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PAN-PACIFIC

ENTOMOLOGIST

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PAN-PACIFIC ENTOMOLOGIST

77(1): 2-8, (2001)

THE GENUS NOTOBITOPSIS BLOTE WITH THE

DESCRIPTION OF TWO NEW SPECIES (HEMIPTERA:

HETEROPTERA: COREIDAE: COREINAE: MICTINI)

HARRY BRAILOVSKY AND ERNESTO BARRERA

Departamento de Zoologia, Instituto de Biologia Universidad Nacional

Aut6noma de México Apdo Postal 70153, México 04510 D.F, México

Abstract—Two new species of Notobitopsis, N. novoguinensis and N. sandaracinus, collected

in New Guinea are described and illustrated. New records for N. limbativentris and N. militaris

are given and a key to the five known species is provided.

Key Words.—Insecta, Hemiptera, Heteroptera, Coreidae, Mictini, Notobitopsis, new species, New

Guinea.

Stal (1863) and Distant (1911a, b) described the three previously known species

of Notobitopsis and included them in the genus Mictis: M. aruana Distant, M.

limbativentris Stal and M. militaris Distant. Bléte (1938) placed limbativentris

and militaris in a new subgenus of Mictis (Notobitopsis). O’Shea and Schaefer

(1980) revised the tribe Mictini in the Asian and Australian region, recognizing

nineteen genera, and including Notobitopsis which they elevated to generic status

and redescribed.

Within the tribe Mictini, the genus Notobitopsis is characterized by the lack of

a tubercle at the apex of scutellum; the cylindrical fore tibia; pronotum not steeply

declivent with humeral angles rounded, not angulate; middle third of posterior

margins of abdominal sternites III and IV not produced posteriorly; abdominal

stemite III with large tubercle on each side; and posterior margin of abdominal

stemite VI without a median tubercle.

This contribution adds two noteworthy records, two new species and a key to

the five known species of Notobitopsis.

NOTOBITOPSIS ARUANA (Distant)

Mictis aruana Distant 1911b: 389-390.

This is the largest known species of Notobitopsis (length over 34 mm) and

readily distinguishable by the following combination of characters: antennal

segment IV and clavus black, and apical margin of corium yellow to orange

yellow.

Distribution.—This species was described from the Aru Islands and is only

known from the type specimen.

Material Examined—1 male; data: Aru Islands. Type deposited in The Natural History Museum,

London.

NOTOBITOPSIS LIMBATIVENTRIS (Stal)

(Figs. 5 and 6)

Mictis limbativentris Stal 1863: 603.

This species is easily identified within the genus by having the clavus and apical

margin of corium black, and the paramere tip stout and almost straight (Figs. 5

2001 BRAILOVSKY & BARRERA: THE GENUS NOTOBITOPSIS 3

and 6). In N. aruana and N. militaris the corial apical margin is yellow to orange

yellow, and the clavus orange yellow or black with reddish reflections. Notobi-

topsis limbativentris is much smaller than the two other species with a total body

length of less than 30.00 mm.

Distribution.—This species was originally described from Dory, New Guinea

(Stal), and subsequently reported from Salawatti, New Guinea (Blote 1938).

Material Examined.—1 female; data: NE New Guinea. Adelbert Mts., Wanuma, 800-1000 m, 23

October 1958, J. L. Gressitt. Deposited in Bernice P. Bishop Museum, Honolulu, Hawaii. 12 males,

11 females; data: Indonesia Irian Jaya. Baliem Valley nr., Jiwika, 5800-6000’, 24 October 1990, P. J.

Clausen. Deposited in University of Minnesota, Insect Collection.

NOTOBITOPSIS MILITARIS (Distant)

(Figs. 7 and 8)

Mictis militaris Distant 191la: 595-596.

This species is similar to N. aruana (Distant) but slightly smaller, less than

33.00 mm, with clavus orange yellow to dull sanguineous. Notobitopsis aruana

is longer than 34.00 mm, with clavus black.

Distribution.—This species previously was known from Wataikwa River and

Hollandia in New Guinea (Distant 1911la, Blote 1938).

Material Examined.—1 female; data: New Guinea. Saidor Apo 321, March—May 1944, O. H. Gra-

ham. Deposited in Bernice P. Bishop Museum, Honolulu, Hawaii. | male, 2 females; datas NE New

Guinea. Kar Kar Is., Kurum Bagiai Crater trail, 0-100 m, August 1968, N. L. H. Krauss. Deposited

Instituto de Biologia, UNAM, México.

NOTOBITOPSIS NOVOGUINENSIS BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 1, 3 and 4, 11)

Types.—Holotype: male; data: Papua New Guinea. Eastern Highlands, Uba-

gubi, 20 mi S Goroka, 6°22’ S, 145°11’ E, June 1986, G. Dodge. Deposited in

Cornell University, Insect Collection, Ithaca, New York. Paratype: 1 male: same

data as holotype. Deposited in the Coleccién Entomolégica del Instituto de Biol-

ogia, UNAM, México.

Description.—Male (holotype). Dorsal coloration: Head, antennal segments I-IV, pronotum, scu-

tellum, and abdominal segments black; head below each postocular tubercle with small yellow dis-

coidal spot; clavus shiny reddish dark; corium shiny reddish dark, with costal margin black; hemelytral

membrane shining metallic blue; connexival segments IJI—VI and anterior half of VII shiny orange;

posterior half of connexival segment VII black. Ventral coloration. Ground color including rostral

segments and legs black; propleura, mesopleura, and metapleura with large carmine-red spot, one on

each segment; anterior and posterior lobe of metathoracic peritreme yellow; pleural margin of abdom-

inal sterna III—-VII orange, with spiracles black. Structure. Head: Rostrum reaching posterior margin

of mesosternum. Pronotum: Lateral margins finely dentate; humeral angles slightly produced. Legs:

Fore and middle femora with two distinct spines near apex; hind femur longer, incrassate, slightly

curved, attenuated at base, densely granulate, without apical spine; fore and middle tibiae sulcate,

cylindrical; hind tibia sulcate, moderately dilated ventrally, with strong spine at inner face before apex.

Abdomen: Abdominal sternite III with small tubercle on each side. Genitalia (Figs. 1, 3 and 4).—

Genital capsule (Fig. 1): Posteroventral edge thickest, scarcely cordiform; body without tubercles.

Parameres (Figs. 3 and 4): Stout, with elongate and almost straight tip. Measurements. Total body

length: 26.90 mm. Head length: 2.04 mm; width across eyes: 2.66 mm; interocular space: 1.36 mm;

interocellar space: 0.68 mm. Antennal segments length: I, 5.70 mm, II, 5.16 mm, III, 4.18 mm, IV,

4.18 mm. Pronotal length: 5.47 mm; width across frontal angles: 2.43 mm; width across humeral

angles: 7.52 mm. Scutellar length: 3.11 mm; width: 2.96 mm.

4 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Figures 1 and 2. Male genital capsule of Notobitopsis spp. Figure 1. N. novoguinensis Brailovsky

and Barrera, NEW SPECIES. Figure 2. N. sandaracinus Brailovsky and Barrera, NEW SPECIES.

Figures 3-10. Parameres of Notobitopsis spp. Figures 3 and 4. N. novoguinensis Brailovsky and

Barrera, NEW SPECIES. Figures 5 and 6. N. limbativentris (Stal). Figures 7 and 8. N. militaris

(Distant). Figures 9 and 10. N. sandaracinus Brailovsky and Barrera, NEW SPECIES.

Female.—Unknown.

Discussion—This handsome species is readily separable by having the clavus

and corium shiny reddish dark, and the hemelytral membrane metallic blue. The

other known species have black or orange yellow clavus, the corium black, and

the hemelytral membrane shiny dark olivaceous or black. The parameres are dis-

tinct (Figs. 3-10).

Distribution.—Only known from New Guinea.

Etymology.—Named for its occurrence in New Guinea.

2001 BRAILOVSKY & BARRERA: THE GENUS NOTOBITOPSIS 5

Figure 11. Dorsal view of Notobitopsis novoguinensis Brailovsky and Barrera, NEW SPECIES.

6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

NOTOBITOPSIS SANDARACINUS BRAILOVSKY AND BARRERA, NEW SPECIES

(Figs. 2, 9 and 10, 12)

Types.—Holotype: male; data: New Guinea. Finschhafen, 16—21 April 1944,

E. S. Ross. Deposited in California Academy of Sciences, San Francisco, Cali-

fomia (CAS). Paratypes: 9 males, 9 females; data: same locality and date as

holotype. Deposited in California Academy of Sciences, San Francisco California

and Coleccién Entomolégica del Instituto de Biologia, UNAM, México.

Description.—Male (holotype). Dorsal coloration: Head black; antennal segments I-III black red-

dish brown, IV shiny orange with basal joint black; pronotum chestnut orange with anterior margin,

lateral margins, and humeral angles black, and lateral border orange; scutellum chestnut orange with

apex orange; clavus chestnut orange; corium chestnut orange with costal margin black, and costal

border yellow; hemelytral membrane dark shining olivaceous, with blue, green and pink metalic iri-

discence; connexival segments dark chestnut orange with upper margin densely granulate with black;

dorsal abdominal segments dark chestnut orange. Ventral coloration. Head and bucculae shiny orange

with the space close to eyes black; rostral segments I-IV black with chestnut orange reflections;

prosternum, mesosternum, and metasternum chestnut orange; propleura, mesopleura, and metapleura

with large and wide carmine-red spots, one on each segment; upper margin of propleura and meso-

pleura black, and upper margin of metapleura reddish orange; acetabulae reddish brown to shiny

orange; anterior and posterior lobe of metathoracic peritreme shiny orange; coxae with inner face

chestnut orange and outer face reddish brown; trochanters reddish brown; femora chestnut orange with

apical joint black; fore and middle tibiae dark reddish brown, with chestnut orange reflections; hind

tibia chestnut orange with subapical spine and apical third reddish brown; tarsi reddish brown; ab-

dominal sterna dark chestnut orange with spiracles, apex of the tubercle of abdominal sternite III, and

posterior margin of abdominal sternite VII black; pleural margin of abdominal sterna II-VI dirty

yellow; genital capsule black. Structure. Head: Rostrum reaching posterior margin of mesosternum.

Pronotum: Lateral margins densely crenulate; humeral angles obtuse, not exposed. Legs: Similar to

N. novoguinensis. Abdomen: Abdominal sternite II with large and prominent tubercle on each side;

upper margin of connexival segments densely granulate. Genitalia (Figs. 2, 9 and 10).—Genital cap-

sule (Fig. 2): Posteroventral edge thickest, simple, with small tubercles close to middle third and

sinuate. Parameres (Figs. 9 and 10): Stout with broad curved tip. Measurements. Total body length:

30.80 mm. Head length: 2.26 mm; width across eyes: 2.96 mm; interocular space: 1.52 mm; intero-

cellar space: 0.82 mm. Antennal segments length: I, 6.30 mm, II, 5.92 mm, III, 4.71 mm, IV, 4.86

mm. Pronotal length: 6.08 mm; width across frontal angles: 2.66 mm; width across humeral angles:

7.82 mm. Scutellar length: 3.42 mm; width: 3.02 mm.

Female——Coloration: Similar to male. Connexival segments VIII and IX dark chestnut orange;

dorsal abdominal segments VIII and IX, and genital plates black. Structure. Legs: Hind femur slightly

incrassate, granulate with distinct spine near apex; hind tibia sulcate, moderately dilated, unarmed.

Measurements. Total body length: 31.30 mm. Head length: 2.35 mm; width across eyes: 2.99 mm;

interocular space: 1.53 mm; interocellar space: 0.91 mm. Antennal segments length: I, 6.46 mm, II,

5.39 mm, III, 4.25 mm, IV, 4.94 mm. Pronotal length: 6.46 mm; width across frontal angles: 2.87

mm; width across humeral angles: 8.05 mm. Scutellar length: 3.64 mm; width: 3.19 mm.

Discussion.—Antennal segment IV of N. sandaracinus, is shiny orange with

basal joint black, and general color of body chestnut orange. In other species,

antennal segment IV is always black, and the general color of the body black or

black with yellow stripes. Additionally, the shape of the parameres of N. sandar-

acinus, N. limbativentris (Stal), and N. militaris (Distant) are distinct (Figs. 5—

10).

Distribution.—Only known from New Guinea.

Etymology.—From the greek sandaracinos, orange colored.

: THE GENUS NOTOBITOPSIS

BRAILOVSKY & BARRERA

2001

Dorsal view of Notobitopsis sandaracinus Brailovsky and Barrera, NEW SPECIES.

Figure 12.

8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

KEY TO SPECIES OF THE GENUS NOTOBITOPSIS

1. Antennal segment IV shiny orange; scutellum chestnut orange; fore and

middle rémora chestnut orange... 6.5 foe. gh eats te teens ee ages oes

se ee N. sandaracinus Brailovsky and Barrera, NEW SPECIES

1’. Antennal segment IV black; scutellum black; fore and middle femora

IDIACKS 2 SE Shee eel ieee cas teien ce tet WELT ced ee Ss vit ee ee Ge ee 2

2. Clavus shiny reddish dark; corium shiny reddish dark with costal margin

black; hemelytral membrane shining metallic blue ....................

Pt, Oa ee Sa N. novoguinensis Brailovsky and Barrera, NEW SPECIES

2’. Clavus black or orange yellow; corium almost entirely black; hemelytral

membrane dark shining olivaceous or black ...................005. 3

3. Apical margin of corium black; body length less than 30.00 mm. ........

FU ever OP exes PP eee ER, cs cs SAMS SS OPIS SS rees SUBS EO N. limbativentris (Stal)

3’. Apical margin of corium yellow to orange yellow; body length longer

PietriS 0300 gritty Me hotles EN ER la cea Mri hela ee cig 4

epee aA SOS ACT Pg ee Er UN Cte ne ee nts See ee Ce Pe N. militaris (Distant)

4’. Clavus black with reddish reflections; body length longer than 34.00 mm.

As os AEN, A PR ean ORLA Re Hm ins ok oe LRN Pee N. aruana (Distant)

ACKNOWLEDGMENT

We thank the following colleagues and institutions for loaning us specimens

and giving assistance relevant to this study: Gordon Nishida (Bemice P. Bishop

Museum, Honolulu, Hawaii); Mick Webb (The Natural History Museum, London,

U.K.); Norman D. Penny and Vincent Lee (Califomia Academy of Sciences, San

Francisco, Califomia); Richard C. Hoebeke (Cornell University, Insect Collection,

Ithaca, New York); P. J. Clausen (University of Minnesota, Insect Collection).

Special thanks to Albino Luna for dorsal view illustrations.

LITERATURE CITED

Blote, H. C. 1938. Catalogue of the Coreidae in the Rijksmuseum van Natuurlijke Historie Part IV.

Coreinae, Third Part. Zool. Meded., 20: 275-508.

Distant, W. L. 1911a. An Enumeration of the Rhynchota collected during the Expedition of the British

Omnithologists Union to Central Dutch New Guinea. Trans. Ent. Soc. London, 3: 591-604.

Distant, W. L. 1911b. Rhynchota from the Aru Islands. Ann. Mag. Nat. Hist. (Series 8), 8: 389-390.

O’Shea, R. & C. W. Schaefer. 1980. A generic revision of the Asian and Australian Mictini (Heter-

optera: Coreidae). Oriental Insects, 14: 221-251.

Stal, C. 1863. Hemipterorum exoticorum generum et specierum nonnullarum novarum descriptiones.

Trans. Ent. Soc. London, 1: 571—603.

Received 15 Nov 1999; Accepted May 31, 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 9-18, (2001)

THREE NEW SPECIES OF HELICOPSYCHE FROM

VIETNAM (TRICHOPTERA: HELICOPSY CHIDAE)

PATRICIA W. SCHEFTER! AND KJELL ARNE JOHANSON?

