Museum of Natural History, Smithsonian Institution, Washington, D.C.

Chilacis typhae becomes the fifth lygaeoid species considered adventive in the

Pacific Northwest; distributions of the four previously recorded adventive ly-

gaeoids in that region, all litter-inhabiting species, were reviewed by A. Asquith

& J. D. Lattin (1991. Pan-Pac. Entomol. 67: 258-271). Western populations of

H. saturejae, whose Old World distribution is more southern than that of C. typhae

(Hoffman & Slater 1995), have been found as far north as Colusa County in

north-central California. Both artheneids are among several Palearctic heteropter-

ans that likely represent separate accidental introductions to eastern and western

North America (Wheeler, A. G. & T. J. Henry. 1992. A synthesis of the Holarctic

Miridae (Heteroptera). Entomological Society of America, Lanham, Maryland).

The possibility exists that at least H. saturejae might represent a long-distance

(aeolian) migrant (Hoffman & Slater 1995).

Typha latifolia is a naturally Holarctic plant, but T. angustifolia, though often

considered Holarctic, might not be a native species (Grace, J. B. & J. S. Harrison.

1986. Can. J. Plant Sci., 66: 361-379). Both lygaeoids, however, should be con-

sidered nonindigenous in the Nearctic Region, based mainly on their recent de-

tection and the absence of indigenous artheneids in the North American fauna

(Wheeler & Fetter 1987, Hoffman & Slater 1995). In addition, neither lygaeoid

shows a Beringian distribution that is typical of many naturally Holarctic heter-

opterans (Wheeler & Henry 1992).

Records.—Chilacis typhae: OREGON: WASHINGTON Co.: Tigard, 29 Sep

1997, A. G. Wheeler, Typha latifolia, old pistillate heads, 11 adults. WASHING-

TON: COLUMBIA Co.: Rt. 30, 8 km W of Clatskanie, 20 May 1998, C. A.

Stoops & A. G. Wheeler, T. latifolia, pistillate heads, 4 adults. GRAYS HARBOR

Co.: McCleary, 20 May 1998, C. A. Stoops & A. G. Wheeler, T. latifolia, pistillate

heads, 3 adults. KING Co.: Issaquah, 22 May 1998, C. A. Stoops & A. G. Wheel-

er, T. latifolia, pistillate heads, 2 adults. KITSAP Co.: Poulsbo, 28 Sep 1997, C.

A. Stoops & A. G. Wheeler, T. latifolia, old pistillate heads, 9 adults. KITTITAS

Co.: Ellensburg, 22 May 1998, C. A. Stoops & A. G. Wheeler, T. latifolia, pis-

tillate heads, 1 adult. LEWIS Co.: E of Napavine, 29 Sep 1997, A. G. Wheeler,

T. latifolia, old pistillate heads, 3 adults. MASON Co.: Shelton, 29 Sep 1997, A.

G. Wheeler, T. latifolia, old pistillate head, 1 adult. PACIFIC Co.: Raymond, 20

May 1998, C. A. Stoops & A. G. Wheeler, T. latifolia, pistillate heads, 4 adults.

SNOHOMISH Co.: Edmunds, 21 May 1998, C. A. Stoops & A. G. Wheeler, T.

latifolia, pistillate heads, 4 adults. THURSTON Co.: Nisqually, 29 Sep 1997, A.

G. Wheeler, T. latifolia, old pistillate heads, 5 adults.

Holcocranum saturejae: ARIZONA: COCHISE Co.: Gray Hawk Ranch, along

San Pedro River, 13 km E of Sierra Vista, 5 Jun 1997, A. G. Wheeler, T. angus-

tifolia, pistillate heads, 16 adults. PIMA Co.: Buenos Aires National Wildlife

Refuge, Arivaca Cienega, E of Arivaca, 28 Jun 1997, A. Smith, Typha sp., pis-

tillate heads, 5 adults. CALIFORNIA: COLUSA Co.: Rt. 20, 1.6 km W of Colusa,

54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

9 Aug 1998, A. G. Wheeler, Typha sp., pistillate heads, 11 adults; Rt. 45, 1.3 km

N of Yolo Co. line SW of Tisdale, 9 Aug 1998, A. G. Wheeler, Typha sp.,

pistillate heads, 6 adults. IMPERIAL Co.: Salton Sea National Wildlife Refuge,

Hazard Unit, NW of Calipatria, 4-5 Dec 1997, K. Sturm & K. Haley, T. latifolia,

pistillate heads, 4 adults. KERN Co.: nr Inyokern, 3 Nov 1997, C. A. Stoops,

Typha sp., old pistillate head, 1 adult; Kern National Wildlife Refuge, Marsh Unit

1, 30 km W of Delano, 21 Nov 1997, J. Allen, Typha sp., pistillate heads, 3

adults. MONTEREY Co.: Monterey, 20 Mar 1998, C. A. Stoops, Typha sp.,

pistillate head, 1 adult. YOLO Co.: Rt. 16, NW of Rumsey, 10 Aug 1998, A. G.

Wheeler, Typha angustifolia, pistillate heads, 5 adults. NEVADA: CLARK Co.,

Floyd Lamb State Park, Las Vegas, 13 Nov 1998, W. K. Reeves, Typha sp.,

pistillate heads, 6 adults. NEW MEXICO: SOCORRO Co.: Bosque del Apache

National Wildlife Refuge, nr Socorro, 15 Sep 1997, M. Oldham, T. latifolia,

pistillate heads, 8 adults. TEXAS: HIDALGO Co.: Santa Ana National Wildlife

Refuge, S of Alamo, 18 Feb 1998, W. A. Jones & W. Warfield, T. domingensis,

pistillate heads, 10 adults.

Acknowledgment——The first collection of either Old World cattail bug in the

West was at Gray Hawk Ranch in Arizona, and we thank naturalist Sandy An-

derson for taking one of us (AGW) to a cattail stand on her property, Noel

McFarland for his hospitality in Arizona, Walker Jones (USDA, ARS, Weslaco,

Texas) for collecting Holcocranum saturejae in Texas, and Will Reeves (Clemson

University) for collecting it in Nevada. We are indebted to personnel (and vol-

unteers) of the National Wildlife Refuge System for making or facilitating col-

lections of H. saturejae: Jack Allen (Kern), Sally Gall (Buenos Aires), Katie

Haley (volunteer, Salton Sea), Alex Smith (volunteer, Buenos Aires), Mike Old-

ham (Bosque del Apache), and Ken Sturm (Salton Sea). We also appreciate at-

tempts to find cattail bugs at other National Wildlife Refuges by David Blankin-

ship at Lower Rio Grande Valley in Texas and Kim Forrest at Humboldt Bay in

California. I. M. Kerzhner (Russian Academy of Sciences) called our attention to

his recent paper, and P. H. Adler (Clemson University) provided useful comments

on a draft of this manuscript.

A. G. Wheeler, Jr., Dept. of Entomology, Clemson University, Clemson, South

Carolina 29634-0365, & Craig A. Stoops, 14990 Growler Ave., Apt. D, Naval

Submarine Base Bangor, Silverdale, Washington 98315-9619.

Received 20 Apr 1998; Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 55-57, (1999)

Scientific Note

FIRST RECORD OF GLYCASPIS BRIMBLECOMBEI

(MOORE) (HOMOPTERA: PSYLLIDAE) IN NORTH

AMERICA: INITIAL OBSERVATIONS AND PREDATOR

ASSOCIATIONS OF A POTENTIALLY SERIOUS NEW

PEST OF EUCALYPTUS IN CALIFORNIA

Eucalyptus spp. were first introduced to California in the mid 1800s and within

a short time were planted throughout the state for fuel wood, windbreaks, cut

foliage, and landscaping. In California, eucalyptus were considered pest free until

the early 1980s when the situation changed with the discovery of the psyllid

Blastopsylla occidentalis Taylor (Taylor, K. L. 1985. J. Aust. Ent. Soc., 24: 17-—

30), and the borer Phoracantha semipunctata Fabr. (Scriven, G. T., Reeves, E. L.

& Luck, R. F 1986. Calif. Agric. 40: 4—6). Subsequently, several other eucalyptus

feeding insects from Australia were discovered in California (Table 1). It is note-

worthy that most of the species in Table 1 are psyllids, and two of them (Cten-

arytaina longicauda Taylor and C. spatulata Taylor) were not described prior to

their discovery in California (Taylor, K. L. 1987. J. Aust. Ent. Soc., 26: 299-233,

Taylor, K. L. 1997. Aust. J. Ent., 36: 113-115). Presently, only the borers (Phor-

acantha spp.) and the blue gum psyllid (C. eucalypti Maskell) are considered

economic pests of eucalyptus in California. University of California biological

control programs introduced parasitoids from Australia which have effectively

controlled C. eucalypti on the ornamental foliage crop E. pulverulenta Sims and

P. semipunctata on several susceptible species (Dahlsten, D. L., Rowney, D. L.,

Copper, W. A., Tassan, R. L., Chaney, W. E., Robb, K. L., Tjosvold, S., Bianchi,

M. & Lane, P. 1998. Calif. Agric., 52: 31-34; Hanks, L. M., Paine, T: D. & Miller,

J. G. 1996. Calif. Agric., 50: 14-16). This note focuses on the initial distribution

and predators in northern California of Glycaspis brimblecombei Moore, a re-

cently recorded and potentially serious eucalyptus pest.

The genus Glycas pis Taylor (Homoptera: Psyllidae) is one of the largest of the

Myrtaceae-feeding psyllids and includes over 120 species which are naturally

distributed from Australia to the Philippine Islands (Moore K. M. 1970 Aust.

Zool., 15: 248-342). We found no previous reports of G. brimblecombei or any

Glycaspis spp. outside their native range. In contrast to the other free-living eu-

calyptus psyllids in California, G. brimblecombei nymphs live individually under

round glabrous, conical coverings called ‘lerps’ (Morgan, E D. 1984. Psylloidea

of South Australia). The presence of the 1-4 mm diameter lerps on both sides of

expanding and fully expanded leaves makes this psyllid highly conspicuous even

at low population densities.

Glycas pis brimblecombei was first found in El Monte, Los Angeles County in

June 1998 on Eucalyptus camaldulensis Dehnh (Cindy Werner, personal com-

munication), and it was found in Fremont, Alameda County in northern California

a month later. By September 1998, G. brimblecombei had been recorded in several

other cities in northern California including Alameda, Hayward, Oakland, Palo

Alto, San Bruno, San Mateo, South San Francisco, and San Francisco. The new

56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(1)

Table 1. Exotic pests of Eucalyptus found in California.

Date found County

Psyllidae

Blastopsylla occidentalis Taylor 1983 Los Angeles*

Ctenarytaina longicauda Taylor 1983 San Diego*

C. eucalypti Maskell 1991 Monterey*

C. spatulata Taylor 1991 Orange*

Cryptoneossa triangula Taylor 1995 Orange*

Glycaspis brimblecombei Moore 1998 Los Angeles*

Coleoptera

Phoracantha semipunctata Fabr. 1984 Orange*

Gonipterus scutellatus Gyllenhal 1994 Ventura*

Phoracantha recurva Newman 1995 Riverside*

Trichomela sloanei Blackburn 1998 Riverside*

Hymenoptera

Aprostocetus sp. 1995 Santa Barbara*

* New record for North America.

1 Gill, R.J. 1998. Cal. Plant Pest Dis. Rpt., 17: 21-24.

psyllid is responsible for severe defoliation and decline of several eucalyptus

Species in northern California, and has become a nuisance in ornamental settings

where the lerps and honey dew coated leaves stick to shoes of pedestrians. We

speculate that the damage caused by G. brimblecombei in California may reflect

the absence of natural enemies and the unusually high rainfall this past winter. In

Australia, outbreaks of several species of lerp forming psyllids are known to

follow years of high rainfall (Moore, K. M. 1961. Proc. Linn. Soc. N. S. W., 86:

185—200, White, T: C. R. 1969. Ecol., 50: 905—909).

In Australia, eight Eucalyptus spp. are known hosts of G. brimblecombei: E.

blakelyi Maiden, E. brassiana Blake, E. bridgesiana Baker, E. camaldulensis, E.

camphora Baker, E. dealbata Cunn. ex Schauer, E. mannifera ssp. maculosa Bak-

er, E. nitens Deane & Maiden, and E. teriticornis Smith (Moore 1970; Carver,

M. 1987. J. Aust. Ent. Soc., 26: 369-372). In contrast, we have found eggs, early

through late stage nymphs, and adults of G. brimblecombei on three other species

including E. diversicolor EK Muell, E. globulus Labill, and E. sideroxylon Cunn.

ex Woolls. We are currently studying the host associations and effects of G.

brimblecombei on additional Eucalyptus spp. in California in an effort to identify

resistant species and monitor the effectiveness of existing predators.

Moore (1961) reported several predators (bell-birds, spiders, mites, and larvae

of Syrphidae (Diptera), Hemerobiidae (Neuroptera), Chrysopidae (Neuroptera),

and Coccinellidae (Coleoptera) and parasitoids (Chalcidoidea: Hymenoptera)) at-

tacking Glycaspis spp. in Australia. At the Fremont site in northern California we

have observed several arthropod predators associated with G. brimblecombei in-

cluding: spiders, mites, Heteroptera (Anthocoris nemoralis Fabr. and Zelus ren-

ardii Kolenati), Coccinellidae (Harmonia axyridis Pallas, Chilocorus bipustulatus

L., Hippodamia convergens Guérin, and Coccinella californica Mannerheim),

Syrphidae, Hemerobiidae, and Chrysopidae. In California, A. nemoralis is an ac-

cidentally introduced predator of the Australian acacia psyllid (Acizzia uncatoides

1999 SCIENTIFIC NOTE 57

Ferris & Klyver) (Hagen & Dreistadt 1990). Furthermore, despite the abundance

of the introduced parasitoid of C. eucalypti, Psyllaephagus pilosus Noyes (Hy-

menoptera: Encyrtidae), in the area, neither this wasp nor others have been ob-

served parasitizing G. brimblecombei. Based on our observations, there is an

apparent inability of the existing predators to control this new psyllid in Califor-

nia, and we conclude that additional biological control organisms will need to be

introduced to effect biological control.

The economic impact of G. brimblecombei may be far more serious and wide-

spread than that of other eucalyptus psyllids in California for several reasons.

First, G. brimblecombei oviposits and successfully completes its life cycles on

both expanding and fully expanded leaves, and may defoliate the host. In contrast,

the other extant psyllid species (C. eucalypti, C. spatulata, B. occidentalis) in

California do not cause defoliation, and their populations are partially controlled

by host phenology because they oviposit and feed almost exclusively in buds and

tender foliage (Brennan, personal observation 1995-1998). Second, G. brimble-

combei appears to have a broader host range than the other psyllids, and, thus,

may impact commercial plantations, and ornamental and forestry plantings of

eucalyptus. Third, of all the Australian psyllid species established in California,

Glycaspis is the only genus within its native range known to exhibit outbreak

populations and cause severe damage to eucalyptus forests and plantings (Moore

1961). Fourth, it is likely that the repeated defoliation caused by G. brimblecombei

may induce stress and thus increase Phoracantha spp. attacks as has been ob-

served with other Glycas pis spp. and beetles (Xyleborus sp.) in Australia (Moore

1961). The ‘Eucalypt Dieback’ syndrome in Australia that has caused the decline

and premature death of eucalytpus in agricultural environments, is thought to be

caused by interactions between weather-induced stress and attacks by several

guilds of insects (Landsberg, J. 1990. Aust. J. Ecol., 15: 89-96).

It is clear that G. brimblecombei has the potential to affect eucalyptus culture

in California. Existing biological controls are not effective at preventing damage

and new ones are needed. Eucalyptus can no longer be considered pest-free in

California, and pest resistance should be considered in selecting species for future

plantings.

Acknowled gment.—The authors thank Cindy Werner of the Los Angeles Coun-

ty Department of Agriculture, Nancy Brownfield and Ira M. Bletz of the East Bay

Regional Parks, Ronnie Eaton and Earl G. Whitaker of the Alameda County

Department of Agriculture, and Carol Sweetapple and Herb Fong of Stanford

University Facilities Operations for their cooperation. This research was partially

funded by a Jastro Shields research award from the Graduate Group in Plant

Biology at U.C. Davis.

Eric B. Brennan!, Raymond J. Gill*, G. Frederic Hrusa? and Steven A. Wein-

baum!, ‘Department of Pomology, University of California, Davis, California

95616; *California Department of Food and Agriculture, Sacramento, California

95832-1448.

Received 17 Sep 1998: Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(1): 58-59, (1999)

Scientific Note

THE PAPER WASP POLISTES DOMINULWUS (CHRIST)

(HYMENOPTERA: VESPIDAE) IN THE STATE OF

WASHINGTON

Polistes dominulus (Christ) is native to the Palearctic region, where it is broadly

distributed (Carpenter 1996. Mem. Mus. Nat. Hist. Not., 173: 135-161). It is a

‘hitchhiker’? species, apparently traveling around the world with human traffic

and commerce. It has been introduced into North America; first reported as Pol-

istes gallicus (L.) in Massachusetts (Hathaway 1981. Psyche, 88: 169-173). Pol-

istes dominulus appears to now be generally distributed in much of the north-

eastern U.S., and westward at least into Ohio and Michigan (Jacobson 1994,

Sphecos 27: 14; Judd & Carpenter 1996. Great Lakes Entomol., 29: 45—46;

Staines & Smith 1995. Proc. Entomol. Soc. Wash., 97: 891). It has not been

reported heretofore in western North America. We report here the establishment

of Polistes dominulus at multiple sites in central Washington (in the Yakima Val-

ley) as well as in Pierce County, west of the Cascade Mountain Range.

In an urban neighborhood of Sunnyside, Yakima County, Washington, on 4

Sep 1998, a cluster of wasps was observed on the front of a travel trailer in the

front yard of a residence. On closer inspection, it was noted that wasps were

entering and exiting a break in the wall of the trailer, with about 30 wasps re-

maining outside within 15 cm of that opening. These wasps were not collected,

but were photographed, and appear to be P. dominulus. Later that day, five Pol-

istes dominulus were collected on a hedge of planted snowberry, Symphoricarpus

albus (L.), heavily visited by golden paper wasps, Polistes aurifer (Saussure).

Polistes dominulus were also observed entering a garage at the same site via a

fallen soffet under an eave and also were entering a gap in a cinder block wall.

On 5 and 6 Sep 1998, five male P. dominulus were netted while they were

flying about an ornamental crabapple tree in a suburban neighborhood 4—8 km

west of the City of Yakima. A small nest (about 40 cells) occupied by eight P.

dominulus wasps was later (18 Oct) observed under an eave on the south side of

a residence at the same location. On 7 Sep 1998, two active colonies were located

near Ahtanum, in the West Valley area of Yakima County. Both nests were under

an eave on the east side of a garage, with one abutting the envelope of a nest of

the aerial yellowjacket Dolichovespula arenaria Fabr. The P. dominulus nest ad-

jacent to the aerial yellowjacket nest was collected. It was 18 X 14 cm in diameter

and contained 322 cells. There were 23 wasp pupae and 7 larvae in the cells of

that nest. Forty-one female and 3 male wasps were collected with the nest, while

2 escaped. On 8 Sep 1998, 11 paper wasp nests were located under eaves of a

house in the city of Yakima. Seven of these nests were occupied by P. dominulus

wasps, and 4 were not occupied. The two largest nests were collected. A nest on

the east side of the house was 17 cm in diameter and contained 412 cells with

27 pupae, 45 larvae and 12 eggs. Many cells appeared to contain honey; probably

concentrated sugars from honeydew (Rau, 1928. Biol. Bull. Marine Biol. Lab.,

Woods Hole, Massachusetts, 24: 503-519). There were 42 female P. dominulus

PAN-PACIFIC ENTOMOLOGIST

75(1): 60, (1999)

Scientific Note

A NEW HOST RECORD OF ORNITHOPHILA GESTROI

(DIPTERA: HIPPOBOSCIDAE) ON THE LESSER

KESTREL (FALCO NAUMANNI FLEISCHER) IN

GALAXIDI, GREECE

Field research on Lesser Kestrels (Falco naumanni) breeding in Greece re-

vealed occurrence of the bird-lousefly, Ornithophila gestroi (Rondani). On 1 Apr

1994 in the town of Nikea, two to four louseflies were observed trailing a few

cms behind Lesser Kestrels in flight. These louseflies easily tracked the aerial

maneuvers of kKestrels, even during rapidly circling courtship flights. It appeared

that individual flies could enter and exit the kestrels in flight.

On 13 Jun 1995, while banding Lesser Kestrels in Galaxidi, a breeding female

captured at 0800, weighing 147 gm, wing chord 244 mm, tail length 161 mm,

tarsus length 37 mm, with #1 and 2 left secondaries absent: Band #000175 had

seven bird-louseflies of which five were captured and preserved in an alcohol

solution of a local spirit called tsipouro. These rapidly moving flies within the

plumage of the female were difficult to capture. When pursued, O. gestroi left

the falcon.

Bird-louseflies were found on young and in nests of Lesser Kestrels. Because

the Lesser Kestrel is a social falcon, nesting colonially in houses and outbuildings

of villages and cities, the bird-lousefly readily moves among the nesting falcons.

The occurrence, however, of this hippoboscid is rare in the falcons of Greece.

Infestations ranged from two to seven flies per falcon; most birds under study

were free of O. gestroi.

T. C. Maa (Pacific Insects Monograph 20: 1—23, 1969) found the bird-lousefly,

O. gestroi to be confined to the Mediterranian subregion and to the Falconiformes:

Falconidae. Within the Ornithomyinae, Ornithophila is an archaic and rare genus

with a single male and four females constituting the type series in Genoa, Flor-

ence, and Harvard Museums. Galita Island near Malta, Crete, and Tangier are the

only localities with host documentation on record, with specimens collected from

Eleanora’s Falcon (fF. eleonorae Gene) and Common Kestrel (F. tinnunculus L.).

The five specimens taken in this study from mainland Greece represent both a

new host in Lesser Kestrels and locality at Galaxidi. Because the Lesser Kestrel

migrates to subsaharan Africa during the non-breeding season, O. gestroi can be

expected on that continent. Collected specimens reside at the California Academy

of Sciences and the J. Gordon Edwards Museum of Entolmology at San Jose

State University, San Jose, CA 95192.

Acknowled gment.—For identification, we thank Dr. Eric M. Fisher of the Cal-

ifornia Department of Food and Agriculture, Plant Pest Diagnostic Laboratory.

Thomas G. Balgooyen', Ben Hallmann?’, and Stanley E. Vaughn.! 'Department

of the Biological Sciences, San Jose State University, San Jose, CA 95192.

?GR-400 O08 Rapsani, Greece.

Received 23 Feb 1998; Accepted 18 May 1998.

1999 SCIENTIFIC NOTE 59

wasps collected with the nest. The other nest collected was on the north side of

the house, was 26 X 23 cm in diameter and contained 480 cells with 5 pupae, 4

larvae and no eggs. Again, many cells appeared to contain honey. Thirty two

wasps were collected with the nest.

On 7 Oct 1998, six P. dominulus females were collected at an urban residence

in the city of Puyallup, in Pierce County, Washington. This collection followed a

complaint of wasps inside of a home, and the wasps were not found with a nest.

These records indicate that P. dominulus is not only established but is probably

common and widespread in Yakima County, Washington. The presence of the

same species in the city of Puyallup indicates widespread distribution within the

state of Washington. Yakima and Pierce counties are east and west of the Cascade

Mountains respectively, which is a major barrier to insect dispersal. Polistes dom-

inulus probably was introduced into the Pacific northwest a number of years ago.

This wasp has a history of preferring nesting sites on or in structures and

enclosures (Hathaway 1981. Psyche, 88: 169-173), making them somewhat syn-

anthropic. Human activities and constructions provide abundant nest sites. Ob-

servations to date of P. dominulus made in Yakima County support this gener-

alization, with nests under eaves of buildings and within any type of cavity. Also,

intensive trapping of wasps in a rural area east of Yakima indicated large popu-

lations of Polistes aurifer but no Polistes dominulus (Landolt 1998. Environ.

Entomol., 27: 129-1234; Landolt 1999. J. Kansas Entomol. Soc., In press).

Voucher specimens of P. dominulus collected in Yakima County, Washington

were deposited in the James Entomological Collection, Department of Entomol-

ogy, Washington State University, Pullman, Washington.

Acknowled gments—Jim Bunch inititially called our attention to the presence

of large numbers of wasps at his apiary in Sunnyside, WA. Thanks to D. Horton

and M. Heidt of Yakima and K. Hobson of Puyallup for access to their properties

to collect wasps. Technical assistance was provided by J. Beauchene and L. Bid-

dick of USDA, ARS, Yakima and K. Hobson, WSU, Puyallup. This study was

supported in part by funds from Sterling International, Veradale, Washington.

Peter J. Landolt! and Arthur Antonelli’, 'U.S. Department of Agriculture, Ag-

ricultural Research Service, 5230 Konnowac Pass Road, Wapato, Washington

98951, *>Washington State University Cooperative Extension, 7612 Pioneer Way,

Puyallup, Washington 98371.

Received 23 Oct 1998; Accepted 7 Dec 1998.

PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for

discussion of this journal's specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English,

but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please maintain a copy

of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing.

Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in,

nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF

ANY FIGURES TO THE 8.5 X 11 IN PAGE. Number pag es as: title page (page 1), abstract and key words page (page 2), text pages

(pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List

the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. The title

must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use five

to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.

Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases

underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet

1988, Dorly et al. 1989), (Burton in press) and (R. F Tray, personal communication). For multiple papers by the same author use:

(Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith

1987: table 3).

Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that

volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are

applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED

(INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author

of each species after its first mention.

Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these

format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before

submitting manuscripts for which these formats are applicable.

Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. F Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide

some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. F Dixon & L. Weismann (eds.). Population structure, genetics and

taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB

Academic Publishing, The Hague, The Netherlands.

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.

Illustrations. — [Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label

letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. Authors

are strongly encouraged to provide illustrations no larger than 8.5 X 11 in for easy handling. Number figures in the order presented.

Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of

the manuscript, and (5) journal. FIGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures.

Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued on

additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter.

Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited

section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156—161.). A short

acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of

the note with the affiliation underscored.

Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for

pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include

reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page

charge fee notice with acknowledgment of initial receipt of manuscripts.

THE PAN-PACIFIC ENTOMOLOGIST

Volume 75 January 1999 Number 1

Contents

YOUNG, D. K.—Transfer of the Taiwanese Pseudopyrochroa umenoi and the Japanese P.

amamiana to Pseudodendroides (Coleoptera: Pyrochroidae: Pyrochroinae) ------------------

GILBERT, A. J. & E G. ANDREWS—Studies on the Chrysomelidae (Coleoptera) of the Baja

California peninsula: A new species of Scelolyperus (Galerucinae), with notes on the

genus in Baja California

DAANE, K. M. & L. E. CALTAGIRONE—A new species of Metaphycus (Hymenoptera:

Encyrtidae) parasitic on Saissetia oleae (Olivier) (Homoptera: Coccidae) —-...--...-.-.----.--

MANLEY, D. G.—Synonymy of Dasymutilla noctura Mickel (Hymenoptera: Mutillidae) ___.-

LATTIN, J.D. & N. L. STANTON—Host records of Braconidae (Hymenoptera) occurring in

Miridae (Hemiptera: Heteroptera) found on lodgepole pine (Pinus contorta) and asso-

Clan <@ayunierape ea Fk, a Seg I eer eee eee re een

MANLEY, D. G.—A synonymy for Pseudomethoca donaeanae (Cockerell and Fox) (Hyme-

MmOptcre: Winer agyS lee oP a. ee 5 Sa Ae oe we a i es De

SALGADO, J. M.—The Leiodidae (Coleoptera) of the Carnegie Museum of Natural History.

New data and description of two new species

MANLEY, D. G. & G. O. POINAR, JR.—A second species of fossil Dasymutilla (Hymenop-

tera: Mutillidae) from Dominican amber

SCIENTIFIC NOTES

WHEELER, JR., A. G. & C. A. STOOPS—Chilacis typhae (Perrin) and Holcocranum saturejae

(Kolenati) (Hemiptera: Lygaroidea: Artheneidae): first western North American records

of two Palearctic cattail bugs

BRENNAN, E. B., R. J. GILL, G. E HRUSA & S. A. WEINBAUM—First record of Glycaspis

brimblecombei (Moore) (Homoptera: Psyllidae) in North America: initial observations

and predator associations of a potentially serious new pest of eucalyptus in California

LANDOLYT, P. J. & A. ANTONELLI—tThe paper wasp Polistes dominulus (Christ) (Hymenop-

tera: Vespidae) in the State of Washington

BALGOOYEN, T. G., B. HALLMANN & S. E. VAUGHN—A new host record of Ornithophila

gestroi (Diptera: Hippoboscidae) on the lesser kestrel (Falco naumanni Fleischer) in

eevee Site: Closes Leben es, eve A eee ue he OM ee Te ie oe ee a

13

PR,

52

35

48

eV

ee

58

60

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 75 April 1999 Number 2

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

EDITORIAL BOARD

R. V. Dowell, Editor R. M. Bohart

Jeffery Y. Honda, Associate Editor J. T; Doyen

R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.

Julieta EF Parinas, Treasurer Warren E. Savary

Published quarterly in January, April, July, and October with Society Proceed-

ings usually appearing in the October issue. All communications regarding non-

receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary;

and financial communications should be addressed to: Julieta EK Parinas, Treasurer;

at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-

emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addizssed to: William Hamersky, Membership Committee chair, Pacific Coast

Entomological Society, Dept. of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118-4599.

Manuscripts, proofs, and all correspondence concerning editorial matters (but

not aspects of publication charges or costs) should be sent to: Dr. Robert V.

Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,

1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-

tributors, and volume 73(4): 248—255, October 1997, for more detailed informa-

tion. Information on format for taxonomic manuscripts can be found in volume

69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer.

The annual dues, paid in advance, are $25.00 for regular members of the So-

ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for

institutional subscriptions or sponsoring members. Members of the Society receive

The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes

are available; see 72(4): 247 for current prices. Make checks payable to the Pacific

Coast Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 1999

Laura A. Irons, President Vincent EK Lee, Managing Secretary

Julieta E Parinas, Treasurer Stanley E. Vaughn, Recording Secretary

THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per

year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate

Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional

mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society,

% California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

This issue mailed 7 July 1999

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

PAN-PACIFIC ENTOMOLOGIST

75(2): 61-67, (1999)

CAENIS CHAMIE, A NEW SPECIES FROM COLOMBIA

(EPHEMEROPTERA: CAENIDAE)

J. ALBA-TERCEDOR! AND S. MOSQUERA?

‘Departamento de Biologia Animal y Ecologia (Unidad deZoologia),

Facultad de Ciencias, Universidad de Granada,

18071-Granada, Spain

*Departamento de Procesos Quimicos y Biolégicos. Fac. de Ingenieria.

Universidad del Valle. Cali. Apartado Aereo 25360. Colombia

Abstract.—The alate stages, nymph and egg of Caenis chamie (Ephemeroptera: Caenidae), NEW

SPECIES, are described and illustrated on the basis of reared material from Colombia. A dis-

cussion of its relationships with other South American species of Caenis is included.

KeyWords.—Insecta, Ephemeroptera, Caenidae, winged stages, nymph, egg, Caenis chamie,

South America, Colombia.

The family Caenidae (Ephemeroptera) in South America is poorly known, with

25 recorded species in four genera (Brachycercus Curtis 1834; Brasilocaenis

Puthz 1975, Caenis Stephens 1835 and Cercobrachys Soldan 1986). The genus

Caenis is the most diverse with 17 species (Navas 1915, 1929a, 1920b, 1922,

1930; Froehlich 1969; Malzacher 1986, 1990; Pereira & Da Silva 1990; Da Silva

1993): 4 species are known from Argentina (C. albata Navas, C. argentina Navas,

C. ludicra Navas and C. nemoralis Navas), 8 from Brazil (C. candelata Mal-

zacher, C. cigana Pereira & Da Silva, 1990; C. cuniana Froehlich, C. fittkaui

Malzacher, C. pfulgfelderi Malzacher, C. quatipuruica Malzacher, C. reissi Mal-

zacher, and C. sigillata Malzacher), 2 from Chile (C. axillata Navas and C. nigella

Navas), and 2 from Paraguay (C. burmeisteri Malzacher and C. pseudamica Mal-

zacher). Berthélemy (1965) reported material from Chile, deposited in the Mu-

seum of Natural History in Paris, labelled by Navas as type of C. chilensis.

However, C. chilensis is a manuscript name and is therefore a nomen nudum.

During a general study of mayflies of southeast Colombia (Mosquera 1995),

specimens of an undescribed species of Caenis were collected. Based on labo-

ratory reared material herein describe all stages of the new species.

The types of the new species are deposited in the following intitutions: Museo

de Entomologia, Universidad del Valle, Cali, Colombia (MEUV) and the senior

author’s collection in the Departmento de Biologia Animal y Ecologia (Zoologia),

Universidad de Granada, Spain (JAC).

CAENIS CHAMIE ALBA-TERCEDOR & MOSQUERA, NEW SPECIES

Types.—Holotype, male imago (genitalia and left wing on slide # 300); data:

COLOMBIA. VALLE DEL CAUCA: Acueducto de Pavas, 1200 m. a.s.1., 76°33’ W,

3°40’ N, September 1994. S. Mosquera leg.; deposited MEUV. Paratypes: 1 male

imago (genitalia mounted for SEM), 4 male subimagos, 5 females, 5 nymphs, 5

exuviae skins (one on slide no. 301) and eggs on slide no. 302, all from the same

locality and date as the holotype; deposited in MEUV (except slides 301 and 302

deposited in JAC).

62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Figs. 1-7. Male imago (1-5): head and pronotum (1); wing (2); genitalia (3); forceps (4); and

prosternum (5). Egg 400 (6) and detail of micropyle, in light microscope 1000 (7).

Etymology—The new species is named after the “‘chamies’’ an ethnic group

that in the past inhabited the Cauca Valley.

Male imago. (in alcohol).—Length body: 2.9-3.7 mm, wings: 2.3—2.9 mm. Head: (Fig. 1): middle

longitudinal white line from occiput to frontal suture; dark band between lateral ocelli; antennae

1999 ALBA-TERCEDOR & MOSQUERA: NEW CAENIS SPECIES 63

Figs. 8-15. Caenis chamie sp.n. nymph: ventral view of female (8) and male abdominal sternites

(9); pronotum (10) and detail of the surface ornamentation <400 (11); fore femur (12) and bristles in

the row on the distal third (13); claws (14) (a: anterior, b: middle, c: posterior); details of distal part

of posterior tibia and claw (15); labial glossa and paraglossa (16).

yellow-white, pedicel length approximately twice that of scape; flagellum basally slightly enlarged;

yellow, progressively paler apically. Thorax: light brown pattern of pronotum as in Fig. 1; prosternum

shaped as in Fig. 5; meso- and metanotum yellow with dark sutures; wings hyaline, costal, subcostal

cells and veins C, Sc and R, shaded with light purple-grey; forefemora brown, with dark longitudinal

margins; tibiae and tarsi white; femur: tibia ratio = 0.54—0.61, fore tarsal formula 1<5<4=3<2.

Abdomen: no projection on second abdominal tergum; posterior 2/3 of each tergum with grey-brown

64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Ty Wy Wy Wy WY WY WY

Wy A

Ay WY OY wy ily WY Wy Yh wy

b

20

21

Figs. 17-20. Caenis chamie sp.n. nymph: lamellar apophysis of middle (17) and hind coxae (18:

a and b correspond to the right and left side); 2nd gill (19); submarginal row of scales on 2nd gill

X400 (20) (a: distal zone, b: middle zone, c: distal zone); dorsal projection on the 2nd abdominal

tergite (21).

pigmentation, terga 1-2 pigmented only in distal third; longitudinal black stripe ventrally at junction

of sterna and pleura. Genitalia (Figs. 3—4, 27-29): Penes anvil shaped, distal margin straight, with

rounded and well-developed lateral lobes, and with central longitudinal incision (Fig. 3). Forceps (Fig.

4) with blunt tips, densely covered with short setae (Figs. 28 and 29). Anterior apophysis of the

transversal sclerite of styliger (after Malzacher 1991) pointed (Fig. 3).

Subimago male. (in alcohol).—Similar to male in color and pattern except obscured by subimaginal

cuticle.

Female. (in alcohol)—Length body: 3.5—3.7 mm, wings: 3.6—3.8 mm.; general color yellow-brown

with pattern similar to that of male.

Egg.—(Figs. 6-7, and 22—26). Length: 159.5-176.2 wm. (* = 166, n = 20); width: 83.3-100 wm

(x = 92.5, n = 20). Egg bean shaped (Figs. 6 and 25). Two epithema (polar caps) present. Micropyle

long, with opening preceded by depression. Chorionic surface with small pores.

Nymph. (in alcohol).—Length body: 3.0—3.5 mm. (male), 4.3-5.3 (female). Head: transverse dark

band between eyes; pedicel length twice that is scape, 4—7 bifid setae % pedicel length; segment 3 of

maxillary palpi subequal in length segment 2, or slightly longer; glossae broad, longer than paraglossae

(Fig. 16). Thorax: Pronotum as in Fig. 10, dorsal surface with reticulate ornamentation (Fig. 11)

(similar ornamentation on vertex and metanotum); forefemora with irregular row of long bifid bristles

(Fig. 12); bristles distally pinnate (Fig. 13); lateral flanges of coxae as in Figs. 17 and 18; claws of

fore and middle legs long and slender, without denticles (Fig. 14a, 14b); hind claws more curved with

row of small denticles (Figs. 14c and 15); distal inner margin of hind tarsi with two rows of strong

and pinnate bristles (Fig. 15), bristles absent on forelegs, sparse and arranged in one row on middle

legs. Abdomen: dorsal projection on tergum 2 as in Fig. 21; postero lateral projections of abdominal

segments as in Fig. 8; abdominal sternite 9 slightly emarginate, sexually dimorphic (compare Figs. 8

and 9); lateral margins of terga 4-8 with long setae; posterior margins of terga 7-8 with long setae;

1999 ALBA-TERCEDOR & MOSQUERA: NEW CAENIS SPECIES 65

Figs. 22-25. SEM of the egg of Caenis chamie sp.n.: general view of egg (22, 25); polar cap

edge (23); micropyle showing the oval chorionic zone before the opening (24).

posterior margin of tergum 9 without setae but with small, somewhat rounded, denticles. Gill 2 as in

Fig. 19, with setae only on mesal fork of triangular ridge; submaginal row of scales simple (Fig. 20).

Discussion.—There is some variability in the shape of the male genitaliafor-

ceps. Thus, in one specimen the left forceps appear as normal (Fig. 29), but the

right forceps is broader (Fig. 28).

Of the 17 named species of Caenis from South America, seven were identified

and inadequately described by Navas. Material of four species (C. argentina, C.

axillata, C. ludicra and C. nemoralis) are deposited in Argentina (Museo Argen-

tino de Ciencias Naturales ‘“‘Bernardino Rivadavia’’, Buenos Aires, and Museo

de la Plata) (Dominguez 1989), and Spain (Museo de Zoologia de Barcelona)

(Alba-Tercedor & Peters 1985). Recently, the senior author has examined the

material deposited in Spain. The only existing specimen of C. axillata is extremely

damaged, lacking the wings, legs, and most of the abdomen, and thus is impos-

sible to redescribe. Malzacher (in litt——29.06.1997) has been studying the type

material of C. argentina and C. ludicra and he is preparing a redescription of

them.

Malzacher (1986) divided the South American species of Caenis into two

groups: the C. fittkaui—group (with long sclerotised and pointed forceps) and the

C. reissi—group (with short, apically rounded forceps). The new species belongs

to the C. reissi—group. This group includes other 7 known species (C. argentina,

C. cigana, C. ludicra, C. pfulgfelderi, C. quatipurunica, C. reissi and C. sigillata).

Diagnosis.—The winged stage of C. chamie sp.n. can be distinguished from

C. argentina because in C. argentina the area between the subcosta and radius

veins is conspicuously shaded dark brown, as was drawn by Navas (1915: fig.

4). Caenis chamie is close to C. pflugfelderi; however it can easily be distin-

66 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Figs. 26-29. SEM of Caenis chamie sp.n.: micropylar rib (26); male genitalia (27) (note that penes

were somewhat retracted after the deshydration process); detail and variability of forceps in a single

specimen (28, 29).

guished by the straight hind margin of penes, pointed and not inwardly curved

apophysis of the styliger sclerite, and broader prosternite. The chorionic surface

of egg has pores (similar to C. reissi, compare Figs. 23, 24 and 25 with Malzacher

1986: figs. 3 and 4) but not honeycombed as in C. pflugfelderi. The nymph is

similar to that of C. pflugfelderi. Both species share middle and hind claws without

denticles, but the new species can be easily distinguished by the emargination of

the hind margin of sternum 9.

ACKNOWLEDGMENT

We are indebted to: Peter Malzacher from Ludwigsburg (Germany) for confir-

mating the new species, useful comments and additional data on Navas’s species;

William and Janice Peters from Florida A&M University, Tallahassee, Florida

(U.S.A.) for their valuable suggestions advise, comments, and editorial work;

Eduardo Dominguez from Tucumén (Argentina) and Mike Hubbard from Talla-

hassee (U.S.A.) for bibliographic help. Support to the senior author to travel to

Colombia (Centro de Investigacién y Control de Contaminantes Ambientales de

la Universidad del Valle, Cali) was provided by the Institute of Environmental

Engineering-Bioengineering (Laussane, Switzerland). Page charges partially offset

by a grant from the C. P. Alexander fund.

LITERATURE CITED

Alba-Tercedor, J. & W. L. Peters. 1985. Types and additional specimens of Ephemeroptera studied by

Longinos Navas in the Museo de Zoologia del Ayuntamiento, Barcelona, Spain. Aquat. Ins.,

7: 215-217.

1999 ALBA-TERCEDOR & MOSQUERA: NEW CAENIS SPECIES 67

Berthélemy, C. 1965. Types of pinned Ephemeroptera deposited at the Natural History Museum of

Paris. Eatonia, 7: 2.

Da Silva, E. R. 1993. Descrigao do imago macho de Caenis cuniana Froehlich, com notas biolégicas

(Ephemeroptera, Caenidae). Revta. Bras. Zool., 10: 413-416.

Dominguez, E. 1989. Tipos de Ephemeroptera de L. Navas depositados en las colecciones entomo-

ldgicas de la Argentina. Rev. Soc. Entomol. Argent., 45: 271-274.

Froehlich, C. G. 1969. Caenis cuniana sp.n., a parthenogenetic mayfly. Beitr. Neotrop. Fauna, 6: 103—

108.

Malzacher, P. 1986. Caenidae aus dem Amazonasgebiet (Insecta, Ephemeroptera). Spixiana, 9: 83-—

103.

Malzacher, P. 1990. Neue Arten der Eintagsfliegen-Familie Caenidae (Insecta, Ephemeroptera) aus

Stidamerika. Stud. Neotrop. Fauna Environm., 25: 31-39.

Malzacher, P. 1991. Genital-morphological features in the Caenidae. pp. 73-85. In J. Alba-Tercedor

& A. Sanchez-Ortega (eds.). Overview and strategies of Ephemeroptera and Plecoptera. Sandhill

Crane Press.

Mosquera de A., S. 1995. Emergencia, formacién de enjambres y distribucién de algunas especies de

Ephemeroptera del sur occidente colombiano. Master’s Thesis, Universidad del Valle, Cali.

Navas, L. 1915. Neurdépteros sudamericanos (2? serie). Brotéria Zool. (Sér. Zool.), 13: 5-13.

Navas, L. 1920a. Insecta nova. VI series —Mem. Pontif. Accad. Rom. Nuovi Lincei, (2) 5: 11-19.

Navas, L. 1920b. Insectos sudamericanos (3? serie). An. Soc. Cient. Argent., 90: 52-72.

Navas, L. 1922. Insectos nuevos 0 poco conocidos. Mem. R. Accad. Rom. Nuovi Lincei, 6: 1-27.

Navas, L. 1932. Insectos de Argentina y Chile (3° serie). Rev. Soc. Entomol. Argent., 5: 79-86.

Pereira, S. M. & E. D. Da Silva. 1990. Nova espécie de Caenis Stephens, 1835 do sudeste do Brasil

(Ephemeroptera, Caenidae). Boletim do Museu Nacional. Nova Série, Rio de Janeiro, Zoologia

(341): 1-8.

Received 16 Oct 1997; Accepted 7 Dec 1998.

PAN-PACIFIC ENTOMOLOGIST

75(2): 68-72, (1999)

MICROCLIMATES ASSOCIATED WITH

CRYPTOTERMES BREVIS

(ISOPTERA: KALOTERMITIDAE)

IN THE URBAN ENVIRONMENT

R. J. WOODROW AND J. K. GRACE

Department of Entomology,

University of Hawaii at Manoa,

3050 Maile Way, Room 310,

Honolulu, Hawaii 96822

Abstract—Thirty-day studies of structural lumber infested by Cryptotermes brevis revealed that

microclimates in Hawaii were fairly uniform with an overall mean wood-core temperature of

24.33° C. The highest wood-core temperature was 43.31° C and the lowest was 13.94° C. The

mean maximum wood core temperature across all sites was 37.04° C. Temperatures varied widely

over a 24 h period with the greatest diurnal variation being 23.72° C. Thermal gradients between

upper and lower locations of infestations were as high as 8.71° C. Ambient relative humidity

was more variable than temperature with values varying as much as 55% RH during a single

month. Monthly mean RH was as high as 75.09% and ranged from 98.2% to 27.2%.

Key Words.—Insecta, West Indian drywood termite, temperature, climate.

Climate studies on termites have been conducted with the Kalotermitidae (Rust

et al. 1979, Williams 1977, Williams 1976) as well as higher termites (Jones &

Nutting 1989, Greaves 1964). Rust et al. (1979) recorded climatic extremes in

the desert habitats of Incisitermes minor (Hagen) and Incisitermes fruticavus Rust,

and found that daytime temperatures in the desert shrub habitat of these two

species routinely reached lethal levels, often exceeding 60° C in exposed soil.

Drywood termite colonies were present in living shrubs, where temperatures were

considerably lower than those of the surrounding soil, and extended from the

coolest basal portions into the peripheral branches where temperatures were much

higher. Additionally, Williams (1976) reported the stress limits for Cryptotermes

brevis (Walker), and Williams (1977) found similarities between observed mac-

roclimatic values and physiological limits established in the laboratory. To date

however, no published works have documented the microclimate, 1.e., the climate

existing within the microhabitat (gallery systems), of C. brevis.