‘Centre for Biodiversity and Conservation Biology, Royal Ontario Museum,

Toronto, Ontario, Canada M5S 2C6

Swedish Museum of Natural History, PO. Box 50007,

S-104 05 Stockholm, Sweden

Abstract.—Three new species of Helicopsyche are described from Vietnam: Helicopsyche khe-

moiensis NEW SPECIES, Helicopsyche azunensis NEW SPECIES and Helicopsyche dackles-

tensis NEW SPECIES. Helicopsyche khemoiensis is most closely related to H. coreana Mey,

the two species comprising the subgenus Galeopsyche. Helico psyche khemoiensis is distinguished

from H. coreana by the shape of the gonocoxite and armature of tergite X. Immature stages of

H. khemoiensis are described. Helicopsyche azunensis and H. dacklestensis, subgenus Helicop-

syche, are most closely related to Helicopsyche chrysothoe (Schmid), H. xenothoe (Schmid) and

other species characterized by four-segmented maxillary palps, absence of mesoscutal warts and

distally hooked gonocoxal secondary branch. Helicopsyche azunensis and H. dacklestensis are

distinguished from other species in that group by the presence on the phallus of large endothecal

sclerites.

Key Words.—Insecta, Trichoptera, Helicopsychidae, Helicopsyche, Vietnam, NEW SPECIES.

Biologists from the Centre for Biodiversity and Conservation Biology at the

Royal Ontario Museum, Toronto, are collecting insects, reptiles and amphibians,

and small mammals from the forests of Vietnam in an effort to determine the

diversity and uniqueness of their faunas. Deforestation is proceeding rapidly and

environmentalists are racing to find areas of high endemicity which should be

protected from further degradation. Collections brought back to the Museum for

identification have yielded a number of species unknown to science. Terrestrial

and aquatic insect collections, including Trichoptera, are particularly rich in un-

described species. This paper describes three new species of Helicopsyche (Tri-

choptera) from streams in Vietnam.

Originally described as a pulmonate snail (Say 1821), the larva of the caddisfly

Helicopsyche builds a dextrally coiled sand grain case within which it grazes

periphyton from rock surfaces. The case which bears an uncanny resemblance to

that of a snail provides protection from predators and its shape enhances the

larva’s ability to maintain position in the current of a freshwater lotic habitat; its

sturdy construction resists crushing, and permits the larva to burrow deeply into

the substrate (Williams et al. 1983).

The family Helicopsychidae occurs in all faunal regions and is most diverse in

the Oriental and Neotropical regions. Of the nearly 180 species which have been

described approximately 150 are in the genus Helicopsyche. Only two species of

Helicopsyche, H. coreana Mey from North Korea and Helicopsyche azwudschgal

Malicky from northern Vietnam were previously known as adults from the East

Asian Subregion (sensu Banarescu 1992) (central and northern Vietnam, China,

Korea, the Japanese archipelago, Sakhalin Island, Taiwan, Hainan); Helicopsyche

yamadai Iwata from Japan was described from larval material and undescribed

larvae of the family were also reported from Hong Kong (Dudgeon 1988). The

10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

three species described in this paper, Helicopsyche khemoiensis Schefter & Jo-

hanson NEW SPECIES, Helicopsyche azunensis Schefter & Johanson NEW SPE-

CIES and Helicopsyche dacklestensis Schefter & Johanson NEW SPECIES sub-

stantially increase the known helicopsychid fauna of the subregion. The neigh-

boring South Asian Subregion (sensu Banarescu 1992) (southwestern Vietnam,

Thailand, Malaysia, Laos, Myanmar, the Indonesian archipelago, southern India

and Sri Lanka) harbours approximately 50 species of Helicopsyche, sensu lato

(Malicky & Chantaramongkol 1993; Schmid 1993; Malicky 1994, 1995; Johanson

1998).

Morphological terms used in this paper are adopted from Johanson (1998).

Types are deposited in the Royal Ontario Museum, Toronto, Canada.

HELICOPSYCHE KHEMOIENSIS SCHEFTER AND JOHANSON, NEW SPECIES

(Figs. 1-26)

Types.—Holotype, male; VIETNAM. NGHE AN: West of Con Cuong, Khe

Moi Forestry Camp, 28 Oct 1994, D. C. Currie, ROM946112, coal oil lantern,

tropical forest, Khe Moi River margin. Paratype: female, same data as holotype.

Paratypes, immatures: VIETNAM. NGHE AN: ca. 25 km SW of Con Cuong, Khe

Moi River Forestry Camp, 6 June 1995, Brad Hubley, ROM956170, small stream

100 m upriver of camp, tropical forest edge, 18°56'N, 104°49’E (2 pupae, 2

larvae).

Description —Male. Head (Fig. 1): postantennal warts small, ovoid; cephalic warts large, ovoid;

eyes large, diameter equal to length of head (Fig. 1); antenna with scape swollen, four and one half

times length of pedicel; pedicel and first flagellomere subequal, subsequent four segments graduated

in length, remainder equal to fifth; tentorium (Fig. 2) with anterior arms subparallel, distally slightly

expanded, short posterior arms and broad tentorial bridge; maxillary palps with segments 1 and 2

subequal, apical segment with long setae dorsally. Pronotum with one pair of setal warts. Legs: spur

formula 1,2,4, apical tibial spur of anterior leg slightly longer than one half length of first tarsal

segment (Fig. 3). Wings (Fig. 4): fore wing length 4.6 mm, forks 1,2 and 3 present; fork 1 originates

on distal one third of Dc; R, subequal to Dc; A, and A, fused basally, A,,, fused with Cu, without

reaching posterior wing margin; hind wing 3.7 mm, with 16 hamuli. Abdomen with small, pointed

ViIth sternal process. Genitalia: segment IX, lateral view (Fig. 5), anteriorly nearly oval, with weakly

developed apodemes; dorsally reduced to a sclerous transverse bridge (Fig. 7); in ventral view broad

and deep (Fig. 9); gonocoxite divided into large primary and short secondary branch; in lateral view

(Fig. 5) primary branch slender, arcuate; dorsal and ventral margins subparallel; secondary branches

fused ventromesally forming shelf-like star shaped plate (Fig. 9); superior appendage inserted mesally,

directed posteriad, clavate in lateral view (Fig. 5); Xth tergum depressed in lateral aspect, apices

hooked dorsally; in dorsal view (Fig. 7) apices divergent, separated by V-shaped notch approximately

one fourth length of segment, with short lobate lightly sclerous lateral processes inserted near base of

tergum; macrosetae absent. Phallus slightly arcuate (Fig. 6), apically expanded, with bifurcate sclerous

process inserted dorsally in endotheca (Figs. 6 and 8).

Female.—Head: eyes smaller than in male; cephalic warts small, round; antenna with scape not

swollen, four and one half times length of pedicel, pecidel shorter than first and subsequent flagellom-

eres. Spur formula 1,2,4; apical tibial spur of foreleg one third length of first tarsal segment. Wings

(Fig. 10): fore wing length 5.6 mm; venation as in male, except Dc longer than M,; A,,, reaches

posterior wing margin at some distance before Cu,; hind wing length 3.7 mm, with 19 hamuli; venation

as in male, apex more acuminate than in male. Abdomen with short VIth sternal process, rounded in

ventral view (Figs. 11 and 15). Sternite VIII with row of stiff, clear setae on apical margin. Genitalia:

in lateral view (Fig. 12) segment IX with small group of ventrolateral setae, deep incision between

the [Xth and Xth segments includes the posterior vaginal opening; Xth segment with setae arranged

along dorsal margin of dorsal branch; dorsal branches broad and slightly curved ventrad; in ventral

view (Fig. 14) ventral branch of segment X very small, external part of gonopods IX narrow, arcuate;

2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES 11

superior

appendage

Figures 1-9. Helicopsyche khemoiensis (male).

Figure 1. Head, dorsal view.

Figure 2. Tentorium, dorsal view.

Figure 3. Fore leg.

Figure 4. Wings.

Figure 5. Genitalia, lateral view.

Figure 6. Phallus, lateral view.

Figure 7. Genitalia, dorsal view.

Figure 8. Phallus, dorsal view.

Figure 9. Genitalia, ventral view.

spermathecal sclerite subquadrate; spermathecal duct with anterad-oriented microtrichia at midsection

(Figs. 12 and 13).

Larva—Head (Figs. 16-18): light tan with six darker spots on lateral surface, three or four minute

setae posterolaterally (Figs. 16 and 17, setae 19,20,21); dorsally with dark spots scattered on surface,

and postocular spot with several long setae (setae 13-16); frontoclypeus with characteristically shaped

dark figure on posterior surface; ventrally unicolorous with pair of dark spots near postoccipital margin

(Fig. 18). Thorax (Fig. 19): pronotum with anterior margin bearing large sharp spine-like setae; dor-

sally with scattered setae each surrounded by a dark spot; propleuron forming right angle, tipped by

stout seta (Fig. 20). Legs: foreleg (Fig. 20) with long setae laterally on coxa, long ventral setae on

trochanter and femur; mid leg (Fig. 21) and hind leg (Fig. 22) with thin setae laterally on coxa and

12 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

13}.

vaginal

sclerite

spermathecal

duct

IXth segment

Xth segment

spermatheca

Figures 10-15. Helicopsyche khemoiensis (female).

Figure 10. Wings.

Figure 11. VIth sternal process of abdomen, lateral view.

Figure 12. Genitalia, lateral view.

Figure 13. Genitalia, dorsal view.

Figure 14. Genitalia, ventral view.

Figure 15. VIth sternal process of abdomen, ventral view.

ventrally on trochanter. Abdomen (Fig. 23) with small group of setae dorsolaterally on segment IX;

long setae dorsad of anal prolegs on segment X; abdominal gills absent; anal gills form pair of

expanded lobes; anal claw minute with many teeth. The larval case is similar to that illustrated by

Wiggins (1996a).

Pupal case: anterior enclosure (Fig. 26) of densely woven silk for 75% of surface; the inner, ventral

quadrant with several open bands bearing transverse strands of silk. Pupa: pupal mandibles and hook-

plates as illustrated by Wiggins (1996b, fig. 17-87). Segment IX with long setae; two pairs dorsolateral,

two pairs lateral and two pairs ventrolateral; anal appendages, lateral view (Fig. 24) short, angled

dorsad, with two pairs long setae posteriorly and six pairs ventrally; in ventral view (Fig. 25) oriented

posteriad; the ventral setae in two longitudinal rows.

Diagnosis.—Males of H. khemoiensis can be separated from closely related H.

coreana by the star-shaped gonocoxal plate formed by the fused secondary

2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES

a

pronieroye |

Figures 16-23. Helicopsyche khemoiensis (larva).

Figure 16. Head, lateral view.

Figure 17. Head, dorsal view.

Figure 18. Head, ventral view.

Figure 19. Thorax, dorsal view.

Figure 20. Propleuron and fore leg, anterior view.

Figure 21. Mid leg, anterior view.

Figure 22. Hind leg, anterior view.

Figure 23. Abdominal segments IX and X, lateral view.

25

Figures 24—26. Helicopsyche khemoiensis (pupa).

Figure 24. Segment IX and anal appendages, lateral view.

Figure 25. Segment IX and anal appendages, ventral view.

Figure 26. Pupal case membrane.

13

14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

branches of the gonocoxite; by the Xth tergum which lacks megasetae, is shal-

lowly bifurcate and divergent apically, hooked dorsally and which bears short

baso-lateral processes. Females are recognized by the acuminate hind wing, the

short rounded VIth sternal process and by the large dorsal process of the Xth

segment which curves ventrad concealing the small ventral branch.

Etymology.—khemoiensis, refers to river from which species was collected.

Material examined.—see Types.

HELICOPSYCHE AZUNENSIS SCHEFTER AND JOHANSON, NEW SPECIES

(Fics. 27—33)

Types.—Holotype, male: VIETNAM. GIA LAI: An Khe Dist; Tram Lap, Azun

R., 2 km NW on trail from forestry building, 17 Jun 1996, D.C. Currie, J. Swann,

ROM 961056, UV light, at rainforest edge/coffee plantation, 14°27’ N, 108°33' E.

Description —Male. Head (Fig. 27): antennae with lobate scape about 2X length of pedicel, 1st

flagellomere as long as pedicel; eyes very large; interantennal warts as long as scape breadth, spherical,

cephalic warts large, ovoid, laterad of elevated, triangular vertex; maxillary palps 4-segmented, the

basal two segments together longer than labial palp. Legs: spur formula 1,2,4; fore leg spur longer

than first tarsal segment. Pronotum with 2 pairs setal warts. Wings (Fig. 28): fore wing length 3.0

mm; forks 1,2,3 and 5 present; fork 1 originating on distal one third of Dc, R, subequal to Dc, A,

and A, form basal loop, A,,, reaches wing margin close to Cu,: hind wing 2.3 mm, with 14 hamuli.

Abdomen with small VIth sternal process. Genitalia: segment IX in lateral view (Fig. 29) subtriangular,

with well developed lateral apodeme; narrow ventrally (Fig. 31); anterior margin ogival in dorsal view

(Fig. 30); gonocoxite with large primary and small secondary branch; primary branch in lateral view

(Fig. 29) with dorsal margin gently concave, apically hatchet-shaped; secondary branch slender, arcuate

with slight apical hook (Fig. 31); superior appendage originating dorsally on segment nine, directed

ventrad, clavate in lateral view; Xth tergum depressed in lateral view, in dorsal view (Fig. 30) bifurcate

for one half length, divided into two tapering processes; macrosetae absent. Phallus thick, slightly

curved ventrad (Fig. 32), dorsoapical membranous part of endotheca bifurcate, lobate, with a pair of

slender sigmoid processes each with a small lateral tooth (Figs. 32 and 33).

Female and immature stages.—Unknown.

Diagnosis.—Helicopsyche azunensis is most closely related to H. chrysothoe

(Schmid) and can be separated from it and other species with 4-segmented max-

illary palps by the following characters: gonocoxite lacking an internal branch

(present in H. chrysothoe); with hatchet-shaped primary branch longer than the

tapered slightly notched secondary branch (in most other species of this group

the secondary branch is strongly notched); lobate Xth tergum lacking macrosetae

or notches (in other species the Xth tergum is acuminate and frequently notched

apically); and phallus with a pair of stout sigmoid sclerous processes each with

a subapical tooth (these do not occur in other species in this group).

Etymology.—azunensis, refers to river at collection site.

Material examined.—see Types.

HELICOPSYCHE DACKLESTENSIS SCHEFTER AND JOHANSON, NEW SPECIES

(Figs. 34—39)

Types.—Holotype, male: VIETNAM: GIA LAI, An Khe Dist., Dacklest River,

5.2 km NE Tram Lap on forest road, 28 Jun 1996, D. C. Currie, J. Swann, ROM

961102, UV light, 200 m upstream bridge, 1° rainforest, 900 m, 14°24’ N,

108°33’ E.

Description.—Male. Head: as in Helicopsyche azunensis, except vertex forming a smaller triangle.

2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES 15

Figures 27-33. Helicopsyche azunensis (male).

Figure 27. Head, lateral view.

Figure 28. Wings.

Figure 29. Genitalia, lateral view.

Figure 30. Genitalia, dorsal view.

Figure 31. Genitalia, ventral view.

Figure 32. Phallus, lateral view.

Figure 33. Phallus, dorsal view.

Legs and pronotum as in H. azunensis. Wings (Fig. 34): fore wing length 2.7 mm, forks 1,2,3 and 5

present, fork 1 originating at distal one third of Dc, R, subequal to Dc, A,,, well separated from wing

margin, does not reach wing margin; hind wing 2.1 mm; with 18 hamuli. Abdomen with well devel-

oped VIth sternal process. Genitalia: segment IX, lateral view (Fig. 35), subtriangular and longer

dorsally, with well developed lateral apodeme. Gonocoxite divided into large primary and small sec-

ondary branch; primary branch in lateral view (Fig. 35) clavate with undulate apical margin; secondary

branch slender, arcuate, notched at apex. Superior appendage originates medially on segment nine,

directed ventrad, clavate in lateral view (Fig. 35); Xth tergum slender, slightly sinuate, strongly de-

pressed in lateral view (Fig. 35), in dorsal view (Fig. 36) deeply bifurcate, each tapering lobe with a

small ventrolateral process (Fig. 37); macrosetae absent. Phallus thick, slightly curved ventrally (Fig.

38) dorsoapical membranous endotheca expanded as posterior lobes with embedded paired slender

sclerous processes curved apicolaterally (Figs. 38 and 39).

Female and immature stages.—Unknown.

16

THE PAN-PACIFIC ENTOMOLOGIST

Figures 34-39. Helicopsyche dacklestensis (male).

Figure 34.

Figure 35.

Figure 36.

Figure 37.

Figure 38.

Figure 39.

Wings.

Genitalia, lateral view.

Genitalia, dorsal view.

Genitalia, ventral view.

Phallus, lateral view.

Phallus, dorsal view.

Vol. 77(1)

Diagnosis.—Helicopsyche dacklestensis is most similar to H. xenothoe

(Schmid) and can be separated from that and other species in this group by the

slender and strongly arched Xth tergum, deeply bifurcate with small mesolateral

processes, and the armature of the phallus.

Etymology.—dacklestensis, refers to river at collecting site.

Material Examined.—see Types.

DISCUSSION

Helicopsyche khemoiensis is most closely related to the North Korean H. co-

reana Mey. Both are restricted to the East Asian subregion (sensu Banarescu

1992), the two species comprising the subgenus Galeopsyche (Johanson 1998).

Characteristics supporting monophyly of this group are (plesiomorphic states in

parenthesis): reduction of the lateral pronotal setal warts (present, unreduced);

2001 SCHEFTER & JOHANSON: NEW HELICOPSYCHE SPECIES 17

Figure 40. Collecting sites of Vietnamese Helicopsyche. A. H. awudschgal Malicky. B. H- khe-

moiensis. C. H: azunensis. D. H. dacklenstensis.

fore wing crossvein R,,;—R,,; directed posterobasad (directed ventrad), and fore

wing A,,, meeting Cu, or fusing with the posterior wing margin close to Cu,

(distant to Cu,). East Asian subgenus Galeopsyche and the South American sub-

genus Cochliopsyche are sister groups (Johanson 1998).

Species described in genus Cochliophylax (Schmid 1993) from northeastern

India and Nepal were redesignated as a derived monophyletic clade within the

subgenus Helicopsyche (Johanson 1998). Helicopsyche azunensis and H. dack-

lestensis are closely related to the many Cochliophylax species described by

Schmid, and together with them have 4-jointed maxillary palps, mesoscutal warts

absent, narrow hind wings without crossvein M-Cu, and the gonocoxal secondary

branch tapered and with an apical hook. Helicopsyche azunensis and H. dackles-

tensis are considered sister species based on the presence of phallic endothecal

sclerites. Their relationship to the remaining species in the group is as yet un-

determined.

Distribution of the four known Vietnamese Helicopsyche species is shown in

Fig. 40.

18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

ACKNOWLEDGMENT

We acknowledge the support of the Entomology division of the Centre for

Biodiversity and Conservation Biology at the Royal Ontario Museum, Toronto,

Ontario, Canada. This constitutes Contribution Number 158 of the Centre for

Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario,

Canada.

LITERATURE CITED

Banarescu, P. 1992. Zoogeography of fresh waters. Volume 2. Distribution and dispersal of freshwater

animals in North America and Eurasia. AULAG-Verlag Wiesbaden.

Dudgeon, D. 1988. Preliminary investigations on the faunistic and ecology of Hong Kong Trichoptera.

pp. 111-117 in Bournaud, M. & H. Tachet (eds), Proc. 5th Int. Symp. Trichoptera. Junk, The

Hague.

Johanson, K. A. 1998. Phylogenetic and biogeographic analysis of the family Helicopsychidae (Insecta:

Trichoptera). Ent. Scand. Suppl., 53: 1-176.

Malicky, H. 1994. Zwei neue Helicopsyche (Helicopsychidae) aus Perak, Malaysia. Braueria, 22: 4.

Malicky, H. 1995. Neue Kocherfliegen (Trichoptera, Insecta) aus Vietnam. Linzer Biol. Beitr. 27: 851-

885.

Malicky, H. & P. Chantaramongkol. 1993. Neue Trichopteren aus Thailand. Teil 2: Rhyacophilidae,

Philopotamidae, Polycentropodidae, Ecnomidae, Psychomyidae, Xiphocentronidae, Helicopsy-

chidae, Odontoceridae (Arbeiten iiber thailandische K6cherfliegen Nr. 12) (Fortsetzung). Linzer

Biol. Beitr, 25: 1137-1187.

Say, T. 1821. Descriptions of univalve shells of the United States. J. Acad. Phil., 2, 1.

Schmid, E 1993. Considérations sur les Hélicopsychidés (Trichoptera, Integripalpia). Beaufortia, 43:

65-100.

Wiggins, G. B. 1996a. Larvae of the North American caddisfly genera (Trichoptera) (2nd ed). Uni-

versity of Toronto Press, Toronto, Buffalo, London.

Wiggins, G. B. 1996b. Trichoptera families. In Merrit, R. W. & K. W. Cummins (eds). An introduction

to the aquatic insects of America (3rd ed). Kendall/Hunt, Dubuque, Iowa.

Williams, D. D., A. T. Read & K. A. Moore. 1983. The biology and zoogeography of Helicopsyche

borealis (Trichoptera: Helicopsychidae): a Nearctic representative of a tropical genus. Can. J.

Zool., 61: 2288-2299.

Received 27 Apr 1999; Accepted 7 Feb 2000.

PAN-PACIFIC ENTOMOLOGIST

77(1): 19-27, (2001)

PREY OF THE SPIDER, DICTYNA COLORADENSIS, ON

APPLE, PEAR, AND WEEDS IN CENTRAL WASHINGTON

(ARANEAE: DICTYNIDAE)

EUGENE R. MILICZKY AND CARROL O. CALKINS

Yakima Agricultural Research Laboratory, United States Department of

Agriculture—Agricultural Research Service, 5230 Konowac Pass Road,

Wapato, Washington 98951

Abstract.—The-cribellate spider, Dictyna coloradensis Chamberlin, constructed webs on the up-

per surface of apple and pear leaves (trees not treated with insecticide), and on weeds in adjacent,

uncultivated ground, at a site in south central Washington. Prey found in D. coloradensis webs

were assigned to one of three categories: pests, predators and parasitoids, or neutral in impact

with respect to fruit trees. Pest taxa comprised 32%, predators and parasitoids 24%, and neutral

groups 44% of 18,314 prey. Most prey were small, winged insects (length < 5 mm). Insects

from 58 families in 10 orders were represented and small spiders in four families were occa-

sionally trapped. Sciaridae and Chironomidae (Diptera) were the most numerous prey and made

up 37% of the total. Most webs contained one or more of these flies and occasionally 25 or

more were trapped. Alate aphids were the most frequently captured pest insects. Other pests

included adults of the white apple leafhopper, the pear psylla, and thrips. Relatively non-mobile

stages of the pests (leafhopper and pear psylla nymphs and apterous aphids) were less commonly

found in the webs. Nineteen percent of all prey were parasitoid wasps, 14 families of which

were identified. Known parasitoids of apple and pear pests were included. The only other pred-

ator or parasitoid taxon that comprised more than 1% of total prey was the Empididae (3%).

Key Words.—Arachnida, Araneae, spider, Dictyna, prey use, apple, pear.

The cribellate spider genus Dictyna is represented in the Nearctic region by

more than 100 species (Roth 1993). Dictyna construct irregular mesh webs in a

variety of situations, at times in considerable numbers and high densities (Cham-

berlin & Gertsch 1958, Heidger & Nentwig 1985). Species of Dictyna have fre-

quently been reported from orchards where they are at times abundant. Muma

(1975) found D. florens Ivie and Barrows common and widespread in Florida

citrus where it constructed webs on leaves of orange and grapefruit. Putman

(1967) reported D. annulipes Blackwall to be a common spider in Ontario, Canada

peach orchards where it constructed webs on areas of rough bark. Also in Ontario,

Hagley & Allen (1989) found D. annulipes to be the most abundant foliage-

inhabiting spider in an apple orchard and they studied its prey utilization by

examination of webs and assay of gut contents.

Dictyna coloradensis Chamberlin occurs throughout much of the central and

northern United States and into the Northwest Territories of Canada. With females

approaching 4 mm in length, it is among the larger species in the genus (Cham-

berlin & Gertsch 1958). Dondale (1956) reported D. coloradensis from apple trees

in Nova Scotia, Canada. During 1997, 1998, and 1999 this spider was very abun-

dant on foliage of apple trees at the USDA-ARS research farm near Yakima,

Washington. Webs were less abundant on pear foliage. Large numbers of D. co-

loradensis also constructed webs on tall, dead stalks of annual weeds in adjacent,

uncultivated ground in the spring, and later in the year utilized the current season’s

growth.

Webs of D. coloradensis were collected during 1997, 1998, and 1999 and their

20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

prey contents identified. Because spiders are considered generalist predators (Wise

1993), we were interested in determining the taxonomic range of prey captured

and the relative proportions that fell into three broad categories. 1) Pests: Included

taxa are generally regarded as plant pests although not all specimens were nec-

essarily pests of apple and pear. 2) Predators and parasitoids: All predatory and

parasitic groups were included although not all were known predators and para-

sitoids of apple and pear pests. 3) Neutral: Taxa in this category probably have

little or no detrimental or beneficial impact on fruit trees.

MATERIALS AND METHODS

This study was conducted at the USDA-ARS research farm, 26 km east of

Yakima, Yakima County, Washington. Several small blocks of fruit trees are plant-

ed at the 130 ha farm. Other crops grown include asparagus and potatoes, but

some ground has never been cultivated and native vegetation, dominated by big

sagebrush, Artemisia tridentata Nuttall (Asteraceae), remains within and sur-

rounds parts of the farm. Uncultivated ground with mixed native and introduced

vegetation partly surrounded some of the tree fruit blocks. Fruit trees were not

treated with insecticides.

Pear and apple leaves and weed stems with D. coloradensis webs were placed

in plastic vials with tight fitting lids and refrigerated until examined. Webs were

immersed in 70% isopropyl alcohol in a petri dish and examined under 6.5 X—

50 for prey identification. Prey were identified to family, if possible, using keys

in Borror, Delong, & Triplehorn (1976) and Goulet & Huber (1993). Exceptions

included the following: Chironomidae and Sciaridae, the most abundant Dipteran

prey, were difficult to distinguish when large numbers were present and specimens

were damaged and entangled in webbing. Many specimens were therefore cate-

gorized as unidentified Nematocera. These two families made up the vast majority

of prey in this category. Cyclorrhaphous Brachycera were categorized as medium

size muscoid flies if larger than Drosophila but smaller than a housefly, and as

small muscoid flies if Drosophila-size or smaller. Many chalcidoid wasps were

identified only to superfamily (Chalcidoidea) and many small, non-chalcidoid,

parasitoid wasps were classified as unidentified parasitoid wasps because small

size, damaged specimens, and entanglement in webbing made identification dif-

ficult.

Webs were collected from three apple varieties (“‘Fuji’’, ‘““Golden Delicious’’,

and ‘Red Delicious’’), pears (mixed “‘Anjou”’ and “‘Bartlett’’ varieties), and dead

weed stems in uncultivated land adjacent to the “‘Fuji’”’ apples. Weeds were pri-

marily tumble mustard, Sisymbrium altissimum Linnaeus (Brassicaceae), an intro-

duced species. A total of 984 webs was examined, distributed among the plant

types as indicated in Table 1. Five to 15 webs were collected from a plant type

per sample date at one to two week intervals. The sampling periods were 22 May

to 16 Oct 1997, 30 Mar to 3 Nov 1998, and 20 Jul to 1 Oct 1999. Each plant

type was not sampled throughout each sample period.

RESULTS

Web Placement.—Most webs on apple and pear were constructed on the upper,

concave surfaces of leaves. The small webs of young spiders covered only 2 or

3 cm? and were usually near the leaf apex. Webs of older, larger individuals often

2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 21

Table 1. Number of sample dates and total number of Dictyna coloradensis webs examined from

different plants during 1997, 1998, and 1999.

No. of samples dates—No. of webs examined

Plant 1997 1998 1999

Apple—“‘Fuji”’ 11-141 19-192 —

Apple—“Golden”’ 10-57 10-56 —

Apple—“‘Red”’ 9-59 — —

Pear 5-17 9-91 11-155

Weeds —_ 18-216 —

covered most of the upper surface of a leaf. Leaves were up to 10 cm long. Prey

accumulated in webs and older, larger webs contained up to 40 or more prey.

Webs were occasionally found in the angle formed by two branches, among flower

petioles, or between a leaf petiole and a branch. They were less visible in these

locations and few were sampled. Webs on dead weed stalks were constructed

among branches on the upper part of a stalk, 0.3—-1.0 m above ground.

Prey Utilization.—Fifty-eight families of insects in 10 orders and four families

of spiders were identified from webs of D. coloradensis (Table 2). Some taxa

were represented by few specimens (two Ephemeroptera among 18,314 prey)

whereas others made up a high proportion of prey in webs from all sources all

three years.

Insects ‘classified as neutral with respect to impact on fruit trees made up the

largest proportion of prey items overall (7981 prey = 44%). Nematocerous Dip-

tera, primarily Sciaridae (dark-winged fungus gnats) and Chironomidae (midges)

were the most abundant prey of any kind (37% overall). Sciarids and chironomids

were present throughout the season and were found in a majority of webs re-

gardless of source. Webs occasionally contained 25 or more of these small insects.

Other taxa of neutral prey rarely comprised more than 1% of the total from a

plant in one year. Small muscoid flies, however, made up 5.4% of prey in webs

from “‘Golden Delicious” in 1997.

Aphids were the most numerous pest insects found in D. coloradensis webs,

and alates were generally much more abundant than apterous forms. Five to 30%

of the total insects found in webs from each plant type each year were aphids.

Aphids were not identified to species because of the large number captured and

their often poor state of preservation (discoloration, damage, dehydration). Several

Species are considered pests of apple in Washington and many appeared to be

green apple aphid, Aphis pomi DeGeer, or the nearly identical spirea aphid, A.

spireacola Patch. Apple/spirea aphid colonies were abundant on developing apple

shoots all three years. Thrips (Thysanoptera) made up 5—11% of total prey in the

samples from fruit trees but were more abundant in webs on weeds (22%). The

western flower thrips, Frankliniella occidentalis (Pergande) is the only species

listed by Beers et al. (1993) as a pest of tree fruit in Washington. A pale, yellowish

insect, its host range includes several fruit trees, alfalfa, potatoes, and numerous

species of weeds (Beers et al. 1993). Thrips were not identified to species due to

small size, entanglement in webbing, and poor preservation. The vast majority,

however, were pale, yellowish insects, in general resembling F. occidentalis.

White apple leafhopper adults, Typhlocyba pomaria McAtee, were most abundant

22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Table 2. Total number of prey items in each taxon found in Dictyna coloradensis webs from apples,

pears, and weeds.