Insects live in dynamic environments (Geiger 1965) and have various adapta-

tions for dealing with environmental extremes. Recent work with Hodotermes

mossambicus (Hagen) established that termites living in warmer habitats have

higher thermotolerances than those from cooler habitats (Mitchell et al. 1993).

Microclimatic extremes are of particular interest with C. brevis because high tem-

peratures are currently being used to control this termite species (Woodrow &

Grace 1997).

Hawaii represents an ideal setting to study C. brevis biology because it is a

common pest throughout the islands. In addition, the climate is relatively constant

throughout the year. In Honolulu, mean temperatures during the warmest and

coolest months vary, on average, less than 4° C while diurnal variation can be on

the order of 5 to 10°; varying more within a typical day than throughout the entire

1999 WOODROW & GRACE: MICROCLIMATES OF CRYPTOTERMES 69

year (Armstrong 1983). Thus, in contrast with more variable North American

conditions, a small time period can be representative of the climate. In the present

study, thirty-day observations were taken of temperature fluctuations in the warm-

est and coolest portions of infested lumber along with ambient temperature and

relative humidity within residential and commercial structures on the island of

Oahu, Hawaii.

MATERIALS AND METHODS

Four study sites were selected on the basis of accessibility. A thorough visual

inspection was performed to determine the location and extent of C. brevis in-

festations prior to placing thermocouples. Thermocouples were placed in the po-

tentially hottest (highest point facing clear sun in the case of an attic infestation),

or the coolest (lowest) locations within a given infested board. Two 1.1 mm (7/

64 in) dia holes were drilled to the center of the infested lumber, one at the highest

and one at the lowest possible sites. Temperatures within the drilled cores were

recorded with Hobo XT temperature loggers (Onset Computer Corp., Pocasset,

Massachusetts) (accuracy: + 0.75° C), which were pre-equipped with thermistor

probes which were inserted into the cores. Ambient relative humidity was re-

corded with a Hobo-RH (Onset Computer Corp., Pocasset, Massachusetts) (ac-

curacy: + 5%) and ambient temperature with a Hobo-Temp (Onset Computer

Corp., Pocasset, Massachusetts) (accuracy: + 0.5° C) at the highest possible site

on a given board. Data loggers were set to record temperature every 24 minutes

for a period of 30 days.

Microclimate surveys took place within four non-air-conditioned structures on

the island of Oahu: a warehouse at Wheeler Army Air Field, Mililani, Hawaii (7

Mar-6 Apr 1996), two single family dwellings, one at Ulua St. (19 Jan—22 Feb

1996) and one at Aina Koa St. (28 Feb—31 Mar 1997), Honolulu, Hawaii, and a

warehouse on the Pearl Harbor Naval Base (14 Apr—9 May 1997). Sampling

periods are purely arbitrary and based on when the infestations were reported by

cooperating pest control professionals, and subsequently, when permission to do

research was granted by the property owners. In the case of both residences, the

readings were taken in roof-rafters; within an attic crawlspace at Ulua St. resi-

dence and in an exposed eve over the entrance to the house at the Aina Koa St.

The Wheeler AAF warehouse measurements were taken in a wall stud, while the

Pearl Harbor warehouse readings were taken within a hardwood shipping-pallet.

RESULTS

Thirty-day studies of the C. brevis microclimates revealed that habitats in Ha-

wail were fairly uniform with an overall mean wood-core temperature of 24.33° C

(SD = 1.36) (Table 1). The highest wood-core temperature recorded across all

sensors was 43.31° C in the Ulua Street residence, and the lowest wood-core

temperature, 13.94° C, was recorded in the Wheeler AAF warehouse. The mean

maximum wood core temperature across all sites was 37.04° C (SD = 4.47, n = 4).

Ambient relative humidity was more variable than temperature with values

deviating as much as 55% RH during a single month (Table 2). Monthly means

ranged from 56.76% for the Ulua St. residence to 75.09% at the Aina Koa St.

70 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Table 1. Summary temperatures (° C) from four sites on the Island of Oahu.

Ht.2 Dimen.*

Location (m) C/AY (cm) Mean + SEM? Min. Max.

Pearl Harbor hae C pallet 25430 2 0:09 18.84 Gots

Aina Koa St. 3 C 3.8 X 8.6 24.76 + 0.08 16.74 34.39

Aina Koa St. 2 C 3.8 X 8.6 24.38 + 0.10 20.79 31.96

Ulua St. 4 GC 3.8 X 8.6 25.91 + 0.14 16.96 43.31

Ulua St. 3 C 3.8 X 8.6 24.94 + 0.12 16.74 5959

Ulua St. 4 A — 26.45 + 0.13 LF coal 41.77

Wheeler AAF 0.1 C 3.8 X 13.3 22.38 + 0.09 14.99 30.98

Wheeler AAF 5 € 3.8 X 13.3 22.56 + 0.10 14.5 S2te

Wheeler AAF 3. A — 21.57 + 0.08 13.94 29.87

“ Height above grade.

’ Core/Ambient temperature probes.

¢ Dimensions of lumber.

¢Thirty day mean, SEM = standard error of the mean.

— No data collected.

residence, and monthly maximums and minimums ranged from 98.2% to 79.2%

and 43.3% to 27.2%, respectively.

Diurnal temperature variation was measured by determining the maximum and

minimum temperatures for each day. Temperatures varied widely over a 24-h

period (Table 3). The greatest amount of diurnal variation occurred at the Ulua

St. residence, which had a mean difference of 14.70° C and a specific difference

as high as 23.72° C. Figure 1 illustrates a typical day in a C. brevis microhabitat.

In almost all cases, the lowest temperatures of all the probes occurred in the early

morning, with temperatures slowly rising, peaking in the late afternoon and then

gradually decreasing through the night. Typically, the upper core temperature rose

first, followed by either the ambient or lower core locations, which tended to trail

behind the upper core probe. Thermal gradients between upper and lower sites

ranged from a monthly maximum of 8.71° C, 5.60° C and 4.48° C for the Aina

Koa St., Wheeler AAF and Ulua St. sites, respectively.

DISCUSSION

The similarity of wood-core temperatures across all sites could be an indication

that these sites represent typical conditions for this species, at least in Hawaii.

The mean (37.04° C) and absolute maximum wood core temperature (43.31° C)

reported here are similar to the macroclimatic values reported previously by Wil-

liams (1977) for C. brevis and other Cryptotermes species. Williams (1977) re-

Table 2. Summary relative humidity (%) data from one month recordings at three sites on the

island of Oahu.

Location Mean + SEM? Minimum Maximum

Wheeler AAF 63.54 + 0.27 39.90 87.20

Ulua St. 56.76 + 0.30 27.20 79.20

Aina Koa St. 75.09 + 0.31 43.30 98.20

@ Standard error of the mean.

1999 WOODROW & GRACE: MICROCLIMATES OF CRYPTOTERMES 71

Table 3. Wood core diurnal variation (° C) of four sites on the Island of Oahu.

Location Mean + SEM? Maximum Minimum

Wheeler AAF lower core 8.83 + 0.45 13.44 4.22

Wheeler AAF upper core 10.69 + 0.51 15.30 Ceol!

Ulua St. upper core 14.70 + 0.95 PacMeSs a0

Ulua St. lower core 12.49 + 0.77 19.87 B19

Aina Koa St. upper core 8.13 + 0.46 13.91 1.75

Pearl Harbor NB pallet core 10.45 + 0.44 15.02 ZA.

@ Standard error of the mean.

ported an absolute maximum of 46° C and a mean high temperature of 38° C for

this species. Although C. brevis can tolerate short exposures to temperatures in

excess of 43° (Woodrow and Grace 1998) extended exposures to 43° C are po-

tentially lethal to this species (Scheffrahn et al. 1997, Williams 1976) and to the

protozoan symbionts (Williams 1977).

In studies with termites in forest and desert habitats it has been reported that

both mound building and tree inhabiting species live in a media that insulate the

colony against high ambient temperatures during hot summer months (Greaves

1964, Rust et al. 1979). In a drywood termite shrub habitat, Rust et al. (1979)

found that wood temperature lagged behind that of the air. Our observations

indicate that the opposite occurs within structures in the urban environment. This

phenomenon is most likely due to radiant heat from the sun heating up the roof

which then in turn conducts the heat to the underlying wood structure, which then

conducts the heat to the surrounding air (Fig. 1). Thus, living shrubs act as in-

sulators to high ambient air temperatures, whereas in the urban environment, wood

within structures may actually be more a conductor of stressful temperatures than

an insulator in some cases.

28

26 —e— Lower core

24 4 —#- Upper core

a —4— Ambient

O 22 een

© |

s 20

—_

y)

® 18

2.

5

hi 16 4

14 4

12 -

10

So = w BAN (op) [oe] wo os = = = = = ie) ie)

S> &. 225 Woe oe er a ee ee a eee UP

= N wow © G = N = B BS OC W = B DW

ow o a = Ni (ss) oOo ai

Time

Figure 1. Twenty-four hour temperature (° C) profile of ambient and wood core temperatures of

a 3.8 X 13.3 cm wall stud in a warehouse at Wheeler Army Air Field.

72 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Although higher termites regulate their colony temperatures (Greaves 1964),

drywood termites may move within their gallery systems according to where

temperatures are most optimal (Rust et al. 1979, Cabrera and Rust 1996). The

most extreme temperature measured in this study, 43.1° C, as discussed previ-

ously, has been shown to be potentially lethal. Thus, our results suggest that C.

brevis might have either behavioral or physiological mechanisms for dealing with

microclimatic extremes that could have implications on the use of extreme tem-

peratures for the control of this termite species.

ACKNOWLEDGEMENT

We are grateful for the assistance of Robin T. Yamamoto, U.S. Army; Mary-

Ann Oshiro, Victor Peters and Stanley Higa, U.S. Navy; James L. Eschle; and

homeowners, Stanley and Florence Miyashiro and Linda and Christopher Carlson;

Carrie H. M. Tome and Robert J. Oshiro provided technical assistance, and Julian

R. Yates III and Amold Hara commented on the manuscript. This research was

supported by USDA-ARS Specific Cooperative Agreement 58-6615-4-037. This

is Journal Series No. 4366 of the Hawaii Institute of Tropical Agriculture and

Human Resources.

LITERATURE CITED

Armstrong, R. W. (ed.). 1983. Atlas of Hawaii. University of Hawaii.

Cabrera, B. J., & M. K. Rust. 1996. The behavioral responses to light and thermal gradients by the

western drywood termite (Isoptera: Kalotermitidae). Environ. Entomol., 25: 436—445.

Geiger, R. 1965. The climate near the ground. Scripta Technica, Inc., translator. Harvard University

Press, Cambridge, Massachusetts.

Greaves, T. 1964. Temperature studies of termite colonies in living trees. Aust. J. Zool., 12: 250-262.

Jones, S. C. & W. L. Nutting. 1989. Foraging ecology of subterranean termites in the Sonoran desert.

pp. 70-106. Jn Schmidt J. O. (ed.). Special biotic relationships in the arid southwest. University

of New Mexico Press, Albuquerque, New Mexico.

Mitchell, J. D., P H. Hewitt & T. Cd. K Van der Linde. 1993. Critical thermal limits and temperature

tolerance in the harvester termite Hodotermes mossambicus (Hagen). J. Insect Physiol., 39:

523-528.

Rust, M. E., D. A. Reierson & R. H. Scheffrahn. 1979. Comparative habits, host utilization and xeric

adaptations of the southwestern drywood termites, Jncisitermes fruticavus Rust and Incisitermes

minor (Hagen) (Isoptera: Kalotermitidae). Sociobiology, 4: 239-255.

Scheffrahn, R. A., G. A. Wheeler & N-Y Su. 1997. Heat tolerance of structure-infesting drywood

termites (Isoptera: Kalotermitidae) of Florida. Sociobiology, 29: 237-245.

Williams, R. M. 1977. The ecology and physiology of structural wood destroying Isoptera. Mater.

Organismen., 12: 111-140.

Williams, R. M. 1976. Factors limiting the distributions of building damaging dry-wood termites

(Isoptera: Cryptotermes spp.) in Africa. Mater. Organismen., 3: 396-403.

Woodrow R. J. & J. K. Grace. 1997. Cooking termites in the Aloha State: the state of thermal pest

eradication in Hawaii. Pest Control, 65: 57, 61-62.

Woodrow R. J. & J. K. Grace. 1998. Laboratory evaluation of the use of high temperatures to control

Cryptotermes brevis (Isoptera: Kalotermitidae). J. Econ. Entomol., 91: 905-909.

Received 15 Jan 1998; Accepted 27 Aug 1998.

PAN-PACIFIC ENTOMOLOGIST

75(2): 73-81, (1999)

CATALOGUE OF PIMPLINAE (HYMENOPTERA:

ICHNEUMONIDAE) FROM PENINSULAR MALAYSIA

A. B. IDRIS

Department of Zoology,

Faculty of Life Sciences,

University Kebangsaan Malaysia,

43600 Bangi, Selangor,

Malaysia

Abstract—A catalogue of Pimplinae (Hymenoptera: Ichneumonidae) from Peninsular Malaysia

which listing 43 species under three tribes and 14 genera is presented. The genus of Xanthopim-

pla (Tribe: Pimplini), Theronia (Tribe: Theroniini) has the greatest number of species (21). A

total of 14 species were described from Peninsular Malaysia with type specimens deposited in

the U.S. National Museum of Natural History—Washington (4), British Natural History Muse-

um—London (3), V. K. Gupta Collection (3), Berlin Museum—Germany (2), and one each in

Munich—Germany and Gainesville—U.S.A.

Key Words.—Insecta, Ichneumonidae, Pimpline, Malaysia, Peninsular Malaysia, Malaya.

Smith (1858) in describing Pimpla punctator Smith from Sarawak (East Ma-

laysia) started the systematic study of the Pimplinae from Malaysia. It was fol-

lowed by Krieger (1897), Cameron (1903) and Tosquinet (1903) who also covered

the West of Malaysia (Peninsular Malaysia). Since then, a number of species had

been described or recorded from Malaysia (Morley 1913, Cushman 1925, Beeson

& Chatterjee 1935, Townes et al. 1961, Gupta 1961, Baltazar 1961, Momoi 1966,

Townes & Chiu 1970, Kamath & Gupta 1972, Gupta & Tikar 1976, Kusigemati

1985).

The present catalogue lists the tribes, genera and species of Pimplinae recorded

from Peninsular Malaysia. Most of the species listed have notes about the location

at which it was collected or recorded. The locations for the type specimens de-

posited are also indicated (capital letters). Species that have no type or lectotype

specimen from Peninsular Malaysia were referred to type or lectotype specimen

recorded from other countries especially from the Indo-Australian region. Note

that the word ‘Malaya’ is also referred to ‘Peninsular Malaysia’ in this catalogue

because Peninsular Malaysia was called ‘Malaya’ before its independent (1957)

and during early years after independent.

Museums in which type or lectotype material is located or deposited are ab-

breviated as follows:

AMSTERDAM. Afdeling Entomologie, Zoologisch Museum, Universiteit van

Amsterdam, Plantage Middenlaan 64, Amsterdam 1004, The Netherlands.

BERLIN. Museum fiir Naturkunde der Humboldt-Universitat zu Berlin, Berlin,

Invaliderstrasse 43, Germany.

BUDAPEST. Termeszettudomanyi Muzeum Allattara (Zoological Department of

the Hungarian Natural History Museum), 1088 Budapest, Baross-Utca 13, Hun-

gary.

COPENHAGEN. Zoologisk Museum, Universitetsparken 15, DK 2100 Kgben-

havn, Denmark.

74 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

EBERSWALDE. Institut f. Pflanzenschutzforschung Kleinmachnow, Abt. Tax-

onomie der Insekten, DDR-13 Eberswalde-Finow 1, Schickler-strasse 5, DDR

Germany.

GAINESVILLE. American Entomological Institute, 3005 S. W. 56th Avenue,

Gainesville, Florida 32608, U.S.A.

GLASGOW. University Museum, University of Glasgow, Glasgow, Scotland,

U.K.

GUPTA. Collection of Virendra Gupta, Located at the American Entomological

Institute, 3005 S. W. 56th Avenue, Gainesville, Florida 32608, U.S.A.

HONOLULU. Bernice P. Bishop Museum, Department of Entomology, Honolulu,

Hawaii 96819, U.S.A.

LEIDEN. Rijksmuseum van Natuurlijke Historie, Postbus 9517, 2300 RA Leiden,

The Netherlands.

LONDON. Department of Entomology, British Museum (Natural History), Crom-

well Road, London SW7, 5BD, England.

MUNICH. Zoologische Staatssammlung, Miinchhausenstrasse 21, D-8000, Miin-

chen 60, FDR Germany.

OXFORD. Hope Entomological Collections, University Museum, Oxford, OX1

3PW, England.

STOCKHOLM. Naturhistoriska Riksmuseet, Sektionen for Entomologi, 10405

Stockholm, Sweden.

WARSAW. Institut Zoologiczny, Polska Akademia Nauk, ul. Wilcza 64, Warsza-

wa, Poland.

WASHINGTON. U.S. National Museum of Natural History, Smithsonian Insti-

tute, Washington, D.C. 20560, U.S.A.

WUFENG. Entomology Collections, Taiwan Agricultural Research.

Other abbreviations are: M = male, F = female, des = description, fig. = figured,

syn = synonyms listed.

TRIBE: PIMPLINI

Pimplaetus mucronis (Gupta & Tikar)

Pimplaetus mucronis. Gupta & Tikar, 1976. Oriental Ins. Monogr., 1: 67. M,

E key, des. Type: EK Malacca & Perak (BERLIN).

Flavopimpla nigromaculata mangae Betrem

Flavopimpla nigromaculata mangae: Gupta & Tikar, 1976. Oriental Ins. Mon-

ogr. 1: 73, E key, des., fig. Selangor: Kenny Hills.

Type: E, Taiwan (EBERSWALDB), Exeristes gracilis Cushman, 1933. Insecta

Matsumurana, 8: 36.; EK India (WASHINGTON), Calliephialtes bimarginatus

Cushman 1937. Indian Forest Rec. (N. S.) Ent. 3: 141.

Camptotypus rugosus (De Geer)

Trichiothecus ruficeps Cameron, 1903. J. Straits Branch Roy. Asiatic Soc., 39:

137. E des. Type: EK Sarawak: Kuching (LONDON).

Hemipimpla rugosus: Morley, 1914. Revision of the Ichneumonidae in the Brit-

ish Museum, 3: 97. M, E key, syn., des. Perak: Tengar.

Camptotypus rugosus: Gupta & Tikar, 1976. Oriental Ins. Monogr., 1: 185, M,

1999 IDRIS: MALAYSIAN PIMPLINAE 75

E key, syn., des., fig. Malacca: Tengah Geh.; N. Sembilan: Seremban; Kedah:

Gajah Mati, Gurun, Bukit Bintang; Selangor: Bukit Kutu—900 m.

Zaglyptus Indicus nigrithorax Gupta

Zaglyptus indicus nigrithorax Gupta, 1961. Indian J. Ent., 22: 250. FE key, des.

Type: EK Pahang: Lubok Tamang, 3500 ft. WASHINGTON).

Echthromorpha agrestoria notulatoria (Fab.)

Echthromorpha maculipes Cameron, 1905. J. Straits Branch Roy. Asiatic Soc.,

44: 121. [F]. des. Lectotype (designated by Townes, Townes & Gupta 1961): E

Sarawak: Kuching (LONDON). Syn. With laeva by Morley 1913.

Echthromorpha notulatoria: Beeson & Chatterjee, 1935. Indian Forest Rec.

(N.S.) Ent., 1: 159. Biol. Malay Peninsular. Host: Hyblaea puera.

Echthromorpha agrestoria notulatoria: Townes, Townes & Gupta, 1961. Mem.

Amer. Ent. Inst., 1: 39. Lectotype design., syn. Malaya.

Xanthopimpla alternans Krieger

Xanthopimpla alternans: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14;

170. M, E key, syn., des., fig. Kuala Lumpur; Penang I: Sungei Siru, Tanjung

Bungah.

Type: Xanthopimpla alternans Kriger, EF Taiwan, Chiai (BERLIN); Xanthopim-

pla genualata Krieger, Indonesia: West Sumatera (BERLIN).

Xanthopimpla clivulus clivulus Townes & Chiu

Xanthopimpla clivulus clivulus: Townes & Chiu, 1970. Mem. Amer. Ent. Inst.,

14: 161. E key, des., Fig. N. Sembilan: Kuala Klawang, Kampai.

Type: FE, Indonesia (GAINESVILLE).

Xanthopimpla curvimaculata pendleburyi Townes & Chiu

Xanthopimpla curvimaculata pendleburyi: Townes & Chiu, 1970. Mem. Amer.

Ent. Inst. 14: 264. key, des., Type: K Selangor: Kuala Lumpur.

Xanthopimpla decurtata detruncata Krieger

Xanthopimpla decurtata detruncata: Townes & Chiu, 1970. Mem. Amer. Ent.

Inst., 14: 143. M, E n. status, key, des., fig. Pahang (4200 ft); Penang I.

Type: Xanthopimpla detruncata: Krieger, 1914. Arch. E Naturgesch., (A) 80

(6); 39, 115. EK Taiwan (BERLIN).

Xanthopimpla ecaudata Krieger

Xanthopimpla hispida: Krieger, 1914. Arch. E Naturgesch., (A) 80 (6): 24. Key;

(A) 80 (7): 130. FE des., fig. Perak.

Xanthopimpla hispida: Townes, Townes & Gupta, 1961. Mem. Amer. Ent. Ints.,

1: 56. Malaya.

Xanthopimpla hispida: Krieger, 1899. Sitzber. Naturf. Gesell. Leipzig, 1897/

98: 70. E key des., Type: K Perak (BERLIN). Syn. By Townes & Chiu 1970.

Xanthopimpla enderleini Krieger

76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Xanthopimpla enderleini: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14:

118. M, E Lectotype design., key, syn., des., fig. Selangor: Ulu Langat.

Type: Xanthopimpla enderleini: Krieger, 1914. Arch. EK Naturgesch., (A) 80

(6): 35. Indonesia: Sumatera (WARSAW).

Xanthopimpla exigua exigua Krieger

Xanthopimpla exigua exigua: Townes & Chiu, 1970. Mem. Amer. Ent. Inst.,

14: 203. M, E n. status, key, syn., des., fig. Selangor: Ulu Langat, 300—390 m.

Xanthopimpla exigua: Krieger, 1914. Arch. EK Naturgesch., (A) 80 (6): 41, 100.

M. key, des., fig. Type: M, Sarawak: Lundu (BERLIN).

Xanthopimpla flavolineata Cameron

Xanthopimpla flavolineata: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14:

114. M, E key, syn., des., fig. Malaya.

Xanthopimpla emaculata: Kusigemati, 1985. Mem. Kagoshima univ. res. Cen-

ter S. Pacific, 5: 127. M, E Pahang: Cameron Highland.

Type: Xanthopimpla flavolineata: Cameron, 1907. Tijdschr. V. Ent., 50; 48. E

key, des. Indonesia: West Irian (AMSTERDAM).

Xanthopimpla gampsura Krieger

Xanthopimpla gampsura: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14:

56. M, E key, des., fig. Selangor: Beting; Kuala lumpur; Teluk Merbau: Penang:

Ginting.

Type: Xanthopimpla gampsura: Krieger, 1914. Arch. E Naturgesch., (A) 80

(6): 44. Indonesia (BERLIN).

Xanthopimpla glaberrima Roman

Xanthopimpla glaberrima: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14:

183. M, E key, syn., des., fig. Penang I.

Type: Xanthopimpla glaberrina: Roman, 1913. Arkiv for Zool., 8 (15): 22.

Philippines (STOCKHOLM); Xanthopimpla sauteri: Krieger, 1914. Arch. EK Na-

turgesch., (A) 80 (6): 31. Lectotype (designated by Townes, Townes & Gupta

1961), Taiwan (BERLIN).

Xanthopimpla incompleta Krieger

Xanthopimpla incompleta: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14:

269. FE key, des., fig. Selangor: Kuala Lumpur (WASHINGTON).

Type: Xanthopimpla incompleta: Krieger, 1914. Arch. E Naturgesch., (A) 80

(6): 23. Key; (A) 80 (7): 145. E des. Indonesia (STOCKHOLM).

Xanthopimpla jacobsoni jacobsoni Krieger

Xanthopimpla jacobsoni jacobsoni: Townes & Chiu, 1970. Mem. Amer. Ent.

Inst., 14: 172. M, E n. status, key, syn., des., fig. Kuala Lumpur; Penang I.; Perak:

Teluk Anson.

Type: Xanthopimpla jacobsoni: Krieger, 1914. Arch. K Naturgesch., (A) 80 (6):

32. Indonesia: Semarang (BUDAPEST). Host: Eulemma versicolor.

Xanthopimpla konowi Krieger

1999 IDRIS: MALAYSIAN PIMPLINAE i,

Xanthopimpla konowi: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14: 48.

M, E key, syn., des., fig. Malaya. Host: Attacus dohertyi, Saturnia pyretorum and

other Saturnidae.

Type: Xanthopimpla konowi: Krieger, 1899. Sitzber. Naturf. Gesell. Lepzig,

1897/98: 87. E key, des., fig. Japan (BERLIN); Xanthopimpla watsoni: Cameron,

1911. Soc. Ent., 26: 46. M. key, des. Type: M, India: Bengal (LONDON).

Xanthopimpla latice ps liturata Townes & Chiu

Xanthopimpla laticeps liturata: Townes & Chiu, 1970. Mem. Amer. Ent. Inst.,

14: 181. E key, des., fig. Type: K Penang I. (WASHINGTON).

Townes & Chiu 1970. Mem. Amer. Ent. Inst., 14: 372 pp.

Townes, Townes & Gupta 1961. Mem. Amer. Ent. Inst., 1: 532 pp.

Xanthopimpla modesta modesta (Smith) (Gupta 1987); Xanthopimpla modesta

(Smith) (Yu & Horstmann 1997).

Xanthopimpla modesta modesta: Townes & Chiu, 1970. Mem. Amer. Ent. Inst.,

14: 106. M, FE n. status, key, Syn., des., fig. Selangor: Kepong.

Xanthopimpla latebalteata: Cameron, 1903. J. Straits Branch Roy. Asiatic Soc.,

39:137. M. des. Lectotype (designated by Townes, Townes & Gupta 1961): E

Sarawak: Kuching (LONDON). Syn. By Townes & Chiu 1970.

Xanthopimpla naenia Morley

Xanthopimpla naenia: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14: 268.

M, E key, syn., des., fig. Kuala Lumpur.

Type: Xanthopimpla naenia: Morley, 1913. Fauna of British India, Hymenop-

tera, 3: 115. E key. des., India (OXFORD).

Xanthopimpla pedator (Fab.)

Xanthopimpla pedator: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14: 39.

M, E key, syn., des., fig. Penang I.

Type: Ichneumon pedator Fab., 1775. Systema Entomologiae, p. 828. EF des.

India (GLASGOW).

Xanthopimpla pulvinaris Townes & Chiu

Xanthopimpla pulvinaris: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14:

271. M, FE key, des., fig. Kuala Lumpur.

Type: EK Taiwan (WUFENG).

Xanthopimpla punctata (Fab.)

Xanthopimpla punctata: Momoi, 1966. Mushi, 40: 4. M, E Malaya

Xanthopimpla punctata: Kusigemati, 1985. Mem Kagoshima Univ. Res. Center

S. Pacific, 5: 127. M, FE des., Perak.

Type: Xanthopimpla lissonota: Cameron, 1906. J. Straits Branch Roy. Asiatic

Soc., 46: 115. E des., EK Sarawak: Kuching (LONDON). Syn. By Townes, Townes

& Gupta 1961.

Xanthopimpla regina Morley

78 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Xanthopimpla regina: Townes & Chiu, 1970. Mem. Amer. Ent. Inst., 14: 43.

M, E key, des., fig. Kuala Lumpur.

Type: Xanthopimpla regina: Morley, 1913. Fauna of British India, Hymenop-

tera, 3: 118. M, FE key, des. Bangladesh: Sylhet (LONDON).

Xanthopimpla walshae walshae Townes & Chiu

Xanthopimpla walshae walshae: Townes & Chiu, 1970. Mem. Amer. Ent. Inst.,

14: 191. E key, des., fig.). Pahang: Fraser’s Hill, 400 ft.

Type: EK Indonesia: Java (GAINESVILLE).

TRIBE: THERONIINI

Theronia clathrata malayensis Gupta: Yu & Horstmann (1997) put it under Pim-

plini

Theronia (Theronia) clathrata malayensis: Gupta, 1962. Pacific Ins. Monogr.,

4: 50. E key, des., Type: EK Pahang: Fraser’s Hill, 1300 m (LONDON).

Theronia nigrivertex Gupta (in Gupta, 1987); Yu & Horstmann (1997) put it under

Pimplini

Theronia (Theronia) nigrivertex: Gupta, 1962. Pacific Ins. Monogr., 4: 45. E

key, des. Type: EK Perak: Jor Camp, 550 m, Batang Padang (WASHINGTON).

Pahang: Cameron Highlands, Terbukui, 1200 m.

Theronia pseudozebra pseudozebra Gupta

Theronia (Theronia) pseudozebra pseudozebra: Gupta, 1962. Pacific Ins. Mon-

ogr., 4: 21. M, E key. des., Pahang: Fraser’s Hills, 1300 m, Kedah: Kedah Pcak,

1200 m. Penang I.

Type: E Sabah: Sandakan (WASHINGTON).

Theronia zebra zebra (Vollenhoven)

Theronia (Theronia) zebra zebra: Gupta, 1961. Pasific Ins. Monogr., 4: 16. M,

E n. status, key, des., fig. Kuala Lumpur.

Type: Pimpla zebra: Vollenhoven, 1879. Stetiner Ent. Ztg., 40: 147. E des.

Indonesia (LEIDEN).

Nomosphecia pyramida pyramida (Gupta) (in Gupta 1987): Neotheronia pyram-

ida Gupta (Tribe: Pimpilini) Gn Yu & Horstmann 1997)

Theronia (Nomosphecia) pyramida pyramida: Gupta, 1962. Pacific Ins. Mon-

ogr., 4: 80. M. key, des.

Type: M, Kedah: Kedah Peak, 1200 m. (GUPTA). Momoi 1968. Kontyu, 36(2);

182-185; Gupta, 1987. Mem. Amer. Ent. Inst., 41 (Part 2). Pp 598-1210.

Nomosphecia zebroides zebroides (Krieger)

Theronia (Nomosphecia) zebroides zebroides: Gupta, 1962. Pacific Ins. Mon-

ogr., 4: 71. En. comb., key, des. Penang I’ Water Fall garden Host: Eumenes

arcuata.

Type: Theronia zebroides: Krieger, 1906. Ztschr. Syste. Hymen. Dipt., 6: 236.

E des. In key. Type: FE Indonesia: Sumatera (BERLIN).

1999 IDRIS: MALAYSIAN PIMPLINAE 79

Epitheronia tomeus (Gupta)

Theronia (Epitheronia) tomeus: Gupta, 1962. Pacific Ins. Monogr., 4: 96. M,

E key, des. Kedah: Langkawi I.

Type: E Indonesia (LEIDEN).

TRIBE: NEOXORIDINI = POEMENIINI

Eugata nigrita Gupta

Eugalta nigrita: Gupta, 1980. Oriental Ins., 14: 106. M, FE key, des., fig. Type:

K Perak (MUNICH), Selangor: Bukit Kutu, 3500 ft.

Eugalta santoshale Gupta

Eugata santoshae: Gupta, 1980. Orietal Ins., 14: 99. FE key, des. Selangor: Bukit

Kutu, 3500 ft. Type: EK Indonesia (GAINESVILLE).

Achorocephalus nigricollis malayanus (Gupta)

Pseudeugalta nigcollis malayanus: Gupta, 1980. Oriental Ins., 14: 120. M. key,

des. Type: M, Perak: Larut Hills, 3700—4000 ft (GUPTA).

Achorocephalus nigricollis malayanus: Gupta, 1985. Oriental Ins., 19: 324. N.

comb.

TRIBE: RHYSSINI

Lytarmes maculipeunis maculipennis (Smith)

Megarhyssa variegata: Mocsary, 1905. Ann. Mus. Natl. Hungarici, 3: 2. M.

des. Lectotype: Perak (BUDAPEST). Syn. By Kamath & Gupta 1972.

Lytarmes maculipennis variegatus: Townes, Townes & Gupta, 1961. Mem.

Amer. Ent. Inst., 1: 84. Lectotype design., n. comb., syn. Malaya.

Lytarmes maculipennis maculipennis: Kamath & Gupta, 1972. Oriental Ins.

Monogr., 2: 23. M, E key, syn., des., fig. Perak; Penag I Penang Hills, Air Hitam,

300 m; Selangor: Ampang Reservoir; Malacca, Mt. Tengah.

Type: Rgyssa varilineata: Cameron, 1907. Ann. & Mag. Nat. Hist., (7) 20; 15.

FE des. Sarawak: Kuching (LONDON). Syn. By Gupta & Kamath 1972.

Sychnostigma binarium binarium Kamath & Gupta

Sychnostigma binarium binarium: Kamath & Gupta, 1972. Oriental Ins. Mon-

ogr., 2: 135. E key, des. Type; K Pahang: Mt. Tahan, 800-1200 m (LONDON).

Sychnostigma flavopictum flavopictum (Smith)

Epirhyssa flavopicta: Morley, 1913. Revision of the Ichneumonidae in the Brit-

ish Museum, 2; 6. FE key, syn. (in part), des. Penag I; Pahang: Mt Thaun, 2500—

3500 ft.

Epirhyssa flavopicta: Morley, 1914. Ann. & Mag. Nat. Hist. (8) 14: 409. Ma-

laysia (in part).

Sychnostigma flavopictum: Townes, Townes & Gupta, 1961. Mem. Amer. Ent.

Inst., 1: 87. N. comb. Penang I.

Epirhyssa curvimaculata: Cameron, 1907. Ann. & Mag. Nat. Hist., (7) 20; 16.

E des. Lectotype (designated by Kamath & Gupta 1971), K Sarawak: Kuching

(LONDON).

80 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Sychnostigma malayanum Kamath & Gupta

Epirhyssa flavobalteata: Morley, 1913. Revision of the Ichneumonidae in the

British Museum, 2: 7, (in part). K Perak: Maxwells’s Hill.

Epirhyssa flavobalteata: Morley, 1913. Fauna of British India, Hymenoptera,

3: 88 (in part).

Sychnostigma malayanum: Kamath & Gupta, 1972. Oriental Ins. Monogr., 2:

173. E key, des., fig. Perak: Maxwell’s Hill.

Type: Pahang: Cameron Highlands, 1500—1600 m (LONDON).

Sychnostigma nigrobalteatum (Cameron)

Sychnostigma nigrobalteatum: Kamath & Gupta, 1972. Oriental Ins. Monogr.,

2: 190. E key, syn., des. Kedah: Kedah Peak, 1100 m.

Epirhyssa nigrobalteata: Cameron, 1903. J. Straits Branch Roy. Asiatic Soc.,

39; 134. EK des. Lectotype (designated by Townes, Townes & Gupta 1961): E

Sarawak: Kuching (LONDON).

Myllenyxis javensis Kamath & Gupta

Myllenyxis javensis: Kamath & Gupta, 1972. Oriental Ins. Monogr., 2: 212. M.

key, des., fig. Selangor: Kuala Lumpur.

Type: M, Java (GAINESVILLE).

Myllenyxis muelleri (Vollenhoven)

Myllenyxis muelleri: Kamath & Gupta, 1972. Oriental Ins. Monogr., 2: 207. F

key, syn., des., fig. Perak.

Epirhyssa nigerrima: Morley, 1913. Revision of the Ichneumonidae in the Brit-

ish Museum, 2: 6. E key, des. Type: EK Sarawak (LONDON). Syn. By Kamath

& Gupta 1972.

ACKNOWLEDGMENT

I would like to express thank you to Associate Professor Mohamed Salleh

Mohamedsaid for encouragement in publishing this catalogue. This work was

supported by Malaysian Government Research Grant IRPA No. 09-02-02-022.

LITERATURE CITED

Baltazar, C. R. 1961. New generic synonyms in parasitics Hymenoptera. Philippine Journal of Science,

90: 391-395.

Beeson, C. EF C. & S. N. Chatterjee. 1935. On the biology of Ichneumonidae (Hymenoptera). Indian

Forest Records, 1: 151-168.

Cameron, P. 1903. Descriptions of new genera and species of Hymenoptera taken by Mr. Robert

Shelford at Sarawak, Borneo. Journal of the Straits Branch of the Royal Asiatic Society, 39:

89-181.

Cushman, R. A. 1925. H. Sauter’s Formosa—collection: Xanthopimpla (Ichneum. Hym.). Entomolo-

gische Mitteilungen, 14: 41-50.

Gauld, I. D. 1984. An introduction to the Ichneumonidae of Australia. British Museum (Natural

History): 413 pp.

Gupta, V. K. 1987. Catalogue of the Indo-Australian Ichneumonidae. Memoirs of the American En-

tomological Institute, 41 (part 1): 17—237.

Gupta, V. K. 1961. A revision of the Oriental species of the genus Zaglyptus (Hymenoptera: Ichneu-

monidae). Indian Journal of Entomology, 22: 244-257.

Gupta, V. K. & D. T. Tikar. 1976. Ichneumonologia Orientalis or a monographic study of Ichneu-

1999 IDRIS: MALAYSIAN PIMPLINAE 81

monidae of the Oriental Region. Part I. The tribe Pimplini (Hymenoptera: Pimplinae). Oriental

Insects Monograph, 1: 1-313.

Kamath, M.K. & V. K. Gupta. 1972. Ichneumonologia Orientalis. Part II. Subfamily Pimplinae, Tribe

Rhyssini (Hymenoptera: Ichneumonidae). Oriental Insect Monograph. Pp. 1—300.

Krieger, R. 1897. Zwei neue Ichneumonidae von Borkum. Entomologische Nachrichen, 23: 7-10.

Kusigemati, K. 1985. Some Ephialtinae of South East Asia, with descriptions of eleven new species

(Hymenoptera: Ichenumonidae). Memoirs of the Kagoshima University, Research Center for

the South Pacific, 5: 126-150.

Momai, S. 1966. Ichneumonidae (Hymenoptera) collected in paddy fields of the Orient with descrip-

tions of new species. Part I. Subfamilies Ephialtinae, Gelinae, Banchinae, Anomalinae and

Mesochorinae. Mushi, 40: 1—11.

Morley, C. 1913. The Fauna of British India including Ceylon and Burma, Hymenoptera. Volume 3.

Ichneumonidae. London. British Museum, 531 pp.

Townes, H. & S-C. Chiu. 1970. The indo-Australian Species of Xanthopimpla (Ichneumonidae). Mem-

oirs of the American Entomological Institute, 14: 372 pp.

Townes, H. K., M. Townes & V. K. Gupta. 1961. A catalogue and reclassification of the Indo-Aus-

tralian Ichneumonidae. Memoirs of the American Entomological Institute, No. 1. 522 pp.

Yu, D. S. & K. Horstmann. 1997. A catalogue of world Ichneumonidae (Hymenoptera). Memoirs of

the American Entomological Institute, 58 (Part 2): 791-876.

Received 28 Mar 1998; Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(2): 82-93, (1999)

DIMORPHOPALPA, A NEW GENUS OF TORTRICID

MOTHS FROM CENTRAL AND SOUTH AMERICA

(LEPIDOPTERA: TORTRICIDAE: EULIIND

JOHN W. BROWN

Systematic Entomology Laboratory, PSI, Agricultural Research Service,

U.S. Department of Agriculture, % National Museum of Natural History,

Washington, D.C. 20560-0168

Abstract——Dimorpho palpa, NEW GENUS, is described from Central and South America. Five

species are recognized: D. albopunctana, NEW SPECIES, from Costa Rica and Venezuela; D.

striatana, NEW SPECIES, from Costa Rica and Venezuela; D. striatanoides, NEW SPECIES,

from Ecuador and Colombia; D. teutoniana, NEW SPECIES, from Brazil (type species); and D.

xestochalca (Meyrick), NEW COMBINATION, from Colombia. Putative synapomorphies for

species of the new genus include the following: 1) sexually dimorphic labial palpi; those of the

male are moderate in length while those of the female are exceedingly elongate; 2) male genitalia

with a pair of unique, sclerotized, ventrally projecting extensions of the tegumen between the

base of the uncus and base of the gnathos; and 3) short, rounded valvae. Dimorpho palpa appears

to be most closely related to Uncicida Razowski, with which it shares similar processes from

the gnathos and a pair of lateral, rounded structures distally on the caulis of the aedeagus that

represent the point of attachment to the juxta.

Key Words.—Insecta, Lepidoptera, Tortricidae, Tortricinae, Euliini, Dimorphopalpa, Neotropics.

Sexual dimorphism in the tortricid tribe Euliini (Tortricinae) usually is restricted

to slight differences in forewing length, subtle differences in intensity and defi-

nition of forewing pattern, and antennal cilia length. In addition, males of many

genera possess a distinctive foreleg hairpencil (Brown 1990a). During continuing

studies on the phylogeny and taxonomy of Neotropical Euliini (Brown 1989,

1990a, b, 1991a, b, 1998; Brown & Powell 1991), I discovered a small, homo-

geneous group of species that exhibit conspicuous sexual dimorphism in the

length of the labial palpi and a slight difference in forewing shape. Males have

moderate, weakly upturned labial palpi and a moderately broad forewing (length

ca. 2.6 times width). In contrast, females have exceptionally elongate, porrect

labial palpi and a slightly more slender forewing (length ca. 2.7—2.8 times width);

females also have a slightly greater forewing length. The sexes can be associated

by forewing pattern, although that of the male is slightly less defined in some

species, and by the sympatry of males and females of similar phenotype. Dimor-

phopalpa, new genus, is described to accommodate this group of species which

includes ‘“‘Tortrix’’? xestochalca Meyrick and four previously undescribed species.

The description of Dimorphopalpa brings to 80 the number of described Neo-

tropical genera in Euliini (Powell, Razowski & Brown 1995). An additional 50

or so described species that lack meaningful generic assignments also are included

in the tribe (Brown 1989, Powell, Razowski & Brown 1995), and a large number

of undescribed species are present in collections worldwide. The tribe may be the

most speciose and diverse group of Neotropical Tortricidae.

MATERIALS AND METHODS

Material for this study was acquired from the following institutions: National

Museum of Natural History (USNM), Smithsonian Institution, Washington, D.C.,

1999 BROWN: DIMORPHOPALPA NEW TORTRICID GENUS 83

3A 1A42A.~—SCCuP

Figure 1. Wing venation of Dimorphopalpa teutoniana.

United States; The Natural History Museum (BMNH), London, England; Essig

Museum of Entomology (UCB), University of California, Berkeley, California,

United States; Instituto Nacional de Biodiversidad (INBio), Santo Domingo, He-

redia, Costa Rica; and Vitor Becker personal collection, Planaltina, Brazil (VBC).

A total of 98 specimens was examined.

Dissection methodology follows that presented in Brown and Powell (1991).

Forewing measurements were made using an ocular micrometer mounted in a

dissecting microscope. Terminology for wing venation and genitalic structures

follows Horak (1984). Abbreviations and symbols are as follows: FW = forewing;

HW = hindwing; DC = discal cell; nm = number of specimens examined or

measured; ca. = circa (approximately); * = mean.

SYSTEMATICS

Dimorphopalpa J. Brown, NEW GENUS

Type Species.—Dimorphopalpa teutoniana J. Brown, NEW SPECIES.

Head.—Frons smooth, sparse-scaled below mid-eye, rough-scaled above; over-

hanging tuft of scales from vertex. Antennal cilia in male ca. 1.25 times width of

flagellomere; cilia in female ca. 0.1 times width of flagellomere; antenna pale tan

with pale yellow scales. Labial palpus (segments II and III combined) ca. 1.5

times horizontal diameter of compound eye in male, ca. 3.0 times horizontal

diameter of compound eye in female; segment II expanded distally by scaling,

slightly curved; segment III about one-third as long as II, partially exposed. Max-

illary palpus rudimentary. Ocelli moderate to small. Chaetosema present. Probos-

cis present, presumably functional. Thorax: Smooth-scaled, without upraised tufts.

Male without foreleg hairpencil. Forewing (Fig. 1): Length ca. 2.6 times width

in male, 2.7—2.8 times width in female; length of DC 0.64—0.65 FW length; width

84 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

of DC 0.13-0.14 its length; CuA, originates 0.54—0.58 along length of DC; all

veins separate beyond DC; R, to costa, R; to termen; CuP present at margin; M-

stem and chorda absent; costal fold absent in male. Hindwing: Sc+R and Rs

separate; Rs and M, closely approximate at base; M, and CuA, connate; CuP

present. Abdomen: Dorsal pits absent; no modified corethrogyne scaling in fe-

male. Male genitalia: Tegumen short, extremely broad, rounded dorsally, with a

pair of slender, elongate, free, ventrally projecting extensions between base of

uncus and base of gnathos; anterodorsal suture of lateral halves of tegumen ob-

solete. Uncus well developed, variable, stout and straight, deflexed, or flared and

enlarged submesally. Socius moderately large, narrow, pendant, with fine long

scales. Gnathos smooth, highly modified; each arm with a large, lateral, distally

attenuate projection and a subapical lobelike process; mesal junction of arms

weakly sclerotized, short, rounded, upturned. Transtilla a simple, narrow, slightly

arched, non-dentate band. Valva short, broad, rounded apically; sacculus weakly

developed; costa not or only weakly sclerotized. Pulvinus absent. Neither hami

nor subscaphium developed. Juxta an irregular, somewhat triangular plate. Ae-

deagus short, stout; caulis with a pair of rounded lateral processes distally; vesica

with lines of small, beadlike scobination; cornuti absent. Female genitalia: Papil-

lae anales simple. Apophyses posteriores ca. 2 times length of apophyses anter-

iores. Sterigma a weakly sclerotized, ventrally bilobed pocket; ostium at saddle

between lobes. Ductus bursae short, without distinct junction between ductus and

corpus. Corpus bursae simple, elongate; spiculae and signa lacking. Ductus sem-

inalis from corpus bursae near junction of corpus and ductus bursae.

Distribution and Biology.—Dimorphopalpa is known from Costa Rica south

to Bolivia, and east to southeastern Brazil. The early stages are unknown.

Diagnosis.—Species of Dimorphopalpa feature a fawn brown ground color

with a variably developed forewing pattern consisting of one or more diagonal

fasciae, sometimes less defined in the male. Dimorphopalpa can be distinguished

from all other genera in Euliini (except Strophotina Brown) by the conspicuously

dimorphic labial palpi, i.e., moderate in length in the male, extremely elongate in

the female. Among previously described genera, only Strophotina Brown has

sexually dimorphic labial palpi (Brown 1998). It is likely that this character has

arisen independently in the two genera because Strophotina and Dimorphopalpa

have little else in common. Although male secondary sex structures must be used

with caution in defining phylogenetic relationships because they may be evolu-

tionarily more labile than other morphological features, dimorphism in the labial

palpi in Dimorphopalpa apparently is consistent within the genus, and as corrob-

orated by other morphological characters cited below, represents one of several

putative apomorphies for the genus.