Prey taxa Apples Pears Weeds

Neutral impact taxa

Chironomidae 195 163 745

Sciaridae 760 238 112

Bibionidae 25 0 94

Psychodidae 6 2 1

Scatopsidae 1 4 1

Simuliidae 1 3 0

Tipulidae 1 1 0

Nematocera—unidentified 2873 1022 590

Stratiomyidae 1 0 0

Therevidae 4 0 0

Bombyliidae 2 0 3

Drosophilidae 175 78 28

Phoridae 38 29 8

Conopidae 0 0 1

Small muscoid flies 375 100 87

Medium muscoid flies 30 26 10

Diptera—unidentified 0 0 5

Halictidae 12 7 0

Chrysididae 1 0 0

Tenthredinidae 0 0 1

Scarabaeidae 2 0 pi

Coleoptera—unidentified 6 1 6

Aleyrodidae 1 0 0

Fulgoroidea 0 0 1

Lygaeidae 6 0 17

Hemiptera—unidentified 2 1 3

Psocoptera 42 18 9

Ephemeroptera 1 1 0

Pest taxa

Aphididae—alate 1549 1011 431

Aphididae—apterous 230 67 5

Typhlocyba pomaria 289 ms) 2D

Other Cicadellidae 18 1 2

Cacopsylla pyricola—adults 3 165 1

Cacopsylla pyricola—nymphs 0 50 0

Phyllonorycter elmaella 193 37 3

Lepidoptera—unidentified 3 0 3

Caterpillar 0 2 0

Mindae—Lygus sp. Z 0 3

Mindae—Campylomma sp.? 12 1 1

Thysanoptera 687 281 777

Acari 3 5 0

Predator and parasitoid taxa

Cecidomyiidae 6 5 4

Empididae 387 132 6

Dolichopodidae 11 4 0

Pipunculidae 25 9 3

Tachinidae 6 6 1

Syrphidae 2 0 0

Braconidae 349 56 81

Ichneumonidae 76 16 18

2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 23

Table 2. Continued.

Prey taxa Apples Pears Weeds

Pnigalio flavipes 177 63 3

Trechnites insidiosus 0 50 0

Mymaridae 641 17 20

Encyrtidae 0 0 1

Chalcidoidea—unidentified 713 373 106

Platygastridae 40 16 21

Scelionidae 25 14 5

Ceraphronidae 73 14 16

Megaspilidae 2 2 1

Proctotrupidae 3 4 1

Dryinidae 1 1 2

Bethylidae 3 0 6

Diapriidae 1 0 3

Cynipoidea 12 3 1

Parasitoids—unidentified® 249 94 123

Sphecidae 8 8 0

Formicidae 15 7 133

Vespidae 1 1 0

Staphylinidae 39 4 35

Carabidae 2 0 3

Coccinellidae PL 1 0

Hemerobiidae 6 2 0

Chrysopidae ys 0 0

Anthocoridae—Orius 27 5 of

Nabidae—Nabis 0 0 1

Lygaeidae—Geocoris 0 0 5

Miridae—Deraeocoris 2 2 0

Salticidae 7 1 1

Liny phiidae—Erigone 13 2. 2

Linyphiidae 10 9 17

Thomisidae 2 0 0

Oxyopidae 6 1 0

Araneae—unidentified 0 2 0

Prey totals 10,493 4243 3578

4 Campylomma also act as predators by feeding on such pests as aphids and mites.

> Category includes only hymenopteran parasitoids.

in webs from “Fuji” apples during 1998 when they comprised nearly 5% of all

prey. Typhlocyba comprised less than 2% of total prey in the other samples, and

nymphs were rarely captured. Adult, western tentiform leafminer, Phyllonorycter

elmaella Doglanar and Mutuura, made up 1—2% of the prey in each tree fruit

sample, but only three of 3578 prey in webs from weeds. Pear psylla, Cacopsylla

pyricola (Foerster), a serious pest of pear in the Pacific Northwest, made up 5.5%

of the prey in D. coloradensis webs from pear during 1997, just under 1% in

1998, and 8% in 1999. Psylla populations at the research farm were low during

1998, probably accounting for their rarity as prey despite the greater number of

webs examined compared to 1997. Psylla numbers were higher during 1999 when

leaves and shoots were often sticky with honeydew, and this was reflected in the

number captured in D. coloradensis webs. Most captured C. pyricola were adults,

although a substantial number of nymphs fell victim in 1999 (50 nymphs, 134

24 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

adults). However, it was difficult, at times, to distinguish psylla nymphs from

their cast skins.

Small, hymenopterous parasitoids were the most abundant predatory and par-

asitoid insects trapped in D. coloradensis webs. They comprised 12% to 28.6%

of total prey in yearly samples from each of the plants. Hosts of many of the

parasitoids were not determined or are unknown, some are probably hyperpar-

asitoids, and some probably occurred only incidentally in the fruit trees. Known

parasitoids of apple and pear pests were, however, captured. Pnigalio flavipes

(Ashmead) (Eulophidae), the most common parasitoid of the western tentiform

leafminer in the Pacific Northwest (Beers et al. 1993), made up 1%—2% of total

prey in samples from fruit trees. It was rarely found in webs on weeds. Mymaridae

(fairyflies) were captured in substantial numbers in webs on “Fuji”? (449 =

11.9%) and “‘Golden Delicious” (132 = 6.6%) during 1998. Both represented

large increases over numbers found in 1997. Mymarids in the genus Anagrus are

important egg parasitoids of the white apple leafhopper and parasitism rates of

up to 70% have been reported in unsprayed orchards (Beers et al. 1993). Many

mymarids found in webs on apple may have been leafhopper egg parasitoids. Two

percent of the insects in webs from pear during 1999 were Trechnites insidiosus

(Crawford) (Encyrtidae), the most important parasitoid of pear psylla in western

North America (Beers et al. 1993).

Other taxa of insect predators and parasitoids rarely comprised more than 1%

of the total prey in webs from a given plant type (Table 2). Empididae, however,

made up 7.5% (233 flies) of the prey found in webs on “Fuji” apple in 1997.

Spiders also were infrequently snared in D. coloradensis webs (< 1% of prey in

samples from any of the plant types) and were small, either immatures or taxa of

small body size.

DISCUSSION

The diversity of prey captured by D. coloradensis is in agreement with the idea

of spiders as generalist predators (Wise 1993). Individual species, however, utilize

a restricted range of available prey depending on factors such as spider size,

hunting strategy, and web size and placement (Marc & Canard 1997). Spiders

generally feed on prey smaller than themselves (Jackson 1977, Nyffeler et al.

1994). This was true of D. coloradensis, the vast majority of whose prey consisted

of insects less than 5 mm in length. The predominant use of small prey has also

been noted in D. segregata (Nyffeler et al. 1988), D. arundinacea (Heidger &

Nentwig 1985), and 11 species, including D. coloradensis, studied by Jackson

(1977). Large and dangerous prey were, however, captured occasionally. Two

worker yellow-jackets (Vespula sp.) were found in 1997 webs and on 26 Apr

2000 a recently captured worker honeybee, Apis mellifera L., was noted in the

web of a female on a dead weed. The spider was feeding on the bee at the time.

Overall, however, few prey larger than a housefly were found in the webs. No

codling moth, Cydia pomonella (L.) or leafrollers, Pandemis pyrusana Kearfott,

were captured. Both are important apple pests in Washington (C. pomonella was

very abundant at the farm in 1997-1998), but at adult lengths of 12 mm or more,

they may be too large to be readily subdued by D. coloradensis webs.

A given species of spider may not utilize all stages in a prey species’ life cycle

to equal degrees (Marc and Canard 1997). This was true for several prey species

2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 23

of D. coloradensis, most or all of whose life cycles are spent on the fruit trees.

Adult white apple leafhopper, adult pear psylla, and alate aphids were common

prey but immature leafhoppers and psylla and apterous aphids fell victim less

frequently. This is probably related to the relative mobility of different stages of

the prey and their locations on the plant. White apple leafhopper and pear psylla

adults are mobile and fly readily when disturbed. They would appear much more

likely to blunder into webs than the more sedentary nymphs. Also, white apple

leafhopper nymphs generally feed on the lower surface of the leaf (Beers et al.

1993) whereas D. coloradensis webs are constructed almost exclusively on the

upper surface. Psylla nymphs, with their flattened body form and leaf-surface

hugging habits were often found alive beneath D. coloradensis webs—apparently

able to avoid entanglement in the silk. Alate aphids, although not strong fliers,

are more mobile than the apterous forms and again must be more likely to come

into contact with webs. Green apple aphid colonies generally develop on succu-

lent, young tissue and are found on growing shoot tips, shoot stems, and the

undersides of leaves (Beers et al. 1993). Thus the sedentary, apterous forms are

less likely to come into contact with D. coloradensis webs.

Dictyna coloradensis webs trapped many small (1—3 mm), hymenopterous par-

asitoids (19.1% of all prey). Some parasitoids were observed crawling over leaf

surfaces, which must often bring them into contact with webbing. This is true of

P. flavipes and T. insidiosus females, both of which search leaf surfaces for hosts

(Beers et al. 1993), and many became entangled in the webs. Nearly the same

number of male P. flavipes (122) were captured as females (118). T. insidiosus

is thelytokous in the western states (Unruh et al. 1995) and males were not found

in the webs. Thirteen percent of the prey of D. arundinacea (L.) consisted of

small parasitoid wasps (Heidger & Nentwig 1985).

Predatory insects, and parasitoids other than Hymenoptera, were infrequently

found in D. coloradensis webs. Deraeocoris spp. (Miridae), for example, are

important predators of pear psylla (Beers et al. 1993) and were abundant on the

pears during 1999. Yet Deraeocoris accounted for only 0.09% of the prey in webs

on pear during 1999. Aphid predators such as Coccinellidae and Chrysopidae,

often abundant in unsprayed orchards with high aphid populations, made up sim-

ilarly low percentages of prey in all samples. Such insects, perhaps because of

size, behavior, and distribution on the plants may not be very vulnerable to en-

tanglement in D. coloradensis webs.

Other spiders were infrequent prey in D. coloradensis webs. The low number

taken (0.4% of total prey) is in accord with Nyffeler’s (1999) findings that web

building spiders are 99% insectivorous whereas hunting types take a higher pro-

portion of other spiders. Jackson (1977), for example, found that 27% of the prey

of Phidippus johnsoni (Peckham and Peckham) (Salticidae) was other spiders.

Capture of beneficial insects and spiders by D. coloradensis (intraguild pre-

dation in a broad sense) was substantial in terms of the overall numbers captured.

However, the hosts of many of the parasitoids may not be orchard pests and their

capture may have little negative impact on orchard ecology from a pest manage-

ment standpoint. Greenstone (1999) noted that the net effect of intraguild pre-

dation can only be determined by examining the system in the presence and

absence of the predator. Although intraguild predation and competitive interac-

tions among predators may in some cases disturb natural pest control, in others

26 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

they may promote greater spider biodiversity and allow spiders to survive periods

of low prey density (Sunderland 1999). |

Several studies have shown that small Diptera are important components in the

diet of Dictyna spiders. Several families of small flies made up 50% of the prey

captured by D. arundinacea in a meadow in Germany (Heidger & Nentwig 1985)

and small Diptera dominated the prey of each of 11 Dictyna species studied by

Jackson (1977). Chironomidae comprised 70.7% of the total prey of D. annulipes

in an apple orchard in Ontario, Canada (Hagley & Allen 1989), and a diverse

array of small Diptera were important components in the diet of D. coloradensis

in Washington. The abundance of these small flies was likely important in sup-

porting the high population of D. coloradensis observed during this study. Also,

their presence during much of the season may help fill in gaps in availability of

other types of prey, as noted by Sunderland (1999) with respect to intraguild

predation.

Clearly, D. coloradensis is a polyphagous predator that includes a wide range

of insects in its diet. Prey selectivity is based primarily on size and activity level—

small, active insects are most heavily utilized. Small, active pests of apple and

pear are taken in substantial numbers, but unfortunately, from a pest management

standpoint, known parasitoids of some pests are also trapped, sometimes in con-

siderable numbers.

ACKNOWLEDGMENT

We thank Matthew H. Greenstone, Thomas R. Unruh, and an anonymous re-

viewer for their constructive comments on the manuscript.

LITERATURE CITED

Beers, E. H., J. EK Brunner, M. J. Willett & G. M. Warner. (eds.). 1993. Orchard pest management: A

Resource book for the Pacific Northwest. Published by Good Fruit Grower, Yakima, Washing-

ton.

Borror, D. J.. D. M. DeLong & C. A. Triplehorn. 1976. An introduction to the study of insects (4th

ed.) Holt, Rinehart, and Winston, New York.

Chamberlin, R. V. & W. J. Gertsch. 1958. The spider family Dictynidae in America north of Mexico.

Bull. Am. Mus. Nat. Hist., 116: 1-152.

Dondale, C. D. 1956. Annotated list of spiders (Araneae) from apple trees in Nova Scotia. Can.

Entomol., 88: 697—700.

Greenstone, M. H. 1999. Spider predation: how and why we study it. J. Arachnol., 27: 333-342.

Goulet, H. & J. T. Huber (eds.). 1993. Hymenoptera of the world: An identification guide to families.

Research Branch, Agriculture Canada. Publication 1894/E.

Hagley, E. A.C. & W.R. Allen. 1989. Prey of the cribellate spider, Dictyna annulipes (Araneae,

Dictynidae), on apple tree foliage. J. Arachnol., 17: 366-367.

Heidger, C. & W. Nentwig. 1985. The prey of Dictyna arundinacea (Araneae: Dictynidae). Zool. Beitr.

N. E 29: 185-192.

Jackson, R. R. 1977a. Comparative studies of Dictyna and Mallos (Araneae, Dictynidae): III. Prey

and predatory behavior. Psyche, 83: 267-280.

Jackson, R. R. 1977b. Prey of the jumping spider Phidippus johnsoni (Araneae: Salticidae). J. Arach-

nol., 5: 145-149.

Marc, P. & A. Canard. 1997. Maintaining spider biodiversity in agroecosystems as a tool in pest

control. Agric. Ecosyst. Environ., 62: 229-235.

Muma, M. H. 1975. Spiders in Florida citrus groves. Fla. Entomol., 58: 83-90.

Nyffeler, M. 1999. Prey selection of spiders in the field. J. Arachnol., 27: 317-324.

Nyffeler, M., D. A. Dean & W. L. Sterling. 1988. Prey records of the web-building spiders Dictyna

segregata, (Dictynidae), Theridion australe (Theridiidae), Tidarren haemorrhoidale (Theridi-

2001 MILICZKY & CALKINS: DICTYNA COLORADENSIS PREY 27

idae), and Frontinella pyramitela (Linyphiidae) in a cotton agroecosystem. Southwestern Nat.,

33: 215-218.

Nyffeler, M., W. L. Sterling & D. A. Dean. 1994. How spiders make a living. Environ. Entomol., 23:

1357-1367.

Putman, W. L. 1967. Prevalence of spiders and their importance as predators in Ontario peach orchards.

Can. Entomol., 99: 160-170.

Roth, V. D. 1993. Spider genera of North America with keys to families and genera and a guide to

literature (3rd ed.). (Distributed by American Arachnological Society). U. of Florida, Gaines-

ville.

Sunderland, K. 1999. Mechanisms underlying the effects of spiders on pest populations. J. Arachnol.,

27: 308-316.

Unruh, T. R., P. H. Westigard & K. S. Hagen. 1995. Pear psylla. Chapter 19. pp. 95-100. In Nechols,

J. R., L. A. Andres, J. W. Beardsley, R. D. Goeden, & C. G. Jackson. 1995. Biological control

in the western United States. Accomplishments and benefits of regional research project W-84,

1964-1989. Univ. Calif. Div. Agric. Nat. Res. Publ. 3361.

Wise, D. H. 1993. Spiders in ecological webs. Cambridge University Press, Cambridge, U.K.

Received 29 Nov 1999; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 28-36, (2001)

OVERWINTERING POTENTIAL IN CALIFORNIA OF

TWO RUSSIAN WHEAT APHID PARASITOIDS

(HYMENOPTERA: APHELINIDAE ET APHIDITDAE)

IMPORTED FROM CENTRAL ASIA

JULIO S. BERNAL!, D. GONZALEZ, & ERNESTO DAVID-DIMARINO

Department of Entomology, University of California,

Riverside, California 92521-041

Abstract.—Aphelinus albipodus Hayat and Fatima and Diaeretiella rapae (M’Intosh) (Hyme-

noptera: Aphelinidae and Aphidiidae) were imported to the USA for biological control of Russian

wheat aphid (Diuraphis noxia Mordwilko) in 1992. Early laboratory studies identified potential

limitations for their successful establishment in northern California where these parasitoids were

extensively colonized and evaluated against Russian wheat aphid. Specifically, those studies

showed that A. albipodus was particularly susceptible to moderately low temperatures (10° C).

These results for A. albipodus were in contradiction with seasonal field temperatures prevalent

in its collection site (Tahcheng, People’s Republic of China) where field temperatures are below

freezing for several months each year. This study sought to reconcile these apparently contra-

dictory results. We examined whether short daylength-induced diapause, i.e. winter diapause,

occurs in A. albipodus and D. rapae, and thus could be used as an overwintering strategy by

these parasitoids. Under laboratory conditions, > % of A. albipodus individuals entered diapause

when daylength was reduced from 14 h to 12 h. In contrast, diapause was not detected in D.

rapae under the same conditions. However, we suggested that diapause may occur in D. rapae

under shorter daylengths because (1) this parasitoid’s development and survivorship are not af-

fected by temperatures associated with 12 h daylengths at its collection site (Wuqia, PRC), and

(11) other studies have demonstrated diapause in this parasitoid at daylengths < 12 h. We discuss

our results in light of A. albipodus’ rapid establishment in northern California, and the use of

our analytical procedure as a component of a process for screening natural enemies in importation

biological control programs.

Key Words.—Insecta, Aphelinus albipodus, Diaeretiella rapae, diapause, biological control, Di-

uraphis noxia, establishment, importation.

Numerous biological control campaigns involving exotic natural enemies have

been successful since the introduction of the vedalia beetle to California more

than 100 years ago (DeBach & Rosen 1991; Greathead & Greathead 1992). Ex-

perience gained during that time and theoretical analyses have led to the identi-

fication of traits that presumably make natural enemies effective biological control

agents (Huffaker et al. 1974, 1977; Beddington et al. 1978; Murdoch et al. 1985;

Luck 1990; Murdoch & Briggs 1996). However, at this time it is uncertain wheth-

er these traits are useful or efficient for identifying the most promising natural

enemies among a suite of available natural enemies (Luck 1990; Gonzalez &

Gilstrap 1992). In contrast, identifying natural enemies that offer promise for

successful establishment in a new area seems a more tenable goal. Characterizing

an exotic natural enemy’s potential for establishment is important because lack of

successful establishment precludes its eventual success in suppressing a pest pop-

ulation.

Climate (excessively cold, hot, or dry seasonal weather) is the single most

‘Current Address: Biological Control Laboratory, Texas A&M University, College Station, Texas

77843-2475.

2001 BERNAL ET AL.: PARASITOID OVERWINTERING 29

frequently cited reason for natural enemy failure in biological control campaigns

(Stiling 1993). Climate can early on limit a natural enemy’s potential for success

by impeding its permanent establishment following colonization. Close matching

between the climates of collection and colonization areas of natural enemies used

in classical biological control campaigns, 1.e., climate-matching, is a long-standing

conceptual “‘rule of thumb” for improving the chances of successful establishment

of exotic natural enemies (Flanders 1940; Messenger 1959, 1971). It is unclear,

however, how much emphasis is commonly placed on climate-matching in the

planning phase of foreign exploration efforts. A comparatively information-inten-

sive approach to climate-matching has been used successfully for forecasting

range expansions of exotic pests (Messenger & Flitters 1954; Meats 1989; Hughes

& Maywald 1990).

Shortly after the discovery of the Russian wheat aphid, Diuraphis noxia Mord-

wilko (Homoptera: Aphididae), in the USA in 1986, a biological control campaign

involving several state, federal, and university agencies was organized against this

pest. Numerous Russian wheat aphid natural enemies were imported to the USA

as a result of this combined effort (Gilstrap et al. 1994). Based on information

available from their collection areas, a limited suite of parasitoids was selected

from among these natural enemies for evaluation in California (Gonzélez et al.,

unpublished data). Among these parasitoids were Aphelinus albipodus Hayat and

Fatima and Diaeretiella rapae (M’Intosh) (Hymenoptera: Aphelinidae and Aphi-

diidae, respectively). A series of studies identified potential limitations for suc-

cessful establishment of these parasitoids in California (Bernal & Gonzalez 1995,

1996, 1997; Bernal et al. 1997). In general, those studies suggested that A. albi-

podus was less tolerant of low temperatures than D. rapae, but the reverse was

true at high temperatures. The implications of these results were that establishment

in California of A. albipodus could be restricted by winter temperatures, but es-

tablishment of D. rapae could be restricted by summer temperatures. However,

these results and their implications were in disagreement with available infor-

mation concerning the climate of each of these parasitoid’s collection areas (ex-

tremely cold and long winters, and moderate summers; see below). This was

particularly true with regard to A. albipodus’ susceptibility to low temperatures

in the laboratory (Bernal & Gonzalez 1996; Bernal et al. 1997), and its rapid

establishment in northern California (Gonzalez et al., unpublished data).

Based on these observations, we examined whether A. albipodus and D. rapae

enter diapause in response to a short daylength, 1.e., winter diapause. Entry into

diapause under short daylength conditions would explain how these parasitoids

survive the low temperatures prevalent during the late fall through early spring

in their collection areas, and would influence their performance against Russian

wheat aphid in California. Based on the results of this and previous studies we

suggest that A. albipodus and D. rapae are able to survive the severe winters

prevalent in their collection areas by entering a short daylength-induced diapause.

In addition, we discuss how diapause and temperature-related developmental re-

strictions can affect the potential for successful establishment and the population

dynamics of these parasitoids under California conditions.

MATERIAL AND METHODS

Parasitoid and Host Cultures—Aphelinus albipodus and D. rapae were col-

lected, respectively, near Tahcheng (46°42’ N, 83°00’ E, ca. 500 m elevation) and

30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Wugia (Ulugqat) (39°05’ N, 74°02’ E, ca. 3000 m elevation), Peoples Republic

of China, by D. Gonzalez in 1992. The procedures for rearing Russian wheat

aphid and both species of parasitoids were described earlier (Bernal & Gonzalez

1993).

Experiments.—Two sets of trials were conducted for each of A. albipodus and

D. rapae. The first set was designed to provide baseline information concerming

emergence rates from mummies, and the maximum egg to adult developmental

times in each of the parasitoids. The second set was designed to detect diapause

if it occurred in either parasitoid species. The methodology for both sets of trials,

and for both parasitoids, was similar, except that the daylength used in the second

set of trials was shorter than in the first set of trials.

Previous studies showed that emergence rates from mummies at 21° C, 50—

70% R. H., and 14 h daylengths were high (= 90%) in both A. albipodus and D.

rapae. Thus, these environmental conditions (hereafter “‘non-diapause condi-

tions’’) were used to obtain baseline information concerning emergence rates and

developmental times in the first set of trials. Small pots of wheat, cv MIT, each

covered with a small cage and containing ca. 40 Russian wheat aphids of instars

If-adult (Bernal & Gonzalez 1997) were exposed to 20 previously mated and

fed, 1—2 day old female A. albipodus or D. rapae. The aphids were exposed to

the female parasitoids for 24 h, after which the parasitoids were removed and the

aphids were incubated under the non-diapause conditions described above. When

aphid mummies formed, they were transferred each to a gelatin capsule (size #0)

and further incubated under non-diapause conditions to allow emergence of adult

parasitoids. The mummies were monitored daily for emergence of adult parasit-

oids, and the number of days from oviposition to emergence of individual adult

parasitoids, the number of mummies formed, and the number of mummies yield-

ing adult parasitoids from each pot were recorded. Fifteen pots with Russian wheat

aphid were prepared for each of A. albipodus and D. rapae.

The second set of trials was similar to the first, except that the daylength was

12 h (hereafter “‘diapause conditions’’) rather than 14 h. In this case, 20 pots with

Russian wheat aphid were used for A. albipodus and 15 pots for D. rapae. The

results of the first set of trials showed that under non-diapause conditions, max-

imum egg to adult developmental times were 28 days in A. albipodus (n = 344),

and 23 days in D. rapae (n = 291). Thus, for the second set of trials, mummies

of each species not yielding adults within their corresponding maximum egg to

adult times were tentatively considered to be in diapause. Emergence rates from

mummies under diapause versus non-diapause conditions were compared via log-

likelihood ratios for 2 X 2 contingency tables, with Yate’s correction for conti-

nuity, for each of A. albipodus and D. rapae (Zar 1996).

Mummies under diapause conditions yielded fewer adult parasitoids than mum-

mies under non-diapause conditions in A. albipodus but not in D. rapae (see

Results). Whether the A. albipodus mummies that did not yield adult parasitoids

under diapause conditions indeed contained diapausing individuals was deter-

mined via dissections. These mummies (n = 242) were divided into 2 groups of

equal size at 28 days; one group was maintained under diapause conditions (i.e.,

12 h daylength), the other was transferred to non-diapause conditions (i.e., 14 h

daylength). Subsamples of mummies from each group were dissected at 48, 55,

and 80 day from oviposition. Mummies containing live last instar parasitoid larvae

2001 BERNAL ET AL.: PARASITOID OVERWINTERING 31

Or pupae were considered to be in diapause. Both groups of mummies were further

incubated if diapausing individuals were detected in the subsamples. The days to

emergence, and the numbers of any parasitoids emerging from both groups of

mummies were recorded.

Finally, the mean monthly temperatures and daylengths of each of the parasit-

oid’s collection sites were compared versus a representative site from northern

California, Tulelake (41°58' N, 121°28' W), where A. albipodus and D. rapae

were colonized. Data for these comparisons were obtained from Walter & Lieth

(1967), Pearce & Smith (1990), and Anonymous (1993).

RESULTS

The emergence rate of adult A. albipodus from mummies was significantly

lower (P < 0.001) under the 12 h daylength relative to the 14 h daylength (Fig.

1). Less than % the proportion of A. albipodus adults emerging from mummies

under the 14 h daylength emerged under the 12 h daylength. In contrast, emer-

gence rates of D. rapae adults were similarly high and not significantly different

(P = 0.439) under both daylengths (Fig. 1).

None of 121 mummies that were maintained at 12 h daylength for up to 84

days from the egg stage yielded A. albipodus adults. Dissection of these mummies

revealed that they contained live last-instar larvae throughout this sampling pe-

riod; 98% contained live last-instar larvae at 48 days from the egg stage (n = 65

mummies), 89% at 55 days (n = 28 mummies), and 93% at 80 days (n = 28

mummies). In contrast, ca. 12% of 121 mummies transferred from 12 h daylength

to 14 h daylength at 28 days from the egg stage yielded A. albipodus adults

within 47—75 days.

Dissection of five of the mummies transferred to 14 h daylength at each of 48,

55, and 80 days from the egg stage indicated that the remaining mummies con-

tained living last-instar larvae (15/15 total mummies dissected contained live last-

instar larvae). Although additional adult emergence from the mummies transferred

to 14 h daylength was plausible given the prevalence of live last instar larvae

inside these mummies after 80 days, these were accidentally destroyed at 84 days

when a mechanical failure caused a sudden increase in the ambient temperature

to >40° C for ca. 20 h.

DISCUSSION

Our results showed that A. albipodus entered diapause when it developed under

a 12 h daylength. The incidence of diapause in A. albipodus at this daylength,

discounting for mummy stage mortality evident under the 14 h daylength (ca.

8%), was ca. 68%, and it occurred in the last larval instar. In addition, our results

showed that diapause could be broken in some A. albipodus individuals by ex-

posing diapausing mummies to a 14 h daylength. In contrast to A. albipodus, D.

rapae did not enter diapause when it developed under a 12 h daylength.

Our findings concerning A. albipodus were not unexpected, whereas those con-

cerning D. rapae were initially puzzling given the climatic conditions prevalent

in their areas of origin (Fig. 2). Mean monthly temperatures at Tahcheng, A.

albipodus’ collection site, vary from —16° C in January to 22° C in July, and

daylengths vary from 8.5 h in December to 15.9 h in June. Temperature conditions

in Wuqia, D. rapae’s collection site, are less extreme than in Tahcheng, ranging

32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

—_

=)

i=)

P << 0.001

Proportion Emerged (+ S.E.)

o

i)

ol

=

o

o

A. albipodus D. rapae

Parasitoid

Temperature (°C)

= ,

om ou

-15 ahcheng, PRC

8 9 10 11 12 13 14 15 16 17

Daylength (hours)

Figure 1. Emergence rates of Aphelinus albipodus and Diaeretiella rapae under two daylengths

(21° C, 50-70% R.H.); hollow columns = 14 h daylength, stippled columns = 12 h daylength.

Numbers above column pairs indicate significance of difference in emergence rates between 14 h and

12 h daylengths: A. albipodus, G = 306.470, 1 df, n = 685; D. rapae, G = 0.666, 1 df, n = 719.

Figure 2. Daylength (on the 15th of each month) and mean monthly temperature variation in:

Tahcheng, People’s Republic of China, A phelinus albipodus’ collection site; Wugia, PRC, Diaeretiella

rapae’s collection site, and; Tulelake, California, a representative northern California colonization site

for A. albipodus and D. rapae.

2001 BERNAL ET AL.: PARASITOID OVERWINTERING 33

from —11° C in January to 13.5° C in July, while daylengths are similar, 9.4 h in

December to 14.9 h in June. Despite differences in temperature, winter conditions

in both Tahcheng and Wuqia would require a mechanism by which A. albipodus

and D. rapae could survive during 5 months of subfreezing mean monthly tem-

peratures.

Diapause provides a mechanism for winter survival in A. albipodus. Mean

monthly temperatures and average daylengths in Tahcheng closely match tem-

perature- and daylength-related developmental restrictions in A. albipodus that are

evident from this and other studies (Bernal & Gonzdélez 1996; Bernal et al. 1997).

Those studies showed that A. albipodus: (1) enters diapause at 12 h daylengths,

and; (4i) cannot develop to adulthood if temperatures are 10° C or below. Aphel-

inus albipodus mortality at 10° C is >80% during egg to mummy development,

and adults do not emerge from mummies that develop at this temperature (Bernal

& Gonzalez 1996; Bernal et al. 1997). Mean temperatures in Tahcheng decrease

to 10° C and below in the fall, and are associated with 12 h and shorter daylengths.

Mean temperatures then increase to 10° C and above in the spring and are asso-

ciated with 14 h and longer daylengths. Thus, A. albipodus may avoid fatally low

temperatures by entering diapause when daylengths decrease to 12 h in the fall,

then resuming development when daylengths increase to 14 h in the spring.

In contrast to A. albipodus, diapause was not detected in D. rapae at 12 h

daylength. Based on our results, however, we cannot discount the possibility of

diapause in this parasitoid. Diaeretiella rapae populations in the Netherlands enter

diapause during the winter (Hafez 1961). Moreover, diapause in these D. rapae

populations peaks in late October (Hafez 1961) when daylengths are between

10.7 and 8.8 h. Thus, diapause may be induced at daylengths shorter than 12 h

in the Wuqia D. rapae population. Average daylengths in Wugqia decrease to 12

h in the early fall and are associated with mean temperatures of ca. 7° C. Later

in the fall, mean temperatures decrease to subfreezing levels, but daylengths by

then decrease to 10.5 h and shorter. Previous studies show that developmental

mortality in D. rapae is low at 10° C, and that its lower developmental threshold

is in the range 2.5—3.9° C (Bernal & Gonzalez 1995, 1997). Thus, diapause at 12

h daylength in the Wuqia D. rapae population may not be necessary given the

associated mean field temperatures (ca. 7 C) in the area. Hence, based on the

results of previous studies (Hafez 1961; Bernal & Gonzdlez 1995, 1997) and on

the lengthy and severe winters prevalent in Wugia, a likely scenario is that dia-

pause in D. rapae is induced by daylengths shorter than 12 h, which are associated

with near- or sub-freezing temperatures.

Our results concerning A. albipodus are consistent with previous reports of

diapause in closely related Aphelinus spp. Yu (1992) found that >50% and >95%

of A. nr. varipes (= A. varipes, see Bernal et al. 1997; J. B. Woolley, personal

communication) collected in southern Alberta (Canada) entered diapause when

they developed under 14 h and 12 h daylengths, respectively. The corresponding

rates for A. varipes collected in Kazakhstan (= A. albipodus, see Bernal et al.