Male genitalia of Dimorphopalpa are characterized by a short, broad, rounded

tegumen and short, rounded valvae. Putative synapomorphies for the included

species include a pair of slender, ventrally projecting extensions of the tegumen

between the base of the uncus and the base of the gnathos, and the short, rounded

valva. Female genitalia are moderately uniform within the genus; the sterigma

always consists of a simple, ventrally bilobed pocket. On the basis of characters

of the male genitalia, Dimorphopalpa appears to be most closely related to Un-

cicida Razowski. Putative synapomorphies for the two genera include the elon-

gate, distally attenuate, lateral projection from the gnathos arm and a pair of

1999 BROWN: DIMORPHOPALPA NEW TORTRICID GENUS 85

lateral, rounded structures distally on the caulis of the aedeagus; both genera lack

a male foreleg hairpencil. The female genitalia of Dimorphopalpa are most similar

to those of Bonagota Razowski and Apotomops Powell in the development of the

sterigma as a simple bilobed pocket. However, Dimorphopalpa differs from the

latter two genera in features of the male genitalia, wing venation, dimorphic palpi,

and length of the antennal cilia. The female of Uncicida is unknown, hence no

comparisons of female genitalic structures or sexuai dimorphism can be made.

The ventrally projecting extensions of the tegumen of Dimorphopalpa have

nothing in common with the hami of Chlidanotinae. The latter structures arise at

or just below the base of the uncus near the dorsum of the tegumen, and are free,

digitate, movable processes. In contrast the structures in Dimorphopalpa are ex-

tensions of the tegumen, and are rigid and inflexible.

Etymology.—The generic name refers to the sexual dimorphism in length of

the labial palpi.

KEY TO THE KNOWN SPECIES OF DIMORPHOPALPA

1. FW with distinct black-brown spot or dash near apex of discal cell (Figs.

PM AUIGLS 8 T- Lea het ONE et eRe SN OEE ee te ee wo eR ee OE eee ene! 2

1’. FW lacking distinct black-brown spot or dash near apex of discal cell

IIe SESE ATI Ay) eee ads 2 SA eee NE tt ic Pegi eee eer a Aen 3

2. FW with small white spot adjacent to black-brown spot near the apex of

discal cell (baer 220 ah Ser Rear ea Mite ee eer cele Mie aa ie eco che eee albopunctana

2’. FW with a slender dark dash near apex of the discal cell (Fig. 5)

Pe de EE ete ee ee ee ON. oe ee a eae teutoniana (in part)

3. FW with three distinct parallel diagonal brown lines extending tornad from

MEAT COSE As (ISLES aa Spahr Sm CN Lert ere Tl a ae tt ee Ae Una aa 4

3'. FW with fewer than three diagonal brown lines (Fig. 5) .............. 5

40 UP Wireround-colorerown: [va tic sont ecw cuter tay Ne ies striatanoides

AL RW sround:-¢olorpaletawitwre.. jas gah te tcse. ett dst. striatana

Bie ees Pee ds. oc Tears cee Beate re ee sae Ws eg oe 6

TA ole) cy [een WO OO ler oe abet nage Se Oe Aare UN ee ae 7

6. FW length less than 8.0 mm; known only from southeastern Brazil (Fig. 5)

ee oh ee ee a RE cere, eerie Bete Seren, Ae cin teutoniana (in part)

6’. FW length greater than 9.0 mm; known only from Colombia .. xestochalca

7. FW with at least one distinct diagonal brown line extending tornad from

LICAT TC OS RA tied Rie pik =e, SORE Eko: ay Sree Both acne MS, ca sot ey, teutoniana (in part)

7'. FW lacking distinct diagonal line(s), usually with a small ill-defined

brown blotch near apex of discal cell; unassociated females from Costa

Rica and Bolivia

Dimorphopalpa albopunctana J. Brown, NEW SPECIES

(Figs. 2, 6, 10)

Types.—Holotype, male; data: VENEZUELA. ARAGUA: Rancho Grande, 1100

m, 16-19 Jan 1966, S. & W. Duckworth; deposited in USNM. Paratypes: COSTA

RICA. CARTAGO PROVINCE: Rio Aquiares, 9 km NW Turrialba, nr. Santa Cruz,

1500 m, 12, 15 May 1985 (J. Powell & P. Opler, UCB). PUNTARENAS PROV-

86 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Figures 2-5. Adults of Dimorphopalpa. 2. D. albopunctana, female paratype, Costa Rica; 3. D.

striatana, male paratype, Cost Rica; 4. D. teutoniana, male paratype, Brazil; 5. D. teutoniana, female

paratype, Brazil.

INCE: Monteverde, 1300 m, 1¢, 17—20 May 1985, blacklight (J. Powell & P.

Opler, UCB); Est. La Casona, 1520 m, 1d, 12, Aug 1992, 1d, Jul 1993, 13

Mar 1991 (N. Obando, INBio); Finca Cafrosa, Est. Las Mellizas, P. N. Amistad,

1300 m, 1¢, Oct 1989, 12 Oct 1990, 15, Nov 1990 (M. Ramirez & G. Mora,

INBio). VENEZUELA. ARAGUA: Rancho Grande, 1100 m, 10, 12, 16—23 Oct

1966 (S. S. & W. D. Duckworth, USNM), 1¢, 1-7 Aug 1967 (R. Poole, USNM),

12, 30-31 Mar 1978 (J. Heppner, USNM).

Description.—Male. FW length 6.5—7.0 mm (x = 6.6; n = 5). Head: Frons pale yellow-tan. Labial

palpus pale whitish tan mesally, tan laterally. Thorax: Yellow-tan. Forewing: Pale brown, dorsum

lighter, fawn brown in basal two-thirds; terminal area with diffuse whitish band extending to costa

just before apex; apex of DC with black-brown spot bordered distally by a white spot roughly equal

in size; faint whitish streak from latter white spot curving to costa ca. two-thirds distance from base

to apex. Fringe pale whitish tan. Hindwing: Uniform pale gray-brown. Fringe pale whitish tan. Gen-

italia: As in Fig. 6 (drawn from JWB slide no. 264, Monteverde, Costa Rica; n = 3). Uncus relatively

stout, with short attenuate tip. Gnathos with distal process oblong, lateral process elongate, spinelike,

the processes together forming a somewhat chelate projection. Socius, transtilla, and valva as described

for genus. Aedeagus short with strongly curved, attenuate phallobase.

Female.—FW length 7.5-8.5 mm (x = 8.0; n = 4). As described for male (Fig. 2). Genitalia: As

1999 BROWN: DIMORPHOPALPA NEW TORTRICID GENUS 87

Figures 6-9. Male genitalia of Dimorphopalpa, valvae spread, aedeagus removed. 6. D. albo-

punctana; 7. D. striatana; 8. D. teutoniana; 9. D. striatanoides. Scale bar = 1.0 mm.

88 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

in Fig. 10 (drawn from JWB slide no. 459, Cartago Province, Costa Rica; n = 3). Sterigma a deeply

bilobed pocket ventrally, with lateral edge of each lobe extending nearly to posterior lip of sterigma

as sclerotized line. Corpus and ductus as described for genus.

Diagnosis.—Dimorphopalpa albopunctana can be distinguished superficially

from its congeners by the small black-brown spot and adjacent white spot near

the apex of the forewing discal cell. The male genitalia are most similar to D.

striatana; in D. albopunctana the uncus is relatively straight rather than deflexed

subapically, and the lateral processed of the gnathos are narrow and spinelike

rather than broad and curved (see Figs. 6 and 7). Autapomorphies for D. albo-

punctana include the shape of the gnathos and the deeply bilobed sterigma.

Etymology.—tThe specific epithet refers to the white spot on the forewing.

Dimorphopalpa striatana J. Brown, NEW SPECIES

(Figs, 3, 7, 11)

Types.—Holotype, male; datas VENEZUELA. ARAGUA: Rancho Grande, 1100

m, 21—25 Jan 1966, S. & W. Duckworth; deposited USNM. Paratypes. COSTA

RICA. CARTAGO PROVINCE: Ref. Nac. Fauna Silv. Tapanti, 1250 m, 2d 4,

12, Nov 1991 (G. Mora, INBio); P[arque] N[acional] Tapanti, A. C. Amistad,

1150 m, 16, Jan 1994 (G. Mora, INBio). PUNTARENAS PROVINCE: Est. Biol.

Las Alturas, Coto Brus, 1500 m, 12, Aug 1991 (M. Ramirez, INBio); Finca

Cafrosa, Est. Las Mellizas, PR N. Amistad, 1¢, Oct 1989 (M. Ramirez & G. Mora,

INBio). VENEZUELA. ARAGUA: Rancho Grande, 1100 m, 1¢, 22 ¢, 8-14 Aug

1967 (R. Poole, USNM).

Description Male. FW length 7.5—7.8 mm (x = 7.6; n = 4). Head: Frons pale yellow-tan. Labial

palpus pale whitish tan mesally, tan laterally. Thorax: Pale tan-yellow. Forewing (Fig. 3): Fawn brown

with faint, indistinct, slightly darker mottling; three parallel diagonal brown fasciae extending from

near costa to near tornus; apex with diffuse brown patch along costa. Fringe yellow-tan. Hindwing:

Uniform light gray-brown. Fringe pale whitish tan. Genitalia: As in Fig. 7 (drawn from INBio no.

331353, Cartago Province, Costa Rica; n = 2). Uncus deflexed dorsally in apical one-fourth, with

attenuate apex. Gnathos with subapical process rounded, lobelike; lateral process broad at base, atten-

uating into elongate curved spine. Socius, transtilla, and valva as described for genus. Aedeagus short,

stout, curved basally.

Female.—FW length 8.2-10.0 mm (* = 9.1; n = 4). As described for male except forewing pattern

more well defined. Genitalia: As in Fig. 11 (drawn from USNM slide no. 69506, Rancho Grande,

Venezuela; n = 2). Sterigma simple, a weakly sclerotized, shallow, ventrally bilobed pocket. Ductus

and corpus as described for genus.

Diagnosis.—Dimorphopalpa striatana is superficially most similar to D. stria-

tanoides in the presence of three parallel diagonal lines of the forewing; however,

it has a considerably lighter ground color. The male genitalia can be distinguished

from D. striatanoides by the broader, more strongly curved lateral process of the

gnathos. The female genitalia of D. striatana can be distinguished from its con-

geners by the extremely shallow bilobed process of the sterigma. Autapomorphies

for D. striatana include the shape of the lateral and subapical processes of the

gnathos and the deflexed tip of the uncus.

Etymology.—The specific epithet refers to the fasciae of the forewing.

Dimorphopalpa striatanoides J. Brown, NEW SPECIES

(Figs. 9 and 13)

Types.—Holotype, male; datas ECUADOR. CARCI: Maldonado, 2200 m, 9—

11 Jan 1993, V. Becker; deposited VBC. Paratypes. COLOMBIA. MAGDALENA:

1999 BROWN: DIMORPHOPALPA NEW TORTRICID GENUS 89

San Pedro de la Sierra, Sierra Nevada de Santa Marta, 1500 m, 1d, 21-23 Aug

1973 (BMNH). ECUADOR. CARCI: Maldonado, 2200 m, 13, 22 ¢, 29-11 Jan

1993 (V. Becker, VBC).

Description—Male. FW length 6.0—7.3 mm (* = 6.5; n = 3). Head: Frons pale yellow-tan. Labial

palpus pale whitish tan mesally, tan laterally. Thorax: Pale tan-yellow. Forewing: Light brown with

three parallel diagonal darker brown fasciae extending from near costa to near tornus; apex with diffuse

brown patch along costa. Fringe yellow-tan. Hindwing: Uniform light gray-brown. Fringe pale whitish

tan. Genitalia: As in Fig. 9 (drawn from JWB slide no. 1096, Ecuador; n = 3). Uncus weakly deflexed

dorsally in apical one-fourth, with attenuate apex. Gnathos with subapical process rounded, lobelike;

lateral process broadest at base, weakly curved, attenuating into elongate thorn. Socius, transtilla, and

valva as described for genus. Aedeagus short, stout, only slightly curved basally.

Female——FW length 8.5-8.7 mm (* = 8.6; n = 2). As described for male. Genitalia: As in Fig.

13 (drawn from JWB slide no. 1097, Ecuador; n = 2). Sterigma with a weakly sclerotized, shallow,

ventrally bilobed pocket; a subrectangular, weakly sclerotized area posterad of ostium. Ductus and

corpus as described for genus.

Diagnosis—Dimorphopalpa striatanoides is most similar to D. striatana; the

two are nearly indistinguishable superficially, although the ground color of the

forewing of D. striatanoides is darker than that of D. striatana. Male genitalia

can be distinguished by features described above in the diagnosis of D. striatana.

The female genitalia of D. striatanoides are unique in the genus in the possession

of a subrectangular sclerotized region of the sterigma immediately postered of the

ostium. Autapomorphies for D. striatanoides include the shape of the lateral pro-

cesses of the gnathos and the sclerotized region of the sterigma.

Remarks.—In the specimen from Colombia the lateral arms of the gnathos are

slightly broader than those in specimens from Ecuador. However, in all other

features, both superficial and genitalic, the specimens are nearly identical.

Etymology.—tThe specific epithet refers to the similarity of this species with D.

striatana.

Dimorphopalpa teutoniana J. Brown, NEW SPECIES

(Figs. 4, 5, 8, 12)

Types.—Holotype, male; data: BRAZIL. SANTA CATARINA: Nova Teutonia,

27°11' S, 52°23’ W, 300-500 m, Sep 1963, E Plaumann; deposited in USNM.

Paratypes. BRAZIL. BAHIA: Camaca, 400-700 m, 2¢ 6, 21-30 Sep 1991, 16,

13-14 Apr 1992 (V. Becker, VBC). MINAS GERAIS: Caraga, 1300, 12, 2—4 Jan

1985, 12, 1-2 Apr 1992, 13, 4 Mar 1993, 24 3, 25 Oct 1994 (V. Becker, VBC),

PARANA: Castro, 13, 1898 (E. Jones, BMNH); Curitiba, 920 m, 1d, 10 Feb

1975, 16, Oct 1975 (V. Becker, VBC); Banhado, Quatro Barras, 800 m, 12, 12

Apr 1970, 22 2, 5 Jun 1970, 12, 6 Jun 1970, 12, 29 Aug 1970, 13, 28 Dec

1970, 16, 12, 22 May 1971 (Becker & Laroca, VBC); Marumbi, Morretes, 500

m, 1d, 21 Nov 1970 (Becker & Laroca, VBC). RIO DE JANEIRO: Itatiaia, 1200

m, 246, 25 Jan 1993, 136, 13 May 1996 (V. Becker, VBC); Nova Friburgo, 800

m, 1d, 22 Jan 1993 (V. Becker, VBC). RIO GRANDE DO SUL: Pinherio, 14, 2

Jan 1989 (A. Camargo, VBC). SANTA CATARINA: Nova Teutonia, 27°11’ S,

52°23' W, 300-500 m, 22 2, Oct 1962, 25d, Aug 1963, 366, Sep 1970 (EF

Plaumann, USNM); Bom Jardim da Serra, 1500 m, 5¢.4, 12, 1-4 Oct 1996 (V.

Becker, VBC); Joinville, 500 m, 22 ¢, 3 Jan 1989 (V. Becker, VBC); Neu [Nova]

Bremen, Rio Laeiss, 12, Aug 1931 (KE Hoffmann, BMNH); Séo Joaquim, 1400

m, 1d, 12, 22—24 Jan 1983 (V. Becker, UCB), 24 36, 222, 22-24 Jan 1983 (V.

90 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

NY

Figures 10-15. Female genitalia of Dimorphopalpa. 10. D. albopunctana; 11. D. striatana; 12.

D. teutoniana; 13. D. striatanoides, 14. D. species (Bolivia); 15. D. species (Costa Rica). Scale bar

= 1.0 mm.

1999 BROWN: DIMORPHOPALPA NEW TORTRICID GENUS 91

Becker, USNM), 72 2, 2 Feb 1993, 4446, 12, 25 Oct 1995 (V. Becker, VBC).

SAO PAULO: Campos do Jordao, 1600 m, 12, 4 May 1995 (V. Becker, VBC);

Sao Paulo, 1000 m, 26 46, 29 Jan 1993 (V. Becker, VBC). Unknown State: Agul-

has Negras, NE Cruceiro, 13, (B. V. Ridout, BMNH). “‘Saunders,”’ “Stn. Coll.

1893-134,” 12 (BMNBH).

Description—Male. FW length 7.0—-8.0 mm (x = 7.2; n = 7). Head: Frons pale yellow-tan, whitish

gray above. Labial palpus pale whitish tan mesally, tan laterally. Thorax: Pale yellow-tan. Forewing

(Fig. 4): Pale cream with light fawn brown reticulation throughout; faint, light brown, diagonal fascia

from costa ca. 0.6 distance from base to apex, extending towards tornus; a diffuse, irregular, light

brown fascia from costa ca. 0.6 distance from base to apex, extending toward base; a faint longitudinal

brown fascia from near base, arching gently toward tornus; flattened, triangular patch along costa in

apical region. Fringe off white. Hindwing: Pale gray-brown. Fringe pale whitish tan. Genitalia: As in

Fig. 8 (drawn from USNM slide no. 68609, Nova Teutonia, Brazil; n = 4). Uncus bent near middle

with broad lateral flange; attenuate apically. Gnathos with subapical process a weakly protruding nub;

lateral process highly variable, truncate or slightly bifurcate terminally. Socius, transtilla, and valva

as described for genus. Aedeagus short, stout, only slightly curved basally.

Female—FW length 8.0-10.0 mm (* = 8.7; n = 9). As described for male but forewing pattern

more defined, fasciae considerably darker than ground color; sometimes with distinct black dash near

distal end of DC (Fig. 5). Genitalia: As in Fig. 12 (drawn from JWB slide no. 358, Nova Teutonia,

Brazil; n = 1). Sterigma with a pair of weakly sclerotized, rounded pockets ventrally.

Diagnosis.—Dimorphopalpa teutoniana is superficially most similar to D. xes-

tochalca in its poorly developed forewing pattern. The absence of the abdomen

of the holotype (and only known specimen) of D. xestochalca prevents genitalic

comparisons between the two. However, geographical and elevational differences

between the type localities (Santa Catarina, Brazil, 500 m vs. Mount Tolima,

Colombia, 1800 m) and the difference in forewing length (greater in D. xesto-

chalca) suggest that the two are not conspecific. Male genitalia of D. teutoniana

can be distinguished from other species in the genus by the conspicuous, broad,

lateral flange of the uncus. Female genitalia have the bilobed pocket of the ste-

rigma deeper than in either D. striatana or D. striatanoides, and shallower than

in D. albopunctana.

Remarks.—The genitalia (USNM slide no. 69364) of one of the females from

Nova Teutonia are remarkably dissimilar to those of all other Dimorphopalpa and

appear to be associated incorrectly with the specimen; this specimen is excluded

from the type series.

Etymology.—The specific epithet is derived from the type locality of Nova

Teutonia, Brazil.

Dimorphopalpa xestochalca (Meyrick), NEW COMBINATION

Tortrix xestochalca Meyrick 1926: 248; Clarke 1958: 256 (figure of adult).

‘“‘Eulia’’ xestochalca; Powell, Razowski & Brown 1995: 146.

Type.—Holotype, male; datas COLOMBIA. Tolima Canyon, 5600’ [1805 m],

Nov 1920; deposited in BMNH.

Description.—Male. FW length 9.1 mm (n = 1). Head: Frons pale whitish tan, above with gray-

tipped, pale whitish tan scales. Labial palpus pale whitish yellow mesally, fawn brown laterally.

Thorax: Gray and dingy white. Forewing: Pale cream with light fawn brown reticulation; a pair of

indistinct brown fasciae from costa ca. 0.5 distance from base to apex, one extending distad half way

to termen and the other extending basad to near base; a faint longitudinal brown fascia from near mid-

base, arching gently toward tornus; flattened triangular-shaped patch along costa in apical region.

92 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Fringe pale gray-yellow. Hindwing: Pale gray-brown. Fringe pale whitish-yellow. Genitalia: Single

specimen lacks abdomen.

Female——Unknown.

Diagnosis.—The holotype of D. xestochalca is nearly identical to males of D.

teutoniana in forewing color, pattern, and shape; size and scaling of the labial

palpus; and length of the antennal cilia, providing strong evidence for the inclu-

sion of xestochalca in Dimorphopalpa. The absence of an abdomen prevents

genitalic comparisons. Although it is possible that D. teutoniana is conspecific

with D. xestochalca, it is unlikely based on geography and elevation. A single

specimen from Colombia (BMNH) also could represent D. xestochalca, but its

forewing pattern matches D. striatanoides and not D. xestochalca; hence, it is

included as a paratype of the former. Female genitalia of unassociated specimens

from Bolivia (Fig. 14) and Costa Rica (Fig. 15) are illustrated. One of these may

represent the opposite sex of the holotype of D. xestochalca; however, both have

a considerably larger forewing length than the latter. Until additional material

becomes available from Colombia, these females are treated as Dimorphopalpa

Spp z

ACKNOWLEDGMENT

I thank the following for allowing me to examine material in their care: the

late J. EK G. Clarke (USNM); J. A. Powell (UCB); K. R. Tuck (BMNH); E. Philips

(INBio); and D. H. Janzen, University of Pennsylvania, whose material is depos-

ited at INBio. In particular, I thank V. O. Becker (VBC), Planaltina, Brazil, whose

material represents over half of the specimens examined. I thank Richard Brown,

Mississippi State University; Jerry A. Powell, University of California, Berkeley;

Marianne Horak, CSIRO, Canberra, Australia; Andrew Jensen, USDA, Systematic

Entomology Laboratory, Beltsville, Maryland; David Smith, USDA, Systematic

Entomology Laboratory, Smithsonian Institution, Washington, D.C.; and one

anonymous reviewer for helpful comments and suggestions on the manuscript.

Susan Escher, Front Royal, Virginia, provided the drawings of the genitalia, except

for Fig. 13.

LITERATURE CITED

Brown, J. W. 1989. Generic reassignments for Neotropical tortricid moths (Tortricidae). J. Lepid. Soc.,

43: 313-322.

Brown, J. W. 1990a. Taxonomic distribution and phylogenetic significance of the male foreleg hair-

pencil in the Tortricinae (Lepidoptera: Tortricidae). Entomol. News, 101: 109-116.

Brown, J. W. 1990b. Review of Hynhamia Razowski (Lepidoptera: Tortricidae) and critique of its

phylogenetic position. Entomol. Scand., 21: 321-328.

Brown, J. W. 1991a. Punctapinella, a new genus for three previously known and three new species

from South America (Lepidoptera: Tortricidae). Contrib. Sci., 423: 1-9.

Brown, J. W. 1991b. Systematic revision of Paraptila Meyrick (Tortricidae). J. Lepid. Soc., 44: 257—

212

Brown, J. W. 1998. Description of Strophotina, new genus, from Central and South America (Lepi-

doptera: Tortricidae). Proc. Entomol. Soc. Washington, 100: 43—49.

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

Tortricidae: Euliini). Univ. Calif. Publ. Entomol., 111. 87 pp. + figs.

Clarke, J. EF G. 1958. Catalogue of the type specimens of Microlepidoptera in the British Museum

(Natural History) described by Edward Meyrick. Volume 3. British Museum (Natural History),

London.

1999 BROWN: DIMORPHOPALPA NEW TORTRICID GENUS 93

Horak, M. 1984. Assessment of taxonomically significant structures in Tortricinae (Lep., Tortricidae).

Mitt. Schweiz. Entomol. Gesell., 57: 3-64.

Meyrick, E. 1926. Exotic Microlepidoptera 3: 289-320. Marlborough, England.

Powell, J. A., J. Razowski & J. W. Brown. 1995. Tortricidae: Tortricinae, Chlidanotinae, pp. 138-151.

In Heppner, J. B. (ed.). Atlas of Neotropical Lepidoptera, Checklist. Part II: Hyblaeoidea -

Pyraloidea - Tortricoidea. Assoc. Trop. Lepid., Gainesville, Florida.

Received 14 Apr 1998; accepted 7 Dec 1998.

PAN-PACIFIC ENTOMOLOGIST

75(2): 94-102, (1999)

IMMATURE STAGES OF OXYPORUS JAPONICUS SHARP

(COLEOPTERA: STAPHYLINIDAE: OXYPORINAE),

WITH NOTES ON PATTERNS OF HOST USE

RODNEY S. HANLEY! AND KEN-ICHI SETSUDA?

‘Snow Entomological Museum, Snow Hall,

University of Kansas, Lawrence, Kansas 66045, U.S.A.

?Gifu Prefectural Museum, Oyana, Seki 501-32, Japan

Abstract.—Eggs, larvae, and pupae of Oriental species Oxyporus japonicus Sharp are described

and illustrated based upon field collected and laboratory reared material. Known aspects of the

life history and habits of O. japonicus are also described. Adults are known to feed on mature

basidiocarps of various fungi, including Pleurotus ostreatus (Fries) Kummer, Lampteromyces

japonicus Singer, Armillaria mellea (Vahl ex Fries) Karsten, Panellus serotinus (Fries) Kiihner,

and Pholiota lenta (Fries) Singer. Morphological comparisons of larval instar III are made to O.

stygicus Gravenhorst. Oxyporus japonicus is hypothesized to exhibit a pattern of overall host

selection that is relatively narrow with a well defined subset of preference, which is similar to

numercus New World species of Oxyporus, including O. stygicus.

Key Words.—Insecta, Coleoptera, Staphylinidae, Oxyporinae, Oxyporus japonicus, mycophagy,

fungus feeding, immature stages, behavior.

All members of the staphylinid subfamily Oxyporinae are included in the single

genus Oxyporus Fabr., a primarily holarctic genus with most species-level diver-

sity in the Nearctic and Oriental regions. Species of Oxyporus are obligate in-

habitants of higher fleshy mushrooms (Ashe 1984; Leschen & Allen 1988; Hanley

& Goodrich 1994a, b, 1995), even though they are typically placed within the

staphylinine lineage of staphylinid subfamilies whose members are mostly pred-

atory and use extra-oral digestion (Lawrence & Newton 1982). Adults and larvae

burrow into and feed on the tertiary mycelia, pileus, and stipe tissue of mushrooms

using various modifications of the mandibles, labial palpi, maxillae, and labrum

(Hanley & Goodrich 1995). The fleshy mushrooms that serve as hosts for Oxy-

porus are members of only three orders within the class Hymenomycetes: Agar-

icales (the gilled mushrooms), Boletales (the bolete mushrooms), and Polyporales

(the polypore mushrooms) (Hanley & Goodrich 1995).

Adults of Oxyporus are characterized by a large prognathous head with large

mandibles and the apically expanded terminal segments of the labial palpi (Fig.

1). Larvae have a distinctive trilobed mala and stout mandibles that are deeply

bifid. Both adults and larvae feed by slicing off bits of host fungi and saturating

the fungal chunks with preoral digestive fluid (Newton 1984, Leschen & Allen

1988, Hanley & Goodrich 1995). Contrary to what was once thought, adults and

larvae are not known to use their prominent mouthparts in a predatory manner.

The immatures of only 7 of the 90 described species of Oxyporus are known

(Paulian 1941, McCabe & Teale 1981, Leschen & Allen 1988, Frank 1991, Han-

ley & Gocdrich 1994a, Goodrich & Hanley 1995), and none are known for any

of the Oriental species. This study describes the immature stages and life history

of O. japonicus Sharp, a common Oriental species. Patterns of host use and other

phenomena, which could be of evolutionary or taxonomic significance, are dis-

cussed.

1999

Figure 1.

HANLEY & SETSUDA: OXYPORUS IMMATURES

Oxyporus japonicus Sharp, adult, dorsal habitus.

95

96 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Table 1. Known fungal hosts of Oxyporus japonicus Sharp.

Number of Number of

Host collections specimens taken

Tricholomataceae

Armillaria mellea (Vahl ex Fries) Karsten 2 2

Lampteromyces japonicus Singer! 8 1106*

Panellus serotinus? (Fries) Kiihner 1 8

Pleurotus ostreatus (Fries) Kummer as ay

Cortinariaceae

Pholiota lenta (Fries) Singer 1 2

* Includes adults and immatures.

' Six records reported by Setsuda (1993, 1994a).

2 One record reported by Suzuki (1986).

MATERIALS AND METHODS

Descriptions of the immature stages of O. japonicus are based on 51 eggs, 116

larvae (36 instar I, 52 instar II, and 28 instar TI), and 2 pupae. The chaetotaxy

used in the description of the immatures of O. japonicus is based on the system

used by Hanley & Goodrich (1994a) in the description of the immatures of O.

stygicus Say.

RESULTS

Host Relationships.—Adults of O. japonicus were found on five fungal hosts

from 3 families of fungi (Table 1), but larvae were found only on Lampteromyces

Japonicus Singer.

Behavior.—Females of O. japonicus were found within cylindrical tunnels ex-

tending from an opening on the undersurface of the fungal cap into the center of

the basidiocarp of L. japonicus. These tunnels likely serve as feeding chambers

for both adults and larvae. An enlarged chamber was typically found at the apical

end of each tunnel within the stipe tissue of young mushroom fruiting bodies.

Within this chamber, eggs were usually found covered with fungal frass. Females

remained within their egg chambers after oviposition and repelled conspecific

female adults and other predacious beetles before and after the eggs hatched. This

behavior has been regarded as subsocial (Setsuda 1994b). Males were less fre-

quently collected within the basidiocarps, but were present. Similar behavior has

been reported for some North American species: O. occipitalis Fauvel (Hanley &

Goodrich 1993), O. stygicus (Hanley & Goodrich 1994a, b), and O. major Grav-

enhorst (Goodrich & Hanley 1995). Oxyporus japonicus, however, is the first

species in which subsocial behavior has been adequately quantified.

Development.—The rapid development of species of Oxyporus is well known,

and O. japonicus fits previously reported patterns. In O. stygicus the develop-

mental time from egg to adult was 16—18 days, with 7—10 days in the pupal stage

(Hanley & Goodrich 1994a); in O. major developmental time was even more

rapid (13-15 days) (Goodrich & Hanley 1995). Development of O. japonicus

from egg to pupa required 12-13 days at room temperature (22—24°C), slightly

longer than for O. stygicus and O. major.

1999 HANLEY & SETSUDA: OXYPORUS IMMATURES 97

IMMATURE STAGES OF OXYPORUS JAPONICUS

Description of Egg—Length 1.2—1.7 mm; white, darkening with age; cylin-

drical without distinct sculpture; mouthparts and appendages visible through cho-

rion in more mature eggs.

Description of Larval Instar ITI.—Length 8.8—12.0 mm. Body elongate, gently

curved, parallel-sided, slightly flattened dorsoventrally. White with thoracic and

abdominal terga brown; head dark yellow to brown. Vestiture length variable,

setae simple. Head cylindrical to oval; ecdysial lines distinct, lateral arms forked,

complete from back of head to bases of antennae; six pigmented stemmata in two

vertical rows on each side (Fig. 2); setal arrangement as in Figs. 2—4. Antenna

3-segmented and inserted anterodorsally near ocelli in membranous socket; seg-

ment I elongate, narrowed toward middle, asetose, length 5X width; segment II

trisetose, 0.6 length of segment I, bearing tubercle-like sensory appendage with

distinct basal collar, single narrower, conical sensory appendage also present; seg-

ment ITI 0.6X length of segment II, bearing inner circle of 3 small, subequal setae

at apex, surrounded by outer circle of 3 longer setae (Fig. 5). Labrum fused to

frons with anterior margin serrate; chaetotaxy with labral marginal and labral

lateral rows of 2 setae each (LI, positioned near FI,), labral dorsal row of 1 seta

(Fig. 6). Adoral surface of labrum (epipharynx) with numerous branched micro-

trichia and a large median furrow. Mandibles broad, flat, bifid apically, stout

basally; margins finely serrate with many fine teeth, lateral margin with 2 small

setae, prostheca absent (Fig. 7). Maxilla with cardo triangular, fused to stipes and

mala, with 1 small seta; mala short, stout, trilobed, inner lobe with 2 non-artic-

ulated and 2 articulated spines, middle lobe with 2 non-articulated spines and no

articulated spines, outer lobe with 1 non-articulated and 2 articulated spines dor-

sally (Fig. 8). Maxillary palpus 3 segmented; segment I asetose, about as long as

wide; segment II bisetose, segment III conical, asetose, length 1.3 length of

segment II, minute sensory structures at apex (Fig. 8). Labium of diamond-shaped

submentum and trapezoidal mentum, ligula absent; labial palpus 2 segmented,

directed ventrally; segment I subequal to length of segment II; segment II elongate

and conical with 3 very minute setae at apex; palpigers fused to form ventral

premental sclerite, bearing 1 pair setae, no campaniform sensilla present (Fig. 9).

Thorax. Pronotum transverse, broadly oval, moderately sclerotized; chaetotaxy

with anterior row of 7 setae, discal row with 3 setae, lateral row of 4 setae each,

posterior row of 5 setae (Fig. 10). Mesonotum transverse, moderately sclerotized;

chaetotaxy with anterior row of 8 setae, posterior row of 6 setae, lateral row of

5 setae, membrane with 5 setae (Fig. 11). Metanotum transverse; chaetotaxy sim-

ilar to mesonotum, except anterior row of 7 setae, and membrane with 3 setae

(Fig. 12). Legs long, each similar in size and configuration; chaetotaxy with 14

setae on coxa, 7 setae on trochanter, 7 setae on femur, 8 setae on tibia, 2 setae

on tarsus (Fig. 13). Abdomen. Tergum I transverse, chaetotaxy with anterior and

posterior rows of 4 setae each, lateral row of 3 setae, laterotergite with 3 setae

each, 1 minute marginal seta present (Fig. 14); tergites and sternites of segments

II—VIII similar in setation. Tergite [IX with 4 pairs of setae, 1 pair minute cam-

paniform sensilla on disc (Fig. 15). Urogomphi 2 segmented; basal segment fused

to tergum IX, with 4 setae on apical half; segment II with 1 small ventral seta

98 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

N

‘ or,

Q ° x; \ A. & pig ee BS

WW O°)

Figures 2-9. Oxyporus japonicus Sharp, larval instar HI. 2. Head, dorsal aspect. 3. Head, lateral

aspect. 4. Head, ventral aspect. 5. Antenna, ventral aspect. 6. Labrum, dorsal aspect. 7. Mandible,

dorsal view. 8. Maxilla, dorsal view. 9. Labium, ventral view. Abbreviations: Ed, epicranial dorsal

setae; El, epicranial lateral setae; Fd, frontal dorsal setae; Fl, frontal lateral setae; P, posterior epicranial

suture; Sa, sensory appendages; T, temporal setae; L, lateral setae; V, ventral setae; VI, ventral lateral

setae.

1999 HANLEY & SETSUDA: OXYPORUS IMMATURES 99

M

bb -ae

ght 4

P2 3.

1.0 mm

Figures 10-15. Oxyporus japonicus Sharp, larval instar III. 10. Pronotum. 11. Mesonotum. 12.

Metanotum. 13. Prothoracic leg, anterior aspect. 14. Abdominal tergum I. 15. Abdominal terga IX—

X. Abbreviations: A, anterior setae; L, lateral setae; M, marginal setae; P, posterior setae.

100 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

WI 0'Z

Figures 16-17. Oxyporus japonicus Sharp, pupa. 16. Dorsal aspect. 17. Ventral aspect.

and 2 small apical setae (Fig. 15). Abdominal segment X slightly tapered from

base to apex, setation composed of 8 setae (Fig. 15).

Variation in Larval Instars.—Overall body lengths of each instar are as follows:

instar I, 1.5—2.5 mm; instar II, 3.5—6.0 mm; instar III, 8.5—12.0 mm. Structurally,

the first instar differs from the second and third as follows: antennae much shorter

and more robust, general reduction in setation over entire body, urogomphi and

abdominal segment X much shorter and more robust. The second instar differs

from the third as follows: antennae much shorter and more robust, a short Ed,

seta is present on the dorsal surface of the head, general reduction in the number

of setae on the dorsal surface of the thorax, legs shorter and more robust, uro-

gomphi shorter and more robust.

Description of Pupa.—Length 5.5—10.0 mm, white, exarate; with 12 pairs non-

articulated projections on head, prothorax, and abdominal areas. Head positioned

ventrally, not completely visible in dorsal view, bearing 2 pairs projections above

each eye and no setae on the eyes. Pronotum bearing 2 prominant setae on anterior

angles, and 2 large setae off midline along posterior margin. Abdominal segments

IJ-VIUI bearing elongate projections along lateral margins; segment IX tapered

apically; segment X bearing 2 elongate projections and 2 cylindrical inner lobes

(Figs. 16 and 17).

Material Examined.—JAPAN. Kyoto, Asyf, 15 Oct 1993 and 19 Oct 1993. K.

Setsuda, from Lampteromyces japonicus on dead Fagus crenata (51 eggs, 36 first

1999 HANLEY & SETSUDA: OXYPORUS IMMATURES 101

instar larvae, 52 second instar larvae, and 28 third instar larvae). JAPAN. Kyoto,

Miyama, Asyf, 30 Oct 1995, K. Setsuda, from L. japonicus (2 pupae). Total

number examined, 169 specimens.

Comments.—The mature larva of O. japonicus is similar to that of other de-

scribed species of Oxyporus, including O. vittatus Gravenhorst (Leschen & Allen

1988), O. stygicus (Hanley & Goodrich 1994a), and O. major (Goodrich & Han-

ley, 1995). Larval instar III of O. japonicus can be differentiated from the previous

species through the combination of the following: 2 setae in epicranial dorsal (Ed)

row on head, 5 setae in anterior (A) row on pronotum, 8 setae in anterior (A)

row on mesonotum.

DISCUSSION

Immature Stages.—Larval instar III of O. japonicus differs from O. stygicus

in the following ways:

Oxyporus japonicus Oxyporus stygicus

Lateral head seta L, absent. Lateral head seta L, present.

2 setae in epicranial dorsal (Ed) row

on head.

3 setae in epicranial dorsal (Ed) row

on head.

3 setae in temporal (T) row on head. 4 setae in temporal (T) row on head.

3 setae in lateral (L) row on head. 4 setae in lateral (L) row on head.

5 setae in anterior (A) row on prono-

tum.

8 setae in posterior (P) row on pro-

notum.

8 setae in anterior (A) row on meso-

notum.

3 setae in anterior (A) row on prono-

tum.

3 setae in posterior (P) row on pro-

notum.

5 setae in anterior (A) row on meso-

notum.

Host Relationships —Hanley & Goodrich (1995) reported five patterns of host

usage in New World Oxyporus based primarily on distributions of adults among

available host mushrooms: 1) overall host selection broad (7 or more families)

with a moderately broad subset of preference (majority of specimens are taken

from about half of the total number of genera), 2) overall host selection moder-

ately broad (many genera from 4—6 families) with a relatively narrow subset of

preference (majority of specimens are taken from less than one third of the total

number of genera), 3) overall host selection relatively narrow (few genera from

2-4 families) with a well defined subset of preference (1—2 genera), 4) overall

host selection relatively narrow (few genera from 2—4 families) with no defined

subset of preference, and 5) host selection is species specific. Based on the avail-

able host records, we hypothesize that O. japonicus fits into pattern 3. Adults are

found on few genera of fungi, typically two to four families with a noticeable

preference towards one or two genera. For O. japonicus, 1125 specimens were

collected from 5 genera in 2 families of fungi. The vast majority of those spec-

imens, 1106 (98%), were collected from the genus Lampteromyces. New World

species of Oxyporus that exhibit this pattern are O. guinquemaculatus LeConte,

O. balli Campbell, O. bierigi Campbell, O. lateralis Gravenhorst, O. lawrencei

102 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Campbell, O. mexicanus Fauvel, O. rufipennis LeConte, and O. stygicus (Hanley

& Goodrich 1995).

Few collection records for species of Oxyporus include fungal hosts for larvae.

From the host data available for the genus, larvae appear to be specialized on one

or two species of related fungi within a small portion of the subset of preferences

exhibited by adults. Large numbers of larvae of O. japonicus were collected from

only one species of fungus, which further supports this pattern. The reasons for

this apparent host specificity are unknown; however, Hanley & Goodrich (1995)

hypothesized that mushrooms within an adult’s subset of fungal preferences that

exhibit fleshy-fibrous context. allow more efficient internal feeding while being

fibrous enough to maintain overall shape of the mushroom.

ACKNOWLEDGMENT

We extend thanks to J. S. Ashe, University of Kansas and K.-J. Ahn, Chungnam

National University, Korea for their critical reviews of this manuscript.

LITERATURE CITED

Ashe, J. S. 1984. Major features of the evolution of relationships between gyrophaenine staphylinid

beetles (Coleoptera: Staphylinidae: Aleocharinae) and fresh mushrooms. pp. 227-255. In

Wheeler, Q. & M. Blackwell (eds.). Fungus-insect relationships: perspectives in ecology and

evolution. Columbia University Press, New York.

Frank, J. H. 1991. Staphylinidae (Staphylinoidea). pp. 341—352. Jn Stehr, FE W. (ed.). Immature insects,

Volume 2. Kendall/Hunt Publishing Co., Dubuque, Iowa.

Goodrich, M. A. & R. S. Hanley. 1995. Biology, development and larval characters of Oxyporus

major (Coleoptera: Staphylinidae). Entomol. News, 106: 161-168.

Hanley, R. S. & M. A. Goodrich. 1993. Biology, life history and fungal hosts of Oxyporus occipitalis

(Coleoptera: Staphylinidae), including a descriptive overview of the genus. Proc. Wash. State

Entomol. Soc., 55: 1003-1007.

Hanley, R. S. & M. A. Goodrich. 1994a. Natural history, development and immature stages of Oxy-

porus Sstygicus Say (Coleoptera: Staphylinidae: Oxyporinae). Coleopts. Bull., 48: 213-225.

Hanley, R. S. & M. A. Goodrich. 1994b. The Oxyporinae (Coleoptera: Staphylinidae) of Illinois. J.

Kansas Entomol. Soc., 67: 394—414.

Hanley, R. S. & M. A. Goodrich. 1995. Review of mycophagy, host relationships and behavior in the

Oxyporinae (Coleoptera: Staphylinidae). Coleopts. Bull., 49: 267-280.

Lawrence, J. EF & A. E Newton, Jr. 1982. Evolution and classification of beetles. Ann. Rev. Ecol.

Syst., 13: 261-290.

Leschen, R. A. B. & R. T. Allen. 1988. Immature stages, life histories and feeding mechanisms of

three Oxyporus spp. (Coleoptera: Staphylinidae: Oxyporinae). Coleopts. Bull. 42: 321-333.

McCabe, T. L. & S. A. Teale. 1981. The biology of Oxyporus lateralis Gravenhorst (Staphylinidae).

Coleopts. Bull., 35: 281-285.

Newton, A. FE, Jr. 1984. Mycophagy in Staphylinoidea (Coleoptera). pp. 302-353. In Wheeler, Q. &

M. Blackwell (eds.). Fungus-insect relationships: perspectives in ecology and evolution. Co-

lumbia University Press, New York.

Paulian, R. 1941. Les premiers états des Staphylinoidea (Coleoptera). Etude de morphologie comparée.

Mém. Mus. nat. Hist. (N.S.) Paris, 15: 1-361.

Setsuda, K. 1993. The component and structure of beetle community inhabiting fruit bodies of wood-

rotting fungi. Akitu, suppl., 1: 1-22.

Setsuda, K. 1994a. Beetles obtained from Lampteromyces japonicus (KAMAM.) SING. (Agaricales:

Tricholomataceae). Bull. Gifu Pref. Mus., 15: 13-16.

Setsuda, K. 1994b. Construction of the egg chamber and protection of the eggs by female Oxyporus

japonicus Sharp (Coleoptera: Staphylinidae: Oxyporinae). Jpn. J. Entomol., 62: 803-809.

Suzuki, K. 1986. Insect fauna of the Kasagatake Mountains, Gifu Pref., Central Japan (Coleoptera).

Bull. Gifu Pref. Mus., 7: 33-25.

Received 11 Aug 1998; Accepted 11 Nov 1998.

PAN-PACIFIC ENTOMOLOGIST

75(2): 103-111, (1999)

REDEFINITION OF THE GENUS CHELANOPS GERVAIS

(PSEUDOSCORPIONIDA: CHERNETIDAE)

WILLIAM B. MUCHMORE

Department of Biology, University of Rochester,

Box 270211, Rochester, New York 14627-0211

Abstract—The type material of Chelifer (Chelanops) coecus Gervais, type species of the genus

Chelanops Gervais, is lost, and the diagnostic characters of the genus are uncertain. A neotype

series from near the original type locality in southern Chile has been studied, and new descrip-

tions of the species and genus are presented.

Key Words.—Arachnida, Pseudoscorpionida, Chernetidae, Chelanops, emended diagnosis, Che-

lano ps coecus, neotype designation, redescription.

The genus Chelanops was established originally as a separate section of Chel-

ifer, with Chelifer coecus as type species (Gervais 1849: 13). The description of

C. coecus was very sketchy by modern standards, and the diagnosis of the section

Chelanops was exceedingly brief. TOmO6svary (1882), without comment, placed

Chelanops in the synonymy of Chernes. In a series of papers from 1890 to 1914,

Banks, using a very broad (unwritten) definition of the genus, described 28 species

of Chelanops from North and South America (see Hoff 1947). However, such

careful workers as Tullgren (1907), With (1908), and Ellingsen (1910) did not

recognize the species or genus in the South American fauna. Joseph (1927) made

some observations on the life history of a species which he identified as Chelan-

ops coecus, but he included no taxonomic data.

Beier (1932: 177, 1933: 538) recognized Chelifer (Chelanops) coecus as the

type species of the genus Chelanops, but he evidently did not see the type spec-

imens, as he reported the species as an “‘unsichere Art’? and gave no description

of it. This did not, however, deter him from giving a diagnosis of Chelanops,

based apparently on four other species, namely Chelifer (Trachychernes) rotun-

dimanus Ellingsen, Chelifer (Trachychernes) altimanus Ellingsen, Chelanops chi-

lensis Beier, and Chelanops costaricensis Beier. Also, he did not include coecus,

the type species, in the key to species of Chelanops because of insufficient de-

scription (“‘ungentigenden Beschreibung’’). As a result, Beier’s working concept

of the genus Chelanops was based not on the type species but on several other

species which he believed were congeneric with coecus (as it turns out, two of

the four species he recognized in 1932-33, altimanus and costaricensis, belong

in other genera—see below).

The true nature of Chelifer (Chelanops) coecus Gervais has never been estab-

lished. Beier (1964b: 307) stated that the original material of Gervais’ Chilean

species had been lost. The present whereabouts of the types are indeed unknown;

they are not in the Muséum national d’ Histoire naturelle, Paris, where they might

be expacted (J. Heurtault, in litt.) or in any other museum I have contacted, and

may be presumed lost.

Fortunately, there are available several series of specimens from near the type

locality which Beier identified as Chelanops coecus (1964b: 367—368) and which

generally conform to the description given by Gervais. I have mounted and stud-

104 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

ied a few of these specimens and from them have selected a neotype. On the

basis of this new type and conspecific specimens, I have prepared a redescription

of the species Chelanops coecus and a redefinition of the genus Chelanops.

MATERIALS AND METHODS

The specimens examined here were collected and preserved in alcohol. Many

were dissected, cleared, and mounted on slides for detailed study under a com-

pound microscope.

Some abbreviations are used in the text, as follows: L = length; L/B = ratio,

length/breadth; L/D = ratio, length/depth; T = tactile seta.

All material dealt with in this study is from the collection of the California

Academy of Sciences, San Francisco, California.

SYSTEMATICS

Family Chernetidae Menge

Chernetidae Menge: Muchmore 1982: 101; Harvey 1991: 534 (complete synon-

ymy to 1989); Harvey 1992: 1427.

Genus Chelanops Gervais

Chelifer (Chelanops) Gervais, 1849: 13. Type species: Chelifer (Chelanops) coe-

cus Gervais.

Chelanops Gervais: Beier 1932: 177; Beier 1933: 538; Hoff 1947: 503; Hoff

1949: 455, 460; Beier 1964b: 370; Harvey 1991: 553.

Stigmachernes Beier 1957: 457; Harvey 1991: 634. NEW SYNONYMY (see

below).

Diagnosis (emended).—A representative of the family Chernetidae (see Harvey

1992: 1427). With the characters of the type species, Chelanops coecus, partic-

ularly the following. Well sclerotized and generally dark in color. Carapace with

surface lightly granulate; 2 transverse furrows; 2 faint eyespots; numerous narrow,

clavodentate setae. Abdominal tergites with 25-30 setae. Male genitalia typical

of the family (see Harvey 1992: 1394). Spermathecae of female delicate and easily

broken, but apparently consisting of 2 slender tubes of uniform diameter. Chelic-

eral hand usually with 7 setae, rarely with 8 or 9; flagellum of 4 setae; galea

moderate in size, with several small rami. Palp robust, more so in male than in

female; surfaces finely to moderately granulate; setae clavodentate to acuminate;

each chelal finger with 45-50 marginal teeth and 7—13 external and internal ac-

cessory teeth; venom apparatus present only in movable finger of chela. Tricho-

bothria as shown in Fig. 4: on fixed finger, ist lies proximad of est; on movable

finger st is closer to t than to sb. Legs typical of the family, moderately slender;

tarsus of leg IV with a long, erect tactile seta distad of middle.

Remarks.—In the possession of four setae in the flagellum of the chelicera,

Chelanops is allied with some 30 other genera in the Chernetidae (tribe Hesper-

ochernetini Beier 1932 ?). Of these, it appears most closely related to Stigmacher-

nes Beier (1957), from the Juan Fernandez Islands, Chile. In most particulars the

two genera are very similar, though the spermathecae of Stigmachernes skotts-

bergi Beier, the only known species of Stigmachernes, have not yet been de-

scribed. Stigmachernes, undoubtedly, is a synonym of Chelanops.

1999 MUCHMORE: CHELANOPS REDEFINED 105

Chelanops is similar to Epactiochernes Muchmore (1974b) and Epichernes

Muchmore (in Muchmore & Hentschel 1982) in general habitus, nature of the

spermathecae, and nature and placement of the tactile seta on the tarsus of leg

IV. Chelanops differs most notably from those two genera in body size, placement

of trichobothria est and ist on the fixed chelal finger, and the number of setae on

the hand of the chelicera.

Semeiochernes Beier (1932) seems to be closely related to Chelanops. Beier

(1954) noted this and decided that the holotype of Chelanops costaricensis, one

of his original Chelanops species, is actually the female of Semeiochernes mili-

taris Beier (1932). Mahnert (1987) also mentioned the similarity of the two gen-

era. Though Semeiochernes has unique processes on the palpal chela of the male

and lacks a tactile seta on the tarsus of leg IV, it is much like Chelanops in most

other respects, including the spermathecae and the placement of trichobothria ist

and est.

Chelanops strongly resembles Dinocheirus Chamberlin (1929) (and see Much-

more 1974a) in body size and shape, but it differs fundamentally from that genus

in the nature of the spermathecae and the relative positions of trichobothria ist

and est on the fixed chelal finger.

Neochelanops Beier (1964b: 370) should no longer be considered a subgenus

of Chelanops. It appears unlikely that Chelifer (Chelanops) patagonicus Tullgren,

type species of the subgenus, is congeneric with Chelanops coecus. In addition

to the differences mentioned by Beier, the two species differ in the size and

location of the tactile seta on the tarsus of leg IV: that of coecus is long and erect,

and located just distad of the middle of the tarsus, but that of patagonicus is

small, “‘pseudotactile,’’ and near the end of the segment. The actual status of

Neochelanops must await restudy of the type material of Chelifer (Chelanops)

patagonicus (if available). The other species presently assigned to Neochelanops

also need to be reevaluated.

Chelanops coecus (Gervais)

(Figs. 1-7)

Chelifer (Chelanops) coecus Gervais, 1849: 13. Type locality: southern Chile,

near Calbuco.

Chelifer coecus Gervais: With 1908: 327.

Chelifer (Trachychernes) rotundimanus Ellingsen, 1910: 379 (synonymized by

Beier 1959: 215).

Chelanops (?) coecus (Gervais): Beier 1932: 179.

Chelanops (Chelanops) coecus (Gervais): Beier 1964b: 367; Harvey 1991: 554

(complete synonymy to 1989).

Chelanops chilensis Beier 1932: 178, fig. 186; Beier 1933: 538, fig. 10; Harvey

1991: 554. NEW SYNONYMY.

Redescription of adults—Males and females similar, but palpal chelae of males more robust. Car-

apace and tergites light brown, palps dark reddish brown, other parts tan. Carapace longer than broad;

surface lightly granulate, with 2 shallow, transverse furrows; no eyes; about 200 narrow clavodentate

setae, 6-10 at anterior and 14-18 at posterior margin. Abdominal tergites 1-11 and sternites 4—10

divided; surfaces of tergites transversely reticulated to scaly, of sternites nearly smooth; interscutal

membranes papillose to scaly; pleural membranes longitudinally papillose; most dorsal setae narrow

clavodentate, ventral setae acuminate to denticulate; sternites with numerous “sense spots’’ scattered

106 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Figures 1-7. Chelanops coecus (Gervais). Figure 1. Setae on 2nd and 3rd sternites of allotype

female (spiracular setae omitted). Figure 2. Spermathecae of allotype female. Figure 3. Right palp of

neotype male, dorsal view. Figure 4. Right palp of allotype female, dorsal view. Figure 5. Left chela

of neotype male, lateral view (setae omitted; darkened areoles are underneath). Figure 6. Left chela

of allotype female, lateral view. Figure 7. Leg IV of neotype male (most setae omitted).

over surfaces. Tergal chaetotaxy of neotype (male) 24:24:27:31:31:30:33:31:29:28:T18T:2, others, both

males and females, similar. Sternal chaetotaxy of neotype (male) 37:[6-5]:(3)35(3):(1)17(1):34:43:40:

34:35:31:T15T:2, other males similar. Sternites 2-6 of females with about 45:(3)25(3):(1)20(1):27:35;

setae on sternite 2 are close-set in a triangular group (Fig. 1). Though the tactile setae on tergite 11

and sternite 11 of the neotype are missing from their areoles, these are clearly present in many other

specimens, both male and female. Internal genitalia of male typical of the Chernetidae, moderate in

size and sclerotization. Spermathecae of female delicate and often destroyed in these preparations, but

occasionally seen as 2 slender tubes of uniform diameter (Fig. 2). Chelicera 0.35—0.4 as long as

carapace; hand usually with 7 setae (rarely 8 or 9), is and /s long, acuminate, es short, acuminate, bs

usually acuminate, others denticulate; flagellum of 4 setae, denticulate distally; serrula of about 26

blades; galea of female long, slender, with 5—6 small rami, that of male a little smaller and with shorter

rami. Palp robust, more so in male than in female (Figs. 3 and 4): L/B of trochanter 1.85—2.0, femur

2.85—3.0, patella 2.4—2.65, and chela (without pedicel) 2.05—2.3; L/D of hand (without pedicel) 1.01—

1.15; movable finger 0.97—1.03 X as long as hand. Surfaces finely to moderately granulate; most setae

on medial sides of trochanter, femur, and patella narrow clavodentate, on lateral sides somewhat longer

and more pointed, those on chela quite long, finely denticulate to acuminate; setae on chelal hand

densely set in male, less so in female. Trichobothriotaxy as shown in Figs. 5 and 6; one paratype male

has an extra trichobothrium eb on fixed finger. Each finger with 45-50 cusped marginal teeth and 9—

13 external and 7-11 internal accessory teeth. Venom apparatus well developed only in movable finger,

nodus ramosus between trichobothria t and st. Legs moderately slender: leg IV with L/D of fe-

mur+patella 3.25—-3.55, tibia 4.45—4.8, tarsus 4.45—4.95. Tarsus of leg IV with a long, erect, tactile

seta about 2/3 length of segment from proximal end (Fig. 7).

Tritonymph.—Much like adults but smaller and more robust. Chelicera with 6 setae on hand; fla-

gellum of 4 setae. Palp with L/B of femur 2.65—2.7; patella 2.15—2.25; chela (without pedicel) 2.6;

1999 MUCHMORE: CHELANOPS REDEFINED 107

L/D of hand (without pedicel) 1.4—1.45; movable finger 0.90 as long as hand. Fixed chelal finger with

7 trichobothria, movable finger with 3. Leg IV with L/D of femur+ patella 3.25-3.4; tibia 3.9—4.1;

tarsus 3.7—4.0; tactile seta on tarsus as in adult.

Deutonymph.—Like tritonymph but smaller. Chelicera with 5 setae on hand; flagellum of 4 setae.

Palp with L/B of femur 2.45; patella 2.3; chela (without pedicel) 2.7; L/D of hand (without pedicel)

1.55; movable finger 0.88 as long as hand. Fixed chelal finger with 6 trichobothria, movable finger

with 2. Leg IV with L/D of femur+patella 3.35; tibia 3.75; tarsus 3.6; tactile seta on tarsus as in

adult.

Measurements (mm), Male.—Figures given first for neotype, followed in parentheses by ranges for

4 paratypes. Body L 4.85 (3.94—4.55). Carapace L 1.48 (1.27—1.40). Chelicera L 0.525 (0.48-0.52).

Palp: trochanter 0.83 (0.74—0.83)/0.43 (0.39-0.415); femur 1.41 (1.33—1.37)/0.495 (0.45—0.48); patella

1.41 (1.26—1.36)/0.55 (0.49-0.52); chela (without pedicel) 2.33 (2.11—2.18)/1.06 (0.94—1.06); hand

(without pedicel) 1.22 (1.11-1.15)/1.21 (0.96—1.13); pedicel L 0.19 (0.16—0.18); movable finger L

1.26 (1.11-1.19). Leg I: femur 0.415 (0.42—0.445)/0.30 (0.265—0.29); patella 0.64 (0.605—0.635)/0.265

(0.245—0.255); tibia 0.755 (0.69—0.725)/0.185 (0.17—0.18); tarsus 0.63 (0.57—0.635)/0.13 (0.12—0.125).

Leg IV: femur+patella 1.26 (1.13-1.21)/0.355 (0.325-0.355); tibia 1.07 (0.94—1.03)/0.22 (0.205-—

0.215); tarsus 0.725 (0.69—0.725)/0.155 (0.14—0.155).

Female.—Ranges for 3 paratypes. Body L 5.29-5.66. Carapace L 1.41—1.52. Chelicera L 0.50-—

0.55. Palp: trochanter 0.77—0.83/0.41-0.43; femur 1.33—1.44/0.45-0.495; patella 1.30—1.37/0.495—

0.55; chela(without pedicel) 2.12—2.22/0.85—0.955; hand(without pedicel) 1.17—1.26/0.83—0.895; ped-

icel L 0.18—0.19; movable finger L 1.11—1.19. Leg I: femur 0.40—0.45/0.28-0.30; patella 0.635—0.665/

0.245-0.26; tibia 0.69—0.74/0.17—0.185; tarsus 0.59—0.62/0.125—0.13. Leg IV: femur+patella 1.19-

1.27/0.355—0.38; tibia 0.96—1.04/0.20—0.215; tarsus 0.67—0.725/0.15-0.16.

Tritonymph.—Ranges for 3 paratypes. Body L 3.77—4.13. Carapace L 1.05—1.13. Chelicera L 0.38—

0.40. Palp: trochanter 0.525—0.55/0.28-0.30; femur 0.87—0.95/0.325-0.35; patella 0.805—0.85/0.36—

0.39; chela (without pedicel) 1.41—1.47/0.54—0.56; hand (without pedicel) 0.785—0.815/0.55—0.585;

pedicel L 0.18—0.19; movable finger L 0.70—0.74. Leg IV: femur+ patella 0.83—0.88/0.25-0.27; tibia

0.63—0.68/0.16—0.175; tarsus 0.49—0.50/0.125—0.14.

Deutonymph.—One specimen. Body L 2.84. Carapace L 0.80. Chelicera L 0.31. Palp: trochanter

0.39/0.215; femur 0.60/0.245; patella 0.57/0.25; chela (without pedicel) 1.00/0.37; hand (without ped-

icel) 0.56/0.36; pedicel L 0.075; movable finger L 0.495. Leg IV: femur+ patella 0.62/0.185; tibia

0.47/0.125; tarsus 0.38/0.105.

Types Examined—CHILE: [DE LOS LAGOS], Los Muermos, forest, 19 Jan

1951, E. S. Ross and A. E. Michelbacher, 9 males, 5 females, 3 tritonymphs, in

2 vials; this material was at some time studied by Beier, and a label was inserted,

reading ‘“‘Chelanops rotundimanus (Ell.) det. Beier 5 2”; 5 males, 3 females, 3

tritonymphs have been mounted on slides. One of the mounted males,

(WM7883.01002), is hereby designated the NEOTYPE; the others are paratypes.

Other Material Examined.—CHILE: [BIJO-BIO], Nuble, 18 km E of San Carlos,

24 Dec 1950, Ross and Michelbacher, 1 male, 1 female, 1 deutonymph, in alcohol;

[DE LA ARAUCANIA], 20 km E of Temuco, 8 Jan 1951, Ross and Michelbacher,

1 male, 1 tritonymph, in alcohol; [DE LA ARAUCANIA], 16 km NE of Pucé6n,

12 Jan 1951, Ross and Michelbacher, 1 male, 4 females, 1 deutonymph, mounted

on slides; “‘Osorno Prov.”? [DE LOS LAGOS], valley forest, 18 km W of Purran-

que, 16 Jan 1951, Ross and Michelbacher, 8 males 2 females, 1 tritonymph, in

alcohol.

Chelanops skottsbergi (Beier), NEW COMBINATION

Stigmachernes skottsbergi Beier 1957: 457, fig. 3; Harvey 1991: 634.

Type locality. Juan Fernandez Islands, Chile.

According to Beier’s (1957) description and figure, this species has all of the

important characters of the genus Chelanops as defined above, even though the

108 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

palps are not as robust as those of C. coecus. Because S. skottsbergi is the type

species of Stigmachernes, that genus becomes a synonym of Chelanops.

DISCUSSION

It is worth noting that although the neotype series was labelled “‘Chelanops

rotundimanus (Ell.),’ Beier (1959) synonymized rotundimanus with coecus, and

in 1964b he treated these very specimens as coecus.

The original type locality of Chelifer (Chelanops) coecus, as given by Gervais,

was near Calbuco, De los Lagos, Chile. The neotype series is from Los Muermos,

also in De los Lagos, less than 40 km NW of Calbuco. The type locality of

Chelifer (Trachychernes) rotundimanus is Philipi, De los Lagos, about the same

distance NE of Los Muermos. The other specimens studied here are from Temuco,

Pucon, and Nuble, some 2—300 km farther north. Chelanops coecus has also been

reported from several other places in Chile and in Argentina (Beier 1959, 1962,

1964a, b, c; Cekalovic 1976), but the status of these records is uncertain.

Chelanops costaricensis, described by Beier in 1932 and one of the four species

of Chelanops recognized by him at that time, was later declared to be a synonym

of Semeiochernes militaris Beier (Beier 1954: 139, Harvey 1991: 632).

Altogether, about 70 species of pseudoscorpions have been assigned to the

genus Chelanops at one time or another. Most have been reassigned subsequently

to other genera, so that at the present time Harvey’s Catalogue (1991: 554-555)

lists 9 species in addition to coecus in the nominate subgenus of Chelanops. The

structure of the spermathecae is not known for any of these species; but, on other

grounds, most of them do not appear to belong in Chelanops as redefined above.

Chelanops chilensis Beier 1932, from Villarica, De la Araucania, southern Chile.

Relying on Beier’s descriptions, figures and keys (1932: 178, fig. 186; 1933:

538, fig. 10), it appears certain that this is a synonym of Chelanops coecus. The

major difference, in Beier’s opinion, seems to have been the absence of tactile

setae from the 11th tergite of chilensis compared to rotundimanus (= coecus);

however, as mentioned above, the tactile setae of both the 11th tergites and 11th

sternites may be lost from their areoles in preserved specimens. All other features

of the two taxa appear to be identical, within the limits described above.

Chelanops affinis Banks 1894, from Florida, USA.

This species certainly belongs in another genus, according to observations I

have made on the types (in the Museum of Comparative Zoology, Cambridge,

Massachusetts) and numerous other specimens. It will be treated in detail in a

later publication.

Chelanops altimanus (Ellingsen 1910), from the Virgin Islands, West Indies.

This species has recently been shown to be a representative of the genus Din-

ocheirus (Muchmore 1997).

Chelanops nigrimanus Banks 1902, from the Galapagos Islands, Pacific Ocean.

This inadequately described species was not mentioned by Beier in 1932 or

1933. It was listed, without comment, as Parachernes (Argentochernes) nigri-

manus (Banks) by Chamberlin (1934) and as Parachernes nigrimanus (Banks)

1999 MUCHMORE: CHELANOPS REDEFINED 109

by Beier (1940); these actions have not been noted by any other author, even by

Harvey (1991), who retained the species in Chelanops. In a footnote in his paper

on ““The species of the pseudoscorpion genus Chelanops described by Banks,”

Hoff stated that C. nigrimanus Banks is a “Species incertae sedis; deposition of

type individuals uncertain, perhaps in the National Museum. Dr. Chapin as yet

has not located the type specimens.”’ (1947: 473). This information may also have

been known to Beier, who in 1948 (p. 472) described a species from Costa Rica

as Parachernes (Argentochernes) nigrimanus. Further, Beier (1977, 1978) de-

scribed three species and one subspecies of Parachernes from the Galapagos

Islands, two from Isabela (= Albemarle), the type locality of C. nigrimanus

Banks, without mention of the latter species. Recently, the holotype of Chelanops

nigrimanus has been found by M. S. Harvey in the J. C. Chamberlin Collection

of Pseudoscorpions, now housed in the Entomology Department of the California

Academy of Sciences; it is a female, mounted on a microscope slide by Cham-

berlin (JC-799.01001; CAS Type No. 17500), and is in good condition. I have

examined the specimen and find that it is, as Chamberlin determined, a represen-

tative of the genus Parachernes Chamberlin. Therefore, Parachernes nigrimanus

Beier, 1948 becomes a junior primary homonym of Parachernes nigrimanus

(Banks, 1902); for that Costa Rican species I propose the replacement name,

Parachernes beieri.

The following species require further study before their status can be known:

Chelanops atlanticus Beier 1955a, from Tristan da Cunha, South Atlantic Ocean.

Chelanops insularis Beier 1955b, from the Juan Fernandez Islands, Pacific Ocean

west of Chile.

Chelanops kuscheli Beier 1955b, from the Juan Fernandez Islands, Pacific Ocean

west of Chile.

Chelanops occultus Beier 1964b, from central Chile.

Chelanops pugil Beier 1964a, from San Ambrosio Island, Pacific Ocean west of

Chile.

They appear to differ in various ways: in the presence or absence of tactile

setae on the 11th tergite, the presence or absence and size and location of a tactile

seta on the 4th tarsus, and the relative positions of trichobothria est and ist on

the palpal chela; and, as mentioned, the spermathecae are not known for any of

them. Reexamination of the types, or of material which is certainly conspecific,

is needed to place these species properly.

The status of the several species presently placed in the subgenus Neochelanops

Beier 1964 (Harvey 1991: 555-556) is uncertain. If the descriptions by Tullgren

(1900) and Beier (1964b) can be relied on, the type species Chelifer (Chelanops)

patagonicus Tullgren is not congeneric with Chelifer (Chelanops) coecus Gervais.

Only restudy of the types can resolve these problems.

ACKNOWLEDGMENT

I am grateful to V. EK Lee and C. E. Griswold for the loan of materials from

the California Academy of Sciences. Many helpful suggestions were provided by

the reviewers.

110 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

LITERATURE CITED

Banks, N. 1894. [description of Chelanops affinis Banks, n. sp.] p. 314. Jn Hubbard, H. G. (ed.). The

insect guests of the Florida land tortoise. Insect Life, 6: 302-315.

Banks, N. 1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898-1899. VII. Ento-

mological results. Arachnida. Part I. Proc. Wash. Acad. Sci., 4: 49-70.

Beier, M. 1932. Pseudoscorpionidea II. Subord. C. Cheliferinea. Tierreich, 58: 1-294.

Beier, M. 1933. Revision der Chernetidae (Pseudoscorp.). Zool. Jahrb., Abt. Syst., Okol. Geogr. Tiere,

64: 509-548.

Beier, M. 1940. Die Pseudoscorpionidenfauna der landfernen Inseln. Zool. Jahrb., Abt. Syst., Okol.

Geogr. Tiere, 74: 161-192.

Beier, M. 1948. Phoresie und Phagophilie bei Pseudoscorpionen. Osterr. Zool. Z., 1: 441-497.

Beier, M. 1954. Eine Pseudoscorpioniden-Ausbeute aus Venezuela. Mem. Mus. Civ. Stor. Nat. Verona,

4: 131-142.

Beier, M. 1955a. Pseudoscorpione von Tristan da Cunha. Results Norw. Sci. Exped. Tristan da Cunha

1937-1938, 35: 7-10.

Beier, M. 1955b. Pseudoscorpione von den Juan-Fernandez-Inseln (Arachnida Pseudoscorpionida).

Rev. Chil. Entomol., 4: 205-220.

Beier, M. 1957. Los Insectos de las Islas Juan Fernandez. 37. Die Pseudoscorpioniden-Fauna der Juan-

Fernandez-Inseln. Rev. Chil. Entomol., 5: 451-464.

Beier, M. 1959. Zur Kenntnis der Pseudoscorpioniden-Fauna des Andengebietes. Beitr. Neotrop. Fauna,

1: 185-228.

Beier, M. 1962. Pseudoscorpionidea. pp. 131-137. Jn Delamere Deboutteville, C. & E. Rapoport (eds.).

Biologie de 1’Amérique Australe, Etudes sur la Faune du Sol. Volume 1.

Beier, M. 1964a. Pseudoscorpione von der Insel San Ambrosio. Ann. Naturhist. Mus. Wien, 67: 303—

306.

Beier, M. 1964b. Die Pseudoscorpioniden-Fauna Chiles. Ann. Naturhist. Mus. Wien, 67: 307-375.

Beier, M. 1964c. The zoological results of Gy. Topal’s collectings in south Argentina. 15. Pseudoscor-

pionidea. Ann. Hist.-Nat. Mus. Natl. Hung., 56: 487-500.

Beier, M. 1977. Pseudoscorpionidea. pp. 93-112. Jn Mission Zoologique Belge aux files Galapagos et

en Ecuador (N. et J. Leleup, 1964-1965). Volume 3.

Beier, M. 1978. Pseudoskorpione von den Galapagos-Inseln. Ann. Naturhist. Mus. Wien, 81: 533-

547.

Cekalovic K., T. 1976. Catalogo de los Arachnida: Scorpiones, Pseudoscorpiones, Opiliones, Acari,

Araneae y Solifugae de la XII Regién de Chile, Magallanes incluyendo la Antartica chilena

(Chile). Gayana, Zool., 37: 1-108.

Chamberlin, J. C. 1929. Dinocheirus tenoch, an hitherto undescribed genus and species of false scor-

pion from Mexico (Arachnida—Chelonethida). Pan-Pacific Entomol., 5: 171-173.

Chamberlin, J. C. 1934. Check list of the false scorpions of Oceania. Occas. Pap. Bernice P. Bishop

Mus., 10: 1-14.

Ellingsen, E. 1910. Die Pseudoskorpione des Berliner Museums. Mitt. Zool. Mus. Berl., 4: 355—423.

Gervais, P. 1849. Aracnidos. II. Quelifereos. pp. 10-13. In Gay, C. (ed.). Fauna Chilena. Volume 4.

Hist. Fis. Pol. Chile, Zool.

Harvey, M. S. 1991. Catalogue of the Pseudoscorpionida. Manchester University Press, Manchester,

England.

Harvey, M. S. 1992. The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arach-

nida). Invertebr. Taxon., 6: 1373-1435.

Hoff, C. C. 1947. The species of the pseudoscorpion genus Chelanops described by Banks. Bull. Mus.

Comp. Zool., 98: 471-550.

Hoff, C. C. 1949. The pseudoscorpions of Illinois. Bull. Ill. Nat. Hist. Surv., 24: 413-498.

Joseph, H. C. 1927. Observaciones sobre el Chelanops coecus Gerv. Rev. Chil. Hist. Nat., 31: 53-56.

Mahnert, V. 1987. Neue oder wenig bekannte, vorwiegend mit Insekten vergesellschaftete Pseudoskor-

pione (Arachnida) aus Siidamerika. Mitt. Schweiz. Entomol. Ges., 60: 403-416.

Muchmore, W. B. 1974a. Clarification of the genera Hesperochernes and Dinocheirus (Pseudoscor-

pionida, Chernetidae). J. Arachnol., 2: 25-36.

Muchmore, W. B. 1974b. Pseudoscorpions from Florida. 3. Epactiochernes, a new genus based upon

Chelanops tumidus Banks (Chernetidae). Fla. Entomol., 57: 397-407.

1999 MUCHMORE: CHELANOPS REDEFINED 111

Muchmore, W. B. 1982. Pseudoscorpionida. pp. 96-102. Jn Parker, S. P. (ed.). Synopsis and classifi-

cation of living organisms. Volume 2. McGraw-Hill, New York.

Muchmore, W. B. 1997. On the status of four old species of pseudoscorpions from Puerto Rico and

the Virgin Islands (Pseudoscorpionida: Olpiidae, Chernetidae, Withiidae). Caribb. J. Sci., 33:

269-280.

Muchmore, W. B. & E. Hentschel. 1982. Epichernes aztecus, a new genus and species of pseudo-

scorpion from Mexico (Pseudoscorpionida, Chernetidae). J. Arachnol., 10: 41-45.

Témésvary, O. 1882. Pseudoscorpiones Faunae Hungaricae. A Magyar Fauna Alskorpidéi. Magyar

Tudomanyos Akad. Math. és Természettud. K6zlemények, 18: 135-256.

Tullgren, A. 1900. Two new species of Chelonethi (Pseudoscorpions) from America. Entomol. Tidskr.,

21: 153-157.

Tullgren, A. 1907. Zur Kenntnis aussereuropdischer Chelonethiden des Naturhistorischen Museums in

Hamburg. Mitt. Naturhist. Mus. Hambg., 24: 21-75.

With, C. J. 1908. An account of the South-American Cheliferinae in the collections of the British and

Copenhagen Museums. Trans. Zool. Soc. Lond., 18: 217-340.

Received 29 Jun 1998; Accepted 28 Dec 1998.

PAN-PACIFIC ENTOMOLOGIST

75(2): 112-120, (1999)

A NEW SPECIES OF CAPITONIUS

(HYMENOPTERA: BRACONIDAE) FROM COSTA RICA

WITH REARING RECORDS

LEENDERT-JAN VAN DER ENT! AND ScoTT R. SHAW?

'Berkenlaan 29, 6581 PM Malden, The Netherlands

Department of Renewable Resources, University of Wyoming,

Laramie, Wyoming 82071-3354

Abstract—Capitonius tricolorvalvus, NEW SPECIES is described from Costa Rica. The species

was reared from a stem gall of Cissus verticillata (L.) Nicholson & C. E. Jarvis and from leaf

petioles of Cecropia spp.

Key Words.—Insecta, Braconidae, Cenocoeliini, Capitonius, Costa Rica, rearing records.

In the past 10 years much effort has been made to sample the insect fauna in

Costa Rica. At least 30 Malaise traps have been operating throughout Costa Rica

to collect specimens used to develop the book ““The Hymenoptera of Costa Rica”’

(Hanson & Gauld 1995). At the same time, INBio (Instituto National de Biodiver-

sidad; see Janzen 1991, Gamez & Gauld 1993) started sampling insects by hand

collecting, Malaise traps, light traps and other trapping devices. As a result, hun-

dreds of specimens of Cenocoeliini were collected, of which most belong to un-

described species (van der Ent & Shaw 1998). Most Cenocoeliini were collected

by Malaise traps and, therefore, host records are lacking. In temperate regions,

Cenocoeliini parasitize wood-boring and bark-boring beetle larvae, mostly be-

longing to the families Cerambycidae and (less commonly) Scolytidae and Bu-

prestidae (Saffer 1982). The only published host record for Cenocoeliini in Costa

Rica is that of Capitonius andirae (Saffer), NEW COMBINATION, parasitizing

seed-infesting curculionid beetle larvae (Saffer 1977). Recently an undescribed

species of Cenocoeliini has been reared from a cerambycid larva which lived in

an epiphytic species of Solanaceae (Quesada, INBio, unpublished data). The spe-

cies described below is commonly encountered in Costa Rica and was reared

from a stem gall and from leaf petioles. These are the first records of a species

of Cenocoeliini from these types of plant substrates. It supports the expectation

(van der Ent & Shaw 1998) that neotropical Cenocoeliini will display a wider

variety of host habitat adaptations than their counterparts in temperate regions.

Capitonius can be identified as member of the tribe Cenocoeliini or subfamily

Cenocoeliinae using the keys of van Achterberg (1993), Sharkey (1997) or Shaw

(1995). The tribe Cenocoeliini is a monophyletic group which traditionally has

been placed within the subfamily Helconinae, but more recently was considered

to form the main tribe in a separate subfamily Cenocoeliinae (Achterberg 1994).

Cenocoeliini are easily recognizable by the high insertion of the metasoma on the

propodeum, a unique character among the non-cyclostome Braconidae. Van Ach-

terberg provided generic keys to the world Cenocoeliinae (1994) and to the New

World Cenocoeliinae (1997). The morphological terminology mostly follows that

used by Sharkey and Wharton (1997) with some additional characters defined by

van Achterberg (1994, 1997). Authorship of this new species is attributed to the

senior author (LJE).

1999 VAN DER ENT & SHAW: NEW CAPITONIUS SPECIES 113

Figure 1. Lateral habitus of Capitonius tricolorvalvus.

CAPITONIUS TRICOLORVALVUS ENT, NEW SPECIES

(Figs. 1-10)

Types.—Holotype, female: COSTA RICA, HEREDIA: 3 km S of Puerto Viejo,

OTS, La Selva, 100 m el, Dec 1992, P. Hanson, Malaise trap; deposited: Insect

Museum, University of Wyoming, Laramie, U.S.A. Paratypes: 22 same data as

holoype; 22 same data except Sep 1992; 12 same data except Oct 1992; 19

same data except Nov 1992; 22 same data except Jan—Feb 1993; 62 same data

except Feb—Mar 1993; 32 same data except Feb—Apr 1993, huertos plots; ALA-

JUELA, 22 Est. Biol. San Ram6n, 900 m el, Jul-Aug 1995; 235 same data except

Jul-Aug 1998 (reared); GUANACASTE, 12 N. P. Guanacaste, Est. Pitilla, 9 km

S of Santa Cecilia, 700 m el, Sep 1993; 1d same data except Jun 1994; LIMON,

12° 4km NE of Bribri, 50 m el, Sep—Nov 1989; 12 same data except Dec 1989-—

Mar 1990; 12 same data except Apr—Jun 1990; 12 same data except Jul—Sep

1990; 12 16 km W of Guapiles, 400 m el, Jul-Sep 1990; 1d sector Cocori, 30

km N de Cariari, Finca E. Rojas, 100 m el, Mar 1994, 12 same data except 15

Dec 1994 (hand collected); 136, 12 Teleférico (Aerial Tram), 500 m el, May 1997

(reared); PUNTARENAS, 2° R. FE Golfo Dulce, 3 km SW of Rincoén, 10 m el,

Oct—Dec 1990; 22 same data except Mar—May 1991; 12 same data except Aug

Figure 2. Capitonius tricolorvalvus. Hind wing.

114 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

=

oe fe >

a oe

i

~ f

.

Figure 3-4. Head of Capitonius tricolorvalvus. Figure 3. Frontal cavity. Postero-dorsal view. Fig-

ure 4. Lower face and clypeus. Anterior view.

1991; 12 same data except Sep 1991; 12 same data except Apr 1993; 1¢ Cerro

de Oro, Rio Rincén, 100 m el, 1-15 May 1995; 12 R. E Golfo Dulce, 24 km W

of Piedras Blancas, 200 m el, Dec 1989—Mar 1990; 12 same data except Nov

1991; 12 same data except June—Aug 1991; 12 R. E Golfo Dulce, 5 km W of

Piedras Blancas, 100 m el, Aug—Sep 1991; 22 Peninsula Osa, Rancho Quernado,

Rio Riyitio, 200 m el, Sep—Oct 1992; 12 Peninsula Osa, 8 km S Rio Rincon

Coopemarti, 30 m el, Feb 1991; 12 PN. Corcovado, Est. Sirena, 50 m el, Apr—

Aug 1989; 12 same except Mar—Jun 1991; 12 San Vito de Coto Brus, Las

Cruces, 1200 m el, 9 Jul-7 Aug 1982; 12 SAN JOSE, Res. Biol. Carara, Est.

Biagual, 500 m el, Jan 1990. Paratypes are deposited at the location of the ho-

lotype, at the Museum de Insectos, Universidad de Costa Rica, Ciudad Univer-

sitaria, San Pedro de Montes de Oca, Costa Rica, at INBio, Santo Domingo de

Heredia, Costa Rica and at the Nationaal Natuurhistorisch Museum, Leiden, The

Netherlands.

Description of Holotype Female.—Body length 3.8 mm; forewing length 3.1 mm (Fig. 1).

1999 VAN DER ENT & SHAW: NEW CAPITONIUS SPECIES 115

i

188m

=

Figure 5—6. Sculpture patterns on mesosoma of Capitonius tricolorvalvus. Figure 5. Left lobe

propodeum. Ventral view. Figure 6. Sternaulus. Lateral view.

Head: antenna with 26 flagellomeres; scapus 3.8X longer than its maximum width; first, middle

and penultimate flagellomere 5.5X, 2.0, 1.3X longer than wide respectively; vertex sparcely punctate

and setose, area behind frontal cavity slightly convex, smooth and bare; distance between lateral ocelli

to diameter of lateral ocellus 3.5X, distance between lateral and median ocellus 2.5X, and distance

between lateral ocellus and eye 0.8X; frontal cavity with strong lateral carinae reaching lateral ocelli

(Fig. 3), median carina of frontal cavity protruding anteriorly (Fig. 3); temple in lateral view 0.35

eye width; eye ovoid with straight antero-ventral margin, 1.2 taller than wide; face densely punctate

and setose; clypeus less densely punctate than face, medio-ventrally of clypeus a distinct tooth (Fig.

4); malar space 0.7 eye height; ventral lobe of mandible less protruding than dorsal lobe.

Mesosoma: mesosoma 1.4 longer than high; propleuron moderately punctate and setose, lateral

carina foveate and complete, median carina with foveae decreasing in size posteriorly (Fig. 5); prono-

tum smooth to weakly punctate, dorsal margin with a row of round foveae extending ventrally at

posterior margin; mesopleuron sparcely punctate with a smooth and bare median area, sternaulus with

large rectangular-oval foveae which are ventrally not carinate and absent in anterior 4 of mesopleuron

(Fig. 6), postpectal and epicnemial carina apparent but not distinctly foveate; mesoscutum sparcely

punctate and setacous, 1.1 broader than long, notauli with large and irregular rectangular foveae,

fusing medio-posteriorly on mesoscutum into a longitudinal carina (Fig. 7), lateral carina on dorsal

part mesoscutum strongly protruding medio-anteriorly to each notaulus; scutellar sulcus with 2 large

foveae; scutellum smooth; metapleuron and propodeum with large irregular aerolate sculpturing (Fig.

116 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

Figure 7-8. Sculpture patterns on mesosoma of Capitonius tricolorvalvus. Figure 7. Notauli, in

part. Dorsal view. Figure 8. Propodeum, in part. Dorsal view.

8); hind femur 3.2 longer than its maximum width; hind tibia 5.4 longer than its maximum width,

hind basitarsus 5.5<X longer than its maximum width; lateral spur of hind tibial 0.4< longer than

basitarsus; tarsal claw with a lobe protruding distally in a tooth; inner side hind coxa with a depression

parallel to anterior margin ending ventrally in % of a circular area (Fig. 9), circular area in ventral

view forms basal % of length of ventral area (Fig. 10).

Wings: forewing; pterostigma 2.5 as long as its maximum width, vein M slightly curved, vein m-

cu interstitial (no Rs+Mb) second submarginal cell 0.45 length of pterostigma; hind wing vein

M+CU 2.7 longer than 1M (Fig. 2); vein 1M 1.3 longer than r-m; vein 1M 1.1X longer than cu-a.

Metasoma: metasoma 2.5 X longer than its maximum width; tergum 1 smooth, 1.5X longer than its

apical width, dorsal carinae parallel and apparent in basal half; dorsope and laterope weakly developed;

terga 2 and 3 smooth; tergum 3 medially 1.3 longer than tergum 2; ovipositor length 4.1 mm,

ovipositor sheath 1.3X longer than forewing.

Color: head black, ventral half of clypeus brown, clypeal tooth black, mandible yellow-brown with

dark brown apex, scapus and pedicel light yellow; antenna dull red-brown, darker towards apex;

mesosoma dull red-brown; legs yellow-brown, slightly darker on apex of middle and hind tibia; tarsi

yellow-white with apical 2 tarsomeres darkened; wings clear, pterostigma dark brown; metasoma

dorsally black and ventrally brown, basal 3/5 of ovipositor sheath light red-brown, darker basally,

remainder black with a long (15-20 longer than wide) yellow-white apex.

Variation in Females—Body length 3.2—4.9 mm; forewing length 2.7—3.9 mm; antenna with 23-—

1999 VAN DER ENT & SHAW: NEW CAPITONIUS SPECIES 117

Figure 9-10. Hind coxa of Capitonius tricolorvalvus. Figure 9. Postero-lateral view with anterior

at inner side a depression. Figure 10. Ventral view with endings depression of inner sides forming

circular area.

28 flagellomeres; dorsal row of foveae on pronotum apparent to weakly developed; antero-medial lobe

on mesoscutum smooth to weakly rugose; notaulus and sternaulus clearly defined to weakly defined

by lacking of carinae, sternaulus equally sized to decreasing in size anteriorly; scutellar sulcus with 2

or 4 (in larger specimens) foveae; pterostigma 2.3—2.7X longer than its maximum width; hind wing

vein M+CU 2.4-3.1X length of vein 1M; vein 1M 1.0—1.2X longer than cu-a and 1.1—1.4x longer

than r-m; dorsal carinae on tergum | apparent from base to midlength; ovipositor length 3.2-5.3 mm;

ovipositor sheaths 1.2—1.4X longer than forewing; ventral half of clypeus brown to black; hind tarsus

with last 2—4 tarsomeres darkened.

Description of Males.—Similar to holotype female; body length 4.1—4.7 mm, forewing length 3.3—

3.6 mm; antenna with 25—28 flagellomeres, more slender and longer then in female; first, middle and

penultimate flagellomere 5.5X, 3.5, 2X longer than wide respectively; hind coxa lacking depression

at inner side.

Remarks.——The key to North American and Mexican Cenocoeliini by Saffer

(1982) identifies C. tricolorvalvus at couplet 18, which includes a group of species

with clear wings, mostly red mesosoma, lack of transverse carinae on the pro-

pleuron, regularly punctate face and clypeus, and ovipositor less than 1.5X longer

than forewing. However, these species are distinctly larger (forewing length >5.0

118 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

mm) than C. tricolorvalvus and have 29 or more flagellomeres, a red metasoma,

sometimes a red head, and uniformly colored ovipositor sheaths. Capitonius tri-

colorvalvus can be distinguished from similarly sized and colored Costa Rican

species that are undescribed by the combination of the following characters: vertex

behind frontal cavity smooth and bare, lobes of propleuron lacking carina or

rugose sculpturing, sternaulus not apparent on anterior % of mesopleuron, hind

wing vein M+CU 2.4-3.1X longer than vein 1M, tricolored ovipositor sheaths

(light red-brown, black, yellow-white), and ovipositor sheaths distinctly longer

than forewing length.

Taxonomy.—The generic position of C. tricolorvalvus is somewhat ambiguous.

Capitonius tricolorvalvus best fits into Promachus in the generic key of van Ach-

terberg (1994) because the hind wing vein M+CU is between 2.1X and 4.7X

longer than vein 1M. The New World genus Promachus was later found to be

pre-occupied by a robberfly genus, and also because intermediates between Pro-

machus and the Old World genus Cenocoelius were found in a Costa Rican sample

of Cenocoeliini, van Achterberg (1995) decided to combine both genera using the

older genus name Cenocoelius. Therefore, C. tricolorvalvus fits into the redefined

Cenocoelius in the generic key to the New World Cenocoeliinae by van Achter-

berg (1997). However, several characters of C. tricolorvalvus other than the size

of hind wing vein M+CU relative to 1M do not fit into the genus description of

Cenocoelius including Promachus. In C. tricolorvalvus the notauli fuse medi-

oposteriorly, not posteriorly as in Cenocoelius; the sternaulus is incomplete versus

complete; hind wing vein 1M is equal to slightly longer than r-m and cu-a versus

shorter to sometimes equal; and the depression at the inner side of the hind coxa

ends about the middle of the coxa versus distad as in several (perhaps all) formerly

Promachus species. These characters do fit into the genus description of Capi-

tonius but described species of Capitonius (van Achterberg, 1994) are larger in

size than C. tricolorvalvus and have a concave vertex behind the frontal cavity.

Recently, we found a morphospecies of Cenocoeliini from Costa Rica with spec-

imens ranging in size of hind wing vein M+CU relative to 1M including both

Capitonius and Cenocoelius character states sensu van Achterberg (1997). There-

fore, the size of hind wing vein M+CU relative to 1M should not be used solely

to distinguish between Capitonius and Cenocoelius. When the size is 2.1 or

less, the wasp likely belongs to Capitonius but with a higher size up to at least

4.0X it could belong to either of the genera and other characters should be taken

in consideration. Because the generic position of C. tricolorvalvus is ambiguous

at present, it was decided to take a parsimonious approach and place it in Capi-

tonius based on a majority of characters in the generic descriptions of van Ach-

terberg (1994).

Distribution—Capitontius tricolorvalvus occurs in Costa Rica from sea-level up

to 1200 m elevation. They are most frequently encountered in tropical lowlands

with moist and wet rain forest types below 500 m altitude. To date, they have

not been collected in the tropical dry forest in NW Costa Rica.

Biology.—A male and a female C. tricolorvalvus have been reared from a stem

gall of the vine Cissus verticillata (L.) Nicholson and C. E. Jarvis, found at

Teleférico (aerial tram). The stem gall was hand collected at ground level and

reared at the University of Costa Rica, San Pedro. It was probably formed by

Cecidomyiidae and several cecidomyiid parasitoids were reared from it. Such stem

1999 VAN DER ENT & SHAW: NEW CAPITONIUS SPECIES 119

galls are often infested by beetles (a more likely host for Cenocoeliini). No traces

were found of which beetle C. tricolorvalvus had parasitized (P. E. Hanson, per-

sonal communication).

Two males C. tricolorvalvus were reared by the senior author from leaf petioles

of Cecropia spp. at the biological field station San Ramon. Also reared from these

leaf petioles were numerous Scolytidae (many species), several Curculionidae (4

spp. of zygopine weevils: Lechriops disparilis Champion, L. rufomaculatus

Champion, 2 undescribed spp. of Pseudolechriops; Hespenheide, personal com-

munication) and the leaf petioles contained a few larval Cerambycidae (probably

a Lasiolepturges sp.; Hespenheide, personal communication). All adult scolytids

were small (<3 mm) and, therefore, unlikely to be attacked by C. tricolorvalvus.

Moreover, most scolytids emerged a few weeks earlier from the leaf petioles than

the parasitoids, the zygopine weevils and the cerambycids. Most likely, based on

size and weight comparisons, C. tricolorvalvus attacked the cerambycid species

inside the leaf petiole of Cecropia spp.

Etymology.—The species was named for its apparent tricolored ovipositor

sheaths.

ACKNOWLEDGMENT

Paul Hanson from the Univerdad de Costa Rica, San Pedro, provided most of

the specimens. Other specimens were (co)-collected by E. Araya, C. Godoy, E.

Quirros, E. Rojas, R. Zuniga and a few other (para)taxonomists from INBio.

Teresa Williams of the Western Research Institute assisted in making SESM pho-

tos. A. Valerio, J. A. Lockwood and two anonymous reviewers made critical

comments on earlier versions of the manuscript.

LITERATURE CITED

Achterberg, C. van. 1993. Illustrated key to the subfamilies of Braconidae (Hymenoptera: Ichneu-

monoidea). Zool. Verh. Leiden, 282: 1-189.

Achterberg, C. van. 1994. Generic revision of the subfamily Cenocoeliinae Szépligeti (Hymenoptera:

Braconidae). Zool. Verh. Leiden, 292: 1—52.

Achterberg, C. van. 1995. New combinations of names for Palaearctic Braconidae (Hymenoptera).