1997; J. B. Woolley, personnel communication) were 0% and >90%. Diapause

was induced by short daylengths both in Yu’s and this study. Short daylengths in

these cases are associated with low winter temperatures that may be particularly

detrimental to Aphelinus species. Developmental thresholds and high mortality at

temperatures between 5 and 10° C are common in Aphelinus spp. (e.g., Force &

34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Messenger 1964; Walker et al. 1988; Trimble et al. 1990; Asante & Danthanarayan

1992; Lajeunesse & Johnson 1992; Yu 1992; Bernal & Gonzdélez 1993, 1996;

Bernal et al. 1997; Lee & Elliot 1998a, b). Thus, Aphelinus spp. appear to be

highly susceptible to low temperatures and to employ diapause as an overwinter-

ing strategy (Trimble et al. 1990; Yu 1992; this study).

Both A. albipodus and D. rapae were imported to Califomia for release against

Russian wheat aphid (Gonzalez et al., unpublished data). Both were extensively

released in northern California and A. albipodus rapidly became established. It is

unclear at present whether D. rapae became established because this species al-

ready occurred in the area. In either case, it is evident that the climate in northern

California is milder during the winter relative to the collection areas of both

parasitoids, and daylength variation during the year is similar between these areas,

or less in the case of D. rapae (Fig. 2). Thus, although diapause may be induced

by short winter daylengths in northern Califomia, it may not be required for winter

survival of A. albipodus and D. rapae. Summer, rather than winter, temperatures

may represent an obstacle for establishment of D. rapae, in particular, in northern

California. Earlier studies showed that D. rapae is more susceptible to high tem-

peratures than A. albipodus (Bernal & Gonzdlez 1995, 1996). This is suggested

also by the seasonal variation in mean temperatures in each of these parasitoid’s

collection sites. Thus, winter diapause and summer survival are two important

factors that may affect the population dynamics of A. albipodus and D. rapae

and their impact on Russian wheat aphid populations in California. Moreover, the

timing of emergence of diapausing parasitoids in the spring will likely influence

their impact on Russian wheat aphid populations. Previously it was suggested that

in annual agroecosystems, early-appearing natural enemies are likely to have a

greater impact on pest populations than those emerging later in the season (Bernal

& Gonzalez 1993; Gilstrap 1997). Our results, and previous studies (Bernal &

Gonzalez 1995, 1996), suggest that A. albipodus is likely to emerge later in the

season than D. rapae.

Differences in climate between collection and colonization areas of exotic nat-

ural areas are reported to contribute substantially to reducing the success rate of

classical biological control (Stiling 1993). Specifically, they hamper biological

control efforts early on in their development by precluding the successful estab-

lishment of exotic natural enemies. A practice that could contribute substantially

to reducing the number of biological control efforts that fail during the coloni-

zation phase is to emphasize closer climate-matching during the planning phase

of foreign exploration efforts. An approach similar to that taken to forecast po-

tential range expansions of pest species (Meats 1989; Hughes & Maywald 1990)

would be desirable, but may seldom be possible with exotic natural enemies.

Unlike pest species, usually little or no information is available concerning an

exotic natural enemy’s response to temperature and other biological parameters

necessary for such an approach. Many times exotic natural enemies are undes-

cribed species and biological information can only be inferred from related spe-

cies; other times, the available information is limited to a taxonomic description.

In both cases, pertinent biological information becomes available only after a

series of studies are conducted. We believe that the analyses presented here, in-

cluding the results from previous studies (Bernal & Gonzalez 1995, 1996, 1997;

Bernal et al. 1997), provide an initial model for assessing the potential for suc-

2001 BERNAL ET AL.: PARASITOID OVERWINTERING 35

cessful establishment of exotic natural enemies early in the development of a

biological control campaign, and a means for improving the success rate of clas-

sical biological control. Ultimately, the success of a classical biological control

campaign is contingent upon the successful establishment of exotic natural ene-

mies.

ACKNOWLEDGMENT

We are grateful to Dr. Bob Wharton and Janet Griset (both at Texas A&M

University) for critically reviewing the manuscript and for suggestions for its

improvement. Marcella Waggoner and her team provided the aphids and parasit-

oids used in the experiments. Our Chinese collaborators, G. Zhang, J. Zhang, W.

Zhang, Y. Min, R. Wang, Du Bing Ren, Wei Zhengming, Geng Shouguang, J.

Wen, X. Liu, and Y. Zhong are gratefully acknowledged for their skilled assistance

during foreign exploration in China.

LITERATURE CITED

Anonymous. 1993. Daily average temperatures and precipitation in California. Statewide IPM Project,

University of California, Davis.

Asante, S. K. & W. Danthanarayana. 1992. Development of Aphelinus mali an endoparasitoid of wooly

apple aphid Eviosoma lanigerum at different temperatures. Entomol. Exp. Appl., 65: 31-37.

Beddington, J. R., C. A. Free & J. H. Lawton. 1978. Characteristics of successful natural enemies in

models of biological control of insect pests. Nature, 273: 513-519.

Bernal, J. S. & D. Gonzalez. 1993. Temperature requirements of four parasites of the Russian wheat

aphid Diuraphis noxia. Entomol. Exp. Appl., 69: 173-182.

Bernal, J. S. & D. Gonzalez. 1995. Thermal requirements of Diaeretiella rapae (M’Intosh) on Russian

wheat aphid (Diuraphis noxia Mordwilko, Hom., Aphididae) hosts. J. Appl. Ent., 119: 273-

277.

Bernal, J. S. & D. Gonzalez. 1996. Thermal requirements of Aphelinus albipodus (Hayat and Fatima)

(Hym., Aphelinidae) on Diuraphis noxia (Mordwilko) (Hom., Aphididae) hosts. J. Appl. Ent.,

120: 631-638.

Bernal, J. S. & D. Gonzalez. 1997. Reproduction of Diaeretiella rapae on Russian wheat aphid hosts

at different temperatures. Entomol. Exp. Appl., 82: 159-166.

Bernal, J. S, M. Waggoner & D. Gonzalez. 1997. Reproduction of Aphelinus albipodus (Hyme-

noptera: Aphelinidae) on Russian wheat aphid (Hemiptera: Aphididae) hosts. Eur. J. Entomol.,

94: 83-96.

DeBach, P. & D. Rosen. 1991. Biological control by natural enemies (2nd ed.). Cambridge University

Press, Cambridge.

Flanders, S. E. 1940. Environmental resistance to the establishment of parasitic Hymenoptera. Ann.

Entomol. Soc. Amer., 33: 245—253.

Force, D. C. & P. S. Messenger. 1964. Duration of development, generation time, and longevity of

three hymenopterous parasites of Therioaphis maculata reared at various constant temperatures.

Ann. Entomol. Soc. Amer., 57: 405—413.

Gilstrap, E E. 1997. Importation biological control in ephemeral crop habitats. Biol. Control, 10:

23-29.

Gilstrap, EF E., D. Gonzalez & L. McKinnon. 1994. Biological control of Russian wheat aphid: a

summary of 1998-1993. pp. 53-61. In Peairs, E B., M. K. Kroening, & C. L. Simmon (comps.).

Proceedings of the sixth Russian wheat aphid workshop. Fort Collins, January 23-25, 1994.

Great Plains Agricultural Council, Fort Collins.

Gonzalez, D. & E E. Gilstrap. 1992. Foreign exploration: assessing and prioritizing natural enemies

and consequences of preintroduction studies. pp. 53-70. Jn Kauffman, W. C. & J. E. Nechols

(eds.). Selection criteria and ecological consequences of importing natural enemies. Entomol.

Soc. Amer., Lanham.

Greathead, D. J. & A. H. Greathead. 1992. Biological control of insect pests by insect parasitoids and

predators: the BIOCAT database. Biocontrol News Info., 13: 61N—68N.

36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Hafez, M. 1961. Seasonal fluctuations of population density of the cabbage aphid, Brevicoryne bras-

sicae (L.) in The Netherlands, and the role of its parasite, Aphidius (Diaeretiella) rapae (Curtis).

T. Pl.-ziekten, 67: 445-548.

Huffaker, C. B., EF J. Simmonds & J. E. Laing. 1974. The theoretical and empirical basis of biological

control. pp. 41-78. Jn Huffaker, C. B. & P. S. Messenger (eds.). Theory and practice of bio-

logical control. Academic Press, New York.

Huffaker, C. B., R. EK Luck & P. S. Messenger. 1977. The ecological basis of biological control. pp.

560-586. In White, D. (ed.). Proceedings, XV International Congress of Entomology. Wash-

ington, D. C., 19-27 August. Entomol. Soc. Amer., College Park.

Hughes, R. D. & G. EK Maywald. 1990. Forecasting the favourableness of the Australian environment

for the Russian wheat aphid, Diuraphis noxia (Homoptera: Aphididae), and its potential impact

on Australian wheat yields. Bull. Entomol. Res., 80: 165-175.

Lajeunesse, S. E. & G. D. Johnson. 1992. Developmental time and host selection by the aphid para-

sitoid Aphelinus sp. nr. varipes (Foerster) (Hymenoptera: Aphelinidae). Can. Ent., 124: 565-

575.

Lee, J. H. & N. C. Elliot. 1998a. Comparison of developmental responses to temperature in Aphelinus

asychis (Walker) from two different geographic regions. Southwest. Entomol., 23: 77-82.

Lee, J. H. & N. C. Elliot. 1998b. Temperature effects on development in Aphelinus albipodus (Hy-

menoptera: Aphelinidae) from two geographic regions. Great Lakes Entomol., 31: 173-179.

Luck, R. F 1990. Evaluation of natural enemies for biological control: a behavioral approach. Trends

Ecol. Evol., 5: 196-199.

Meats, A. 1989. Bioclimatic potential. pp. 241-252. In Robinson A. S. & G. Hooper (eds.). Fruit flies:

their biology, natural enemies and control. Volume 3B. Elsevier, Amsterdam.

Messenger, P. S. 1959. Bioclimatic studies with insects. Ann. Rev. Entomol., 4: 183-206.

Messenger, P. S. 1971. Climatic limitations to biological control. Proc. Tall Timbers Conf. Ecol. Animal

Control Habitat Mgmt., 3: 97-114.

Messenger, P. S. & N. E. Flitters. 1954. Bioclimatic studies of three species of fruit flies in Hawaii.

J. Econ. Entomol., 47: 756-765.

Murdoch, W. W. & C. J. Briggs. 1996. Theory for biological control: recent developments. Ecology,

77: 2001-2013.

Murdoch, W. W., J. Chesson & P. L. Chesson. 1985. Biological control in theory and practice. Am.

Nat., 125: 344-366.

Pearce, E. A. & G. Smith. 1990. The Times books world weather guide. Times Books-Random House,

New York.

Stiling, P. 1993. Why do natural enemies fail in classical biological control programs? Amer. Entomol.,

39: 31-37.

Trimble, R. M., L. R. H. Blommers & H. H. M. Helsen. 1990. Diapause termination and thermal

requirements for postdiapause development in Aphelinus mali at constant and fluctuating tem-

peratures. Entomol. Exp. Appl., 56: 61-69.

Walker, J. T. S., S. C. Hoyt, D. P. Carroll & G. V. Tangren. 1988. Influence of constant and alternating

temperatures on wooly apple aphid (Homoptera: Eriosomatidae) and its parasitoid Aphelinus

mali (Haldeman) (Hymenoptera: Aphelinidae). Melanderia, 46: 36-42.

Walter, H. & H. Lieth. 1967. Klimadiagram Weltatlas. VEB Gustav Fischer Verlag, Stuttgart.

Yu, D. S. 1992. Effects of photoperiod and temperature on diapause of two Aphelinus spp. (Hyme-

noptera: Aphelinidae) parasitizing the Russian wheat aphid. Can. Ent., 124: 853-860.

Zar, J. H. 1996. Biostatistical analysis. Prentice Hall, Englewood Cliffs.

Received 8 Feb 2000; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 37-38, (2001)

A NEW SPECIES OF CORTICARINA FROM ARIZONA

(LATRIDITDAE: CORTICARIIND)

FRED G. ANDREWS

California Department of Food and Agriculture, 3294 Meadowview Road,

Sacramento, California 95832-1448

Abstract.—Corticarina arizonensis is described as new. It is compared to other North American

species.

Key Words.—Insecta, Coleoptera, Latridiidae, Corticarina, Arizona.

In 1994 two specimens of Corticarina were swept from an oak tree at the base

of a talus slope at approximately 9200 feet in the Barfoot campground in the

Chiricahua mountains of Southeastern Arizona. Dissection of the lone male dem-

onstrated a new species based on the unknown aedeagal shape. Recent collecting

(1998) in the same spot yielded an additional twenty-one specimens. The same

aedeagal shape was found when males from this collection were dissected. All

specimens were collected within 100 feet of the original collection, but this time

all were swept from conifers. Additional specimens were collected on Mt. Lem-

mon, Arizona at an elevation of 7500 feet from the flower heads of dead cow

parsnip (Heracleum lanatum Michaux) in the company of numerous Cortinicara

gibbosa Gyllenhal. Numerous collections of Corticarina cavicollis Mannerheim

and Corticarina fuscula Gyllenhal have been made in the Chiricahua Mountains

and Southeastern Arizona at lower elevations without evidence of this new spe-

cies. It is likely that this species is restricted to the higher mountain elevations.

Corticarina arizonensis can be differentiated from all other species of Corticarina

by the shape of the male aedeagus (Figs. 1 and 2). Corticarina arizonensis is the

only North American species in which both the dorsal and ventral lobes of the

aedeagus are sharply pointed. The aedeagus of the other North American Corti-

carina are illustrated by Andrews (1985, 1992).

CORTICARINA ARIZONENSIS Andrews, NEW SPECIES

Description.—Length 1.33—-1.59 mm. Width 0.63—0.74 mm. Pronotal length

0.30—0.37 mm. Pronotal width 0.37—-0.44 mm. n = 43. Color dark brown to dark

reddish brown; legs, antennal segments 2—6 lighter brown; surface shiny. Anten-

nae 0.45—0.51 mm long, segment 8 longer than wide, segment 10 wider than long.

Pronotum 1.11 to 1.47 times as wide as long; lateral margin slightly sigmoid in

anterior one-half, widest in anterior one-half; postmedian depression weakly de-

veloped; lateral depression narrow, weakly developed, limited to posterior one-

half; dorsally evenly rounded from side to side; lateral margins smoothly arcuate,

minutely serrate. Pronotal surface distinctly punctured, punctures generally sep-

arated by approximately a puncture width. Elytra narrowly inflated, 1.13 to 1.75

times as long as wide. Elytral striae well defined; setae arranged in straight lines,

each raised medially and generally terminating in next puncture; humeral callus

distinct. Fully winged. Eyes fully developed, 77 facets in single eye examined.

Male: protibia toothed on inner surface, tooth 45/100 from apex. Male genitalia

as in Figs. 1 and 2.

38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Figures 1-2. Corticarina arizonensis aedeagus. Figure 1. Ventral view. Figure 2. Lateral view.

Types.—HOLOTYPE (Male): ARIZONA. Pima Co: Santa Catalina Mts., Mt.

Lemon, Summerhaven, [X-17-1998, 32°26.26' N, 110°45.52' W, F Andrews & T.

Eichlin. Deposited in the collection of the Califomia Academy of Sciences. PAR-

ATYPES (23). Same data as Holotype (11); ARIZONA. Cochise Co. Barfoot

Park, VIII-2-1994, EF Andrews & T. Eichlin (2); 10.6 mi. NW Southwest Research

Station, Barfoot Cpg., [X-14-1998, 31°55.11' N, 109°16.47' W, E Andrews & T.

Eichlin (10).

LITERATURE CITED

Andrews, E G. 1985. Corticarina scissa (LeConte), a valid species (Coleoptera: Lathridiidae). Coleop.

Bull. 39: 147-149.

Andrews, E G. 1992. Two new species of Corticarina from coastal California (Coleoptera: Lathridi-

idae: Corticariini) with notes on J. L. LeConte types. the Coleop. Bull., 46: 274-280.