Zool. Med. Leiden, 69: 131-138.

Achterberg, C. van. 1997. Subfamily Cenocoeliinae. pp. 185-192. Jn Wharton, R. A., P. M. Marsh &

M. J. Sharkey (eds.). Manual of the New World genera of the family Braconidae (Hymenoptera).

Special publication of the International Society of Hymenopterists. Number 1. Allen Press,

Lawrence, Kansas.

Ent, L. J. van der & S. R. Shaw. 1998. Species richness of Costa Rican Cenocoeliini (Hymenoptera:

Braconidae): a latitudinal and altitudinal search for anomalous diversity. J. Hym. Res., 7: 15-24.

Gamez, R. & I. D. Gauld. 1993. Costa Rica: an innovative approach to the study of tropical biodi-

versity. pp. 329-336. In LaSalle, J. & I. D. Gauld (eds.). Hymenoptera and biodiversity. CAB

International, Wallingford, Oxon, U. K.

Hanson, P. E. & I. D. Gauld (eds.). 1995. The Hymenoptera of Costa Rica. The Natural History

Museum, London. Oxford University Press.

Janzen, D. H. 1991. How to save tropical biodiversity. Americ. Entom., 37: 157-171.

Saffer, B. 1977. A new species of Cenocoelius from Costa Rica (Hymenoptera: Braconidae). Proc.

Entomol. Soc. Wash., 79: 593-596.

Saffer, B. 1982. A systematic revision of the genus Cenocoelius (Hymenoptera: Braconidae) in North

America including Mexico. Polski Pismo Ent., 52: 73-167.

Sharkey, M. J. 1997. Key to the New World subfamilies of the family Braconidae. pp. 39-63. In

Wharton, R. A., PR M. Marsh & M. J. Sharkey (eds.). Manual of the New World genera of the

120 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(2)

family Braconidae (Hymenoptera). Special publication of the International Society of Hymen-

opterists. Number 1. Allen Press, Lawrence, Kansas.

Sharkey, M. J. & R. A. Wharton. 1997. Morphology and terminology. pp 19-38. Jn Wharton, R. A.,

P.M. Marsh & M. J. Sharkey (eds.). Manual of the New World genera of the family Braconidae

(Hymenoptera). Special publication of the International Society of Hymenopterists. Number 1.

Allen Press, Lawrence, Kansas.

Shaw, S. R. 1995. Braconidae. pp. 431—463. Jn Hanson, P. E. & I. D. Gauld (eds.). The Hymenoptera

of Costa Rica. The Natural History Museum, London. Oxford University Press.

Received 28 May 1998; Accepted 28 Dec 1998.

PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for

discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English,

but foreign language summaries are permitted. Manuscnpts not meeting the format guidelines may be returned. Please maintain a copy

of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing.

Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in,

nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF

ANY FIGURES TO THE 8.5 X 11 IN PAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text pages

(pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List

the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. The title

must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words: use tive

to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.

Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases

underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet

1988, Dorly et al. 1989), (Burton in press) and (R. FE Tray, personal communication). For multiple papers by the same author use:

(Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith

1987: table 3).

Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that

volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are

applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED

(INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author

of each species after its first mention.

Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these

format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before

submitting manuscripts for which these formats are applicable.

Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. F Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide

some answers? pp. 233—238. Jn Holman, J., J. Pelikan, A. G. F Dixon & L. Weismann (eds.). Population structure, genetics and

taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB

Academic Publishing, The Hague, The Netherlands.

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.

Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label

letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. Authors

are strongly encouraged to provide illustrations no larger than 8.5 X 11 in for easy handling. Number figures in the order presented.

Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of

the manuscript, and (5) journal. FFGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures.

Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued on

additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter.

Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited

section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156—-161.). A short

acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of

the note with the affiliation underscored.

Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for

pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include

reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page

charge fee notice with acknowledgment of initial receipt of manuscripts.

THE PAN-PACIFIC ENTOMOLOGIST

Volume 75 April 1999 Number 2

Contents

ALBA-TERCEDOR, J. & S. MOSQUERA—Caenis chamie, a new species from Colombia

(Ephemeroptera: Caenidae)

WOODROW, R. J. & J. K. GRACE—Microclimates associated with Cryptotermes brevis (Isop-

tera: Kalotermitidae) in the urban environment

IDRIS, A. B.—Catalogue of Pimplinae (Hymenoptera: Ichneumonidae) from Peninsular Ma-

laysia

BROWN, J. W.—Dimorphopalpa, a new genus of Tortricid moths from Central and South

America (Lepidoptera: Tortricidae: Euliin1)

HANLEY, R. S. & K.-I. SETSUDA—Immature stages of Oxyporus japonicus Sharp (Coleop-

tera: Staphylinidae: Oxyporinae), with notes on patterns of host use

MUCHMORE, W. B.—Redefinition of the genus Chelanops Gervais (Pseudoscorpionida: Cher-

netidae)

ENT, L.-J. VAN DER & S. R. SHAW—A new species of Capitonius (Hymenoptera: Bracon-

idae) from Costa Rica with rearing records

61

68

73

82

94

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 75 July 1999 Number 3

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

EDITORIAL BOARD

R. V. Dowell, Editor R. M. Bohart

Jeffery Y. Honda, Associate Editor J. T. Doyen

R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.

Julieta EF Parinas, Treasurer Warren E. Savary

Published quarterly in January, April, July, and October with Society Proceed-

ings usually appearing in the October issue. All communications regarding non-

receipt of numbers should be addressed to: Vincent EK Lee, Managing Secretary;

and financial communications should be addressed to: Julieta EK Parinas, Treasurer;

at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-

emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addressed to: William Hamersky, Membership Committee chair, Pacific Coast

Entomological Society, Dept. of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118-4599.

Manuscripts, proofs, and all correspondence concerning editorial matters (but

not aspects of publication charges or costs) should be sent to: Dr. Robert V.

Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,

1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-

tributors, and volume 73(4): 248—255, October 1997, for more detailed informa-

tion. Information on format for taxonomic manuscripts can be found in volume

69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer.

The annual dues, paid in advance, are $25.00 for regular members of the So-

ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for

institutional subscriptions or sponsoring members. Members of the Society receive

The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes

are available; see 72(4): 247 for current prices. Make checks payable to the Pacific

Coast Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 1999

Laura A. Irons, President Vincent FE Lee, Managing Secretary

Julieta EF Parinas, Treasurer Stanley E. Vaughn, Recording Secretary

THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per

year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate

Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional

mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society,

% California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

This issue mailed 30 July 1999

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

PAN-PACIFIC ENTOMOLOGIST

75(3): 121-129, (1999)

Obituary: Vincent D. Roth (1924-1997)

DARRELL UBICK

Department of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118,

dubick @calacademy.org, (415) 750-7235

Figure 1. Vincent D. Roth in 1979 at the Southwestern Research Station, Arizona.

Abstract—A chronology of the life of Vincent D. Roth, one of the founding members of the

American Arachnological Society and member of the Pacific Coast Entomological Society from

10 December 1965 until his retirement in 1986, is presented along with a list of his publications

and taxa named in his honor.

It is with sadness that I report the death of Vince Roth, after an illness, on 27

July 1997 at his home in Portal, Arizona. To me, Vince was a mentor and friend

for many years; as he was to numerous others in the Arachnological community.

If the value of a person is to be measured in terms of the vacancy left behind,

122 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

then Vince’s passing is indeed a great loss. No more his myriad notes and spec-

imens scattered to numerous biologists around the globe; nor his newsletters de-

scribing travels to exotic places.

Vince was born as Vincent Daniel Roth on 12 February 1924 in Portland,

Oregon, where he grew up with one sibling, his younger sister, Wilma (Willie).

His father, Frank Xavier Roth, worked for the railroads, was born in WinbGring,

Alt6ting, Bavaria (as Francis Seraph Rott), of Bavarian and Bohemian ancestry,

and immigrated to the United States as a child, in 1893. His mother, Mary Wied-

meyer, was from Richfield, Wisconsin, born to a German father and a Menomonee

Indian mother. The earliest indication of Vince’s entomological bent is from a

photo, showing him at two or three years old dressed as a bumble bee. He attended

the local elementary schools and was a good student but, in response to the illness

and premature death of his mother, failed and had to repeat the 7th grade. From

1938 he attended Benson Polytechical High School and completed 2.5 years of

automotive and trade courses.

In 1941 Vince joined the U.S. Navy and was in Pearl Harbor on 7 December

1941 during the Japanese bombing. He remained in the Navy until the end of the

war, completing machinist school and studying steam and diesel engines, even-

tually attaining the rank of Machinist Mate Ist Class. The Navy gave him an

opportunity to travel throughout the Pacific and he visited ports from Korea to

the Ryukyus, Philippines, New Guinea, New Caledonia, Galapagos, and Hawaii.

This, no doubt, catalyzed his love of travel (despite the fact that Vince never

overcame seasickness) which remained with him for the rest of his life. This was

also the time that he read a book, E. W. Teale’s Grassroot Jungles, which triggered

his interest in entomology.

After the war, in 1946, he enrolled in Oregon State College, Corvallis, Oregon,

and received a B.S. (1949) and M.S. (1951) in Entomology. From the onset he

gravitated to entomology and soon became Curator of the insect collection; a job

he kept throughout his stay at the college. In the summers he took employment

in agricultural entomology, working on cranberry insects (under Dr..J. B. Rosen-

stiel) and surveying mosquitos (under Dr. A. Lindquist). Among his social activ-

ities, he was elected to the Phi Sigma Biological Society and the Sigma Xi Honor

Society, joined the Entomological Society of America and the Society of System-

atic Zoology, served as secretary of the Oregon Entomological Society, and be-

came the president of the Square Dance Club on campus.

During this period Vince became interested in spiders. Unable to identify what

turned out to be a black widow, he contacted Dr. Willis J. Gertsch of the American

Museum of Natural History, who suggested he study Comstock’s The Spider Book

and possibly introduced him to Dr. Harriet Exline (Frizzell) of the University of

Washington. Years later he spoke to me of his appreciation of her generosity, not

only in the sharing of information and providing literature but, as was important

to Vince, teaching him the pronounciation of the many elusive names in arach-

nology. Her influence is reflected in the fact that he followed up her studies of

the western Agelenidae (s. 1.). He collected avidly in Oregon, finding many new

species, some of which he described in his thesis (on the Oregon species of

Cybaeus) and in other papers. From that point on, agelenoid spiders became

Vince’s primary arachnological interest.

In 1952 Vince married Jean Louise Lamb in Milwauke, Oregon, and the fol-

1999 UBICK: ROTH OBITUARY 123

Figures 2-7. Vincent D. Roth. Figure 2. With parents and sister in Oregon in 1935; Figure 3.

Dressed as bumble bee in 1928; Figure 4. With wife, Jean, and daughter, Susan, at Salem, Oregon,

during 1954-1955; Figure 5. With wife, Bobbie, and daughters, Kristin and Kim, at the Southwestern

Research Station, Arizona, in 1969; Figure 6. With Willis Gertsch and Darrell Ubick at Tucson,

Arizona, in 1996; Figure 7. With wife, Barbara, in front of ““E] Ghosto Blanco” at Portal, Arizona,

in 1996.

124 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

lowing year they moved to Berkeley, California, where he enrolled at the Uni-

versity of California with the goal of continuing his graduate studies. These were

curtailed shortly after the birth of their daughter, Susan Elizabeth (in 1954), when

the family moved back to Salem, Oregon. Here Vince started employment with

the Oregon State Department of Agriculture as a Survey Entomologist, where he

sampled for insect pests, assembled collections, and promoted 4-H programs in

entomology. The following year, however, a better position became available at

the University of Arizona Experiment Station. Leaving his family in Salem, Vince

moved to Yuma and, as Assistant Entomologist, worked on the ‘‘Yellow Clover

Aphid Problem”? to develop a control for the Spotted Alfalfa Aphid. Although he

regularly visited his family in Oregon, the arrangement proved unsatisfactory and

the following year Vince and Jean were divorced.

In 1956 Vince began his long field association with Dr. Willis J. Gertsch, of

the American Museum of Natural History, by assisting him on a collecting trip

through the western United States and Mexico. As Vince was a keen collector,

he kept a steady stream of interesting spiders flowing to Willis; on one occasion

donating a particularly large collection of Oregon spiders, for which he received

Life Membership to the AMNH. He also accompanied Willis on three expeditions

to California, in 1958, 1959, and 1960 (the last joined by Wilton Ivie), which

probably represent the first major collection of spiders from this region.

A major turning point in Vince’s career came in 1962, when he became the

Resident Director of the Southwestern Research Station. The station, or SWRS,

located in the scenic and biologically rich Chiricahua Mountains of southeastern

Arizona, was founded in 1955 and directed by Dr. Mont Cazier for the AMNH.

Cazier’s sudden retirement in mid-1962 left a vacancy which was temporarily

filled by Willis Gertsch, who in turn encouraged Vince to take the post. This he

did eagerly, taking up residence at the station’s ‘“‘Log Cabin” in 1963 with his

wife, Dorothea “‘Bobbie’”? Ann Thompson from El Centro, California, whom he

married in 1961. They had a daughter, Kristin Ann, and some years later (1970)

adopted Kim Lee (who was born as Hai Sook Jin in Seoul, Korea; in 1964, the

same year as Kristin). After Vince’s second marriage dissolved (1972), the girls

continued living at the station until they moved to Tucson to attend high school

(1975).

Apart from running SWRS, Vince continued to spend much time exploring the

desert, and took numerous trips throughout the Chiricahuas and Sonora, Mexico,

keeping detailed field notes of the places visited. Most of the time it was in his

old white pick-up, which he named “El Ghosto Blanco’’, and which he (mirac-

ulously) maintained in working order. El Ghosto was always kept well stocked

with survival and collecting gear to be available for field work at a moment’s

notice, which was often the case. Although spiders were his main love, Vince

maintained a generalist’s interest in biology and created the synoptic collection

of the local fauna and flora at SWRS. He enjoyed inventing and created a mini-

ature herbarium and plant press to facilitate plant identification in the field (Roth

1972). His passion for solving biological puzzles is well illustrated in “‘the case

of the lost lizard’’, as he called it (1997). The horned lizard, Phrynosoma ditmarsi

Stejneger, was known from only a few museum specimens, all collected at the

turn of the century at an unknown locality in Sonora. Vince eventually discovered

the locality by closely examining those parts of it that the lizards themselves had

1999 UBICK: ROTH OBITUARY 125

collected, that is, ingested (Roth 1971, Lowe et al. 1971). Vince was also an

outspoken conservationist and expended much effort to preserve the natural en-

vironment in the Chiricahuas. He actively worked to stop overgrazing, much to

the consternation of the local ranchers, and his battle against noxious weeds was

equally legendary; he always carried a shovel in El Ghosto for eradicating the

latest incursion of horehound or Johnsongrass.

In his spare time, he continued taxonomic work on spiders. In 1962 he received

a National Science Foundation Grant for a taxonomic study of the South American

Agelenidae, at the California Academy of Sciences. He published several addi-

tional papers on agelenids during this period, along with several lists, including

Galapagos spiders (with P. R. Craig 1970), nearctic Gnaphosidae (with D. Ubick

1973), Chiricahua Mountain spiders (with A. K. S. Jung 1974), and Yuma Co.,

Arizona, jumping spiders (with D. Richman 1976). His discovery of a new species

of intertidal spider in Sonora (Desidae: Paratheuma interaesta (Roth & Brown))

resulted in a series of papers on spiders from this unusual habitat, mostly with

W. L. Brown from 1975-1980.

In 1968, his close friend and mentor, Willis Gertsch, retired from the AMNH

and moved to the nearby village of Portal. Soon other arachnologists retired and

moved to this area, as did Dr. Findley Russell and Dr. Martin H. Muma, and

many more visited each summer to where the Chiricahuas became a veritable

Mecca for Arachnologists. Appropriately, the first meeting of the American Ar-

achnological Society, co-hosted by Willis and Vince in August 1972, was held

there. Vince continued being active in the AAS throughout his life, serving as

Vice President (from 1973-1975), Director (1980-1982), member of the Editorial

Board (1985-1986), and Archivist (1986-1997). At the 1993 meeting he intro-

duced the Auction, which immediately became a popular and entertaining way of

raising money for the society, as well as redistributing literature (much of which

was donated by Vince). In 1995 the AAS honored Vince with a commemorative

plaque, in recognition of his contributions to Arachnology and to the Society, of

which he was a charter member.

A most important trip for Vince was in the winter of 1975-1976 to Costa Rica,

where he met Barbara Maria Emmanuela Schropfer, a midwife from Cham, Ba-

varia. The following year they were married in an elaborate ceremony at SWRS.

Barbara assisted Vince in running the station, was a hostess to the many visitors,

and helped Vince in the field and laboratory. In 1977 Vince acquired some prop-

erty in Portal from Willis. The following year he and Barbara, with the help of

an elderly carpenter, built a large and attractive house (dubbed Number One Spi-

der Lane), to which they retired in 1986: In October 1996, they had twins, Daniel

and Taran, one month prior to Vince’s death.

After Vince’s retirement, they embarked upon an ambitious program of world

travel, visiting the following: Spain-Morocco-Egypt-Israel-Germany (September

1986—July 1987); South America (March 1988—March 1989); India-SE Asia (Oc-

tober 1989—April 1990); southern Africa (October 1990—April 1991); Madagascar

(March—September 1992); eastern Africa (October-November 1992); Ecuador

(January—December 1994); Siberia (Kiril Islands) (July—September 1995); Hawaii

(1977, 1982, 1984, 1990, 1996). These trips produced rich collections of spiders,

now mostly at the California Academy of Sciences, some at the Museum of

Comparative Zoology. In addition to collecting, Vince and Barbara volunteered

126 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

their services in a number of ways. In Ecuador they spent an entire year running

a field station, Cabafias Alifahui (now known as Butterfly Lodge), located on the

Rio Napo; and in Madagascar, three months conducting a spider survey in Ran-

omafana National Park. They routinely curated spiders at the various museums

visited around the globe. Vince was appointed Research Associate at both the

California Academy of Sciences and the Bernice P. Bishop Museum.

During this period Vince’s spider work was devoted largely to compiling keys

to the Nearctic genera, an ambitious goal given our limited knowledge of many

families, and which he published as Handbook for Spider Identification (1982)

and later as Spider Genera of North America (1985, 1993). For this, he interacted

with many colleagues, eagerly gathering the latest taxonomic information and

corrections for the next edition. This project was typical Vince: a collaborative

effort to produce a usable and up-to-date key, available at a modest cost to a

broad audience, and with the profits going to support arachnological research.

Vince is survived by his wife Barbara, sons Taran and Daniel, daughters Kim,

Kristin, and Suzie, grandchildren Ajelina and Dylan, and sister Wilma.

TAXA NAMED AFTER VINCENT AND BARBARA ROTH

ARANEAE

Aphonopelma rothi Smith, 1995 (Theraphosidae)

Callobius rothi Leech, 1972 (Amaurobiidae)

Castianeira rothi Reiskind, 1969 (Corinnidae)

Cesonia rothi Platnick & Shadab, 1980 (Gnaphosidae)

Euagrus rothi Coyle, 1988 (Dipluridae)

Hesperocranum rothi Ubick & Platnick 1991 (Liocranidae)

Indothele rothi Coyle, 1995 (Dipluridae)

Legendrena rothi Platnick, 1995 (Gallieniellidae)

Loxosceles barbara Gertsch & Ennik, 1983 (Sicariidae)

Loxosceles rothi Gertsch & Ennik, 1983 (Sicariidae)

Metazygia rothi Levi, 1995 (Araneidae)

Misumenops rothi Schick, 1965 (Thomisidae)

Neotama rothorum Baehr & Baehr, 1993 (Hersiliidae)

Rhoicinus rothi Exline, 1960 (Trechaleidae)

Theridion rothi Levi, 1959 (Theridiidae)

Tibellus rothi Schick, 1965 (Philodromidae)

Tricholathys rothi Chamberlin & Gertsch, 1958 (Dictynidae)

Zimiromus rothi Platnick & Shadab, 1981 (Gnaphosidae)

SOLIFUGAE

Therobates rothi Muma, 1962 (now Eremochelis, Eremobatidae)

CHELONETHIDA

Diplotemnus rothi Muchmore, 1975 (Miratemnidae)

OPILIONES

Dalquestia rothorum Cokendolpher & Stockwell, 1986 (Sclerosomatidae)

1999 UBICK: ROTH OBITUARY 127

ISOPODA

Caucasonethes rothi Vandel, 1953 (Trichoniscidae)

DIPLOPODA

Bdellozonium rothi Chamberlin, 1950 (Polyzoniidae)

CHILOPODA

Stenophilus rothi Chamberlin, 1953 (Geophilidae)

COLLEMBOLA

Friesea rothi Christiansen & Bellinger, 1988 (Poduridae)

ORTHOPTERA

Grylloblatta rothi Gurney, 1953 (Grylloblattidae)

TRICHOPTERA

Limnephilus rothi Denning, 1966 (Limnephilidae)

Ochrotrichia rothi Denning & Blickle, 1972 (Hydroptilidae)

COLEOPTERA

Alaocybites rothi Gilbert, 1956 (Curculionidae)

Anilloferonia rothi Hatch, 1951 (Carabidae)

Bryothinusa rothi Moore & Legner, 1975 (Staphylinidae)

Catopocerus rothi Hatch, 1957 (Leiodidae)

Endeodes rothi Moore, 1975 (Melyridae)

Malthinus rothi Fender, 1972 (Cantharidae)

Pselaptrichus rothi Park in Hatch, 1962 (Pselaphidae)

Pseudohaida rothi Hatch, 1957 (Staphylinidae)

Rothium Moore & Legner, 1977 (Staphylinidae)

Scydmaenus rothi Marsh in Hatch, 1957 (Scydmaenidae)

Trigonoscuta rothi Pierce, 1975 (Curculionidae)

DIPTERA

Eupary phus rothi James, 1973 (Stratiomyidae)

HYMENOPTERA

Gryon rothi Masner, 1979 (Scelionidae)

Playaspalangia rothi Yoshimoto, 1976 (Pteromalidae)

SCIENTIFIC PUBLICATIONS OF V. D. ROTH

Bogert, C. M. & V. D. Roth. 1966. Ritualistic combat of male gopher snakes. Pituophis melanoleucus

afinis (Reptila, Colubridae). Amer. Mus. Novit., 2245: 1-27.

Cooke, A. L., V. D. Roth & FE H. Miller. 1972. The urticating hairs of theraphosid spiders. Amer.

Mus. Novit., 2498: 1—43.

Jung, A. K. S. & V. D. Roth. 1974. Spiders of the Chiricahua Mountain area, Cochise Co., Arizona.

J. Ariz. Acad. Sci., 9: 29-34.

Lowe, C. W., M. D. Robinson & V. D. Roth. 1971. A population of Phrynosoma ditmarsi from Sonora,

Mexico. J. Ariz. Acad. Sci., 6: 275-277.

128 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Nesom, G. L. & V. D. Roth. 1981. Erigeron scopulinus (Compositae), an endemic from the south-

western United States. J. Ariz.-Nev. Acad. Sci., 16: 39-42.

Richman, D. B. & V. D. Roth. 1976. A revised list of the jumping spiders (Araneae: Salticidae) of

Yuma County, Arizona. SW Natur., 21: 199-202.

Roth, V. D. 1951. New records for Streblidae and Nycteribiidae. Pan-Pac. Entomol., 27: 96.

Roth, V. D. 1952a. Method of storing spiders and insects in alcohol. Entomol. News, 63: 45.

Roth, V. D. 1952b. Notes and a new species in Cybaeina. Pan-Pac. Entomol., 42: 195-199.

Roth, V. D. 1952c. A review of the genus Tegenaria in North America. J. Wash. Acad. Sci., 42: 283-—

288.

Roth, V. D. 1952d. The genus Cybaeus in Oregon. Ann. Entomol. Soc. Amer., 45: 205-219.

Roth, V. D. 1954. Review of the spider subgenus Barronopsis (Arachnida, Agelenidae, Agelenopsis).

Amer. Mus. Novit. 1678: 1-7.

Roth, V. D. 1955. Entomology leaders handbook and leaflets. Oregon State College Extension Service.

Roth, V. D. 1956a. Revision of the genus Yorima Chamberlin and Ivie (Arachnida, Agelenidae). Amer.

Mus. Novit. 1773: 1-10.

Roth, V. D. 1956b. Taxonomic changes in the Agelenidae. Pan-Pac. Entomol., 32: 175-180.

Roth, V. D. 1959. Alfalfa seed treatments for Spotted Alfalfa Aphid control in southeastern Arizona.

J. Econ. Entomol., 52: 654-658.

Roth, V. D. 1960. Insect immigrants. Farm Bureau Monthly, Dec: 6.

Roth, V. D. 1961. The beneficial spider. Farm Bureau Monthly, Jul-Aug: 8-9.

Roth, V. D. 1962a. Germination of alfalfa seed treated with dry and liquid formulations of di-syston

and phorate. J. Econ. Entomol., 55: 1-2.

Roth, V. D. 1962b. Teaching the classification of Arthropoda to non-scientific groups. Turtox News,

40: 46

Roth, V. D. 1963. The familial affiliation of the spider genus Textrix. System. Zool., 12: 173-174.

Roth, V. D. 1964. The taxonomic significance of the spider trochanter. Ann. Entomol. Soc. Amer., 57:

759-766.

Roth, V. D. 1965. Genera erroneously placed in the spider families Agelenidae and Pisauridae (Ara-

neida: Arachnida). Ann. Entomol. Soc. Amer., 58: 289-292.

Roth, V. D. 1967a. A review of the South American spiders of the family Agelenidae (Arachnida,

Araneida). Bull. Amer. Mus. Nat. Hist., 134: 299-345.

Roth, V. D. 1967b. Redescription of Tegenaria longimana Simon (Araneae, Agelenidae). J. Ariz. Acad.

Sci., 4: 197-198.

Roth, V. D. 1967c. A redescription of the spider genus Mizaga Simon (Agelenidae), with new syn-

onymy. Amer. Mus. Novit., 2291: 1-7.

Roth, V. D. 1967d. Descriptions of the spider families Desidae and Argyronetidae. Amer. Mus. Novit.,

2292: 1-9.

Roth, V. D. 1968. The spider genus Tegenaria in the Western Hemisphere (Agelenidae). Amer. Mus.

Novit., 2323: 1-33.

Roth, V. D. 1970. A new genus of spiders (Agelenidae) from the Santa Catalina Mountains. J. Ariz.

Acad. Sci., 6: 114-116.

Roth, V. D. 1971. Food habits of Ditmars’ Horned Lizard with speculation on its type locality. J. Ariz.

Acad. Sci., 6: 278-281.

Roth, V. D. 1972a. Taxonomic changes in the Agelenidae, II. Notes Arachnol. Southwest, 3: 10-12.

Roth, V. D. 1972b. Proposal for a Nearctic spider catalog. Amer. Arachnol., 8: 6-9.

Roth, V. D. 1972c. A miniaturized plant press and herbarium. Amer. Biol. Teacher, 34: 535-536.

Roth, V. D. 1973. A miniaturized plant press and herbarium. Saguaroland Bull. Desert Bot. Garden

Ariz., 27: 82-84.

Roth, V. D. 1976. XVI. Rediscription of the marine spider Desis galapagoensis (Desidae). Mission

zoologique belge aux iles Galapagos et en Ecuador (N. et J. Leleup, 1964-1965), 3: 11-22.

Roth, V. D. 1977. Speed key to clubionid genera of America north of Mexico. Amer. Arachnol., 17: 12.

Roth, V. D. 1980. Chapter 23. Arthropoda: Arachnida. pp. 347-355. In R. C. Brusca (ed.). Common

intertidal invertebrates of the Gulf of California. Univ. Ariz. Press, Tucson.

Roth, V. D. 1981la. A new genus of spider (Agelenidae) from California exhibiting a third type of leg

autospasy. Bull. Amer. Mus. Nat. Hist., 170: 101-105.

Roth, V. D. 1981b. A miniaturized plant press and herbarium. N Nev. Native Plant Soc. Newsl. 7: 6-8.

Roth, V. D. 1982. Handbook for spider identification. Privately published.

1999 UBICK: ROTH OBITUARY 129

Roth, V. D. 1984a. The spider family Homalonychidae (Arachnida, Araneae). Amer. Mus. Novit.,

2790: 1-11.

Roth, V. D. 1984b. A miniaturized plant press and herbarium. Plants in the Laboratory. Nat. Assoc.

Biol. Teachers, Mono. series 1: 1-2.

Roth, V. D. 1985. Spider genera of North America. Arachnol. Soc. Amer., Gainesville.

Roth, V. D. 1988. American Agelenidae and some misidentified spiders (Clubionidae, Oonopidae, and

Sparassidae) of. E. Simon in the Museum national D’Histoire naturelle. Bull. Mus. natn. Hist.

nat., Paris, 4: 25-37.

Roth, V. D. 1993. Spider genera of North America with keys to families and genera and a guide to

literature (3rd ed.). 1-203.

Roth, V. D. 1997. Ditmar’s Horned Lizard (Phrynosoma ditmarsi) or the case of the lost lizard. Sonoran

Herpet., 10: 2-6.

Roth, V. D. & P L. Brahme. 1972. Nearctic genera of the spider family Agelenidae (Arachnida,

Araneida). Amer. Mus. Novit. 2505: 1—52.

Roth, V. D. & W. L. Brown. 1973. Illustrated key to the Nearctic Gnaphosidae with a speed key

supplement. Amer. Arachnol., 9: 18-22.

Roth, V. D. & W. L Brown. 1975a. Comments on the spider Saltonia incerta Banks (Agelenidae?). J.

Arachnol., 3: 53-56.

Roth, V. D. & W. L. Brown. 1975b. A new genus of Mexican intertidal zone spider (Desidae) with

biological and behavioral notes. Amer. Mus. Novit., 2568: 1-7.

Roth, V. D. & W. L. Brown. 1976a. Chapter 6. Other intertidal air-breathing arthropods. pp. 119-150.

In L. Cheng (ed.). Marine insects. North-Holland Publishing Co., Amsterdam.

Roth, V. D. & W. L Brown. 1976b. A new wolf spider from the Gulf of California beaches. J. Ariz.

Acad. Sci., 11: 61-63.

Roth, V. D. & W. L. Brown. 1986. Catalog of Nearctic Agelenidae. Occ. Pap. Mus. Texas Tech Univ.,

99:1-21.

Roth, V. D. & P. R. Craig. 1970. VII. Arachnida of the Galapagos Islands (Excluding Acarina). Mission

zoologique belge aux iles Galapagos et en Ecuador. Musee Royal de I’ Afrique Centrale, 2:

107-124.

Roth, V. D. & B. M. Roth. 1984. A review of appendotomy in spiders and other arachnids. Bull. Brit.

Arachnol. Soc., 6: 137-146.

Schonhorst, M. H., V. D. Roth & K. W. Nielson. 1959. Breeding Alfalfa to have more resistance to

parasites. Progressive Agriculture, Summer: 11.

Tuttle, D. M., O. L. Barnes, M. W. Nielson, V. D. Roth & M.H. Schonhorst. 1958. The Spotted

Alfalfa Aphid in Arizona. Univ. Ariz. Agric. Exp. Stat., Bull. 294: 1-10.

Ubick, D. & V. D. Roth. 1973a. Nearctic Gnaphosidae including species from adjacent Mexican states

with an index to synonymy and invalid names. Amer. Arachnol., 9: 23-40.

Ubick, D. & V. D. Roth. 1973b. Gnaphosidae of Mexico not listed in the Nearctic catalog. Amer.

Arachnol., 9: 41—43.

van den Bosch, R., L. H. Dawson, V. D. Roth & V. W. Brown. 1961. Promising new parasite of the

Egyptian Alfalfa Weevil imported from southern Iran. Calif. Agric., 15: 11.

Walters, R.D. & V. D. Roth. 1950. Faunal nest study of the woodrat, Neotoma fuscipes monochroura

Rhoads. J. Mammal., 31: 290-292.

ACKNOWLEDGMENT

Thanks to Barbara Roth for her hospitality and help in gathering information

on Vince’s life, and Kristin and Bobbie Roth for making some photographs avail-

able. Patrick Craig and Warren Savary assisted with photography and provided

good companionship on trips to Arizona. Anne Gondor helped with the text and

prepared the plates, and Suzanne Prinsloo assisted with library tasks. Paul Arnaud

and Charles Griswold reviewed the manuscript and gave many useful suggestions.

Finally, thanks go out to the many individuals, too numerous to mention, who

shared with me their recollections about Vince.

Received 13 Nov 1998; Accepted 10 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 130-146, (1999)

AN ANALYSIS OF THE GENUS SALAPIA STAL WITH

DESCRIPTION OF SIX NEW SPECIES, AND SOME

TAXONOMIC REARRANGEMENTS (HEMIPTERA:

HETEROPTERA: COREIDAE: ACANTHOCEPHALINI)

HARRY BRAILOVSKY AND ERNESTO BARRERA

Instituto de Biologia, UNAM, Depto. de Zoologia, Apdo Postal No. 70-153,

México 04510, D.E México

Abstract.—Six new species of Salapia Stal from Brazil, Ecuador, and Peru are described and

illustrated. Salapia humeralis (Burmeister) is redescribed. Salapia guttifera (Stal) is transferred

to the genus Stenometapodus, resulting in the new combination Stenometapodus guttifer (Stal).

A key to the known species of Salapia is included.

Key Words.—Insecta, Hemiptera, Heteroptera, Coreidae, Acanthocephalini, Salapia, new species,

Brazil, Ecuador, Peru.:

Brailovsky (1992) revised the genus Salapia Stal, described four new spe-

cies, and transferred Laminiceps haenschi Breddin (1901) to Salapia. In the

same contribution he added new records for some species, and included a key

to the known taxa, except S. humeralis (Burmeister) and S. guttifera (Stal)

because the types were not located and the original description was too short

to find good characters to separate from the other known species. However,

the discovery of the types of S. humeralis and S. guttifera in the Museum der

Humboldt Universitat zu Berlin, six undescribed species, and some taxonom-

ical problems made the present analysis, which includes a key to the known

species necessary.

The genus Salapia is characterized by an elongate body; cylindrical hind tibiae

that is not expanded; rostrum never extending to abdomen with segment IV the

shortest, segment II the longest, and segment III longer than I; scutellum longer

than wide, humeral angles acute, and metathoracic peritreme with two auricles.

The closely related genus Laminiceps Costa is similar to Salapia except for the

body robust, nearly oval, never parallel-sided, the scutellum wider than long, and

the humeral angles obtuse, not acute.

Previously, twelve species of Salapia (S. abdominalis (Dallas), S. baraquini

(Signoret), S. dimidiata (Dallas), S. guttifera (Stal), S. haenschi (Breddin), S.

humeralis (Burmeister), S. luteola Brailovsky, S. nigra Brailovsky, S. pallida Brai-

lovsky, S. pretiosa Bléte, S. selecta Brailovsky, and S. signata (Dallas)) were

known. In this paper we add six new species collected in Brazil, Ecuador and

Peru, and one species, S. guttifera, is transferred to the genus Stenometapodus,

forming the new combination Stenometapodus guttifer.

With this contribution, the current number of species known in Salapia is 17.

SALAPIA CAUCALANDIA BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 1, 9)

Types.—Holotype: female; data: BRAZIL. Rondonia, vic. Caucalandia,

10°32'S—62°48'W, 160-350 m, 30 October 1991, J. MacDonald. Deposited in

Mississippi Entomological Museum, Mississippi State. Paratype: 1 female: BRA-

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 131

Figures 1-8. Pronotum of Salapia spp. Figure 1. S. caucalandia Brailovsky and Barrera, NEW

SPECIES. Figure 2. S. deckerti Brailovsky and Barrera, NEW SPECIES. Figure 3. S. vanduzeei

Brailovsky and Barrera, NEW SPECIES. Figure 4. S. egeri Brailovsky and Barrera, NEW SPECIES.

Figure 5. S. humeralis (Burmeister). Figure 6. S. kondratieffi Brailovsky and Barrera, NEW SPECIES.

Figure 7. S. onorei Brailovsky and Barrera, NEW SPECIES. Figure 8. S. signata (Dallas).

132 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Figure 9. Dorsal view of Salapia caucalandia Brailovsky and Barrera, NEW SPECIES.

ZIL. Rondonia ‘Austin Trail’’ (linea C-13) off B-65, 2 km. N. Caucalandia, 21—

23 March 1991, Kondratieff and Welch. Deposited in the ““Coleccién Entomo-

lé6gica del Instituto de Biologia, UNAM, México.”

Description—Female (holotype). Dorsal coloration. Head including antennal segments I to IV

black; pronotum yellow with black discoidal spot on middle third; calli yellow with narrow black

longitudinal stripe running on middle third; scutellum black with apex yellow; clavus and corium

yellow with claval vein, apical margin of corium, and elongate stripe on endocorium black; hemelytral

membrane black; connexival segments with upper margin orange, and inner margin black; abdominal

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 133

segments black. Ventral coloration. Head dull dark hazel with dirty yellow stripe contiguous to inner

face of eye; buccula and rostral segments I to IV shiny black to reddish brown; thorax and legs dull

dark hazel, tinged with orange, and following areas shiny reddish brown: acetabulae, anterior and

posterior margin of propleura, and posterior margin of mesopleura and metapleura; anterior and pos-

terior lobe of metathoracic peritreme black; abdominal sterna bright reddish brown with hazel reflec-

tions; pleural margin of abdominal sterna III to VIII and external face of gonocoxae I bright to dull

orange. Structure. Head: Rostrum reaching anterior third of metasternum. Pronotum: Humeral angles

expanded, apically acute; posterolateral margin denticulate; triangular process shorter (Fig. 1). Mea-

surements: Head length including the tylus: 1.45 mm; width across eyes: 2.53 mm; interocular space:

1.00 mm; interocellar space: 0.42 mm; preocular distance: 0.88 mm; length of antennal segments: I,

3.64 mm; II, 3.00 mm; III, 2.44 mm; IV, 6.68 mm. Pronotal length: 4.10 mm; width across frontal

angles: 1.80 mm; width across humeral angles: 6.60 mm. Scutellar length: 2.44 mm; width: 2.24 mm.

Total body length: 19.10 mm.

Discussion.—Like S. nigra Brailovsky with antennal segments IJ and III, scu-

tellum, buccula, and abdominal sterna (except pleural margins III to VII orange)

black to reddish brown. Salapia caucalandia, can be easily distinguished by the

antennal segment IV, and abdominal segments III to V entirely black, and the

pronotum yellow with discoidal black spot near middle third. On S. nigra the

antennal segment I'V is black with discoidal ring yellow, the abdominal segments

III to V orange, and the pronotum mostly black.

Distribution.—Only known from Brazil.

Etymology.—Named for its occurrence near Caucalandia.

SALAPIA DECKERTI BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 2, 10)

Types.—Holotype: female; datas ECUADOR. Camelus, FE V. Feyer. Deposited

in the Museum der Humboldt Universitat zu Berlin. Paratype: 1 female: same

data as holotype. Deposited in the “‘Coleccién Entomologica del Instituto de Bio-

logia, UNAM, México.”’

Description—Female (holotype). Dorsal coloration: Head, pronotum, and scutellum orange; an-

tennal segment I reddish brown with inner face ochraceus to orange or entirely dirty orange; antennal

segments II and III reddish brown and IV yellow with basal and apical third reddish brown; clavus

orange to yellow with broad longitudinal stripe black; corium orange to yellow with 5 black maculae

between veins; hemelytral membrane dark ambarine with basal angle, and inner veins darker; con-

nexival segments III to VI reddish with upper border, posterior joint, and inner margin pale orange;

connexival segments VII to IX, and dorsal abdominal segments II to IX pale orange. Ventral color-

ation: Head, buccula, thorax, coxae, trochanters, femora, and abdomen pale orange yellow; rostral

segments reddish brown with apical third of segments I to III yellow; tibiae reddish brown with basal

joint orange yellow; tarsi reddish brown; anterior and posterior lobe of metathoracic peritreme black.

Structure. Head: Rostrum reaching anterior border of metasternum. Pronotum: Humeral angles ex-

panded, apically acute; posterolateral margin denticulate; triangular process shorter (Fig. 2). Measure-

ments. Head length including the tylus: 1.43 mm; width across eyes: 2.60 mm; interocular space: 1.04

mm); interocellar space: 0.44 mm; preocular distance: 0.90 mm; length of antennal segments: I, 4.08

mm; II, 3.16 mm; III, 2.48 mm; IV, 6.80 mm. Pronotal length: 3.96 mm; width across frontal angles:

1.68 mm; width across humeral angles: 6.80 mm. Scutellar length: 2.52 mm; width: 2.32 mm. Total

body length: 18.80 mm.

Discussion.—Close to S. luteola Brailovsky with antennal segments IJ and III

black to reddish brown, and pronotum and scutellum entirely yellow to orange.

In S. deckerti, the clavus is black with margin yellow to orange, the corium is

yellow to orange with 5 black maculae between veins, and ventrally entirely

134 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

AQ

‘Figure 10. Dorsal view of Salapia deckerti Brailovsky and Barrera, NEW SPECIES.

yellow to orange. In S. luteola the clavus, corium, and ventral region are black

to reddish brown, except the orange pleural margin of abdominal sterna III to VI.

Salapia pretiosa Blote also has the antennal segments II and III black to reddish

brown, and the scutellum mostly yellow to orange (apical third black), but the

clavus is entirely black, the corium black with transverse and straight orange to

yellow stripe, and antennal segment IV black to reddish brown and not bicolorous

like in S.. deckerti.

Distribution.—Only known from Ecuador.

Etymology.—Named for Dr. Jurgen Deckert, distinguished German hemipterist.

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 135

Figure 11. Dorsal view of Salapia vanduzeei Brailovsky and Barrera, NEW SPECIES.

SALAPIA VANDUZEEI BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 3, 11)

Type.—Holotype: female; data: PERU. Putumayo District, E] Encanto, 25 Au-

gust 1920, Cornell Univ. Expedition, lot 569. Deposited in Cornell University,

Insect Collection, Ithaca, New York.

Description—Female (holoty pe). Dorsal coloration: Head, pronotum, scutellum, connexival segments

and abdominal segments orange; antennal segment I reddish brown with inner face dirty orange; segments

II and III reddish brown and IV yellow with basal and apical third reddish brown; tylus reddish brown

136 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

with orange reflections; clavus black with basal angle orange; corium black with yellow irregular fascia;

hemelytral membrane smoky black, with basal angle darker. Ventral coloration: Head, buccula, thorax,

coxae, and abdominal sterna orange; rostral segments I to IV, trochanters, femora, tibiae, and tarsi reddish

brown; anterior and posterior lobe of metathoracic peritreme black. Structure. Head: Rostrum reaching

anterior border of metasternum. Pronotum: Humeral angles expanded, apically acute; posterolateral mar-

gin denticulate; triangular process shorter (Fig. 3). Measurements. Head length including the tylus: 1.45

mm; width across eyes: 2.60 mm; interocular space: 1.00 mm; interocellar space: 0.41 mm; preocular

distance: 0.90 mm; length of antennal segments: I, 3.88 mm; II, 3.04 mm; III, 2.44 mm; IV, 6.92 mm.

Pronotal length: 3.76 mm; width across frontal angles: 1. 66 mm; width across humeral angles: 6.48

mm. Scutellar length: 2.44 mm; width: 2.36 mm. Total body length: 18.50 mm.

Discussion.—Related with S. luteola Brailovsky, in having antennal segments

II and III and legs reddish brown to black, and pronotum and scutellum orange.

Salapia vanduzeei, has the head, thorax, coxae and abdomen orange, in S. luteola

these structures are reddish brown to black, except the connexival segments and

pleural abdominal sterna are orange to yellow.

Distribution.—Only known from Peru.

Etymology.Named for the late Dr. Edward P. Van Duzee, in recognition of

his many fundamental contributions to hemipteran systematics.

SALAPIA EGERI BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 4, 12)

Type.—Holotype: female; data: PERU. Sicuani. Deposited in the ‘‘Coleccién

Entomoldgica del Instituto de Biologia, UNAM, México.”’

Description—Female (holotype). Dorsal coloration: Head pale orange yellow; antennal segments

I to III yellow with apical joint reddish brown; segment IV yellow with apical third reddish brown;

pronotum pale orange yellow with elongate macula orange brown near to middle third; scutellum

orange brown with basal discoidal spot, lateral margins, and apex yellow; clavus pale orange yellow

with inner border reddish brown; corium pale orange yellow with 3 reddish brown maculae between

veins; hemelytral membrane dark ambarine with basal angle darker; connexival segments III to VI

orange brown and VII pale orange; abdominal segments black with creamy yellow longitudinal stripe

running from II to posterior margin of segment VII. Ventral coloration: Head, buccula, rostral seg-

ments I to IV (apex of IV reddish brown), and legs (spines black to reddish brown) pale yellow to

orange; thorax pale orange yellow with following areas orange brown: rectangular spot on propleura;

outer margin of mesopleura, and posterior margin of metapleura; anterior and posterior lobe of meta-

thoracic peritreme black; abdominal sterna shiny orange hazel, with creamy yellow discoidal spot

lateral to middle line on sterna III to VI. Structure. Head: Rostrum reaching posterior border of

mesosternum. Pronotum: Humeral angles expanded, broad, apically with short and broad, spine pos-

terolateral margin denticulate; triangular process broad, well developed (Fig. 4). Measurements. Head

length including the tylus: 2.00 mm; width across eyes: 2.52 mm; interocular space: 0.92 mm; inter-

ocellar space: 0.50 mm; preocular distance: 1.00 mm; length of antennal segments: I, 3.60 mm; II,

2.76 mm; III, 2.20 mm; IV, 6.28 mm. Pronotal length: 3.72 mm; width across frontal angles: 2.20

mm; width across humeral angles: 6.84 mm. Scutellar length: 2.28 mm; width: 2.20 mm. Total body

length: 18.62 mm.

Discussion.—This species is similar in color and general appearance to S. sig-

nata (Dallas) in having the head orange yellow and antennal segments I to III

yellow to orange with apical joint reddish brown to black. In S. egeri, the ace-

tabulae are entirely orange, the pronotal disc, and scutellum without creamy yel-

low longitudinal stripe, the corium without transverse yellow fascia and humeral

angles are broadened, and apically have a short broad spine (Fig. 4). In S. signata

the acetabulae are creamy yellow, and contrast with the orange surface, the pron-

137

NEW SALAPIA SPECIES

BRAILOVSKY & BARRERA

1999

lovsky and Barrera, NEW SPECIES.

Dorsal view of Salapia egeri Brai

Figure 12.