Received 3 Feb 2000; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 39-44, (2001)

MELANOTRICHUS BOYDI, A NEW SPECIES OF PLANT

BUG (HETEROPTERA: MIRIDAE: ORTHOTYLIND

RESTRICTED TO THE NICKEL HYPERACCUMULATOR

STREPTANTHUS POLYGALOIDES (BRASSICACEAE)

MICHAEL D. SCHWARTZ! AND MICHAEL A. WALL?

'% Biological Resources Program, ECORC, Agriculture and Agri-Food Canada,

Ottawa, Ontario K1A OC6 Canada

Department of Botany and Microbiology, Auburn University,

101 Rouse Life Sciences Building, Auburn, Alabama 36849-5407*

Abstract——Melanotrichus boydi, NEW SPECIES is described from the western foothills of the

Sierra Nevada Mountains in California. The nickel hyperaccumulating species, Streptanthus po-

lygaloides (Gray), Brassicaceae, an endemic to serpentine soils, is its only known host plant.

Key Words.—Insecta, Miridae, plant bugs, Melanotrichus, new species, nickel hyperaccumula-

tors, serpentine soil.

As a consequence of its size, exceedingly diverse topography, climate, and

geologic history, California supports a remarkably large and interesting flora

(Munz & Keck 1973). Despite the pioneering work of E. P. Van Duzee and recent

students of North American Muiridae, the plant bug fauna of California is still

poorly known. Current studies by the junior author in serpentine soil exposures

of California have revealed a new species of plant bug restricted to a nickel

hyperaccumulator species of crucifer. In this paper we describe Melanotrichus

boydi Schwartz & Wall, NEW SPECIES, provide illustrations of the male geni-

talia, photomicrographs of the head, pretarsus, ostiolar peritreme, scalelike setae,

a distribution map and discuss the first known example of bioaccumulation for a

native herbivore on a metal hyperaccumulator. All measurements are based on

ten specimens with the mean and range presented.

MELANOTRICHUS BOYDI Schwartz & Wall, New Species

(Figs. 1-14)

Types.—Holotype, male: U.S.A. CALIFORNIA. ELDORADO Co.: NE of Col-

oma, [900 m], 22 Jun 1998, M. A. Wall, ex Streptanthus poly[galoides|., M. Wall

98-105; deposited: California Academy of Sciences, San Francisco (CAS). Par-

atypes: 1d, 12 same data as holotype except no host and M. Wall 1; MARIPOSA

Co.: 16, 12 NW of Coulterville, [700 m], 23 Jun 1998, M. Wall (2); 36, 59

Lake McClure at hwy 49 [bridge], [900 m], 23 Jun 1998, M. Wall (9); 13, 52

NW of Mariposa, 29 Jun 1998, M. Wall (7); PLACER Co.: 16, 3¢ S of Sugar

Pine Res[ervoir], [1270 m] 28 Jun 1998, M. A. Wall; TUVOLUMNE Co.: 436,42

S of Mocassin on hwy 49, [700 m], 11 Jun 1999, M. A. Wall; 4¢, 92 SW of

Chinese Camp, Red Hills Rec Area, [470 m], 28 May 1999, M. A. Wall; 12

Chinese Camp, [470 m], 10 Jun 1997, M. A. Wall; 13¢ S of Chinese Camp in

Red Hills Rec Area, [470 m], 9-13 Jun 1996, M. A. Wall. Paratypes deposited

* Currently Department of Ecology and Evolutionary Biology, University of Connecticut, Torrey

Life Science Building, 75 North Eagleville Road, U-43, Storrs, Connecticut 06269-3043.

40 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Figures 1-4. Scanning electron micrographs of Melanotrichus boydi. Figure 1. Head, lateral view.

Figure 2. Ostiolar peritreme, lateral view. Figure 3. Scalelike setae on corium. Figure 4. Pretarsus,

apical view.

at the American Museum of Natural History, New York, CAS, the Canadian

National Collection of Insects, Ottawa and the National Museum of Natural His-

tory, Smithsonian Institution, Washington, D.C.

Description of Males.—Orthotylinae: Orthotylini. Length from clypeus to apex of membrane 5.74

(5.20—6.15) mm, maximum width across hemelytra 1.88 (1.60—2.10) mm. Head: width 0.93 (0.88—

0.99) mm, vertex width 0.39 (0.36—0.41) mm. Labium: length 1.14 (1.08—-1.23) mm, reaching apex of

mesosternum, sometimes just extending to slightly beyond middle of mesosternum, but labrum usually

reaching apex of mesosternum. Antenna: segment 1, length, 0.55 (0.50—0.60) mm; 2, 1.86 (1.73-2.15)

mm; 3, 1.60 (1.45—1.83) mm; 4, 0.47 (0.45—0.49) mm. Pronotum: length 0.70 (0.63—0.79) mm, basal

width 1.34 (1.23—-1.45) mm.

Generally green with greenish-yellow to orange-yellow on embolium, cuneus, mesoscutum, some-

times scutellum, pronotum on calli and anterior of calli, head, antennal segment 1, extreme base of

antennal segment 2, legs, and venter; black on antennal segments 2—4, apex of labial segment 4,

tarsus, pretarsus; membrane of hemelytra infuscate with greenish-yellow veins; dorsum with inter-

2001 SCHWARTZ & WALL: NEW PLANT BUG SPECIES 41

Figures 5-13. Male genitalia of Melanotrichus boydi. Figures 5—7. Left paramere. Figure 5. Apex

of shaft, apical view. Figure 6. Sensory lobe, apical view. Figure 7. Sensory lobe, lateral view. Figures

8 and 9. Right paramere. Figure 8. Dorsal view. Figure 9. Lateral view. Figure 10. Phallotheca, dorsal

view. Figure 11. Detail of ductus seminis & secondary gonopore, lateral view. Figure 12. Detail of

spiculum, lateral view. Figure 13. Ventral processes of genital segment, apical view.

mixed vestiture of moderately distributed, shining white, reclining simple setae and head, anterior of

pronotum and hemelytra with scattered shining, silvery, sericeous scalelike, setae (Fig. 3).

Head broadly subtriangular; clypeus moderately produced and slightly rounded in dorsal aspect,

anteriorly flattened with slightly rounded ventral margin in lateral aspect (Fig. 1); posterior margin of

head wider than, and slightly removed from, anterior margin of pronotum; basal carina broad, gently

rounded; eyes large, height greater than head height in lateral aspect; posterior margin gently curved

anteriorly in dorsal aspect; vertex width equal to 1.5 width of eye. Antennal segment 1 with mod-

erately distributed, reclining, black simple setae and two subapical black bristles; diameter about 1.5

as wide as segment 2; segment 3 & 4 slightly thinner than segment 2. Pronotum trapeziform, calli

slightly swollen. Mesoscutum moderately broadly exposed. Hemelytra subparallel-sided, widest at

apex of embolium; length of cuneus about 3X width. Venter unmarked; ostiolar peritreme (Fig. 2).

Legs long; tibia with dark brown to black bristles, unmarked at base; claw (Fig. 4) gently curved with

minute pulvillus and apically converging parempodium.

Genitalia: Left paramere (Figs. 5-7) broad, C-shaped, sensory lobe with truncate, broadly serrate

apex; shaft broadly curved, gradually attenuate. Right paramere (Figs. 8 and 9) L-shaped, broad

basally, strongly attenuate distally. Phallotheca (Fig. 10) strongly sclerotized, convoluted, aperture on

right aspect. Vesica with narrow elongate ductus seminis (Fig. 11) and one basally thickened, distally

attenuate spiculum (Fig. 12); apex of spiculum subequal to apex of secondary gonopore (Figs. 11 and

12). Ventral processes of genital segment (Fig. 13); right process larger than left process, strongly

sclerotized with flattened dorsal flange, which protrudes posteriorly beyond margin of genital aperture.

Description of Females.—Similar to males except, eye smaller, head and vertex wider, and heme-

lytral margin more rounded, length from clypeus to apex of membrane 5.86 (5.20—6.50) mm, maxi-

mum width across hemelytra 2.01 (1.80—2.29) mm. Head: width 1.01 (0.98-1.15) mm, vertex width

0.49 (0.45-0.53) mm. Labium: length 1.26 (1.20—1.38) mm, reaching apex of mesosternum, sometimes

just extending to slightly beyond middle of mesosternum, but labrum usually reaching apex of me-

42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Nevada

El Dorado

Madera

Figure 14. Distribution of Melanotrichus boydi. Areas filled in black represent distribution of

serpentine soil within the foothills of the Sierra Nevada. M. boydi was either observed or collected at

all numbered sites. Site numbers correspond with information presented in Table 1 (modified from

Wall 1999).

sosternum. Antenna: segment 1, length, 0.58 (0.53—0.64) mm; 2, 1.85 (1.66—2.06) mm; 3, 1.49 (1.33-

1.58) mm; 4, 0.46 (0.45—0.48) mm. Pronotum: length 0.81 (0.71-0.91) mm, basal width 1.58 (1.35-—

1.75) mm.

Diagnosis.—Melanotrichus boydi NEW SPECIES is most similar to M. mistus

(Knight) and M. stanleyaea (Knight) in size (almost all other species of Melan-

otrichus are less than 5 mm in length) but is distinguished by the white simple

setae on the dorsum, the black antennal segments 2—4, and the structure of male

genitalia, especially the narrow apex of the right paramere and pointed vesical

spiculum. Melanotrichus mistus has conspicuous black setae on the dorsum, yel-

low to orange antenna, and male genitalia with the apices of the right paramere

and vesical spiculum blunt. Melanotrichus stanleyaea has similar pale vestiture

to M. boydi but the antenna is pale, and the apex of the right paramere is longer

and more acutely attenuated.

Taxonomy.—Schuh (1995) and Kerzhner and Josifov (1999) treat Melanotri-

chus Reuter as a subgenus of Orthotylus Fieber. However, given the absence of

a phylogenetic analysis of the genus we choose to follow North American authors

and describe our new species in the genus Melanotrichus. In Henry (1991: 455)

M. boydi will key to M. concolor (Kirschbaum). In Knight (1968) this species

will key to M. stanleyaea Knight, under Dichaetocoris Knight (cf., Knight 1968:

2001 SCHWARTZ & WALL: NEW PLANT BUG SPECIES 43

Table 1. Concentration of Ni found in sample of M. boydi collected at six sites throughout Cali-

fornia (see Fig. 14). Concentrations represent values obtained from samples consisting of 15—20 in-

dividuals of M. boydi from each site analyzed via atomic absorption spectrophotometry.

Site Ni content in ppm on

location dry weight basis

Site 1 735

Site 3 ail

Site 4 789

Site 6 777

Site 7 718

Site 10 751

114) or as M. wileyae Knight (cf. Knight 1968: 117). The black antennal segments

2—4, distribution, and host plant are sufficient to distinguish M. boydi from these

species.

Hostplants.—Streptanthus polygaloides (Gray), Brassicaceae, a winter annual,

endemic to serpentine soils from the foothills of the western slopes of the Sierra

Nevada Mountains in California is the host of both immature and adult stages.

Wall and Boyd (in press) provided evidence to indicate that M. boydi is probably

monophagic. At ten localities where this species was collected on S. polygaloides

they sampled other locally abundant plants—a conifer, three woody dicots, four

herbaceous dicots, including other species of Brassicaceae and another species of

Streptanthus, and a monocot—for the presence of M. boydi and did not collect

any.

Distribution.—Figure 14. In addition to the localities listed under Types, M.

boydi was also collected from the following sites in late-May to mid-July: BUTTE

Co.: N of Magalia, 800 m; CALAVERAS Co.: N of San Andreas, 300 m; FRESNO

Co.: NW of Pine Flat Lake, 400 m; PLACER Co.: S of Washington, 1330 m. In

the western foothills of the Sierra Nevada M. boydi is associated throughout the

range of S. polygaloides from Butte County in the north to Fresno County in the

south encompassing a wide range of elevation (330 m to 1330 m) within the

foothills woodland and the yellow pine forest plant communities.

Etymology—Named to honor Dr. R. S. Boyd, Department of Botany and Mi-

crobiology, Auburn University, who provided insight and encouragement to the

junior author during his Master’s degree, and for his leadership in the study of

hyperaccumulation ecology.

Discussion.—Melantrichus boydi is unique in several ways. Not only is it the

only species of insect reported to specialize on a Ni hyperaccumulator, but it

appears to accumulate Ni at levels one to two orders of magnitude higher than

other insects found feeding on S. polygaloides (Wall 1999). This high level of Ni

accumulation is concordant across the known range of M. boydi (Table 1). Bot-

anists have long used a qualitative colorimetric test for identifying plants in the

field that contain high levels of Ni. In this test, plant material is crushed onto

filter paper permeated with the colorless chemical, dimethylglioxime, which reacts

with the Ni in the plant and changes to a various shades of red (Reeves 1992).

This same test consistently gives a positive result when M. boydi is crushed onto

the filter paper. While perhaps only having novelty status, this colorimetric test

adds another interesting element to the taxonomist’s arsenal for identifying this

44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

species. At the very least, the presence of high levels of Ni in M. boydi is con-

sidered an autapomorphy for the species.

Including M. boydi n. sp., five North American mirids apparently specialize on

brassicaceous hosts, all are members of the genus Melanotrichus. Melanotrichus

albocostatus Knight is known from Cardaria costatus, Descurainia sophia (L.)

Webb, and Sisymbrium irio L. Webb (Henry 1991), M. leonardi Kerzhner and

Schuh is known from D. sophia (Kelton 1980), M. stanleyaea is known from

Stanleya pinnata (Pursh) Britton (Knight 1968), and M. whiteheadi Henry is

known from D. pinnata pinnata (Walt.) Britt. (Henry 1991). Interestingly, S. pin-

nata is a hyperaccumulator of Selenium (Brooks 1998). Determining whether or

not host preference for Brassicaceae has evolved in multiple lineages, or if these

five species represent a monophyletic group within Melanotrichus requires a de-

tailed phylogenetic analysis, which is beyond the scope of this present study.

ACKNOWLEDGMENT

The Pacific and Agri-Food Research Centre, Agassiz, British Columbia pro-

vided support for MDS to study type specimens of E. P. Van Duzee, housed in

the CAS. Support for this study was also provided by the Alabama Academy of

Sciences to Michael Wall and by NSERC and FRBC grants to G. G. E. Scudder,

Department of Zoology, The University of British Columbia, Vancouver, Canada.

Two anonymous reviewers made critical comments on an earlier version of the

manuscript.

LITERATURE CITED

Brooks, R. R. 1998. Geobotany and hyperaccumulators. Chapter 3. pp. 55—94. In Brooks, R. R. (ed.).

Plants that Hyperaccumulate Heavy Metals. CAB International, Wallingford.

Henry, T. J. 1991. Melanotrichus whiteheadi, a new crucifer-feeding plant bug from the southeastern

United States, with new records for the genus and Key to the species of eastern North America

(Heteroptera: Miridae: Orthotylinae). Proc. Entomol. Soc. Wash., 93: 449-456.

Kelton, L. A. 1980. Two new species of Melanotrichus Reuter from western Canada and a description

of the male of M. atriplicis (Heteroptera: Miridae). Can. Entomol. 112: 337-339.

Kerzhner, I. M. & M. Josifov. 1999. Cimicomorpha IT, Miridae. Volume 3. Jn Aukema, B & C. Rieger

(eds.). Catalogue of the Heteroptera of the Palaearctic. The Netherlands Entomological Society,

Wageningen.

Knight, H. H. 1968. Taxonomic review: Mirdae of the Nevada Test Site and the western United States.

Brigham Young Univ. Sci. Bull., Biol. Ser., 9: 1-282.

Munz, P. A. & D. D. Keck. 1973. A California flora and supplement. University of California Press,

Berkeley.

Schuh, R. T. 1995. Plant bugs of the world (Insecta: Heteroptera: Miridae). New York Entomol. Soc.,

New York.