138 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Figure 13. Dorsal view of Salapia kondratieffi Brailovsky and Barrera, NEW SPECIES.

otal disc, scutellum and corium with creamy yellow marks and humeral angles

are narrowed, exposed, and apically with long, acute spine (Fig. 8).

Distribution Only known from Peru.

Etymology.—Named for Dr. J. E. Eger, distinguished American hemipterist.

SALAPIA KONDRATIEFFI BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 6, 13)

Types.—Holotype: female; data: BRAZIL. Rondonia, 62 km. SW, Ariquemes,

nr. Fazenda Rancho Grande, 6-15 December 1990, D. A. Rider and J. E. Eger.

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 139

Deposited in the “‘Colecci6n Entomolégica del Instituto de Biologia, UNAM,

México’’. Paratype: 1 female; data: BRAZIL. Rondonia, Linea C-2.5, off B-65,

12.5 km. S. Caucalandia, 17 March 1991, Kondratieff and Welch. Deposited in

Colorado State University, Fort Collins, Colorado.

Description—Female (holotype). Dorsal coloration: Head dark orange hazel; antennal segments I to

Ill yellow orange with apical joint reddish brown; segment IV reddish brown with pale yellow ring near

middle third; pronotum dark orange hazel with narrow longitudinal stripe pale yellow orange; scutellum

reddish brown with narrow longitudinal stripe pale yellow orange; clavus dark orange with apical third

black; cortum dark orange with 5 to 6 black maculae between apical veins; hemelytral membrane yellow

ambarine with basal angle and space between veins brown; connexival segments reddish brown with

upper margin orange; abdominal segments orange. Ventral coloration: Head including buccula dirty

yellow with middle third pale brown; rostral segments I and II dirty orange hazel with apical join darker;

segments III and IV reddish brown; thorax dark orange hazel with posterior margin of metapleura reddish

brown; coxae and trochanters ochraceus to orange hazel with outer margin reddish brown; femora dark

orange hazel; tibiae and tarsi pale orange yellow; anterior and posterior lobe of metathoracic peritreme

black; abdominal sterna dark orange hazel. Structure. Head: Rostrum reaching anterior border of meta-

sternum. Pronotum: Humeral angles expanded, apically acute; posterolateral margin denticulate; trian-

gular process shorter (Fig. 6). Measurements. Head length including the tylus: 1.45 mm; width across

eyes: 2.50 mm; interocular space: 0.98 mm; interocellar space: 0.44 mm; preocular distance: 0.88 mm;

length of antennal segments: I, 3.68 mm; II, 2.96 mm; III, 2.44 mm; IV, 6.52 mm. Pronotal length: 3.60

mm; width across frontal angles: 1.68 mm; width across humeral angles: 6.00 mm. Scutellar length:

2.20 mm; width: 1.92 mm. Total body length: 17.70 mm.

Discussion.—Like S. egeri with antennal segments I to III yellow or orange

and apically black to reddish brown, the pronotal disc, corium, and acetabulae

without creamy yellow marks, and tibiae and tarsi pale yellow. Salapia kondra-

tieffi, differs in having the abdominal sterna III to VI without pale yellow discoidal

spots, humeral angles narrow with an acute longer spine (Fig. 6) and antennal

segment IV black to reddish brown with yellow ring near middle third. In S. egeri

the abdominal sterna III to VI have pale yellowish spots, the humeral angles are

broadened with a short, broad apical spine (Fig. 4), and antennal segment IV is

yellow with apical third black to reddish brown.

Distribution.—Known only from Brazil.

Etymology.—Named for Dr. Boris C. Kondratieff, distinguished American Ple-

copterist.

SALAPIA ONOREI BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 7, 14)

Type.—Holotype: female; data: Ecuador. Sucumbios, San Pablo, Rio Aguarico,

October 1995, E Nischk. Deposited in the “‘Departamento de Entomologia, Pon-

tificia Universidad Catélica del Ecuador.”’

Description.—Female (holoty pe). Dorsal coloration: Head and corium dark orange hazel; antennal

segment I dark orange hazel, II and II yellow orange, and IV pale yellow; pronotum dark orange

hazel with humeral angles reddish brown, and a black triangular spot on middle third with the base

behind calli, and the apex close to posterior border; scutellum black to reddish brown; clavus dark

orange hazel with claval commissure brown; hemelytral membrane ambarine, with basal angle and

each margin dark brown; connexival segments dark orange hazel with anterior and posterior joint

black; dorsal abdominal segments black with lateral margins and a longitudinal stripe running from

II to VI segment yellow. Ventral coloration: Head pale orange hazel with middle third brown; rostral

segment I, and basal third of II pale orange hazel, middle and posterior third of segment II and basal

half of III reddish brown, and apical half of III and IV dark orange hazel; buccula pale orange hazel;

thorax pale orange yellow with following areas reddish brown: acetabulae (outer edge pale orange

140 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Figure 14. Dorsal view of Salapia onorei Brailovsky and Barrera, NEW SPECIES.

yellow), posterior and outer margin of propleura, mesopleura and metapleura, and prosternum, me-

sosternum, and metasternum; anterior and posterior lobe of metathoracic peritreme black; coxae dark

orange hazel; trochanters, femora, tibiae, and tarsi pale yellow with femoral spines black (hind femora

pale orange yellow); abdominal sterna dark orange hazel with posterior margin of sterna HI to VII,

anterior and posterior joint of pleural sterna III to VII, gonocoxae I (except external angle), and outer

margin of paratergite VIII and IX black. Structure. Head: Rostrum reaching posterior border of me-

sosternum. Pronotum: Humeral angles markedly expanded, with the spine large and robust; postero-

lateral margin conspicuously denticulate; triangular process robust, well developed (Fig. 7). Measure-

ments. Head length including the tylus: 2.12 mm; width across eyes: 2.64 mm; interocular space: 1.18

mm; interocellar space: 0.53 mm; preocular distance: 1.28 mm; length of antennal segments: I, 3.88

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 141

mm; II, 3.00 mm; III, 2.41 mm; IV, 7.64 mm. Pronotal length: 5.07 mm; width across frontal angles:

1.88 mm; width across humeral angles: 8.04 mm. Scutellar length: 2.60 mm; width: 2.54 mm. Total

body length: 19.82 mm.

Discussion.—This new species superficially resembles S. signata (Dallas) in

size and shape. S. onorei is, however, readily distinguishable by the humeral

angles markedly expanded, posterolateral margins conspicuously denticulate, with

triangular process well developed (Fig. 7), pronotal disc and scutellum without

creamy yellow longitudinal stripe, and corium without transverse creamy yellow

vitta, all of them present in S. signata.

Like S. humeralis (Burmeister), the antennal segments II and III are entirely

yellow to orange. In S. humeralis the humeral angles are scarcely expanded, the

spine shorter and the triangular process quite small (Fig. 5), the pronotal disc is

shiny orange with humeral angles pale yellow, and the acetabulae and posterior

margin of propleura, mesopleura, and metapleura creamy yellow. S. onorei differ

in the shape of the humeral angles, the length of the triangular process (Fig. 7),

and on the general coloration.

In S. egeri, the triangular process of the pronotum are broad, and markedly

exposed, both the antennal segments II and HI are yellow with apical third black,

and the humeral angles are broadened, with the spine shorter and robust (Fig. 4).

Distribution.—Known only from Ecuador.

Etymology.—Named for Dr. Giovanni Onore, distinguished Ecuadorian en-

tomologist.

SALAPIA HUMERALIS (BURMEISTER)

(Figs. 5, 15)

Paryphes humeralis Burmeister 1835: 336.

Salapia humeralis, Stal 1868: 50.

Types.—Lectotype (here designated): female; data: BRAZIL, Bahia, Gomez nr

1157. Deposited in Museum der Humboldt Universitat zu Berlin, Germany. Para-

type: 1 female; data: same data as lectotype. Deposited in Museum der Humboldt

Universitat zu Berlin, Germany.

Redescription.—Female (lectotype). Dorsal coloration: Head shiny orange hazel with tylus and

antenniferous tubercles yellow; antennal segment I reddish orange with apical joint yellow; segments

II and HI yellow; segment IV mutilated; pronotum reddish orange with calli orange hazel, and humeral

angles yellow; scutellum reddish orange with apex yellow; corium and clavus yellow; hemelytral

membrane dark ambarine, with basal angle, veins, and external edge darker; connexival segments III

to VI with upper margin yellow and inner margin reddish orange hazel, and segments VII to IX orange

hazel; dorsal abdominal segments orange hazel with black longitudinal and irregular stripe running

from segment II to VI. Ventral coloration. Head including the buccula and rostral segments I to IV

shiny orange hazel; thorax shiny orange hazel with acetabulae, posterior margin of propleura, and

small spot on posterior margin of mesopleura creamy yellow, with following areas shiny reddish

brown: middle third of propleura, anterior angle of acetabulae, and great portion of mesopleura and

metapleura; coxae shiny orange hazel with external face reddish brown; trochanters with outer face

reddish brown, and inner face shiny orange hazel; femora reddish brown; tibiae with two reddish

brown stripes and two yellow stripes; tarsi yellow; abdominal sterna shiny orange hazel with pleural

margins III to VII, and a broad rectangular spot on posterior margin of each sterna and lateral to

middle line orange; genital plates shiny orange hazel. Structure. Head: Rostrum reaching posterior

border of mesosternum. Pronotum: Humeral angles slightly expanded, apically with short and robust

spine; posterolateral margin scarcely denticulate; triangular process conspicuously reduced (Fig. 5).

Measurements. Head length including the tylus: 2.00 mm; width across eyes: 2.80 mm; interocular

142 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Figure 15. Dorsal view of Salapia humeralis (Burmeister).

space: 1.20 mm; interocellar space: 0.54 mm; preocular distance: 0.92 mm; length of antennal seg-

ments: I, 3.32 mm; II, 2.64 mm; III, 2.24 mm; IV, mutilated. Pronotal length: 3.36 mm; width across

frontal angles: 1.80 mm; width across humeral angles: 5.60 mm. Scutellar length: 2.20 mm; width:

2.08 mm. Total body length: 17.96 mm.

Discussion.—Pary phes humeralis Burmeister (1835) is known only from the

female lectotype and female paralectotype, here designated, which bears the labels

“Brazil, Bahia, Gomez, nr. 1157’’ and deposited in the Museum der Humboldt

Universitat zu Berlin, Germany. It was referred to the genus Salapia by Stal

(1868).

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 143

Figure 16. Dorsal view of Stenometapodus guttifer (Stal).

This species is distinguished from the other species of the genus by the follow-

ing characters: antennal segments II and III plus clavus and corium entirely yel-

low; pronotal disc reddish orange with calli orange hazel and humeral angles

yellow; abdominal sterna with broad rectangular spot orange yellow occupying

the posterior margin of sterna III to VII and lateral to middle line; width across

eyes conspicuously developed, longer than 2.70 mm; humeral angles of pronotum

slightly expanded, and apically with short and robust spine (Fig. 5).

Distribution.—Only known from Brazil.

144 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

STENOMETAPODUS GUTTIFER (STAL), NEW COMBINATION

(Fig. 16)

Petalops guttifer Stal 1859: 456-457.

Salapia guttifera, Stal 1868: 50.

Petalops guttifer Stal (1859) is known only from the male lectotype, here des-

ignated, which bears the labels “‘Brazil, Rio Janeiro, Berol, nr. 1476”’ and depos-

ited in Museum der Humboldt Universitat zu Berlin, Germany. It was referred to

the genus Salapia by Stal (1868), and here transferred to the genus Stenometa-

podus Breddin (1903) with the binomius Stenometapodus guttifer, NEW COM-

BINATION.

This species clearly belongs in the widespread South American genus Steno-

metapodus (Brailovsky 1984) by the following characters: hind tibiae expanded,

and longer than abdomen, posterior angles of pronotum acute, scutellum usually

with erect setae, and posterior angle of connexival segments V and VI spined. In

Salapia the hind tibiae are simple and cylindrical, and never expanded.

KEY TO SALAPIA SPECIES

1. Antennal segments II and III yellow to orange, with or without apical

[HORl ca aE Yo) Qn nn ee ee 8 eee ee ee ee eee ON eee 2,

1’. Antennal segments II and III black to reddish brown ................ 6

2. Antennal segments II and III yellow to orange, with apical third black

Be Ais TE oN eae Se Ses. guaeaee Cae A Aaah ge NC Lan, IS Deon ean y ee 3

2'. Antennal segments II and III entirely yellow to orange .............. 5

3. Pronotal disc and scutellum pale orange hazel with wide longitudinal vitta

creamy yellow; acetabulae pale creamy yellow ...... S. signata (Dallas)

3'. Pronotal disc and scutellum pale or dark orange hazel without creamy

yellow longitudinal vitta; acetabulae orange hazel, and unicolorous

WE Me LM OLAX mee CPS tape ncn nee eee eee 22 POL ee 4

4. Abdominal sterna III to VI with large pale yellow discoidal spot, laterally

to midline; humeral angles broad, with short, robust spine (Fig. 4);

triangular process broad, well developed (Fig. 4) ..............

See Re UN 22 US S. egeri Brailovsky and Barrera, NEW SPECIES

4’. Abdominal sterna III to VI without pale yellow discoidal spot; humeral

angles narrowed, with acute, longer spine (Fig. 6); triangular process

smaller, scarcely developed (Fig. 6) ..................0- 22 eeee

ee eee hee Ye S. kondratieffi Brailovsky and Barrera, NEW SPECIES

5. Humeral angles markedly expanded, with the spine longer and acute (Fig.

7); triangular process of pronotum conspicuously developed (Fig. 7);

acetabulae reddish brown; posterior margin of propleura, mesopleura

and: metaploura-reddish -browin- 0. 0.0 s Pees ee ee

Bead Wag eee aD Pee: UP S. onorei Brailovsky and Barrera, NEW SPECIES

5'. Humeral angles scarcely expanded, with the spine shorter (Fig. 5); tri-

angular process of pronotum smaller (Fig. 5); acetabulae creamy yel-

low; posterior margin of propleura, mesopleura, and metapleura

ereamy Velow ac 2 5 Rel ce eee S. humeralis (Burmeister)

6. Scutellum yellow to orange, with or without apex black ............. 7

6’. Scutellum black to reddish brown ..............000 0c c eee 10

1999 BRAILOVSKY & BARRERA: NEW SALAPIA SPECIES 145

7. Head and buccula black; thorax black to reddish brown

7’. Head and buccula yellow to orange; thorax pale orange .............. 9

8. Femora black to reddish brown; pronotal disc entirely yellow

Ee PI es, ta hi RA OR aan ich ee at S. luteola Brailovsky

8’. Femora yellow to orange; pronotal disc not entirely yellow

Ah, eRe rn POU NG, LAD al actin We SPER. MS Se S. pretiosa Blote

9. Femora orange; clavus yellow to orange with broad longitudinal stripe

black; corium yellow to orange with five black maculae between veins

i, SE NE Er ey, S. deckerti Brailovsky and Barrera, NEW SPECIES

9’. Femora reddish brown; clavus black with basal third orange; corium

black with yellow irregular transverse fascia ..................

Me PRPR ean rb 3 Bo oa S. vanduzeei Brailovsky and Barrera, NEW SPECIES

10. Buccula black; thorax black to reddish brown ..................... 11

10’. Buccula yellow; thorax yellow to orange .............. 000 c eee eae 15

11. Abdominal sterna black, with pleural margin yellow to orange ...... 12

11’. Abdominal sterna yellow to orange ............... 00 cee ee eee eee 13

12. Antennal segment IV black; dorsal abdominal segments III to V black;

anterolateral margin and anterior lobe of pronotum yellow. ....

et ee ae S. caucalandia Brailovsky and Barrera, NEW SPECIES

12’. Antennal segment IV yellow with basal and apical third reddish brown;

dorsal abdominal segments III to V orange; anterolateral margin and

anterior lobe of pronotum black .................. S. nigra Brailovsky

13. Corium yellow to orange, with apical margin and elongate longitudinal

stripe on endocorium black ....................4. S. dimidiata (Dallas)

13’. Corium black, with or without transverse vitta yellow to orange ..... 14

14. Corium entirely black; antennal segment IV with small yellow ring ...

dee eee i Ch ne, Bae, Ott na S. abdominalis (Dallas)

14’. Corium black with transversal vitta yellow to orange; antennal segment

IV with broad yellow ring. ................... S. baraquini (Signoret)

15. Pleural margin of abdominal sterna IV to VII bicolorous; clavus black

to reddish brown with posterior third yellow; corium with yellow and

broad transverse vitta .......... 0.20.2 e eee eee S. selecta Brailovsky

15’. Pleural margin of abdominal sterna IV to VII unicolorous; clavus black

or yellow, on the last condition with only the vein black; corium with

slender or without yellow transverse vitta

16. Clavus black or reddish brown; corium black, with ochre, slender and

irregular transversal vitta ............0..000000- S. haenschi (Breddin)

16’. Clavus yellow with the vein black; corium yellow to orange with the

apical margin and elongate stripe on endocorium black .......

ere a re ee ee ee ts ee S. pallida Brailovsky

ACKNOWLEDGMENT

We are indebted to the following individuals and institutions for the loan of

specimens and other assistance relevant to this study: James K. Liebherr and E.

Richard Hoebeke (Cornell University, Ithaca, New York); Boris Kondratieff (Col-

orado State University, Fort Collins, Colorado); R. L. Brown (Mississippi State

University, State University, Mississippi); Giovanni Onore (Pontificia Universidad

146 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Catélica del Ecuador, Quito); Jurgen Deckert (Museum der Humboldt Universitat

zu Berlin, Germany). Special thanks are extended to Jesus Contreras and Cristina

Urbina for the preparation of dorsal view illustrations.

LITERATURE CITED

Brailovsky, H. 1984. Una nueva especie del Género Stenometapodus Breddin y algunas notas acerca

de Empedocles tenuicornis (Westwood) (Hemiptera-Heteroptera-Coreidae-Acanthocephalin1).

Anales Inst. Biol. Univ. Nal. Autén. México (1983), Ser. Zool., 54: 63-68.

Brailovsky, H. 1992. El Género Salapia con descripcién de cuatro especies nuevas (Hemiptera-Het-

eroptera-Coreidae-Acanthocephalini). Anales Inst. Biol. Univ. Nal. Autén. México, Ser. Zool.,

63: 47-59.

Breddin, G. 1901. Neue neotropische Wanzen. Soc. Ent., 16: 41-42.

Breddin, G. 1903. Beitrage zur Hemipteren fauna der Anden. Sitz. Ges. Nat. Freunde Berlin., 1903:

366-383.

Burmeister, H. 1835. Handbuch der Entomologie Zweiter Band: IJ Ordnung Rhynchota. Berlin: Theod.

Ehr. Friedr. Enslin, i-iv: 1—400.

Stal, C. 1859. Till kannedomen om Coreidae. Ofvers. Kong]. Vet.-Akad. Foérh., 16: 449-475.

Stal, C. 1868. Hemiptera Fabriciana I. K. Svenska Vet.-Akad. Handl., 7: 1-148.

Received 9 Sep 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 147-152, (1999)

SUNLIGHT AVOIDANCE COMPARED BETWEEN

HESPEROPSIS GRACIELAE (MACNEILL) (LEPIDOPTERA:

HESPERIIDAE) AND BREPHIDIUM EXILIS (BOISDUVAL)

(LEPIDOPTERA: LYCAENIDAE)

W. D. WIESENBORN

U.S. Bureau of Reclamation, RO. Box 61470,

Boulder City, Nevada 89006-1470

Abstract—tTolerance of solar radiation by the sympatric butterfly species, Hesperopsis gracielae

(MacNeill) and Brephidium exilis (Boisduval), was compared. Adults in varying air temperatures

(30-—40° C) were exposed to different intensities of direct sunlight (13.8-110 kilolux), and the

elapsed response times were recorded when butterflies avoided continued exposure by flying to

shade. Avoidance response times (transformed log [Y + 1]) were shorter in H. gracielae (re-

transformed mean = 44 sec) than B. exilis (102 sec) across all air temperatures and light inten-

sities. Air temperature (affecting the body-temperature increase needed to stimulate flight) and

light intensity (affecting the rate of heating) independently influenced the species’ tolerance of

sunlight. Avoidance response times decreased linearly with increasing air temperature and hy-

perbolically with increasing light intensity. Rates of decrease did not differ between species.

Brephidium exilis’s more prolonged exposure to sunlight contradicts its smaller size and larger

ratio of body surface-area: volume (0.99) compared with H. gracielae (0.78). Hesperopsis gra-

cielae appears physiologically less adapted than B. exilis to radiation exposure and more readily

exploits shade from its hostplant to maintain a lower body temperature.

Key Words.—Insecta, Lepidoptera, comparative thermoregulation, Hesperiidae, Hesperopsis gra-

cielae, Lycaenidae, Brephidium exilis.

MacNeill’s sootywing, Hesperopsis gracielae (MacNeill), is a small (wing-

spread ~ 23 mm) dark-brown butterfly (Lepidoptera: Hesperiidae) found along

the lower Colorado River and near the river along its tributaries in southeastern

California, western Arizona, southern Nevada, and southern Utah (Scott 1986).

Two or three generations of H. gracielae occur from April to October (Emmel &

Emmel 1973, Austin & Austin 1980). Larvae of H. gracielae feed only on Atri-

plex lentiformis (Torrey) (Chenopodiaceae), a shrub found in dense clumps along

lower Colorado River drainages (Emmel & Emmel 1973). Hesperopsis gracielae

is more rare than the distribution of its hostplant (Austin & Austin 1980). In

Nevada, the butterfly’s rarity has afforded the species the conservation ranks of

‘G?S1’, signifying an unknown global (G) rarity and a state (S) rarity of critically

imperiled (<< 6 viable occurrences) (K. Goodwin, Nev. Nat. Heritage Program,

Carson City, personal communication; also see Master 1991).

Hesperopsis gracielae’s distinctive, characteristic tendency of flying within ri-

parian shrubs (MacNeill 1970) suggests the species may limit exposure to direct

sunlight (solar radiation) to prevent overheating in the high insolation and summer

air temperatures within its range (Wiesenborn 1998). The present study further

tests this hypothesis by comparing H. gracielae’s avoidance of sunlight with that

of the pigmy blue, Brephidium exilis (Boisduval). Brephidium exilis (here ssp.

exilis) is a smaller (wingspread ~ 16 mm) brown and white butterfly (Lepidoptera:

Lycaenidae) sympatric with H. gracielae (Scott 1986). Although H. gracielae and

B. exilis larvae both feed on A. lentiformis and are found along and near the lower

148 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Colorado River, B. exilis is less specialized, occurring in other low-altitude, al-

kaline habitats and feeding on other Chenopodiaceae (Scott 1986).

MATERIALS AND METHODS

The study site was located on the eastern edge of the Muddy River floodplain

at an elevation of 450 m near Bowman’s Reservoir, Clark County, Nevada. Av-

erage daily maximum air temperatures at Logandale near the site during 1968—

1992 in April, May, June, July, August, and September were 26.5, 31.8, 37.9,

40.7, 39.1, and 35.5° C, respectively (Nat. Oceanic & Atmospheric Admin., West-

ern Regional Climate Center, Reno, Nevada). The site supported a narrow, linear

band of A. lentiformis with a lesser amount of Pluchea sericea (Nuttall). Both B.

exilis and H. gracielae were frequently observed flying among the A. lentiformis

shrubs throughout the day.

The species’ avoidance of direct sunlight was determined similar to the method

used previously (Wiesenborn 1998). Insects were individually captured with an

aerial net and placed into a shaded 31 cm X 31 cm X 31 cm aluminum-frame

cage. The cage was covered on the bottom and on two sides with 13-mesh/cm

plastic screen, on one side with aluminum, on the top with clear vinyl, and on

one side with a cloth sleeve for inside access. The insect was allowed to acclimate

for 5 min, and the cage was repositioned with its aluminum side shaded and direct

sunlight transmitted through the top to illuminate one-half of the cage bottom. A

10-cm long A. lentiformis branch with 4—5 leaves was placed under the insect at

the beginning of each observation. The insect was allowed to walk or fly onto

the branch and placed in shade on the cage bottom for 1 min. Insects that flew

from the branch before the 1-min shading period had elapsed were placed back

onto the branch and the 1-min period repeated. The branch then was picked-up

and the insect exposed to direct sunlight passing through 8-mesh/cm organdy cloth

laid atop the cage. By using organdy cloth in layers (1, 2, 4, 8, or 16 layers),

light intensity striking the insect was varied without altering the sunlight spectrum

transmitted. Differences in basking posture required B. exilis to be exposed lat-

erally and H. gracielae dorsally, and the former’s walking required the branch to

be continually moved to maintain a constant lateral exposure. The time was re-

corded when the exposure was begun and when the insect flew from the branch.

Subtracting the former from the latter calculated the elapsed avoidance response

time in seconds. Observations were stopped after 20 min if flight did not occur

(2 of 70 observations, both of B. exilis shaded by the 16-layer treatment). Flights

from the branch always were to shade.

Each trial consisted of each light-intensity treatment tested once in random

order on an insect. Species were tested in random order with both species tested

once on 22 April and twice on 23, 24, and 28 April 1998. Trials were performed

under 0-—5% cloud cover between 10:14 PDT and 15:35 PDT and lasted 15—68

min each. Relative humidity was 18-38% and wind speed O—5 kmph. Light in-

tensity (measured with a Sekonic L-398 light meter) inside the cage was 4.1—10

kilolux (kIx) in shade and 103-120 klx in sunlight without shading by organdy

cloth. Air temperature in shade and light intensity beneath the organdy cloth were

measured inside the cage at the beginning of each observation.

Two H. gracielae and all seven B. exilis were collected after being tested.

Thoracic and abdominal widths (at midpoints) and lengths were measured with

E99 WIESENBORN: BUTTERFLIES AVOID SUNLIGHT 149

an ocular micrometer and used (assuming a cylindrical shape) to calculate thoracic

and abdominal surface areas (excluding cylinder ends), volumes, and area: volume

ratios. Ratios likely represent the effect of body size on heating rate, because

surface area would affect the amount of radiation absorbed, and volume would

affect the mass being heated. Collected specimens were verified as to species and

deposited as vouchers (G. Austin, Nev. St. Mus., Las Vegas, personal commu-

nication).

Avoidance response times were transformed log (Y + 1) and analyzed by mul-

tiple regression with cage air temperature, light intensity, and species as indepen-

dent variables (the latter as an categorical variable, Myers 1986). The regression

was improved (r? maximized and residuals most-randomly scattered) by trans-

forming light intensity 1/X. Cage air temperature was not related (F = 1.66; df

= 1,68; P = 0.20) to light intensity. The interactions species X light intensity,

species X air temperature, and air temperature X light intensity were individually

added to the regression model and tested for significance (P < 0.05). For presen-

tation, transformed avoidance response times were adjusted (Sokal & Rohlf 1981)

for transformed light intensity, then retransformed and plotted on logarithmic scale

against air temperature. Similarly, transformed avoidance response times were

adjusted for air temperature, then retransformed and plotted on logarithmic scale

against light intensity. Regression lines were fitted to transformed data for each

species and plotted after retransformation.

RESULTS

Combined thoracic and abdominal surface areas were 22 + 2.5 mm? (mean +

SD; n = 7) in B. exilis and 48 + 4.8 mm? (n = 2) in H. gracielae, and combined

thoracic and abdominal volumes were 22 + 4.2 mm}? in B. exilis and 61 + 7.2

mm?in#H. gracielae. Area: volume ratios were 0.99 + 0.096 in B. exilis and 0.78

+ 0.014 in H. gracielae, 1.3 times greater in B. exilis than in H. gracielae.

Avoidance response time was related to air temperature (F = 34.4; df = 1,66;

P < 0.001), light intensity (F = 99.2; df = 1,66; P < 0.001), and species (F =

30.0; df = 1,66; P < 0.001). Hesperopsis gracielae sought shade earlier (retrans-

formed mean = 44 sec) than B. exilis (retransformed mean = 102 sec), and both

species sought shade earlier with increasing air temperature (Fig. 1A) and light

intensity (Fig. 1B). After accounting for these variables, interactions were not

evident between species and air temperature (F = 0.327; df = 1,65; P = 0.57)

or species and light intensity (F = 2.52; df = 1,65; P = 0.12). The regression

lines in each plot (Fig. 1) therefore do not statistically diverge from parallel; the

two species did not differ in their rate of decrease in avoidance response time as

air temperature or light intensity increased. Air temperature and light intensity

also did not interact (F = 0.26; df = 1,65; P = 0.61), indicating independent

effects on avoidance response time. Regression (F = 46.3; df = 3,66; P < 0.001;

r?> = 0.68) of avoidance response time on air temperature, light intensity, and

species (B. exilis = 1, H. gracielae = 2) produced the following equation (co-

efficients = SEs):

log (s + 1) = 4.9 + 0.5 — 0.086 + 0.015 (@ C)

+ 29 + 3 (1/klx) — 0.44 + 0.08 (species)

150 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

10°

Avoidance Response Time (s)

ry

10

30 32 34 36 38 40

Air Temperature (°C) Light Intensity (klx)

Figure 1. Avoidance of sunlight by Brephidium exilis (circles and broken lines) and Hesperopsis

gracielae (X’s and solid lines): avoidance response time in seconds elapsed (A) adjusted for light

intensity (transformed 1/X) and plotted against air temperature and (B) adjusted for air temperature

and plotted against light intensity. Avoidance response times, plotted on logarithmic scales, were

retransformed to original scale after adjusting transformed log (Y + 1).

DISCUSSION

Flight from sunlight to shade by H. gracielae and B. exilis likely was a be-

havioral response to prevent overheating (Wiesenborn 1998); continual exposure

to sunlight elevated body temperatures until a threshold was reached stimulating

flight to shade. Although body temperatures in butterflies increase as the time of

exposure to sunlight increases, the rate of temperature increase diminishes as body

temperatures asymptote near an upper limit (Heinrich 1972, 1986; Wasserthal

1975). The curvilinear, asymptotic relationship described in these studies suggests

body temperature increases linearly with a proportional increase in exposure time,

a function linearized by transforming exposure time log X (Sokal & Rohlf 1981).

The log (Y + 1) transformation of avoidance response time (exposure time until

flight to shade) in the present study therefore agrees with the diminishing rate of

body temperature increase as previously determined.

The hyperbolic relationship between transformed avoidance response time and

light intensity (Fig. 1B) likely resulted from the latter being a rate (quantity per

time), equal to light energy per area illuminated per time. Rates frequently plot

as hyperbolic curves that are straightened by reciprocal transformation (Sokal &

Rohlf 1981).

The independent effects of air temperature and light intensity on avoidance

response time indicate these variables acted on the butterflies by different mech-

anisms. Air temperature, approximating the initial body temperature prior to ex-

posure to sunlight, contributed additively towards the body temperature increase

required to reach the threshold to stimulate flight. Light intensity, approximating

solar radiation, provided the sole energy influx driving body temperature upward.

Within species, higher light intensity resulted in greater rate of energy absorbance

and greater rate of temperature increase towards the flight threshold. Insect species

with greater rates of radiation absorbance heat more quickly as radiation intensity

increases (Digby 1955). Hesperopsis gracielae and B. exilis appear not to differ

1999 WIESENBORN: BUTTERFLIES AVOID SUNLIGHT 151

in radiation absorbance, because their rates of decrease in avoidance response

time did not differ with increasing radiation intensity.

Earlier flight to shade by H. gracielae due to greater energy absorbance and

body heating rate also is contradictory to the species’ larger body size, as larger

insects exposed to radiation typically heat more slowly (Heinrich 1986). Brephi-

dium exilis’s greater area: volume ratio expectedly would have caused it to heat

1.3 times faster than H. gracielae. Instead, B. exilis remained, while exposed to

sunlight, 2.7 times longer than H. gracielae. Because heating rate does not ap-

preciably differ between lateral and dorsal basking (Heinrich 1986), it is unlikely

B. exilis’s greater tolerance of direct sunlight is due to this behavioral difference.

Hesperopsis gracielae’s earlier flight to shade may have been due to its darker

coloration, increasing radiation absorbance. However, coloration incompletely in-

dicates the proportion of radiation absorbed, because visible light reflected off the

insect does not include the near-infrared, part of the spectrum that can contribute

significant warming (Heinrich 1972).

Equivalent heating rates between the two species would require H. gracielae

to have a lower flight threshold, or temperature tolerance, compared with B. exilis.

The difference between the species’ flight-threshold temperatures can be estimated

from the plot of avoidance response time against air temperature (Fig. 1A). The

mean avoidance response time (retransformed = 67 sec) for both species corre-

sponds to an air temperature of 33° C for H. gracielae and 38° C for B. exilis.

The two species would have responded at the same time if subjected to these two

air temperatures and exposed to the same light intensity. The 5° C air temperature

difference between species therefore estimates the difference between flight-

threshold temperatures; B. exilis tolerated body temperatures 5° C higher than H.

gracielae assuming equivalent heating rates. Brephidium exilis’s tolerance of high

body temperatures resembles that found in dragonflies, where desert species tol-

erate body temperatures 4—9° C higher than species found in cooler regions (Pol-

cyn 1994).

Hesperopsis gracielae is less able to tolerate direct sunlight and therefore less

adapted to the high insolation and air temperatures of its environment. Rather

than tolerating high body temperatures, H. gracielae appears to maintain lower

body temperatures by flying within the shade of its hostplant, A. lentiformis. Thus

H. gracielae’s specialization on A. lentiformis may in part be due to its need for

a foodplant providing adequate canopy cover. This concept is supported by con-

sidering Hesperopsis alpheus (Edwards), a species closely related to H. gracielae

that is more widely-distributed and feeds on Atriplex canescens (Pursh) Nuttall

(MacNeill 1970). Of the two insect species, only H. gracielae inhabits the lower

Colorado River, while H. alpheus is limited to higher elevations (> 1500 m) and

cooler climates (Emmel & Emmel 1973; G. Pratt, UC Riverside, personal com-

munication). Both Atriplex species are found in the lower Colorado River habitats

of H. gracielae (Turner et al. 1995). Atriplex lentiformis is up to 1 m taller than

A. canescens and provides a more dome-shaped canopy (Turner et al. 1995); H.

gracielae’s exploitation of A. lentiformis’s greater cover likely allows the insect

to inhabit an otherwise inhospitable climate. Indeed, H. alpheus does not exhibit

H. gracielae’s habit of flying for prolonged periods within shrubs (MacNeill

1970).

In contrast to H. gracielae, B. exilis’s wide host range does not allow it to

152 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

consistently utilize host shade as a means of thermoregulation, instead requiring

the species to be physiologically better adapted to desert climate. For example,

one of B. exilis’s most-used hosts is Atriplex semibaccata Robert Brown (Emmel

& Emmel 1973), an exotic plant common at low elevations whose prostrate

growth form (Munz 1974) would offer butterflies little protection from sunlight.

Tolerance of direct sunlight expectedly also is required by B. exilis’s migratory

behavior (Scott 1986), reducing the species’ ability to remain sheltered within

plants.

Conservation activities intended to benefit H. gracielae should consider the

species’ requirement for cover. In addition to furnishing its other life requisites,

such as nectar sources for adults (Wiesenborn 1997), restored or preserved ripar-

ian habitat should provide A. lentiformis patches large enough, and contiguous

enough, to allow prolonged flight within host canopies. It is unclear if neighbor-

ing, alternative plant species of adequate canopy would by themselves satisfy the

insect’s shade requirement, because oviposition behavior and plant cover may be

interrelated. Regardless of larval suitability, A. lentiformis shrubs offering inad-

equate canopy may not be selected by ovipositing females.

ACKNOWLEDGMENT

The author is grateful to G. E Pratt of the Entomology Department at UC

Riverside for his helpful comments on the manuscript.

LITERATURE CITED

Austin, G. T. & A. T. Austin. 1980. Butterflies of Clark County, Nevada. J. Res. Lepid., 19: 1-63.

Digby, P. S. B. 1955. Factors affecting the temperature excess of insects in sunshine. Exp. Biol., 32:

279-298.

Emmel, T. C. & J. EK Emmel. 1973. The butterflies of southern California. Nat. His. Mus. Los Angeles

Co., Sci. Series, 26: 1-148.

Heinrich, B. 1972. Thoracic temperatures of butterflies in the field near the equator. Comp. Biochem.

Physiol., 43A: 459-467.

Heinrich, B. 1986. Comparative thermoregulation of four montane butterflies of different mass. Phy-

siol. Zool. 59: 616-626.

Master, L. L. 1991. Assessing threats and setting priorities for conservation. Conserv. Biol., 5: 559—

563.

MacNeill, C. D. 1970. A new Pholisora with notes on P. alpheus (Edw.) (Lepidoptera: Hesperiidae).

Entomol. News, 81: 177-184.

Munz, P. A. 1974. A flora of southern California. Univ. of Calif. Press, Berkeley.

Myers, R. H. 1986. Classical and modern regression with applications. Duxbury Press, Boston.

Polcyn, D. M. 1994. Thermoregulation during summer activity in Mojave Desert dragonflies (Odonata:

Anisoptera). Functional Ecol., 8:441—449.

Scott, J. A. 1986. The butterflies of North America: a natural history and field guide. Stanford Univ.

Press, Stanford, California.

Sokal, R. R. & FE J. Rohlf. 1981. Biometry (2nd ed.). W. H. Freeman & Co., New York.

Turner, R. M., J. E. Bowers & T. L. Burgess. 1995. Sonoran desert plants: an ecological atlas. Univ.

Ariz. Press, Tucson.

Wasserthal, L. T. 1975. The role of butterfly wings in regulation of body temperature. J. Insect Physiol.,

21: 1921-1930.

Wiesenborn, W. D. 1997. Hesperopsis graciliae (MacNeill) (Lepidoptera: Hesperiidae) flight between

hostplants and Prosopis glandulosa Torrey. Pan-Pacific Entomol., 73: 186-189.

Wiesenborn, W. D. 1998. Avoidance of direct sunlight by adult Hesperopsis gracielae (MacNeill)

(Lepidoptera: Hesperiidae). Pan-Pacific Entomol., 74: 157-162.

Received 13 Nov 1998; Accepted 20 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 153-158, (1999)

DESCRIPTION OF IMMATURE STAGES OF

PHY LLOPHAGA (TRIODONYX) LALANZA SAYLOR

(COLEOPTERA: MELOLONTHIDAE, MELOLONTHINAE)

MIGUEL-ANGEL MorRON!, SALVADOR HERNANDEZ-RODRIGUEZ?, &

AGUSTIN RAMi{REZ-CAMPOS?

‘Departamento de Entomologia, Instituto de Ecologia, A.C. Km 2.5 antigua

carretera a Coatepec, A.P. 63, Xalapa, Veracruz 91000, México

*Departamento Técnico de Campo, Ingenio de Puga, S.A. (AGA-Aziicar).

Domicilio conocido Francisco I. Madero, Nayarit 63000, México.

Abstract —Three larval stages and pupa of Phyllophaga (Triodonyx) lalanza Saylor are described

based on a large series of samples obtained in sugar cane fields from Tepic, Nayarit, Mexico,

where the larvae cause severe damage to the root system. Fully developed third-stage larvae

(58-64 mm length) of this species are some of the biggest root-feeding white grubs in Latin

America. Illustrations of diagnostic structures of third-stage larva and pupa are included.

Key Words.—Insecta, white grubs, morphology, taxonomy, sugar cane.

Restimen.—Se describen los tres estados larvarios y la pupa de Phyllophaga (Triodonyx) lalanza

Saylor, con base en series grandes de muestras obtenidas en los cafiaverales de la regidn de

Tepic, Nayarit, México, donde las larvas causan dafios severos al sistema radicular. Las larvas

de tercer estadio completamente desarrolladas alcanzan entre 58 y 64 mm de longitud, por lo

cual representan una de las larvas edaficolas riz6fagas mas grandes descritas hasta el momento

en América Latina. Se ilustran las estructuras diagnésticas para la identificacidn de la larva de

tercer estado y la pupa.

In recent years (1993-1996) reports of damage by white grubs in sugar cane

fields, black bean, chili pepper, and corn crops have increased in the state of

Nayarit, Mexico. Larvae have seriously damaged nearly 1000 ha of sugar cane

around Tepic city. Nearly 90% of the samples of larvae and adults obtained and

studied by the authors were determined as Phyllophaga (Triodonyx) lalanza Say-

lor (Mor6én et al. 1996). In this paper, the third instar larva and pupa of this

species are described, including the taxonomic characters and measures of the

first and second instar larvae, and the third instar larva is briefly compared with

the larvae of other species of Phyllophaga. Technical terms used are those of

Ritcher (1966) and Moron (1986). The descriptions are based on 160 third instar

larvae, 64 second instar larvae, 16 first instar larvae, two cast skins of third instar

larvae and 8 pupae, reared from eggs or directly collected in the field (see Material

Examined). Specimens studied are deposited in the following Mexican collections:

Instituto Nacional de Investigaciones Forestales, Agricolas y Pecuarias, SAGAR,

Celaya, Guanajuato; Instituto de Biologia, U.N.A.M. Mexico City; Departamento

de Investigaciédn en Ciencias Agricolas, Benemérita Universidad Aut6énoma de

Puebla, Puebla; Instituto de Ecologia, A.C. Xalapa, Veracruz; Departamento Téc-

nico de Campo, Ingenio de Puga, S.A. Tepic, Nayarit; the private collection of

M.A. Moron, Xalapa, Veracruz; and in the collections of California Academy of

Sciences, San Francisco; U.S. National Museum, Smithsonian Institution, Wash-

ington, D.C.; University of Nebraska State Museum, Lincoln; and Biosystematics

Research Centre, Ottawa, Canada.

154 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Figure 1. Phyllophaga lalanza third stage larva, lateral aspect. Chaetotaxy simplified. Scale line

is 5 mm.

PHYLLOPHAGA (TRIODONYX) LALANZA SAYLOR, 1941

(Figs. 1-15)

Third Instar Larva.—Head. Maximum width of head capsule 7.4—8.0 mm. Surface of cranium finely

and densely rugopunctate, reddish yellow. Frons (Figs. 1 and 2) with only 1 exterior frontal seta on

each side; 26—30 anterior frontal setae; 30—54 posterior frontal setae, laterally mixed with 8—12 exterior

frontal setae on each side; each anterior angle of frons with 2 setae; remaining cranial surface with

3—4 dorso-epicranial setae, 3 epicranial setae, 18—20 para-ocellar setae on each side. Clypeus (Fig. 2)

with 4 lateral setae. Labrum slightly asymmetrical, lateral margins rounded, 12 posterior setae, 5—7

central setae, and 2-3 lateral setae. Eye spots absent. Epipharynx (Fig. 3) without zygum and epizy-

gum; haptomerum with group of 30-35 heli; each plegmatium with 13 to 18 short plegmata. Pro-

plegmatia absent. Dexiophoba and laeophoba large, extending forward from sense cone for more than

one half distance between sense cone and heli. Laeotorma with 2-3 anterior, short processes. Hapto-

lachus with few microsensillae. Sclerotized plate not developed, but with 2 macrosensillae between

sense cone and dexiotorma. Crepis well sclerotized with both ends bifurcated. Each acanthopariae

with 14-16 curvate, spine-like setae. Chaetoparia moderately developed, with few microsensilla among

the setae. Mandibles (Figs. 4-7) with ventral stridulatory areas absent. Scissorial area of left mandible

with distal blade-like portion separated from proximal tooth by scissorial notch; inner margin without

tooth; molar area with well developed distal lobe (M1); dorsomolar setae absent; acia long, slightly

acute; brustia multisetose. Scissorial area of right mandible formed by 3 short teeth; inner margin

without tooth; molar area with 2 irregular lobes; calx enlarged; brustia multisetose. Galea with 1 well

developed uncus (Fig. 8); lacinia with 3 terminal unci in a longitudinal row, fused at bases, and each

side with 4—6 stout heli in a longitudinal line (Fig. 9). Maxillary stridulatory area with 16—19 sharp-

pointed, anteriorly directed teeth, without anterior process (Fig. 10). Hypopharyngeal sclerome asym-

metrical, produced on right side into broadened process (Fig. 8). Dorsal surface of last antennal

segment with 1 large, oval sensory spot (Fig. 11). Thorax. Thoracic spiracles 0.8—0.9 mm measured

dorsoventrally; respiratory plate reddish yellow, regularly shaped as a closed ‘“‘C’’; distance between

2 lobes of respiratory plate less than dorsoventral diameter of bulla; spiracular bulla rounded, slightly

1999 MORON ET AL.: PHYLLOPHAGA IMMATURES 155

Me 3

DRey Mee

ce

key Te bs ate es gi Ps

fs a Wy : PF! a om

Seu

yw

Figures 2-7. Phyllophaga lalanza Saylor third-stage larva. Figure 2. Frontal view of cranium.

Figure 3. Epipharynx. Figures 4 and 5. Dorsal aspect of left and right mandibles, respectively. Figures

6 and 7. Ventral aspect of right and left mandibles, respectively. Scale lines are 1 mm.

prominent. Pronotum with 2 well-marked, yellowish, lateral scleromes at each side (Fig. 1). Dorsa of

pro-, meso- and metathorax with sparse, slender setae, without short spinelike setae. Tarsal claws of

prothoracic and mesothoracic legs similar, large, each claw bearing 2 setae (Fig. 1). Claws of meta-

thoracic legs reduced in size (Fig. 1). Abdomen. Abdominal spiracles of segments 1 to 4 nearly similar

in size, 0.8-0.9 mm measured dorsoventrally; respiratory plate reddish yellow, regularly shaped as a

closed ‘‘C’’; distance between 2 lobes of respiratory plate much less than dorso-ventral diameter of

bulla; spiracular bulla rounded, slightly prominent (Fig. 12). Respiratory plate with a maximum of

about 18 to 24 oval “‘holes” along any diameter; “holes” not in definite rows. Spiracles of segments

5 to 7 reduced in size, 0.65—0.70 mm; distance between 2 lobes of respiratory plate slightly less than

dorso-ventral diameter of bulla. Spiracles of segment 8 noticeably smaller, 0.50-0.52 mm; distance

between 2 lobes of respiratory plate equal to dorsoventral diameter of the bulla (Fig. 1). Dorsum of

abdominal segments 2 to 7 with large number of spinelike setae and some sparse, long, slender setae;

dorsum of segments 8 to 10 with some paired, sparse, long, slender setae (Fig. 1). Venter of segments

1 to 9 with paired, sparse, slender setae. Raster with pair of longitudinal palidia which are slightly

Vol. 75(3)

THE PAN-PACIFIC ENTOMOLOGIST

156

Ge ..

= o

o

o

oc

vt Gd

"aN gel

ae eee Ree

— 5 we 7

Pe FE — To

x, ~. on Ce

a id Vas y “ae

1a~ 2 ST

~ ~ N ~~’ 7 eS

> ae

iy ~y ». ee

> “— a3 4 yY ao US

q nN N\ nd i

“ o> eles

a rpPAe Ce |

~ a* am: 4% ~

14. nS

ale a

awe thf

Figures 8-14. Phyllophaga lalanza third-stage larva. Figure 8. Dorsal aspect of left maxilla and

labium. Figure 9. Apex of right lacinia, mesial view. Figure 10. Stridulatory area of right maxilla.

Figure 11. Dorsal aspect of apical antennal segment. Figure 12. Fourth abdominal spiracle. Figure 13.

Tarsal claw of fore leg. Figure 14. Palidia and teges. Scale lines are 1 mm.

1999 MORON ET AL.: PHYLLOPHAGA IMMATURES 157

Figure 15. Phyllophaga lalanza male pupa, lateral aspect. Note the humeral peg-like projection

on the elytral tecus (hp), the dioneiform organs (do) and genital ampulla (ga). Scale line is 5 mm.

convergent anteriorly and posteriorly; each palidium formed by 27 to 37 short, spinelike pali (Fig.

14). Teges formed by 65 to 75 hooked, short setae. Campus with 4 long, slender setae. Barbula dense,

with long setae. Dorsal anal lobe with large number of short setae. Lower anal lobe divided by sagittal

cleft, with some sparse short setae. Anal slit ““Y’’-shaped, stem of “‘Y’’ shorter than arms of “‘Y”’.

Approximate total body length: 58-64 mm.

Second Instar Larva——Similar to third instar except as follows: maximum width of head capsule:

4.70—5.25 mm; frons with 30-52 posterior frontal setae; dorso-ventral diameter of prothoracic spiracles

0.40-0.55 mm; length of metacoxae: 3.0—3.5 mm; each palidium with 29-38 pali.

First Instar Larva—Similar to second instar except as follows: maximum width of head capsule:

2.9-3.3 mm; frons with 15-26 posterior frontal setae; respiratory plates of thoracic and abdominal

spiracles kidney shaped; dorso-ventral diameter of prothoracic spiracles 0.25 mm; one thin eclosion

spine at each side of metanotum; length of metacoxae 1.85—2.00 mm; each palidium with 27-39 pali.

Pupa—Head. Glabrous, strongly reflexed downward; antennae and mouth parts clearly differenti-

ated, labrum much exposed, ocular canthi wide, compound eyes sunken (Fig. 15). Thorax. Pronotum

convex, with shallow depressions at sides and near middle, anterior lateral angles clearly projected.

Meso- and metanotum well differentiated. Elytral tecae with shallow, longitudinal sulci and each

humeral corner with short, tubercle-like, sclerotized callus. Wing tecae slightly longer than elytra.

Protibiae with 3 short processes on external borders. Mesotibiae with vague keels. Metatibiae without

keels and apical spurs well differentiated. Abdomen. Segments I-VI clearly wider than distal segments.

Two pairs of poorly developed dioneiform organs between tergites [V—VI. Pleural lobes rounded and

prominent. Spiracle I simple, ovate, with fine peritreme; spiracles II-IV rounded, with wide, prominent

peritreme and adjacent sclerotized, small dorsal plate; spiracles V—VI closed, vague; spiracles VII-—

VIII closed, vague, and surrounded by longitudinal rugae (Fig. 15). Last abdominal tergite with 2

long, sharply pointed, divergent urogomphi. Last abdominal “‘sternite” in males with smooth, prom-

inent genital ampulla; in females, flattened and deeply striated. Body length, 30-35 mm.

Remarks.—Except for its large size (58—64 mm length) and the high number

of long frontal setae, third instars of P. lalanza do not have any definite taxonomic

character that separates the larvae of the subgenus Triodonyx from the known

larvae of other subgenera of continental American Phyllophaga. The high number

of long frontal setae is useful to separate the first and second instars of this species

from other small species that live in the Nayarit area. Using the key to species

of Phyllophaga larvae from the U.S. (Ritcher 1966: 87), P. lalanza vaguely keyed

to near P. quercus Knoch and P. tristis Fab. because of the absence of propleg-

matia in the epipharynx, but it has a raster with closely set palidia formed by

more than 13 pali. Using the key to larvae from the Antillean and exotic Melo-

158 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

lonthini (B6ving 1942: 173), the larvae of the Antillean subgenera Clemora Saylor

or Cnemarachis Saylor show an irregular or dispersed arrangement of pali; Cle-

mora does not have proplegmata, and Cnemarachis has numerous proplegmata.

Some Asiatic species of Ancylonycha Dejean have regular pali and do not have

proplegmatia but have round, black spots on the dorsal side of the cardo, coxae,

and near the spiracles. So, the possible relationships of larvae of Phyllophaga

(Triodonyx) species remains obscure. One interesting character observed in the

pupa of P. lalanza is the presence of humeral, sclerotized, spine-like, elytral calla,

but, unfortunately, no other pupae of the genus are described in detail so that a

comparison can be made.

Material Examined——MEXICO: STATE OF NAYARIT, Tepic Municipality, Ejido FE I. Madero, 16

April 1994, S. Hernandez & M. A. Moron, soil under sugar cane roots, 760 m, 2 cast skins of third

instars associated with immature adults and 4 third instars; 14 third instars, same data except: 15

November 1994, S. Hernandez & A. Ramirez; 12 third instars, same data except: Ejido Pochotitan,

10 December 1994, 910 m; 22 third instars, same data except: 18 January 95, 910 m; 63 third instars,

same data except: Ejido El Refugio, 3 November 1994, 990 m; 10 third instars, same data except: 10

November 1994; 35 third instars, same data except: 5 December 1994. Two second instars reared from

eggs deposited by females collected in Ejido E I. Madero, 20 July 1994; 6 second instars from Ejido

Pochotitan, 17 October 1994, A. Ramirez & S. Hernandez, under sugar cane roots, 910 m; 31 second

instars, same data except: Ejido El Refugio, 17 August 1994; 25 second instars, same data except: 14

September 1994. Five first instars reared from eggs deposited by females collected in Ejido EF I.

Madero, 20 July 1994, 760 m, A. Ramirez; 11 first instars from Ejido El Refugio, 17 August 1994,

A. Ramirez & S. Hernandez, under sugar cane roots, 910 m. Four male and 4 female pupae collected

in Ejido Pochotitan, Potrero La Mesa, 23 April 1994, S. Hernandez, A. Ramirez & M. A. Moron,

inside pupal chamber located under sugar cane roots, 910 m.

ACKNOWLEDGMENT

We thank M. A. Gutierrez, EF Morales, M. J. de la Paz, E. Vallarta, R. Aguilar,

A. & I. Alvarez, J. & M. Carrillo, E. Correa, L. Flores, L. Gil, H. Hernandez, R.

Juarez, J. M. Lara, J. J. Medina, L. Nifiez, M. Panuco, M. Soto and R. Verdin

for their valuable aid during the field and laboratory work. Brett C. Ratcliffe

(University of Nebraska State Museum, Lincoln) critically reviewed an early draft

of the manuscript and helped with the English translation. Financial support for

the publication of this paper was provided by the Instituto de Ecologia, A.C.

(account 902-02).

LITERATURE CITED

Boving, A. G. 1942. Descriptions of the larvae of some West Indian Melolonthine beetles and a key

to the known larvae of the tribe. Proc. U.S. Nat. Mus., 92 (No. 3146): 167-176, plates 18-19.

Morén, M. A. 1986. El género Phyllophaga en México. Morfologia, distribucién y sistematica su-

praespecifica (Insecta: Coleoptera). Publ. No. 20. Instituto de Ecologia, México.

Moron, M. A., S. Hernandez-Rodriguez & A. Ramirez-Campos. 1996. El complejo “‘gallina ciega”’

(Coleoptera: Melolonthidae) asociado con la cafia de azticar en Nayarit, México. Folia Entomol.

Mex., 98: 1-44.

Ritcher, P.O. 1966. White grubs and their allies: a study of North American scarabaeoid larvae. Studies

in Entomology No. 4. Oregon State University Press, Corvallis.

Received 14 Jan 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 159-164, (1999)

TWO NEW SPECIES OF GIULIANIUM MOORE FROM

THE PACIFIC COASTS OF ALASKA AND CALIFORNIA

(COLEOPTERA: STAPHYLINIDAE: OMALIINAE)

KEE-JEONG AHN! AND JAMES S. ASHE?

‘Department of Biology, Chungnam National University, Daejeon City 305-764,

Republic of Korea

*Division of Entomology, Natural History Museum, Snow Hall,

University of Kansas, Lawrence, Kansas 66045, U.S.A.

Abstract—A systematic review of the omaliine genus Giulianium Moore is presented. Diagnosis

of Giulianium is presented, and three species are recognized, two of which are described as new

(G. alaskanum, NEW SPECIES, and G. newtoni, NEW SPECIES). Types and paratypes of the

two new species are designated. A key is provided for separation of known species of Giulianium

and illustrations of diagnostic features are provided.

Key Words.—Insecta, Coleoptera, Staphylinidae, Omaliinae, Giulianium, new species, intertidal.

While taking collecting trips to Alaska and Hokkaido (Japan) and studying

Field Museum of Natural History (Chicago, Illinois) collections, we discovered

two remarkable new species of intertidal staphylinid beetles. We concluded after

detailed study of these specimens that these species represented two new species

of Omaliine genus Giulianium Moore. This genus was first described and char-

acterized by Moore (1976) based on the new species G. campbelli from California,

U.S.A. He tentatively placed Giulianium in the subfamily Phloeocharinae. How-

ever, Newton & Thayer (1995) proposed that Giulianium should be placed in the

Omaliinae (tribe Aphaenostemmini Peyerimhoff) because members of Giulianium

lack the derived characters of the Phloeocharinae and have a well-developed Om-

aliine-type gland system (Thayer 1987). Their phylogenetic analysis indicated that

Giulianium showed a sister group relationship with Aphaenostemmus Peyerimhoff

(see Newton & Thayer 1995 for more detailed discussions).

Our objectives in this paper are to redescribe Giulianium, to describe two new

species (G. alaskanum and G. newtoni), and to provide a key for separation of

known species.

GIULIANIUM MOORE

Giulianium Moore, 1976: 56.

Type Species.—Giulianium campbelli Moore 1976, by original designation.

Diagnosis —Moore (1976) has provided a complete description of this genus. This diagnosis con-

denses and updates that description to include other species, corrects mistakes and adds characters not

mentioned by Moore. Members of Giulianium can be recognized by the following combination of

characteristics: small, length 2.1 mm-—3.0 mm; body uniformly dark brown to light brown (teneral

specimens often dark rufus with darker abdomens); body form slender, parallel-sided or abdomen

slightly wider than very slender head thorax and elytra, flattened; body densely pubescent with very

short and fine microsetae; integument moderately densely reticulate throughout. Head orbicular to

slightly elongate (Figs. 8-10), narrowed at base to broad neck, nuchal constriction faint to absent

dorsally. Gular sutures separated, diverged to base of head. Infraorbital carina absent. Eyes very small,

several setae between facets. Tempora long. Antennae long, reaching to posterior fourth or to beyond

apex of elytra when extended posteriorly, all antennomeres elongate (Figs. 11-13). Labrum (Fig. 1)

160 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Figures 1-7. G. alaskanum. Figure 1. Labrum, dorsal aspect; Figure 2. Epipharynx, ventral aspect;

Figure 3. Mandible, ventral aspect; Figure 4. Maxilla, dorsal aspect; Figure 5. Labium, dorsal aspect;

)

Figure 6. Hypopharynx, dorsal aspect; Figure 7. Hind tarsus, lateral aspect. Scales 0.1 mm.

1999 AHN & ASHE: NEW GIJULIANIUM SPECIES 161

8 9 10

Ie ae ig ky Le GE

Figures 8-10. Body shape, dorsal aspect and Figs. 11-13. Antenna, lateral aspect. Figure 8. G.

campbelli; Figure 9. G. alaskanum, Figure 10. G. newtoni; Figure 11. G. campbelli; Figure 12. G.

alaskanum, Figure 13. G. newtoni. Scales 0.1 mm.

deeply and broadly emarginate anteriorly, appearing bilobed, anterior margin with diffuse fringe of

fine setae. Epipharynx as in Fig. 2. Mandible (Fig. 3) with acute apex, and finely ciliate prostheca,

without enlarged molar region. Maxilla as in Fig. 4; lacinia with 2 large spines apically and numerous

spinose setae and smaller setae along inner margin; galea with numerous moderately long setae at

apex, 4—6 widely separated large setae on inner margin and 1 large seta externally near base of densely

setose area. Maxilla with palpomere 4 as long or longer than 2+3, somewhat bulbous at base and

narrowed apically with a distinctive recurved apex (Fig. 4). Labium (Fig. 5) with 3-articled palpi, 2"4

wider than 3", narrower than 1, ligula very broad and divided to base into 2 broad lobes. Hypo-

pharynx as in Fig. 6. Pronotum subquadrate to slightly elongate (Figs. 8-10). Tibiae (Fig. 7) with a

few scattered spines, tarsal claws falcate. All tarsi 5-articled, tarsomere 1—4 short, subequal, article 5

longer. Hind wings absent. Abdominal sternite HI very deeply sinuate anteriorly, more or less U-

shaped. Abdomen IIJ—VII without paratergites. Abdominal tergite VII slightly longer than VI, abdom-

162 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

inal tergites without transverse basal impressions. Anterior margin of abdominal sternite VIII modified

into broad lobe in association with sternal gland and gland reservoir.

Intertidal, found under stones and boards (and probably other debris) below the high-tide mark on

beaches of the Pacific Coast of North America from Northern California to Alaska and Hokkaido.

Larvae unknown.

Distribution.—Pacific Coasts of California, Alaska, Hokkaido (Japan).

KEY FOR IDENTIFICATION OF KNOWN SPECIES OF GIULIANIUM

1. Larger species, 3.0 mm in length; antenna very elongate, nearly attain-

ing apex of elytra when extended posteriorly, antennal articles 3—10

very elongate, 2.0—1.5 times longer than wide (Fig. 11); aedeagus as

Hi Be Ae IEE at apn ge OER EM ee Bb ete npcy Sot ated cia ee Pe te AP rh pe ET G. campbelli

ie Smaller species, 2.1—2.7 mm in length; antenna less elongate, attaining

middle to apical third of elytra when extended posteriorly, antennal

articles 3-10 less elongate 1.7—1.1 times longer than wide (Figs. 12

and. 13);saedeagus nots AbOVe:.t ti.ioct. . S282 ee ee eee 2

2(1’). Smaller species, 2.1—2.2 mm in length, mature specimens uniformly

light brown; head distinctly elongate, 1.1—1.2 times longer than wide

(Fig. 10); pronotum slightly, but distinctly elongate, about 1.1 times

longer than wide (Fig. 10); aedeagus as in Fig. 16; known from

NOKMETO CaMLOrnig sw. oxeie.c4 8 oa Mee en ns ee meses he G. newtoni

Page Larger species, 2.4—2.7 mm in length, mature specimens uniformly

dark brown; head subquadrate, about as long as wide (Fig. 9); prono-

tum about as long as wide to slightly longer than wide (up to 1.05

times longer than wide) (Fig. 9); aedeagus as in Fig. 15.; known

trom<Adaska-and: Japan 2) sya. oie he cite deen tes G. alaskanum

GIULIANIUM CAMPBELLI MoOorE 1976

(Figs. 8, 11, 14)

Giulianium campbelli Moore, 1976: 57.

Redescription—Length 3.0 mm. Body uniformly dark brown throughout, uniformly covered with

dense vestiture of very fine pubescence, slender, parallel-sided. Head slightly wider than long (1.1—

1.2 times wider than long) (Fig. 8); antenna (Fig. 11) very long and slender, nearly attaining apex of

elytra when extended posteriorly, all antennomeres very elongate, antennomere 3—10 about 2.0-1.5

times longer than wide, with apical articles relatively shorter than more basal articles. Pronotum (Fig.

8) about as wide as long (Moore 1976 says that the pronotum is ¥% longer than wide but our mea-

surements of the holotype and one paratype did not confirm this). Aedeagus as in Fig. 14 (dorsal

aspect only, lateral aspect of parameres not available because the aedeagus is permanently mounted

in mounting medium and could not be reoriented).

Distribution.—California.

Material Examined —Holotype, labeled as follows: ‘Bear Harbor, Humboldt County, California,

June 1964, Derham Giuliani; Holotype, Giulianium campbelli Moore’ deposited in the Entomological

Research Institute, Ottawa, Canada. Paratype 1, same data as holotype.

GIULIANIUM ALASKANUM, NEW SPECIES

(Figs. 9, 12, 15, 16)

Types.—Holotype, labeled as follows: ‘U.S.A.: Alaska. Seward, 24 V 1994, K.

J. Ahn, ex., under boulder in low tide; Holotype, Giulianium alaskanum Ahn and

1999 AHN & ASHE: NEW GIULIANIUM SPECIES 163

Figures 14-18. Aedeagus. Figure 14. G. campbelli, dorsal aspect; Figure 15. G. alaskanum, lateral

aspect; Figure 16. Dorsal aspect; Figure 17. G. newtoni, lateral aspect; Figure 18. Dorsal aspect. Scales

0.1 mm.

Ashe, Desig. K.-J. Ahn and J. S. Ashe 1998.’ deposited in the Snow Entomolog-

ical Museum, University of Kansas, Lawrence, Kansas. Paratypes, 28; 2 labeled

as follows: ‘Japan: Hokkaido. Akkeshi, Tokotan, 15 VI 1994, ex., under beach

ground’ 26 same data as type (18, Snow Entomological Museum; 8, Chungnam

National University Insect Collection, Daejeon, Korea; 2, Field Museum of Nat-

ural History, Chicago, Illinois).

Description—Length 2.4—2.7 mm. Body uniformly dark brown throughout, uniformly covered with

dense vestiture of very fine pubescence, slender, parallel-sided. Head about as wide as long (Fig. 9);

antenna (Fig. 11) long and slender, attaining apical third of elytra when extended posteriorly, all

antennal articles elongate, articles 3-10 about 1.7—1.3 times longer than wide, with apical articles

relatively shorter than more basal articles. Pronotum (Fig. 9) subquadrate to slightly longer than wide

(less than 1.1 times longer than wide). Aedeagus as in Figs. 15 and 16; parameres more or less straight

in lateral aspect, not bent upwards with respect to the medial lobe, lateral edge narrowed in apical

fifth (Fig. 15), dorsal aspect more or less uniformly divergent from base to apex (Fig. 16).

Distribution.—Alaska and Japan (Hokkaido).

GIULIANIUM NEWTONI, NEW SPECIES

(Figs. 10, 13, 17, 18)

Types.—Holotype, labeled as follows: ‘U.S.A.: California, San Mateo Co. Moss

Beach, 27 II 1952, under boards below high tide, O. Bryant, Field Museum Nat

164 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

Hist; Holotype, Giulianium newtoni Ahn and Ashe, Desig. K.-J. Ahn and J. S.

Ashe 1998.’ deposited in the Field Museum of Natural History, Chicago. Para-

types, 5; same data as type (3, Field Museum of Natural History; 2, Snow En-

tomological Museum).

Description—Length 2.1—2.2 mm. Body uniformly light brown throughout, uniformly covered with

dense vestiture of very fine pubescence, very slender, parallel-sided. Head distinctly longer than wide

(Fig. 10); antenna (Fig. 13) long and slender, attaining apical half of elytra when extended posteriorly,

all antennomeres elongate, antennomeres 3—10 about 1.5-1.1 times longer than wide, with apical

articles relatively shorter than more basal articles. Pronotum (Fig. 10) distinctly longer than wide,

about 1.1 times longer than wide. Aedeagus as in Figs. 17 and 18; parameres distinctly bent upwards

with respect to the median lobe, lateral edge not narrowed in apical fifth (Fig. 17), dorsal aspect with

parameres diverging in basal half and converging toward midline from near middle to apex (Fig. 18).

Distribution.— California.

Discussion.—The three known species are superficially very similar, but they

can be distinguished by the characters in the key, and by the slight, but consistent,

differences in the parameres noted in the descriptions (see also Figs. 14-18). To

facilitate comparisons between these similar species, the illustrations of compa-

rable structures of all three species are drawn to the same scale. The differences

in relative size and proportions of the head, pronotum and antennae, and the

differences in the parameres of the aedeagi, are readily apparent in these illustra-

tions. !

ACKNOWLEDGMENT

We extend special thanks to Dr. A. Smetana (Entomological Research Institute,

Ottawa, Canada) and Dr. A. E Newton, Jr. (Field Museum of Natural History,

Chicago) for loan of specimens. This research was partially supported by Chung-

nam National University Research Grant to K.-J. Ahn.

LITERATURE CITED

Moore, I. 1976. Giulianium campbelli, a new genus and species of marine beetle from California

(Coleoptera: Staphylinidae). Pan-Pacif. Entomol., 52: 56-59.

Newton, A. F & M. K. Thayer. 1995. Protopselaphinae new subfamily for Protopselaphus new genus

from Malaysia, with a phylogenetic analysis and review of the Omaliine Group of Staphylinidae

including Pselaphidae (Coleoptera). pp. 219-320. In J. Pakaluk & S. A. Slipinski (eds.). Biol-

ogy, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80" Birthday of Roy

A. Crowson.

Thayer, M. K. 1987. Biology and phylogenetic relationships of Neophonus bruchi, an anomalous south

Andean staphylinid (Coleoptera). Syst. Entomol., 12: 389-404.

Received 9 Sep 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 165-169, (1999)

ESTABLISHMENT OF AN EXOTIC PLASTER BAGWORM

IN CALIFORNIA (LEPIDOPTERA: TINEIDAE)

HANIF GULMAHAMAD

Terminix International, 1501 Harris Court, Anaheim, California 92806

Abstract—This paper reports the first North American and California records of the plaster

bagworm, Phereoeca praecox Gozmany and Vari. Distributional records for California are re-

ported. A case history of P. praecox as a fabric pest in southern California is described. Food

sources of Phereoeca are summarized. Dissection of live larval cases reveals that in southern

California, P. praecox is essentially chitinophagous and keratinophagous.

Key Words.—Insecta, Lepidoptera, Tineidae, plaster bagworm, fabric pest, food sources, Pher-

e0eca praecox.

The genus Phereoeca (Lepidoptera: Tineidae) was created in 1956 in order to

separate a group of flat, case-bearing moths from case-making moths of the genus

Tineola (Hinton & Bradley 1956). Phereoeca spp. are commonly referred to as

wall bagworms, plaster bagworms, and household case-making moths. The larva

live in, feed from and pupate inside a characteristic watermelon-shaped, flattened,

broadly spindle-shaped case lined internally with silk. The outside of the case is

usually covered with sand, soil particles, brick dust, and other miscellaneous de-

bris found within the larva’s habitat. The genus Phereoeca occurs throughout the

wet tropics of the Old and New World. Robinson & Nielson (1993) stated that

they are aware of six Phereoeca spp. worldwide of which four are named. The

nomenclature and taxonomy of the genus Phereoeca has been, and remains con-

fusing.

Prior to this paper, the only other Phereoeca species recorded from North

America was Phereoeca uterella (Walsingham) which is reported from Florida,

Louisiana, Mississippi, and North Carolina (Hetrick 1957). For many years, this

species was reported as Tineola walsinghami (Busck) (Villanueva-Jimenez 1996,

Katz 1997). Tineola walsinghami was put in synonymy in 1984 (Davis 1984).

The species in Florida was then referred to as Phereoeca dubitatrix (Meyrick)

(Villanueva-Jimenez 1996). Phereoeca dubitatrix was put in synonymy in 1993

(Robinson & Nielsen 1993). The Phereoeca from Florida is now designated as

P. uterella (Walsingham). The name changes associated with this single species

testifies to the confusion regarding the identity and nomenclature of the genus

Phereoeca.

Phereoeca praecox Gozmany and Vari in California.—The first report of a

plaster bagworm occurring in California was on 28 January 1986 when a number

of larval cases were submitted by a pest control operator to Orange County Ag-

riculture Commissioner for identification (Nick Nisson, personal communication).

Adults reared from this sample were sent to the National Museum of Natural

History for identification. A species determination could not be made at that time

because the specimens submitted were in poor condition preventing proper iden-

tification. Adults of a Phereoeca sp. captured on 2 April 1997 from inside a

residence on Granville Drive, Newport Beach, Orange County, California, and

additional moths reared from larval cases retrieved from the above address were

166 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

sent to Dr. Davis of the NMNH for a species determination. Davis subsequently

fowarded illustrations of his genitalia dissections of this species to Gaden Rob-

inson of the British Museum (Natural History). Robinson identified the male as

P. praecox, but he was uncertain if the female represented the same species.

Phereoeca specimens reared from larval cases collected on 27 August 1998 from

Gardena, Los Angeles County, California, were also submitted to Davis for iden-

tification. Males from this sample were found to be consistent with P. praecox.

P. praecox was originally described in 1973 (Gozmany & Vari 1973). The male

holotype was reported to have been taken on 30 August 1928 in Njala, Sierra

Leone. P. praecox has since been reported from Australia (Robinson & Nielson

1993):

This paper reports the first North American and California records of P. prae-

cox. Other species of Phereoeca are not known to occur in California.

Since 1989, I have encountered plaster bagworms in southern California as-

sociated with structures during the course of my work as a structural pest man-

agement professional. Specimens collected by the author, and others submitted to

him for identification, have always been empty larval cases. Empty cases are

easily recognizable because the pupal exuviae are usually found partially pro-

truding from one end of the case. I have collected cases from inside garages, in

substructural areas, under exterior stairwells, on exterior walls, under eaves of

structures, on walls and ceilings of entryways, under patio covers and at patio/

building junction, and on bathroom and laundry room walls. On one occasion, a

case with a live larva was found on the asphalt parking lot of a commercial office

building about 10 m from the structure.

Distribution Records of P. praecox in Southern California —U.S.A. CALIFORNIA. LOS ANGELES

Co.: Los Angeles, 26 August 1987. California Dept. of Food & Agriculture, Plant Pest Diagnostic

Branch record. Los Angeles, 20 July 1996. H. Gulmahamad. Gardena, 20 January 1998, R. Arias.

Redondo Beach, 27 August 1998, S. Howard. Gardena, 27 August 1998, H. Gulmahamad. Gardena,

15 October 1998. D. Jimbo. Long Beach, 30 October 1998, S. Howard. Redondo Beach, 4 November

1998, S. Howard. San Pedro, 10 November 1998, S. Howard. ORANGE Co.: Westminster, 28 January

1986. Nick Nisson. Anaheim, 20 March 1989. H. Gulmahamad. Aliso Viejo, 15 June 1996. V. Herrera.

Newport Beach, 2 April 1997. H. Gulmahamad. Irvine, 17 May 1997. D. Kern. Placentia, 22 May

1997. R. Lagana. Yorba Linda, 30 May 1997. P. Palamara. Anaheim Hills, 12 July 1997. D. Simkin.

Mission Viejo, 4 August 1997. M. Tassinari. Anaheim, 31 October 1997. H. Gulmahamad. San Cle-

mente. 16 February 1998. D. Eschevarria. Costa Mesa, 2 March 1998. B. Griffin. Costa Mesa, 11

March 1998. B. Smallwood. Newport Beach, 4 April 1998. H. Gulmahamad. Anaheim, 1 May 1998.

H. Gulmahamad. Corona del Mar, 22 July 1998. V. Lucero. Newport Beach, 23 July 1998. V. Lucero.

Irvine, 16 March 1999. H. Gulmahamad. RIVERSIDE Co.: Beaumont, 2 April 1993. Tracy. SAN

BERNARDINO Co.: Chino, 17 September 1997. R. Lampman. SAN DIEGO Co.: San Diego, 12

December 1988. R. Skelly. San Diego, 19 March 1992. Glassford. Spring Valley, 27 March 1994.

Taylor. SANTA BARBARA Co.: Santa Barbara, 25 February 1988. CDFA Plant Pest Diagnostic Branch

record. YOLO Co.: Sacramento, 20 April 1993. L. Allen.

Food Sources of the Genus Phereoeca.—There is much disagreement regarding

natural food sources of the larvae of the various species of Phereoeca. Cited food

sources include insect parts, fur, flannel, wool, spider webs, bat and bird drop-

pings, and other fabrics (Meyrick 1905; Walsingham 1914; Kea 1933; Busck

1933; Watson 1939, 1946; Mallis 1954; Hinton 1956; Hetrick 1957; NPCA 1977;

Zimmerman 1978; Aiello 1979; Robinson & Nielson 1993; Koehler & Castner

1994). Dissection of 15 cases of P. praecox containing live larvae which were

taken from three different locations in southern California, revealed that P. prae-

1999 GULMAHAMAD: PHEREOECA IN CALIFORNIA 167

Figure 1. Top. Plaster bagworm cases hanging at wall/ceiling junction of garage. Ninety cases

were counted in this area.

Figure 2. Bottom. Carpet below zebra skin rug showing plaster bagworm damage. Note more

pronounced damage at periphery of rug.

168 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

- ris

wt ap WP oy

rh tyes "

_ |

hey’

» 7 »*

ey a

a

\ BCE Cae ba ee tae Stes es OPIATE Ei yk

é aaa

Figure 3. Photograph shows damage caused by plaster bagworm to edge of carpet adjacent to a

sliding glass door.

cox fed on dead insects, insect parts and fragments, human and animal hairs, and

bird feathers. This is the first record of a Phereoeca species feeding on bird

feathers.

A Case History of P. praecox as a Fabric Pest in California. —On 26 March

1997, the owner of a residence on Granville Drive, Newport Beach, Orange Coun-

ty, California called Terminix International and requested assistance with an un-

usual pest problem. On 2 April 1997, I interviewed the owner and conducted an

inspection of the premises. In the detached garage, I found numerous cases of P.

praecox on the walls and at the wall/ceiling junction. Many empty cases were

hanging in spider webs at the wall/ceiling junction (Fig. 1).

The area of concern to the homeowner was a portion of the carpet in front of

the fireplace which was covered with a natural zebra skin rug. The carpet was a

two-year-old 100% looped wool Berber carpet portions of which under the zebra

skin was eaten to the base (Fig. 2). No larval cases were present at this time.

However, interviewing the maid, I discovered that, as part of her regular cleaning

routine, the zebra skin rug is picked up at least once every two weeks and the

area underneath it vacuumed. The maid claimed that it was during one of the

these bimonthly cleaning in March 1997 that she discovered the damage and about

30 larval cases of a plaster bagworm under the rug. She maintained that the

damage occurred within a two week period. Damage to the carpet was also found

at several other locations in the living room, primarily at the floor/wall junction

(Fig. 3). At some of these areas, empty larval cases were found under the edges

of the carpet. I conducted a meticulous inspection of the premises for other fabric

pests. None were found. A number of live adult moths were found in one dimly

1999 GULMAHAMAD: PHEREOECA IN CALIFORNIA 169

lit corner of the living room. Additional larval cases were taken from: in and on

the fireplace, in the chimney, under a sofa, under a sofa cushion, under a chair,

under and behind a china cabinet, behind baseboards, under the edges of the

carpet, on walls and ceilings of closets, on the living room walls particularly in

corners where two walls meet. Larval cases were also found on the stucco and

brick walls on the exterior of the entryway to the home. The maid claimed that,

at one time, plaster bagworm cases were so numerous on the exterior walls of

the entryway that she had to wash them off with a water hose.

ACKNOWLEDGMENT

I thank Stoy Hedges, Ken Hobbs, Don Davis, and Harry Katz for their con-

structive review of the manuscript, and Don Davis and Gaden Robinson for iden-

tifying the California plaster bagworm.

LITERATURE CITED

Aiello, A. 1979. Life history and behavior of the case-bearer Phereoeca allutella (Lepidoptera: Ti-

neidae). Psyche, 86: 125-136.

Busck, A. 1933. Microlepidoptera of Cuba. Entomologica Americana 13: 151-203.

Davis, D. R. 1984. Tineidae. Chapter 15. pp. 19-24. In Heppner, J. B. (ed.). Atlas of Neotropical

Lepidoptera. Checklist. Part 1. Microterigoidea-Immoidea. W. Junk Publishers, The Hague, The

Netherlands.

Gozmany, L. A. & L. Vari. 1973. The Tineidae of the Ethiopian region. Mem. Transv. Mus., 18: 1—

238.

Hetrick, L. A. 1957. Some observations on the plaster bagworm, Tineola walsinghami Busck (Lepi-

doptera: Tineidae). Fla. Entomol., 40: 145-146.

Hinton, H. E. 1956. The larvae of the species of Tineidae of economic importance. Bull. Entomol.

Res., 47: 251-346.

Hinton, H. E. & J. D. Bradley. 1956. Observations on species of Lepidoptera infesting stored products.

XVI: two new genera of clothes moths (Tineidae). The Entomologist, 89: 42—47.

Katz, H. 1997. Clothes moths. Chapter 10. pp. 427-462. In Moreland, D. (ed.). Handbook of pest

control. Mallis Handbook & Technical training company. Cleveland, Ohio.

Kea, J. W. 1933. Food habits of Tineola uterella. Fla. Entomol., 17: 66.

Koehler, P G. & J. L. Castner. 1994. Pests that occasionally invade structures. Univ. of Florida. Institute

of Food and Agricultural Sciences. Dept. of Entomology and Nematology. SP 148.

Mallis, A. 1954. Handbook of pest control (1st ed.). Mac Nair-Dorland Company, New York.

Meyrick, E. 1905. Descriptions of Indian Micro-Lepidoptera. J. Bombay Nat. History Soc., 16: 580-

619.

NPCA. 1977. Fabric pests. NPCA Technical release, ESPC 031101—031141. NPCA. Dunn Loring,

Virginia.

Robinson, G. S. & E. S. Nielsen. 1993. Tineid genera of Australia (Lepidoptera). Monographs of

Australian Lepidoptera. Volume 2. CSIRO Publications, East Melbourne, Victoria. Australia.

Villanueva-Jimenez, J. A. 1996. Plaster bagworm Phereoeca dubitatrix (Meyrick) Lepidoptera: Ti-

neidae. http://guv.ifas.ufl.edu/~insect/urban/occus/P-Bagworm.htm.

Walsingham, T. De. G. L. 1914. Family 20. Tineidae. Biologia Cent.-Am. Insecta Lepid. Heterocera,

4: 344-375.

Watson, J. R. 1939. Control of four household pests. Fla. Agr. Exp. Sta. Press Bull., 536.

Watson, J. R. 1946. Control of three household insects. Fla. Agr. Exp. Sta. Press Bull., 619.

Zimmerman, E. C. 1978. Insects of Hawaii. Volume 9. Microlepidoptera. The University Press of

Hawaii, Honolulu. Hawaii.

Received 19 Nov 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 170-177, (1999)

PUPAL DIAPAUSE OF COLORADIA PANDORA BLAKE

(LEPIDOPTERA: SATURNIIDAE)

ELIZABETH A. GERSON!, RICK G. KELSEY!, & DARRELL W. Ross?

'USDA Forest Service, PNW Research Station, 3200 SW Jefferson Way,

Corvallis, Oregon 97331

*Department of Forest Science, Oregon State University,

Corvallis, Oregon 97331

Abstract.—Pupae of the pandora moth, Coloradia pandora Blake, were collected in central

Oregon and stored at 5° C for 8 to 24 weeks, then incubated at 25° C. The minimum cold storage

time required to break diapause was 12 weeks, but emergence rates were highest (87.5%) for

14-18 weeks. In a separate experiment, 1000 pupae were maintained in field enclosures for 3

years while soil temperature was monitored. Seventy-two percent of these pupae emerged in

Year 1. Soil temperature fell below 5° C for 21.7, 22.9, and 25.1 weeks over the three consecutive

winters, and the minimum soil T was —2° C. In the lab study, =22 weeks at 5° C limited

emergence to <40%, therefore mortality from duration of cold could be considerable in winters

such as Year 3. Prolonged (extended) diapause was observed in only 0.6% of the sample pop-

ulation.

Key Words.—Insecta, Coloradia pandora, pandora moth, pupae, diapause, phenology, rearing,

adult emergence.

The pandora moth, Coloradia pandora Blake, a native defoliator of Pinus spp.

in the western U.S., has a biennial life cycle throughout most of its range (Tuskes

et al. 1996). The pupal stage generally lasts 12 to 13 months (June through July

the following summer in central Oregon) (Ross 1995, 1996), although pupal dia-

pause as long as 6 year may occur (Carolin 1971). The proportion of the popu-

lation remaining in diapause beyond 1 yr has been estimated at <5% (Massey

1940) up to ‘‘a substantial part of the generation” in some areas (Carolin & Knopf

1968). A recent outbreak of C. pandora in central Oregon was characterized by

alternating summers of severe defoliation, and summers with dense moth popu-

lations. If a substantial proportion of an epidemic population remained in extended

diapause, then conspicuous defoliation and moth flights during off-years would

be expected, however, this was not observed.

The life histories of abundant, episodic forest insects can influence ecological

processes such as predator-prey relationships and host tree physiology. In the case

of C. pandora, a high incidence of extended diapause would diminish the biennial

nature of its life cycle, possibly increasing defoliation stress on host pine trees

and enhancing the availability of C. pandora pupae as prey for insectivores.

Therefore, one objective of this study was to quantify the occurrence of extended

diapause in central Oregon. To do this, we tracked emergence of a single cohort

of pupae for three years under natural conditions.

The second objective of this study was to determine the length of cold storage

required for successful pupation of C. pandora. We were interested in minimizing

pupal diapause for the purpose of rearing adult moths in the laboratory. Massey

(1940) attempted to break diapause by exposing pupae to low temperatures in

December, then incubating the pupae at 21—27° C. He did not specify the cold

storage temperature or duration of exposure, and his pupae required 4 months

1999 GERSON ET AL.: DIAPAUSE IN COLORADIA 171

exposure to high temperatures before any development was noted. Time to emer-

gence generally decreases with the duration of exposure to cold, provided the

temperatures are within an acceptable range for the species (Danks 1987). Our

cold storage experiment, in conjunction with the extended diapause study, pro-

vided some insight about environmental conditions that could promote extended

diapause in C. pandora, and conditions that could cause mortality in the pupal

stage.

METHODS AND MATERIALS

Cold Storage.—Pupae were collected 1 November 1992 in an area of the Des-

chutes National Forest, Oregon, that had been heavily defoliated the previous

spring based on USDA Forest Service Region 6 aerial surveys. The pupae were

covered with sandy soil and refrigerated at 2 to 5° C for a minimum of eight

weeks. After eight weeks, eight female and eight male pupae [sexed according to

characteristics of the fourth and fifth abdominal segments (Tuskes et al. 1996)]

were transferred to 4 liter paper buckets with screen tops, covered minimally with

sand and incubated at 25° C in a rearing room with a 16:8 light : dark photoperiod.

Another 8 female and 8 male pupae were removed from cold storage at 12, 14,

16, 18, 20, 22 and 24 weeks. Soil in the refrigerator and sand in the rearing room

were misted with water several times weekly. Female and male emergence was

monitored daily.

Extended Diapause.—Pupae were collected on 15 June 1995 from 2 locations

in Pringle Falls Experimental Forest, about 45 km SW of Bend, Oregon. These

sites had been moderately to heavily defoliated by the current generation, but not

the previous generation of C. pandora based on USDA Forest Service Region 6

aerial surveys, so the pupae were unlikely to be more than 1 year old. Two

hundred pupae were laid in the bottom of each of 5 screen enclosures (0.9 x 0.9

x 0.9 m) and covered with ~6 cm top soil. Three of the enclosures were located

within the Pringle Falls Research Natural Area in a mature stand of ponderosa

pine, and the other 2 enclosures were placed 4 km distant at the Pringle Falls

Research Forest Headquarters under a similar canopy. Adult emergence was tal-

lied by gender for each enclosure on 3 or 4 dates in July and August from 1995

through 1997 (Year 1—Year 3). A soil and air temperature monitor (Omnidata

Datapod® model DP-212, Logan, Utah) was installed in November 1994, to doc-

ument exposure to cold temperatures in the first winter of pupal diapause prior

to collecting the pupae, and for the remainder of the experiment. The soil tem-

perature probe was buried at the same depth as the pupae (6 cm), and the air

temperature probe was placed next to one of the enclosures at the Headquarters

location. The Datapod® recorded maximum, minimum, and average daily tem-

peratures.

RESULTS AND DISCUSSION

Cold Storage—No adults emerged from the cohort receiving 8-weeks cold

storage, even after 177 days at 25° C. Successful completion of diapause was first

observed for pupae in the 12-week treatment group (Fig. 1). Optimal survival

rates (87.5%) were achieved with 14 to 16 weeks at 5° C for males and 16 to 18

weeks for females. Female emergence lagged behind that of males, indicating

protandry (Fig. 1). The length of time to emergence at 25° C declined with in-

172 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

—y female

% Emergence

8 10 12 14 16 18 20 22 24

Duration of Cold Storage (weeks at 5 °C)

Figure 1. Percent C. pandora adult emergence relative to length of cold storage at 5° C. n = 8

female and 8 male pupae per time period.

creasing time in cold storage (Fig. 2). Pupae stored for 24 weeks at 5° C required

1 month less time at 25° C to complete diapause compared to pupae in cold

storage for 12 weeks. Linear regression of time to emergence on the length of

cold storage revealed that, for each additional week of cold storage, the incubation

time declined by 3 days. Thus in a laboratory situation, additional cold storage

beyond the minimum required to break diapause would not be time efficient,

except that the rate of successful emergence was not constant (Fig. 1). The uni-

100

Oo females

—~ 90 A males

Oo —— y= 116.5 - 3.0(x)

fs P= 0.82

N

= «80-4

”

o

Sn]

CO)

e)

d

mo 604

O)

=

LL

g 0-4

o

=

aa ii

0 “it T T 2 T T T T =~] (ee ae eee

8 10 12 14 16 18 20 22 24

Duration of Cold Storage (weeks at 5 °C)

Figure 2. Incubation time at 25° C needed for C. pandora adult emergence relative to duration of

5° C cold storage period of pupae.

1999 GERSON ET AL.: DIAPAUSE IN COLORADIA 173

Table 1. Emergence rates of Coloradia pandora adults for 5 enclosures, each containing 200 pupae.

Year 1 Year 2 Year 3 Total

Enclosure % % % %

A 74.0 0.5 0 74.5

B 70.5 0.5 0 71.0

Cc #35 0.5 0 74.0

D 75.0 0 0 75.0

E 66.5 1.5 0 68.0

Mean 71.9 0.6 0 G25

form incubation temperature of 25° C was selected to favor rapid development,

rather than to mimic field conditions. Soil temperatures measured in the field

during the extended diapause experiment never exceeded 23° C.

There is a cold duration threshold between 8 and 12 week below which C.

pandora pupae from this population are physiologically unable to complete dia-

pause. Half of the pupae in the 8-week cold treatment group were dissected fol-

lowing the experiment and were found to be undifferentiated and apparently still

viable. Because this experiment did not incorporate cold storage treatments be-

tween 8 and 12 week, we cannot determine the threshold more precisely. Twelve

weeks should be considered minimal because the pupae in this experiment may

have had some prior exposure to temperatures <5° C. They were collected from

the field on 1 November 1992, and subsequent monitoring in 1995 and 1996 for

the extended diapause experiment indicated that soil temperatures may drop below

5° C in the later half of October. Therefore, the actual length of cold treatment

these pupae received could have been several weeks longer than treatments im-

posed in the lab.

Cold exposure in the pupal phase may not be prerequisite to adult emergence

throughout the range of C. pandora. There are three geographically disjunct pop-

ulations of the nominate subspecies (Tuskes et al. 1996). Central Oregon is the

northern extent of the western distribution which extends south into California

and Nevada. By several accounts (Aldrich 1921, Tuskes 1984), some adults in

the isolated population of southern California have been observed emerging after

just 2 months of pupation. Significant latitudinal variation in pupal diapause is

common (Danks 1987, Tuskes et al. 1996). For example, the saturnid Automeris

io (Fabr.) produces both diapausing and non-diapausing pupae in the lower lati-

tudes of its range (Manley 1993).

Extended Diapause——In Year 1, 67 to 75% of the pupae in the 5 enclosures

successfully completed diapause (Table 1). An additional 0 to 2% emerged in

Year 2, and none emerged the 3rd year following pupation, indicating extended

diapause was rare in the sample population. Most of the pupae remaining in the

enclosures after 3 years were desiccated, a few were moldy, and 4 appeared

potentially viable upon dissection. Total emergence rates in the field enclosures

(grand mean = 72.5%, Table 1) fell below the maximum successful pupation

rates of the cold storage experiment (87.5%, Fig. 2), but this may be explained

by soil temperatures. Soil temperatures averaged =5° C for at least 152 days (21.7

weeks in Year 1 (Fig. 3A), so based on results from the cold storage experiment,

we might expect successful pupation rates as low as 20—40% (Fig. 1). If the

174 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

A--Pupal Year 1

Soil Temperature (°C)

5

1 Sep 94 15 Jan 95 31 May 95

B--Pupal Year 2

Soil Temperature (°C)

5

1 Sep 95 15 Jan 96 31 May 96

C--Pupal Year 3

Soil Temperature (°C)

5

1 Sep 96 15 Jan 97 31 May 97

Date

Figure 3. Daily soil temperatures at Pringle Falls Headquarters, Deschutes National Forest,

Oregon. Temperature probe at 6 cm depth. (A) Ist year, (B) 2nd year, (C) 3rd year of pupal diapause.

pupae used for the cold storage experiment actually did have 1—2 weeks additional

exposure in the field prior to collection, then the expected successful pupation

rate for 22 weeks at 5° C would be 63-75%, which corresponds well with the

emergence rate from the extended diapause experiment. During the winter of Year

2 there were 22.9 weeks with soil temperatures <5° C, and in Year 3 there were

25.1 weeks =5° C (Fig. 3B, 3C). These data, along with the cold storage results,

1999 GERSON ET AL.: DIAPAUSE IN COLORADIA 175

suggest that the duration of cold soil temperatures in central Oregon probably

restricts the maximum successful pupation rates for C. pandora, perhaps as much

as 60% over a long winter.

Winter mortality in other moth species is a function of degree, as well as

duration, of cold (Turnock et al. 1983). Diapausing A. io pupae tolerate air tem-

peratures in excess of —18° C, but freezing temperatures extend the duration of

diapause (Manley 1993). The degree of sensitivity of C. pandora to freezing

temperatures is unknown. In the first year of the extended diapause experiment,

the minimum recorded soil temperature was —1° C and 72.5% of the pupae sur-

vived. Over 3 winters, soil temperatures hovered around 0° C for extended periods

(Fig. 3) despite marked fluctuations in air temperatures, thus it seems likely that

duration, rather than degree of cold, would cause winter mortality under typical

field conditions. Coloradia pandora pupate about 6 cm below the soil surface in

central Oregon which buffers exposure to cold. Snow cover can dramatically

reduce winter mortality from freezing temperatures (Turnock et al. 1983), but

persistence of the snow pack in spring could lengthen pupal exposure to cold by

slowing soil warming. The insulating effects of leaf cover and snow have been

shown to influence pupal development of A. io (Manley 1993). An open forest

canopy that intercepts less snow in winter and allows for rapid snow melting in

spring might be favorable to C. pandora pupation. Decreased canopy cover (Ross

1995), lower elevation and southerly aspect (Schmid 1984), all of which hasten

melting of snow and possibly reduce the duration of cold soil, are known to

promote earlier emergence of adult C. pandora. There is also some evidence that

water from snowmelt may trigger developmental processes in diapausing insects

(Danks 1987).

Synchronized emergence and protandry are characteristics of the temporal

emergence patterns of lepidopteran populations that favor successful mating (Tus-

kes et al. 1996). Coloradia pandora appears to utilize both strategies to some

extent. Moths emerge throughout a 30- to 40-day period, but the peak emergence

period lasts about 7 days (Schmid 1984, Ross 1996). Overall emergence in our

field enclosures was fairly coincident in Year 1 as 95% of the moths which

emerged did so within a 2-week period (Fig. 4). More frequent observations may

have narrowed this time frame. Adult C. pandora may live 7—10 days (Schmid

1984, personal observation) so this peak period is relatively broad compared to

some shorter lived saturniids having synchronized emergence within several days

(Tuskes et al. 1996). Protandry slightly offsets the synchrony of emergence toward

males first, which may benefit the population by increasing the likelihood that

females are able to mate and lay eggs soon after emergence (Tuskes et al. 1996).

In Year 1 of the extended diapause study, 85% of male emergence and 70% of

female emergence occurred by July 28 (Fig. 4). Protandry also was observed in

the cold storage experiment (Fig. 1) and in other C. pandora studies (Massey

1940, Schmid 1984). A total of 299 females and 420 males emerged in Year 1,

suggesting the sex ratio in the population may have been biased in favor of males.

However, the one-thousand pupae in the enclosures initially were not sexed, so it

remains possible that higher mortality rates in female pupae caused fewer adult

females to emerge.

Coloradia pandora pupae are food for golden-mantled ground squirrels (Sper-

mophilus lateralis Say), white-footed deer mice (Peromyscus maniculatus Wag-

176 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

100

—e— male

—w— female

80 -

60 -

40

20

Cumulative % of Adults Emerged

+ fas

5 Jul 17 Jul 28 Jul 3 Aug 26 Aug

Pupal Year 1

Figure 4. Cumulative emergence pattern for adult C. pandora moths. Percentages are based on

the total number of males and females emerged in the Ist year of pupal diapause.

ner), and possibly other wildlife in central Oregon (unpublished data). Extended

diapause in a biennial insect would provide a more reliable prey base for insec-

tivores utilizing only the pupal stage. Fecundity and survivorship of wildlife with

short lifespans (1—5 year), such as deer mice and ground squirrels, may be influ-

enced by annually fluctuating food resources. In our study, less than 2% of pupae

remained viable (i.e., edible) beyond 1 year of development, hence they would

constitute a biennial food source in central Oregon. Carolin’s (1971) paper on

extended diapause has been interpreted as indicating that C. pandora provided an

annual food base for aboriginal Paiute tribes in California, despite the biennial

cycle of C. pandora (Weaver & Basgall 1986; but see Fowler & Walter 1985,

Blake & Wagner 1987). In areas with milder winters (< 12 weeks below 5°C),

C. pandora pupae could possibly hold over a second year in order to acquire the

requisite cold period to complete diapause; but it is also possible that univoltinism

created an annual supply of larvae in southern California (Tuskes et al. 1996).

Geographical differences in environmental regimes may explain Carolin’s

(1971) observations of extended diapause. He also collected pupae in central

Oregon, but he held them over winters in Portland, Oregon which has a relatively

mild, maritime climate. Carolin’s data showed 18% of pupae emerged in their

2nd year, and another 37% emerged the 3rd year. However, even more important

is the fact that his initial population of pupae were culled from unemerged pupae

remaining after a large flight year. Therefore, the extended emergence rates Car-

olin reported are not applicable to whole populations.

ACKNOWLEDGMENT

We thank Gladwin Joseph (Oregon State University, Corvallis) for his enthu-

siastic help in collecting C. pandora pupae and setting up the field enclosures,

and Jocelyn G. Millar (University of California, Riverside) and Paul C. Hammond

(OSU) for helpful reviews of the manuscript. The use of trade names is for the

1999 GERSON ET AL.: DIAPAUSE IN COLORADIA EET

information and convenience of the reader and does not constitute official en-

dorsement or approval by the U.S. Department of Agriculture.

LITERATURE CITED

Aldrich, J. M. 1921. Coloradia pandora Blake, the moth of which the caterpillar is used as food by

Mono Lake Indians. Ann. Entomol. Soc. Amer., 14: 36-38.

Blake, E. A. & M. R. Wagner. 1987. Collection and consumption of pandora moth, Coloradia pandora

lindseyi (Lepidoptera: Saturniidae), larvae by Owens Valley and Mono Lake Paiutes. Bull.

Entomol. Soc. Amer., 33: 22—27.

Carolin, V. M. 1971. Extended diapause in Coloradia pandora Blake. Pan-Pacif. Entomol., 47: 19-23.

Carolin, V. M. & J. A. E. Knopf. 1968. The pandora moth. U.S. Dept. Agric., Forest Serv. Forest Pest

Leaflet, 114.

Danks, H. V. 1987. Insect dormancy: an ecological perspective. Biological Survey of Canada, Ottawa,

Canada.

Fowler, C. S. & N. P. Walter. 1985. Harvesting pandora moth larvae with the Owens Valley Paiute.

J. Calif. Great Basin Anthropol., 7: 155-165.

Manley, T. R. 1993. Diapause, voltinism, and foodplants of Automeris io (Saturniidae) in the south-

eastern United States. J. Lepid. Soc., 47: 303-321.

Massey, C. L. 1940. The pandora moth (Coloradia pandora Blake), a defoliator of lodgepole pine in

Colorado. M. A. Thesis, Duke University, Raleigh, North Carolina.

Ross, D. W. 1995. Short-term impacts of thinning ponderosa pine on pandora moth densities, pupal

weights, and phenology. West. J. Appl. For, 10: 91-94.

Ross, D. W. 1996. Phenology of pandora moth (Lepidoptera: Saturniidae) adult emergence and egg

eclosion in central Oregon. Pan-Pacif. Entomol., 72: 1—4.

Schmid, J. M. 1984. Emergence of adult pandora moths in Arizona. Great Basin Naturalist, 44: 161-

165.

Turnock, W. J., R. J. Lamb & R. P. Bodnaryk. 1983. Effects of cold stress during pupal diapause on

the survival and development of Mamestra configurata (Lepidoptera: Noctuidae). Oecologia,

56: 185-192.

Tuskes, P. M. 1984. The biology and distribution of California Hemileucinae (Saturniidae). J. Lepid.

Soc., 38: 281-309.

Tuskes, P M., J. P. Tuttle & M. M. Collins. 1996. The wild silk moths of North America: a natural

history of the Saturniidae of the United States and Canada. Cornell University Press, Ithaca,

New York.

Weaver, R. A. & M. E. Basgall. 1986. Aboriginal exploitation of pandora moth larvae in east-central

California. J. Calif. Great Basin Anthropol., 8: 161-179.

Received 28 Dec 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(3): 178-180, (1999)

Scientific Note

NEW HOST RECORDS FOR THE BLACK POLYCAON

The black polycaon, Polycaon stoutii (LeConte) (Coleoptera: Bostrichidae) is

a wood-boring beetle which occurs in British Columbia, Washington, Oregon,

California, and Arizona (Ebeling, W. 1975. Urban entomology. Univ. of Calif.

Press, Berkeley, California; Fisher, W. S. 1950. USDA Misc. Publ., 698). It has

been reported to attack alder, curing eucalyptus logs, and three-ply panels which

are used for making desks and other furniture (Doane, R. W. et al. 1936. Forest

insects. McGraw-Hill Book Company, New York). Other host records include

maple, oak, fruit trees, California laurel, madrone, manzanita, sycamore, and other

native California trees (Essig, E. O. 1958. Insects and mites of western north

America. The Macmillan Company, New York). The black polycaon has also

been reported from white ash, Fraxinus americana L. an eastern hardwood species

(Seybold, S. J. & D. L. Wood. 1993. Pan-Pacific Entomol., 69:33-—35).

This paper reports two new host records for the black polycaon and describes

a case history of an infestation in a hardwood warehouse in southern California.

On 30 October 1998, portions of infested hardwoods from a hardwood lumber

warehouse in Santa Ana, Orange County, California, were brought to my office

for identification purposes. Examination of the wood revealed the presence of

adult beetle emergence holes in a few pieces of lumber. Further cutting of the

wood revealed tunnels which were loosely packed with coarse frass (Fig. 1). Two

live larvae, one full grown, were retrieved when portions of infested wood were

dissected. The larvae and the type of damage present in these lumber are consis-

tent with the activity of the black polycaon. An adult P. stoutii subsequently

emerged from infested wood which was held at room temperature in an insect

rearing cage confirming that this infestation was indeed that of the black polycaon.

On 2 November 1998, I visited the lumber yard in question and inspected

several piles of hardwood lumber which were of concern to the owners. Piles of

bostrichid frass were found in several areas of the stacks of lumber. The infested

woods were American cherry (fruitwood), Prunus serotina Ehrhart and North

American black walnut, Juglans nigra L. This is the first record of P. stoutii from

these two eastern hardwood species. The principal source of both of these hard-

woods is the eastern United States (Paxton, E 1987. Beautiful hardwoods. Frank

Paxton Company, Kansas City, Missouri).

According to the lumber company’s records, the American cherry and North

American black walnut in question originated from Bellplaine, Iowa. The infested

lumber had been in storage in their warehouse in Santa Ana since September

1997. The records also show that this lumber was steamed and kiln-dried in Iowa

prior to shipment to California. Upon completion of these processes, the moisture

content of the woods was reported to be in the range of 6 to 8%. These procedures

would have certainly killed any wood-boring insects which may have been present

in the raw lumber. The fact that the black polycaon is not known to occur in Iowa

and the lumber was kiln-dried prior to shipment ruled out the possibility that this

infestation originated at the point of shipment. The owners of this lumber ware-

1999 SCIENTIFIC NOTE 179

Figure 1. Black polycaon damage to hardwood taken from a lumber yard in Santa Ana, California.

house were keenly interested in the origins of this infestation because they were

hoping that the shipper would be liable for the cost of eradicating the infestation.

The owners of this lumber warehouse were concerned about the 22 species of

hardwoods held in storage with an inventory value of millions of dollars. The

source of this black polycaon infestation was from indigenous populations of this

beetle. This bostrichid can complete a generation in one year (Ebeling 1975;

Furniss, R. L. & V. M. Carolin, 1977. Western forest insects. USDA Misc. Publ.,

1339). The infested wood in this incident had been in storage in Santa Ana for

more than one year. This is certainly enough time to allow for adult emergence

and other external manifestations of the infestation. It should be noted here that

moisture content readings taken from both species of hardwoods on 2 November

1998 were 10.1%.

The black polycaon flies at night (Ebeling 1975), and it is attracted to bright

lights particularly mercury vapor lamps. Over the years, I have encountered two

cases where adult beetles were drawn to structures by external lights. In August

1990, on Fourth Street in Rancho Cucamonga, San Bernardino County, California,

black polycaon were attracted by lights to a large defense contractor building,

which was surrounded by wine grape vineyards, in such numbers that they created

a nuisance. In August 1997, P. stoutii was found on and around a large glass-

type structure located on Irvine Center Drive, Irvine, Orange County, California.

This building is illuminated at night by four large banks of light which are located

in concrete pits in the ground at the four corners of the structure. Bright beams

of light are projected upwards on and towards the top of the structure. The prop-

erty management company stated that the illuminated structure served as a good

source of publicity because it attracts the attention of nighttime commuters trav-

eling on the nearby freeway. These lights also attracted large number of other

insects to the structure creating a nuisance.

It is recommended that hardwood lumber yards and lumber warehouses do not

180 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(3)

use exterior mercury vapor lamps. These lamps will attract wood-boring beetles

to the area creating a situation which is conducive to infestations of stored lumber.

Acknowledgment.—I thank Stoy Hedges and Ken Hobbs for reviewing the

manuscript and offering suggestions for its improvement.

Hanif Gulmahamad, Terminix International, 1501 Harris Court, Anaheim, Cal-

ifornia 92806.

Received 8 Dec 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for

discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English,

but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please maintain a copy

of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing.

Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in,

nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF

ANY FIGURES TO THE 8.5 X 11 IN PAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text pages

(pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List

the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right corner. The title

must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use five

to seven words or concise phrases aa KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.

Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases

underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet

1988, Dorly et al. 1989), (Burton in press) and (R. FE Tray, personal communication). For multiple papers by the same author use:

(Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith

1987: table 3).

Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that

volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are

applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED

(INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author

of each species after its first mention.

Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these

format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor. before

submitting manuscripts for which these formats are applicable.

Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. FE Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide

some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. F Dixon & L. Weismann (eds.). Population structure, genetics and

taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB

Academic Publishing, The Hague, The Netherlands.

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.

Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label

letter sizes of at least 1 mm; this reduction must also allow for space below the illustrations for the typeset figure captions. Authors

are strongly encouraged to provide illustrations no larger than 8.5 X 11 in for easy handling. Number figures in the order presented.

Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of

the manuscript, and (5) journal. FIGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures.

Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued on

additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter.

Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited

section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156—161.). A short

acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of

the note with the affiliation underscored.

Page Charges. — PCES members are charged $35.00 per page, for the first 20 (cumulative) pages per volume and full galley costs for

pages thereafter. Nonmembers should contact the Treasurer for current nonmember page charge rates. Page charges do not include

reprint costs, or charges for author changes to manuscripts after they are sent to the printer. Contributing authors will be sent a page

charge fee notice with acknowledgment of initial receipt of manuscripts.

THE PAN-PACIFIC ENTOMOLOGIST

Volume 75 July 1999 Number 3

Contents

UBICK.):—Obituary: Vincent D: Retin: (USDA oF yo a 121

BRAILOVSKY, H. & E. BARRERA—An analysis of the genus Salapia Stal with description

of six new species, and some taxonomic rearrangements (Hemiptera: Heteroptera:

CAOF il is he attr ee AUT) 9 TY aos a a ee eS ee Eee 130

WIESENBORN, W. D.—Sunlight avoidance compared between Hesperopsis gracielae

(MacNeill) (Lepidoptera: Hesperiidae) and Brephidium exilis (Biosduval) (Lepidoptera:

Le VOR CRINCION. oe Bee a 92k be I No Sg ete eek SI ily ee oe mes RR ee 147

MORON, M.-A., S. HERNANDEZ-RODRIGUEZ & A. RAMIREZ-CAMPOS—Description of

immature stages of Phyllophaga (Triodonyx) lalanza Saylor (Coleoptera: Melolonthidae,

eT OV Gti Gh) A es A ene RI a i, 2g td EN Si Slee cn ae 153

AHN, K-J., & J. S. ASHE—Two new species of Giulianium Moore from the Pacific Coasts of

Alaska and California (Coleoptera: Staphylinidae: Omaliinae) —-- 159

GULMAHAMAD, H.—Establishment of an exotic plaster bagworm in California (Lepidoptera:

SSH GVEICORYS) ake Gv wrath! nines A en Aen sO eels: OMA Wile eee PRE Aine uci e se A Dil ee yt le 165

GERSON, E. A., R. G. KELSEY & D. W. ROSS—Pupal diapause of Coloradia pandora Blake

Clistidipherariacurminc ae nce ed a tet) el es oe Ne a ee ee 170

SCIENTIFIC NOTE

GULMAHAMAD, H.—New host records for the black polycaon _ nnn 178

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 75 October 1999 Number 4

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

EDITORIAL BOARD

R. V. Dowell, Editor R. M. Bohart

Jeffery Y. Honda, Associate Editor J. T. Doyen

R. E. Somerby, Book Review Editor J. E. Hafernik, Jr.

Julieta F. Parinas, Treasurer Warren E. Savary

Published quarterly in January, April, July, and October with Society Proceed-

ings usually appearing in the October issue. All communications regarding non-

receipt of numbers should be addressed to: Vincent F. Lee, Managing Secretary;

and financial communications should be addressed to: Julieta F. Parinas, Treasurer;

at: Pacific Coast Entomological Society, Dept. of Entomology, California Acad-

emy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addressed to: William Hamersky, Membership Committee chair, Pacific Coast

Entomological Society, Dept. of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118-4599.

Manuscripts, proofs, and all correspondence concerning editorial matters (but

not aspects of publication charges or costs) should be sent to: Dr. Robert V.

Dowell, Editor, Pan-Pacific Entomologist, California Dept. of Food & Agriculture,

1220 N St., Sacramento, CA 95814. See the back cover for Information-to-Con-

tributors, and volume 73(4): 248—255, October 1997, for more detailed informa-

tion. Information on format for taxonomic manuscripts can be found in volume

69(2): 194-198. Refer inquiries for publication charges and costs to the Treasurer.

The annual dues, paid in advance, are $25.00 for regular members of the So-

ciety, $26.00 for family memberships, $12.50 for student members, or $40.00 for

institutional subscriptions or sponsoring members. Members of the Society receive

The Pan-Pacific Entomologist. Single copies of recent numbers or entire volumes

are available; see 72(4): 247 for current prices. Make checks payable to the Pacific

Coast Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 1999

Laura A. Irons, President Vincent F. Lee, Managing Secretary

Julieta F. Parinas, Treasurer Stanley E. Vaughn, Recording Secretary

THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly for $40.00 per

year by the Pacific Coast Entomological Society, % California Academy of Sciences, Golden Gate

Park, San Francisco, CA 94118-4599. Periodicals postage is paid at San Francisco, CA, and additional

mailing offices. POSTMASTER: Send address changes to the Pacific Coast Entomological Society,

% California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

This issue mailed 30 December 1999

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

The paper used in this publication meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

PAN-PACIFIC ENTOMOLOGIST

75(4): 181-182, (1999)

Obituary: David W. Moss, Jr. (1947-1997)

LARRY G. BEZARK! AND CLIFFORD Y. KITAYAMA2

‘California Department of Food and Agriculture, Sacramento, CA 95814

*Scientific Methods Inc., RO. Box 599, Durham, CA 95938

Dave Moss: That name brings back a lot of memories for those of us who

attended at San Jose State University with him. Although he was well known as

the “party man,’ a moniker not unwarranted, Dave represented the enthusiasm

of a core of young students in the Biology Department hungry for knowledge.

We shared medical entomology and larval taxonomy classes and along with the

encouragement and enthusiasm of J. Gordon Edwards (‘Doc’), our interest in the

field of entomology intensified. Entomology was no longer a simply a branch of

biology to be studied in the laboratory but it became a way of life. It became fun

to collect, study and learn about insects. It became a passion and there were people

who shared that passion and enthusiasm. Dave Moss was one of those individuals

who had that passion. He desired to learn as much as he could about insects and

that meant spending time in the field as well as in the laboratory. Dave loved to

be outdoors and collecting insects was a way to learn about them that transcended

the textbook.

We fondly remember the many great collecting trips with Dave to local haunts

or far away places. The trips up to Mt. Hamilton to collect Rhagium inquistor

(Linnaeus) and trips to the local cherry tree orchard to pick up Synaphaeta quexi

(LeConte), or walking to Williams Street Park, and pulling bark off dead trees in

search of beetles. Weekend trips to blacklight bugs over the hill to Frank Raines

Park, and adventures seeking the cicada-killer wasp, Sphecius. Whittaker’s Forest

trips to collect everything and anything. Walking the creeks of Castle Rock Park

and Uvas creek at Sveadal chasing Amphizoa. Week long trips to Arizona during

spring break or during the summer, driving all night, sleeping in tents or our cars,

running the blacklights all night long and dividing up the catch after we got home.

And while field trips offered a wide array of experiences outside of the class-

room, we spent countless hours with Dave in the laboratory after trips were over,

sorting, sharing and curating specimens, eating Togos sandwiches while poring

over keys trying to figure out the identities of our catch, comparing material with

identified specimens in the collection and wondering why our specimens didn’t

fit the keys.

Dave, like many of Doc’s students gravitated towards the Coleoptera and even-

tually he settled on the elateroids as his group. Dave amassed a sizable collection

of click beetles and developed considerable literature resources with the help of

modern technology. Correspondence with specialists in other countries provided

Dave with specimens and a broader understanding of the group. He was always

willing to provide identifications and many of the click beetles in the San Jose

State University entomology collection bear Dave’s identification labels.

Not being solely devoted to beetles, Dave also had a significant interest in

agricultural entomology and parlayed that interest into a long tenure at Zoecon,

a research firm in the bay area, where he worked in the insectary rearing large

numbers of tobacco hornworms and other species of insects that were used for

182 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

research studies on economic pest problems. Some of the techniques he acquired

in the insectary were shared with fellow students and at times the labs at school

had several rearing projects in progress. One such project involved attempting to

rear larvae of Brachypsectra fulva LeConte, the Texas beetle. The larvae were

collected from under the bark of Eucalyptus trees in the Imperial Valley, and

despite a steady supply of Manduca eggs which Dave provided, these fascinating

immatures fed little and only a few pupated.

Dave Moss was a passionate man who possessed great enthusiasm for the field

of entomology. His enthusiasm was contagious and many fellow students were

affected. He organized numerous collecting trips which turned out to be great

adventures for all those involved. He never let his battle with juvenile diabetes

affect his enthusiasm or his desire to learn. Although Dave’s accomplishments in

entomology did not result in publications, it is appropriate that as a member of

the Pacific Coast Entomological Society (from 1975 through 1982) and faithful

attendee of its meetings at the Califomia Academy of Sciences during his active

entomological career, that he be recognized for his accomplishments. Dave earned

his Bachelors degree in entomology in 1973 and he was an integral part of a time

at San Jose State when entomology was special. Students present during this

period of time shared a common bond, studied together, collected together and

even those who graduated during different eras still share the common bond of

an entomological education from San Jose State. You can find specimens from

his many collecting efforts in San Jose State’s J. Gordon Edwards insect collec-

tion.

What else can you say about him? He always had a grin on his face and always

rolled with the punches. You could always walk into his house and not feel like

you were intruding, in fact most of the time you were likely to be given a meal.

If you were looking for a good time, he was always willing to help you look for

it. If you needed help, Dave was there. He loved clickers.

Dave is survived by his wife Mercy Moss, and their three children; Priscilla,

Tacy and David W. Moss, the third.

Received 26 Oct 1998; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

75(4): 183-199, (1999)

LATE HOLOCENE SONORAN DESERT ARTHROPOD

REMAINS FROM A PACKRAT MIDDEN, CATAVINA,

BAJA CALIFORNIA NORTE, MEXICO

WILLIAM H. CLARK! AND JULIA T. SANKEY?

‘Orma J. Smith Museum of Natural History, Albertson College of Idaho,

Caldwell, Idaho 83605 e-mail: bclark @acofi.edu and

Museo de Entomologia Departamento de Ecologia,

Centro de Investigacion Cientifica y de Educacion Superior de Ensenada,

Apartado Postal 2732, 22830 Ensenada, Baja California, México

?Royal Tyrrell Museum of Paleontology, Box 7500,

Drumheller, Alberta TOJOYO Canada

e-mail: jsankey @ unix1.sncc.lsu.edu

Abstract—Arthropods are reported for the first time from a late Holocene-aged (1770 + 60

radiocarbon years B.P.) packrat (Neotoma) midden located in a granite boulder field in the

Catavifia region of the Central Desert, Baja California, México. More than 23 arthropod taxa

have been identified from 315 fragments found in a subsample of the midden. Ten of these taxa

have not previously been reported from Neotoma midden assemblages. Coleoptera (beetles) and

Hymenoptera, Formicidae (ants) are the dominant forms recovered, possibly reflecting a tapho-

nomic bias. We compared the taxa found in the midden assemblage with that found at the site

today and found little difference, however a study of the midden plants has revealed that the

modern vegetation and climate are drier than ~1770 years ago.

Key Words.—Insecta Neotoma (packrat) midden, Central Desert, Baja California, Mexico, Ho-

locene, arthropods, beetles, ants.

Resumen.—Se reportan por primera vez artr6podos provenientes de un depésito de Neotoma del

Holoceno tardfo (1770 + 60 afios de antigiiedad, fechado por C'*) localizado en un 4rea de rocas

graniticas en la regidn de Catavifia, en el Desierto Central de Baja California, México. Se

identificaron mas de 23 taxa de artr6podos de 315 fragmentos encontrados en una submuestra

del depésito. Diez de estos taxa no habjan sido reportados previamente para los depositos de

Neotoma. Las formas dominantes recuperadas fueron Coleoptera (escarabajos) e Hymenoptera

(hormigas, Familia Formicidae), posiblemente reflejando un sesgo tafonédmico. Comparamos los

taxa encontrados en el depdsito con los que se encuentran actualmente en el sitio y hallamos

poca diferencia, sin embargo un estudio de las plantas del depdsito revel6 que la vegetacion y

el clima modernos son mas secos que hace ~1770 afios.

Fossil arthropods have received recent attention (Carpenter 1992a, b; Buckland

& Coope 1991) especially from packrat (Neotoma spp.) middens (Ashworth 1973,

1976; Elias 1987, 1990, 1994; Elias & Van Devender 1990; Elias et al. 1992;

Hall et al. 1988, 1989, 1990; Hebda et al. 1990; Morgan et al. 1983; Spilman

1976; Van Devender & Hall 1994). The objectives of this paper are to describe

the arthropod assemblage from a midden and to compare it with that presently

found in the area. Neotoma fecal pellets in this midden have been dated at 1770

+ 60 radiocarbon years B.P. Only two other packrat (Neotoma) middens have

been reported in the literature from the peninsula of Baja California, México. The

first is located near San Fernando and was discovered by Philip Wells (Axelrod

1979a, b; Van Devender et al. 1987; Wells 1969, 1976). These reports mention

only two plants (Juniperus and Prunus) and three radiocarbon dates of the midden

(10,000 years BP). The second midden was collected by a College of Idaho field

184 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

expedition during January 1984 to the Catavinia study site (Wells 1986). The

midden contained Juniperus, and was radiocarbon dated at nearly 18,000 years

BP. No detailed description of the middens are given and no mention of arthropods

were reported in these publications. The present study expands this base of in-

formation and documents the first arthropods from a packrat midden in the pen-

insula of Baja California, México.

The packrat currently inhabiting the Catavifia area is Neotoma lepida Thomas

(Huey 1964, Hall 1981). We have live-trapped numerous WN. lepida at this study

Site.

There are a variety of reasons arthropods occur in packrat nests/dens: 1) sea-

sonal inhabitants; 2) transported in on plants; 3) incidental visitors; and 4) those

that are peculiar to the site (Davis 1934). Aalbu & Andrews (1992) have shown

that at least one species of beetle feeds on Neotoma pellets. It may be difficult to

determine exactly how some arthropods ended up in the midden but regardless

the samples make a good datable record of past species and conditions (Van

Devender & Hall 1994). We compared taxonomic groups found in the midden

with those reported from stored food products (Olsen et al. 1996) because there

may be similarities in the microhabitats of the two.

Many other Neotoma middens have been found at the Catavifia site, including

several from the same boulder area. We report here the results of examination of

the youngest midden found to date.

MATERIALS AND METHODS

Location and Habitat—This midden is located 9 km northwest of Rancho

Santa Inés, Baja Califomia, México (Lat. 29°46’ N, Long. 114°46’ W, elevation

550 m). It is a mid-peninsular locality of the Central Desert in the Catavinia region

(Bratz 1976) (Figs. 1 and 2). The area is characterized by the shrubs Larrea

tridentata (Sessé and Mocifio ex Décandolle) Coville, Ambrosia dumosa (A.

Gray) Payne and Ambrosia chenopodifolia (Bentham) Payne, and by the cacti

Opuntia cholla Weber, Opuntia molesta Brandegee, Pachycereus pringlei (S. Wat-

son) Britton and Rose, Lophocereus schottii (Engelm.) Britton and Rose, Fero-

cactus gracilis Gates, and the boojum, Fouquieria columnaris Kellogg (Blom &

Clark 1984). The cholla cactus Opuntia ganderi Rebman & Pinkava is common

at the site (Rebman 1995). Mesquite, Prosopis glandulosa Torrey var. torreyana

(L. Benson) M. C. Johnston is found in the Central Desert (Wiggins 1980) and

at the site and is the dominant plant found in this midden (Sankey et al., unpub-

lished data).

The mean annual precipitation for the Catavifia area is about 96 mm (3.8 in)

with a mean annual temperature of 18.4° C (nm = 14 years, 1956-1967) (Hastings

1964, Hastings & Humphrey 1969). Using 24 years of additional data, Garcia

(1981) reports the mean annual precipitation for the area as 101.7 mm (4 in) and

temperature 19.0° C. Fifty percent of the precipitation occurs in the three winter

months (Dec—Feb) and about another thirty percent of the precipitation occurs

during the fall (Sep—Nov) (Garcia 1981). Spring and summer receive little pre-

cipitation. Blom & Clark (1984) reported 46 mm of precipitation at the site from

9 Jul 1981 through 4 Jan 1982.

Precipitation measured at the site since that time appears to be less than that

reported by Garcia (1981) except for the 1990-1991 winter season (Clark, un-

1999 CLARK & SANKEY: NEOTOMA MIDDEN ARTHROPODS 185

T +

110° !

UNITED STATES

ee

Ly

O

29

wi

RE

Figure 1. Map of the peninsula of Baja California, México, showing general location of the Ca-

tavifia region and an outline of the Sonoran Desert (cross hatched).

El Rosario

Figure 2. Map of the site showing the main areas of granite boulder outcrops (stipled) in the

Central Desert, Baja California, México and the Catavifia site and the location of midden #3.

186 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

published data). Clark et al. (1993) reported 430 mm (16.9 in) of precipitation at

this site from April 1985 through July 1991 for an average of about 61 mm per

year. Mean summer temperatures are 25.8° C and mean winter temperatures are

13.2° C (n = 24 years) (Garcia 1981). The above publications only list monthly

averages of temperature. Through the use of maximum-minimum thermometers

and individual observations during some winter months in some years Clark (un-

published data) has recorded freezing temperatures in this area. During the 1989—

90 winter max-min thermometers in two different areas (one near granite boulders

and one in the open) recorded lows of —3° C and —9° C, respectively.

The weathered Cretaceous granite rocks of the Jaraguay block (Gastil et al.

1975) form ideal areas for the preservation of water soluble middens. The midden

(#3) reported here is located in a boulder pile approximately 10 m in elevation

above the surrounding sandy wash (Fig. 3). The midden is wedged into a crack

of large granitic rock, and is about 3 m below the top of the boulders (Fig. 4).

The front of the midden is exposed to light from an opening in the boulders above

but is protected from direct precipitation.

Midden.—The midden is 2 m in length and 1 m in width (Fig. 4). It is stratified,

with the clearest stratification between the top 0.5 cm of unindurated, loose pale

yellow plant material (subsample a) and the remaining 20—25 cm deep section of

indurated, compacted, and stratified plant material. The radiocarbon date (Beta

Analytic Inc. sample #30453) was obtained from a sample of Neotoma fecal

pellets from the midden.

Material Examined.—A 652 g sample was soaked in distilled water for four days, until it was

completely disaggregated. The arthropod and plant material was recovered after passing the liquid

through a number 20 mesh (0.84 mm) soil sieve. After washing, 237 g of material remained (36.3%).

The liquid was saved for pollen analysis (Sankey et al., unpublished data).

The dry material (237.0 g) remaining was separated into arthropods and vegetation under a dis-

secting microscope using low power. Individual arthropods and arthropod fragments were mounted on

standard entomological card points with water soluble white glue (Borrer et al. 1989).

The arthropod material was then identified using entomological keys (Amett 1985, Borrer et al.

1989) and by consulting specialists. Extant arthropods have been collected at this site since 1976

(Blom & Clark 1980, 1984, 1988; Hovore 1988; Papp 1989; Shook 1989; Leschen 1996; Triplehorn

1996) using pitfall traps (Clark & Blom 1992), mailaise traps, UV light traps, bait traps, pan traps,

and a variety of hand collection techniques (Clark & Blom 1979, 1989, 1992) and these specimens

were used to compare with the arthropod fragments found in the midden.

Voucher specimens of the arthropods, plants, and Neotoma lepida (vertebrate catalog numbers 46,

654, 655, and 656) are located in the Orma J. Smith Museum of Natural History, Albertson College

of Idaho, Caldwell (ALBRCIDA).

RESULTS AND DISCUSSION

The following is a description of the identified arthropod remains from the

subsample (315 arthropod fragments) of this Neotoma midden (Table 1). Taxa

identified are briefly described with their known occurrence in Neotoma middens,

known present occurrence in the Catavifia area (taken from the literature and our

museum collection), and a brief account of their ecological status.

Archnida: Chelonethida (Chernetidae)

Material.—Two palpal chelae. These are from two individuals, one adult and

one a tritonymph.

1999 CLARK & SANKEY: NEOTOMA MIDDEN ARTHROPODS 187

Figure 3. Photographs of the Catavifia site boulder area containing midden #3: a) boulder area

from a distance showing shrubs, including mesquite on the left in the sandy wash; b) close-up of the

boulder area showing no mesquite in the boulders.

188 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

Figure 4. Photograph of the Catavifia site, midden #3 in a granite rock crack. The midden is

approximately one meter in width and two meters in length. The edge of the midden is indicated by

the white arrow.

Occurrence in Packrat Middens.—Pseudoscorpions have previously been re-

ported from middens (Hall et al. 1989, 1990).

Occurrence in the Cataviiia Area.—Several pseudoscorpions of the family

Chernetidae have been collected at this site in pitfall traps and were also found

to be phoretic on Cerambycidae (Coleoptera).

Discussion.—These were the only arachnids found in the midden. The chelae

are very chitinous and might be the only parts of the pseudoscorpions to survive.

Pseudoscorpions often live around rock cracks and in litter and other organic

deposits (Weygoldt 1969, Muchmore 1990) and stored-food products (Olsen et

al. 1996) thus find good habitat in Neotoma nests.

Insecta

The majority of the arthropods collected in this midden are insects and most

of these insects are beetles (Coleoptera) (Table 1). The fact that Coleoptera form

the majority of the assemblage is not surprising considering they are “‘hard-bod-

ied’’ (Elias 1990).

Coleoptera, Anobiidae

Material.—This family is only represented by one fragment.

Occurrence in packrat middens—Only one midden literature record could be

found (Hall et al. 1990).

1999 CLARK & SANKEY: NEOTOMA MIDDEN ARTHROPODS 189

Table 1. Arthropods from 1770 B.P. Neotoma midden sample, Catavifia area, Baja California,

México. *“‘Body, bodies” refer to head, thorax, and abdomen, usually with legs and antennae missing.

Taxa

Material Number

Arachnida

Chelonethida

Chernetidae pedipalpal chela 2

Diplopoda

unidentified segment

Insecta

Coleoptera

Anobiidae body fragment

Bruchidae

Algarobius prosopis (LeConte) bodies* 2

body fragments 31

Carabidae

Harpalus sp. elytra

Lathridiidae

Metophthalmus rudis Fall body

body fragments 10

Ptinidae

Niptus ventriculus LeConte bodies, abdomens 5

Niptus sp. bodies, abdomens 10

Ptinus verticalis (?) bodies, abdomens 17

Tenebrionidae

Blapstinus sp. head

unidentified body fragments o

unidentified Coleoptera body fragments 57

Diptera

Syrphidae (7?) puparium fragment

Hemiptera

unidentified heads Z

Hymenoptera

Apoidea head

Chalcidoidea

Pteromalidae (7?) head

Formicidae

Ecitoninae

Neivamyrmex nigrescens (Cresson) head

Myrmicinae

Pheidole grallipes Wheeler heads 3

Solenopsis xyloni McCook gasters, heads, thoraces 60

unidentified Formicidae heads, gasters, thoraces 26

unidentified Hymenoptera head 7

Lepidoptera

unidentified caterpillar heads 4

unidentified insects body fragments 68

Totals: Total taxa 23+ Total specimens 315

190 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

Occurrence in the Cataviiia Area.—Anobiids are known to occur in this area

and have been collected, but have not yet been determined to species.

Discussion.—Anobiids live in dry plant materials (Amett 1985, Borror et al.

1989) and could thus be expected to be found in a Neotoma den. They are also

found in stored food products (Madenjian 1996).

Coleoptera, Bruchidae, Algarobius prosopis (LeConte)

Material.—Thirty three bodies and body fragments (Fig. 5a).

Occurrence in Packrat Middens.—Literature records for Bruchidae in Neo-

toma middens include Hall et al. (1989, 1990) and Elias & Van Devender

(1990). Algarobius prosopis was one of the bruchids reported by Hall et al.

(1989).

Occurrence in the Cataviiia Area.—Modern Bruchidae recorded for this site

include A. prosopis and Mimosestes protractus (Horn). Algarobius prosopis oc-

curs in Texas, Arizona, and in México: Sonora, Sinaloa, Chihuahua and Baja

California (Kingsolver 1986). Kingsolver (1986) notes that this beetle is the most

common Prosopis bruchid in its range, but does not show it as known from the

Central Desert area.

Discussion.—The dominant identifiable beetle in the midden is the mesquite

beetle A. prosopis (Table 1, Fig. 5a). The Bruchidae or seed beetles are common

in plant seeds especially those of leguminous species (Amett 1985, Borror et al.

1989). Algarobius prosopis is a common seed predator of P. glandulosa Torrey

(Kingsolver et al. 1977, Kingsolver 1986). Because Prosopis glandulosa is the

dominant plant found in the midden (Sankey et al. unpublished data) the beetles

were probably transported into the den by Neotoma carrying in the mesquite.

Bruchids are found in stored food products (Madenjian 1996).

Coleoptera, Carabidae cf. Harpalus

Material.—One elytron (Fig. 5b).

Occurrence in Packrat Middens.—Carabids have been reported from middens

(Elias 1987; Elias & Van Devender 1990; Hall et al. 1989, 1990).

Occurrence in the Catavitia Area.—Many species of Carabidae have been col-

lected and identified at this site, but Harpalus sp. has not been among them to

date.

Discussion.—The ground beetles are the third largest family of beetles and

are common in debris, under objects and on the ground surface (Arnett 1985,

Borror et al. 1989). Carabids are rarely found in stored food products (Madenjian

1996).

Coleoptera, Lathridiidae, Metophthalmus rudis Fall

Material.—One body (head, thorax, abdomen) (Fig. 5c).

Occurrence in Packrat Middens.—Lathnidiidae have been reported from mid-

dens (Elias & Van Devender 1990, Van Devender & Hall 1994). Metophthalmus

rudis has not previously been reported from midden habitats.

Occurrence in the Cataviiia Area.—Modern species of Latharidiidae collected

at this site include Corticarina scissa (LeConte), Dienerella ruthae Andrews, M.

rudius, and Metophthalmus trux Fall.

Discussion.—The family consists of minute beetles which inhabit moldy ma-

1999 CLARK & SANKEY: NEOTOMA MIDDEN ARTHROPODS 191

terials and debris (Amett 1985, Borror et al. 1989). Metophthalmus live in ground

litter and feed on fungi (Andrews 1976). M. rudis has been reported from recent

Neotoma nests in a variety of locations in the coastal area of Central California

(Andrews 1976). Lathridiids are commonly found in stored food products (Mad-

enjian 1996).

Coleoptera, Ptinidae, Niptus ventriculus LeConte and Ptinus verticalis LeConte

Material.—Five bodies and abdomens of N. ventriculus 17 bodies and abdo-

mens of P. verticalis (Fig. 5d).

Occurrence in Packrat Middens.—Ptinidae are common in midden samples

(Elias 1987; Hall et al. 1989, 1990).

Occurrence in the Catavina Area.—Ptinidae are common at this site and are

commonly caught in pitfall traps, but have not yet been identified to species.

Ptinus verticalis is known only from southern California (Papp & Okumura 1959,

Papp 1962). Niptus ventriculus is known from Texas to California south through

Mexico (Papp & Okumura 1959, Papp 1962) and Baja California, including the

Central Desert (Aalbu & Andrews 1992).

Discussion.—The family Ptinidae live in dried plant and animal material and

in animal nests (Amett 1985, Borror et al. 1989). N. ventriculus is known from

Neotoma nests and N. arcanus feeds on Neotoma pellets (Aalbu & Andrews

1992). Ptinids are common in stored food products (Madenjian 1996).

Coleoptera, Tenebrionidae, Blapstinus

Material.——One head.

Occurrence in Packrat Middens.—Tenebrionidae are also common in midden

samples (Elias 1987; Elias & Van Devender 1990; Hall et al. 1989, 1990), and

Blapstinus has previously been reported from packrat midden samples from So-

nora (Hall et al. 1988).

Occurrence in the Catavina Area.—Blapstinus histricus Casey occurs at the

site and Blaisdell (1943) lists an additional seven species of Blapstinus from Baja

California.

Discussion.—Tenebrionidae generally feed on plant materials of some sort

(Borror et al. 1989). The family Tenebrionidae are richly represented in stored

food products (Madenjian 1996).

Diptera, Syrphidae

Material.—One puparium fragment.

Occurrence in Packrat Middens.—Syrphidae have not been reported from mid-

den samples although Stratiomyidae are known from such samples (Hall et al.

1989, 1990).

Occurrence in the Cataviia Area.—Few of the Syrphidae collected at this site

have been identified but we have found Volucella isabellina Williston.

Discussion.—The family contains many species and may be found in a wide

variety of habitats including decaying vegetation (Amett 1985, Borror et al. 1989).

These flies are found in stored food products (Olsen 1996a).

192 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

Figure 5. Photographs of selected arthropods from the Baja California, México, Central Desert,

Catavifia site, midden #3. a) Coleoptera, Bruchidae, Algarobius prosopis (LeConte), entire beetle.

Scale: length is 3.5 mm. b) Coleoptera, Carabidae, cf. Harpalus, elytra. Scale: length is 8 mm. c)

Coleoptera, Lathridiidae, Metophthalmus rudis Fall, entire beetle. Scale: Length is 1.25 mm. d) Co-

leoptera, Ptinidae, Ptinus verticalis LeConte, entire beetle. Scale: Length is 3 mm.

1999 CLARK & SANKEY: NEOTOMA MIDDEN ARTHROPODS 193

Figure 5. (Continued.) e) Hymenoptera, Formicidae, Neivamyrmex nigrescens (Cresson), head and

thorax. Scale: Length is 1.5 mm. f) Hymenoptera, Formicidae, Solenopsis xyloni McCook, petiole,

postpetiole, and gaster. Scale: Length is 1 mm. g) Lepidoptera, unidentified caterpillar, head Scale:

Length is 2 mm.

Hemiptera

Material.—Two unidentified heads.

Occurrence in Packrat Middens.—Hemiptera have been reported previously

from middens (Elias 1987, Hall et al. 1989, 1990).

194 THE PAN-PACIFIC ENTOMOLOGIST Vol. 75(4)

Occurrence in the Cataviia Area.—Many species of Hemiptera are found at

this site today.

Discussion.—Hemiptera are plant feeders and are very common on vegetation

(Amett 1985, Borror et al. 1989).

Hymenoptera, Apoidea

Material.—One head.

Occurrence in Packrat Middens.—Apoidea have not previously been reported

from packrat middens.

Occurrence in the Cataviiia Area.—There are such a variety of species of bees

known from the site that not much can be interpreted by its presence.

Discussion.—Bees are common on flowers where they feed on nectar and act

as pollinators (Amett 1985, Borror et al. 1989).

Hymenoptera, Pteromalidae

Material.—One head.

Occurrence in Packrat Middens.—The family has not previously been reported

from packrat middens.

Occurrence in the Cataviia Area—Pteromalidae have been collected in the

area but remain unidentified.

Discussion.—Pteromalidae are a large family of parasitic wasps (Amett 1985,

Borror et al. 1989), which parasitize a large variety of insect groups and may

have entered the midden within another insect. They are known to parasitize

insects found in stored food products (Avaritt & Richter 1996).

Hymenoptera, unidentified Formicidae

Material.—Twenty six heads, gasters, and thoraces.

Occurrence in Packrat Middens.—Ants are a common component of midden

faunas (Elias 1987; Hall et al. 1989, 1990).

Occurrence in the Catavina Area.—Clark (1980) reported 25 species of ants

from this area representing four subfamilies: Ecitoninae, Pseudomyrmecinae,

Myrmicinae, and Formicinae.

Discussion.—Ants made up the second most abundant group found in the mid-

den (Table 1). The vast majority of identifiable Hymenoptera from the midden

are ants. Avaritt and Richter (1996) report ants from stored food products.

Hymenoptera, Formicidae, Ecitoninae, Neivamyrmex nigrescens (Cresson)

Material.—One head and thorax (Fig. 5e).

Occurrence in Packrat Middens.—The species has been reported from packrat

midden samples twice (Hall et al. 1989, 1990).

Occurrence in the Cataviia Area.—This species occurs at the site (Clark &

Blom 1988). Watkins (1982) records four species of Neivamyrmex for the state

of Baja California Norte, including N. nigrescens.

Discussion.—The ant forages for insect prey (mostly ant brood) at night and

may have strayed into the packrat den.

1999 CLARK & SANKEY: NEOTOMA MIDDEN ARTHROPODS 195

Hymenoptera, Formicidae, Myrmicinae, Pheidole grallipes Wheeler

Material.—Three heads.

Occurrence in Packrat Middens.—Pheidole has been reported previously from

midden collections (Elias 1987; Hall et al. 1989, 1990).

Occurrence in the Cataviiia Area.—Pheidole grallipes is known from this area.

Discussion.—Pheidole grallipes inhabits granitic crevices at this site (Clark et

al. 1986) and is one of four species of this seed harvesting genus to be found

here (Blom & Clark 1980, Clark, unpublished data). It also appears to feed on

other arthropods (Clark et al. 1986). Pheidole is a very large genus of mostly

tropical species (Naves 1985). There are no fossil Pheidole known previous to

the Miocene (Brown 1973).

Hymenoptera, Formicidae, Myrmicinae, Solenopsis xyloni McCook

Material.—Sixty gasters (Fig. 5f), heads, and thoraces.

Occurrence in Packrat middens.—The Solenopsis aurea Wheeler has been re-

ported from middens (Hall et al. 1989, 1990).

Occurrence in the Catavina Area.—Solenopsis xyloni is the only species of the

genus found at the site now (Blom & Clark 1980).

Discussion.—This is the native fire ant and it feeds primarily on other insects