Reeves, R. D. 1992. The hyperaccumulation of nickel by serpentine plants. pp. 253-277. In A. J. M.

Baker, J. Proctor, and R. D. Reeves (eds.). The Ecology of Ultramafic (serpentine) Soils. Inter-

cept, Andover.

Wall, M. A. 1999. Nickel accumulation in serpentine arthropods with emphasis on a species of Me-

lanotrichus (Heteroptera: Miridae). M.S. Thesis, Auburn University, Alabama.

Received 29 Mar 2000; Accepted 5 May 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 45-46, (2001)

Scientific Note

ALTITUDINAL DISTRIBUTION AND PHENOLOGY OF

THREE SPECIES OF CARRION BEETLES

(COLEOPTERA: SILPHIDAE) FROM NEVADO DE

COLIMA, JALISCO, MEXICO

The taxonomy of carrion beetles (Coleoptera: Silphidae) from México was

revised by Peck, S. B. & R. S. Anderson (1985. Quaest. Ent., 21(3): 247-317)

who recorded eleven species belonging to four genera. Other contributions, es-

pecially for Jalisco State include: Volcan de Tequila (Navarrete-Heredia, J. L.

1995. Dugesiana, 2(2): 11—26); Sierra de Manantlan (Rivera-Cervantes, L. E. &

E. Garcia-Real. 1998. Dugesiana, 5(1): 11-22), and La Primavera and Barranca

del Rio Santiago (Navarrete-Heredia, J. L. & H. E. Fierros-Lépez. 1998. Duge-

siana, 5(1): 49-50). This study was done in Jalisco by staff members from the

Center for Zoological Research, University of Guadalajara to determine carrion

beetle distribution. In this note, we describe our trapping results from the National

Park Nevado de Colima, Jalisco, México.

Field work was done on the NW slope of the National Park Nevado de Colima,

in the locality El Floripondio, Cerro Las Viboras, San Gabriel County, between

2200—3000 meters above sea level (m). We used carrion traps (model NTP-80)

designed by Moré6én, M. A. & R. Terrén [1984. Acta Zool. Mex. n.s., (3): 1-47].

Six sites were selected: Cupressus forest (2300 m), oak-pine forest (2400 m),

cloud forest (2600 m), Abies forest (2840 m), disturbed Abies forest (2920 m),

and Abies-grass association (2950 m). One carrion trap was used for each site

during one month, starting in April and ending in October 1998. Rotting squid

was used as bait.

Three species of carrion beetles were collected: Oxelytrum discicolle (Brullé),

24 2,840 om ‘ 82Q Te

2,600 IHL F ae £0 we 2

& yi pO Bek

2.900 04

Nicrophorus |

; - i

"mexicans

trophioris

olds

a sits

liscicolle

ie | Abies a distinbed

Cirpressus | oakcpine | i Wie ke,

“# tly forest —s forest «== Ae

forest forest

SS

association —

Figure 1. Altitudinal distribution of three Silphidae species from El Floripondio, Jalisco, collected

with carrion traps.

46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

250 -—

200 :

150

‘oO

iD)

oq,

NM

= 100

a

—

50

BR ROLES

0 "ANSI NN crac

@ Nicrophorus mexicanus Nicrophorus olidus

Figure 2. Abundance of Nicrophorus mexicanus and N. olidus from El Floripondio, Jalisco, col-

lected with carrion traps between April and October, 1998.

Nicrophorus mexicanus (Matthews) and Nicrophorus olidus (Matthews). These

species were represented by 743 specimens. Nicrophorus mexicanus was the com-

monest species (628 specimens), and was collected monthly and distributed be-

tween 2300—2950 m. Nicrophorus olidus was the second most common species,

represented by 107 specimens, found between 2300—2600 m, but collected only

during May—August (Fig. 1). Finally, only eight specimens of O. discicolle were

collected in the Cupressus forest in June. Other Mexican localities demonstrating

similar silphid abundance and distribution include: Volcan de Tequila, Jalisco

(Navarrete-Heredia, 1995); Sierra de Manantlan (Rivera-Cervantes & Garcia-

Real, 1998) and Cofre de Perote, Veracruz (Arellano, L. 1998. Dugesiana, 5(2):

1-16). In all of them, Nicrophorus species are well represented, mostly at higher

elevations, whereas O. discicolle is less abundant and restricted to lower places.

Acknowledgment.—Dr. Rodolfo Novelo for review of this note, and Dr. Mar-

garet K. Thayer helpful comments.

Hugo E. Fierros-Lé6pez and José Luis Navarrete-Heredia, Entomologia, Centro

de Estudios en Zoologia, Universidad de Guadalajara, Apdo. Postal 234, 45100

Zapopan, Jalisco, México.

Received Nov 30, 1999; Accepted May 31, 2000.

PAN-PACIFIC ENTOMOLOGIST

77(1): 47-50, (2001)

FIRST RECORDS OF LEPTOPODIDAE IN WASHINGTON

STATE (HEMIPTERA: HETEROPTERA) WITH

NOTES ON HABITAT

R. S. ZAcK, C. N. LOONEY, M. E. HITcHcox, & J. P. STRANGE

Department of Entomology, Washington State University,

Pullman, Washington 99164-6382

Abstract.—Patapius spinosus (Rossi) was found in several locations in Benton and Whitman

Counties, Washington State. Adults were found from late January through late October with

mating pairs and immatures being found only in the fall of the year. The bug was associated

with cobblestone and basalt that was being used as roadgrade stabilization or in piles of such

rock that had been left after construction activities.

Key Words.—Insecta, Leptopodidae, Hanford Site, Patapius spinosus (Rossi), Washington State.

Extant leptopodid bugs are primarily Old World in distribution with fossil ev-

idence known from Mexico and Ecuador (Froeschner 1988). The first recording

of a leptopodid from the United States was in California (Arbuckle, Colusa Coun-

ty) where a single individual was found during the examination of tree protectors

used to trap peach twig borer larvae on almond trees (Usinger 1941). Subsequent

to this first finding, the species in question, Patapius spinosus (Rossi) has been

found in Nevada, and Idaho (Brothers 1979). Patapius spinosus is currently

known from California, Nevada, and Idaho in the United States, the Canary Is-

lands, Europe, North Africa, and has been introduced into Chile (Froeschner

1988). Both Brothers (1979) and Froeschner (1988) provide excellent illustrations

of the bug. Our finding of this species at several widely separated locations in

Washington State is a significant extension of its previously known range and

may indicate that the species is relatively widespread in the western United States.

Patapius spinosus appears to be rather generalized in its habitat selection. Both

Usinger (1941) and Brothers (1971) found the species in areas removed from

water sources; Brothers (1971) found specimens on the undersides of cobblestone

in a mine-tailings dump. The sites from which we obtained specimens are com-

parable to a mine tailings area except that our sites are located near water (which

the tailings site may have been). Our first finding occurred at the Hanford Nuclear

Site (Benton County) in southcentral Washington State. The Hanford Site is an

area of native shrub-steppe vegetation with semi-arid climatic conditions that in-

clude hot and dry summers and cold winters. Annual precipitation is less than 12

cm. Temperatures range from an average of 3° C in Jan to 33° C in July; tem-

peratures of 30° C or above occur an average of 56 days per year (ERDA 1975).

Specimens were found on the undersides of cobblestones that had been depos-

ited along the margins of a built-up gravel road that ran parallel to an alkaline

pond (West Lake-—46°36.06’ N, 119°32.78’ W) located at 150 m in elevation.

West Lake is the only naturally occurring pond on the Hanford Site. The pond is

surrounded by an alkaline crust with no emergent macrovegetation along the

shoreline. However, large areas of bullrush (Scirpus sp.) as well as various grasses

and other vegetation occur in the area surrounding the pond. The size of the pond

is a direct function of ground water elevation and fluctuates throughout long and

48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

short term periods depending on climate and seasonal weather conditions. On

average, the pond encompasses 4 hectares. The first specimens were discovered

while examining the undersides of the cobblestone on 15 Jan 1999. Subsequent

to this finding, a more concerted effort was undertaken on 29 Jan during which

several hundred rocks were turned and 30 adult specimens located. Again, on 12

Feb, several specimens were found; we continued to find specimens, sporadically,

throughout the summer and into early September when the study, at this site, was

discontinued. Only material from the first two collecting dates were taken and

processed. Additionally, a single specimen was collected in a pitfall trap (14—26

Aug 1998) located in vegetation surrounding West Lake. Although no effort was

made at these times to quantify our collections, we noticed that specimens were

never found on the section of rock that was in contact with the soil (moist) and

that there appeared to be a certain clumping phenomenon to their discovery.

Although few rocks harbored specimens, multiple specimens were sometimes dis-

covered under individual rocks. Specimens are deposited in the M. T. James En-

tomological Collection, Washington State University.

In an examination of the M. T: James Entomological Collection, Washington

State University, we located two specimens that had been collected along the

Snake River at the McCauley Ranch that was located (area since flooded) between

Almota and Penwawa Canyons (Whitman County) on 20 Mar 1971. A descriptive

label on these specimens stated that they had been collected under rocks, as had

our Hanford Site specimens. We therefore expanded our survey activities to sev-

eral, apparently likely sites adjacent to the Snake River. These sites were all

located in Whitman County, along the Snake River: Wawawai River road, ap-

proximately 19 km SW of Pullman (46°36.91’ N, 117°22.51’ W); corner of Wa-

wawai Grade road and Wawawai River road, approximately 17 km SW of Pullman

(46°36.05’ N, 117°22.65' W); Almota, (Fig. 1) approximately 24 km W of Pull-

man (46°42.22’ N, 117°27.98' W) and; Boyer Park (Fig. 2) (at Lower Granite

Dam) approximately 22 km WSW of Pullman (46°40.64' N, 117°26.74’ W).

On 28 Feb 1999, sampling was conducted at the above-delineated Snake River

sites. Records were kept of the number of rocks turned, the number of times

leptopodids were found, and in what size “‘groups.’’ A total of 1610 rocks were

turned with 94 leptopodids found. Specimens were found singly and in groups of

up to 10 on the underside of a single rock. While no samples were taken during

the summer, on 21 Oct 1999 the Snake River sites were again surveyed. At this

time, no count of bugs was maintained and only a small number were collected

and processed. However, the number of leptopodid adults was high and, indeed,

appeared higher than during our winter sampling periods. Additionally, this was

the only time that we found mating pairs and immatures. Again, leptopodids were

found in small piles of cobblestone and basalt rock that had been used as road-

grade support or had been left in small piles (often of several hundred rocks) after

completion of road or walking path construction. These latter piles were often

scattered and had been overgrown with grasses (Fig. 2). It is interesting to note

that not every pile of rock contained the bugs but, when they were found, nu-

merous rocks within the pile harbored them.

Rocks with leptopodids underneath always contained some open space below

and were almost never wet. In the few instances where rocks with leptopodids

below were wet, all of the surrounding rocks in the pile were also wet. In most

Figure 2. Pile of basalt rock at Boyer Park (Whitman County).

50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

areas, when wet and dry rocks were adjacent, it appeared that the insects favored

the dry rocks. Rocks with leptopodids on them varied extremely in color and

arrangement. There was no apparent pattern in size or color of rock and the

number of specimens occurring on the underside of the rock. Some piles of rocks

were embedded in grasses; others were on bare dirt. In no cases were leptopodids

found on the steep piles of basalt rubble adjacent to roadwork or railroads. When

the insects were found near railways and road cuts, they were always on the

undersides of rocks arranged in a flatter and less congested manner.

It is interesting to speculate on how P. spinosus was introduced into Washing-

ton. At both Hanford and the Snake River sites, the bugs were found under

cobblestone, river rock that is used for stabilization of roadways. In all instances,

the rock had been brought to the site from other areas that could not be discerned

by us. However, at Hanford, the rock may have come from the adjacent Columbia

River (approximately 3 km from West Lake) and, at the Whitman County sites,

rock may have come from the adjacent Snake River. It may be that the bugs are

associated with cobble habitat and are distributed as this rock is moved and used

in construction efforts. Unfortunately, besides our minimal observations, we have

no evidence of such movement.

ACKNOWLEDGMENT

This project was founded by The Nature Conservancy with awards from the

U.S. Department of Energy, The Nature Conservancy of Washington State, and

the Bullitt Foundation. We thank Dara Nunn and Jennifer Strange for aid in

collecting field data.

LITERATURE CITED

Brothers, D. R. 1979. First record of Patapius spinosus in Idaho and Nevada (Hemiptera: Leptopod-

idae). Great Basin Nat., 39: 195-196.

Energy Research and Development Administration (ERDA). 1975. Final Environmental Statement,

Waste Management Operations. ERDA-1538. Volume 2. Energy Research Development Ad-

ministration, Richland, Washington.

Froeschner, R. C. 1988. Family Leptopodidae Amyot and Serville, 1843. p. 166. Jn Henry, T: J. and

R. C. Froeschner (eds.). Catalog of the Heteroptera, or True Bugs, of Canada and the Continental

United States. E. J. Brill, Leiden.

Usinger, R. L. 1941. A remarkable immigrant leptopodid in California. Bull. Brooklyn Entomol. Soc.,

36: 164-65.

Received 24 Nov 1999; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 51-53, (2001)

Scientific Note

ERNOBIUS MOLLIS (L.) (COLEOPTERA: ANOBITDAE): AN

EXOTIC BEETLE COLONIZES MONTEREY PINE,

PINUS RADIATA D. DON, IN NORTHERN CALIFORNIA

In 1990, the first collection of the bark anobiid, Ernobius mollis (L.), west of

Texas was made in Oakland, Alameda County, California from a living Norway

spruce, Picea abies (L.) Karsten, planted as an ornamental tree (Seybold, S. J. &

J. L. Tupy, 1993. Pan-Pacif. Entomol., 69: 36—40.). Ernobius mollis is native to

northern Europe, but has been introduced into North America (Craighead, F C.

1950. Insect Enemies of Eastern Forests, USDA For. Serv. Misc. Pub. 657.) and

the southern hemisphere (Casimir, J. M. 1958. Div. Wood Tech., For. Comm. New

South Wales Tech. Notes, 2: 24—27 [Australia, New Zealand, and South Africa]).

Seybold and Tupy (1993) speculated that E. mollis, known to colonize Monterey

pine, Pinus radiata D. Don, in the southern hemisphere, might soon colonize the

abundant urban plantings of P. radiata in the San Francisco Bay Area.

On 20 Sep 1993, a homeowner in Albany, Alameda County, California (Talbot

Street) reported that she had observed insect damage in a variety of wooden

articles in her home. On examination of her residence, none of the wooden articles

of concern to the homeowner appeared to harbor active infestations of wood-

destroying insects. However, examination of several pieces of barked P. radiata

firewood stacked within the structure indicated that the small branch sections were

infested with a wood-boring insect. These P. radiata branches had emergence

holes through the bark surface. Peeling the already loose bark from the xylem

revealed live larvae and pelleted frass in the bark-xylem interface as well as larval

insect galleries etched in the xylem surface. The homeowner reported that the P.

radiata branches had been cut approximately six months earlier (1.e., Mar 1993)

from a standing tree in Fremont, Alameda County, California on Bud Court near

the Highway 880 Mowry Avenue Exit. The branches were placed in a laboratory

cage at the University of California at Berkeley at ambient indoor temperature

(16° C to 29° C) and larval and adult E. mollis were periodically collected from

the logs from Sep 1993 to Jun 1996. By Jun 1996 the bark-xylem interface had

been completely obliterated and the remaining paper thin bark formed an easily

damaged shell over the xylem.

This collection documents the presence of E. mollis in urban plantings of P.

radiata in the San Francisco Bay Area. Although this collection record was from

cut branches, future northern California reports of infestations from moribund P.

radiata and perhaps other ornamental conifers should be expected. Through its

association with moribund tree tissue, E. mollis may also play a role in the dis-

semination of the pitch canker fungus, Fusarium circinatum Nirenberg &

O’Donnell (Nirenberg, H. I & K. O’Donnell. 1998. Mycologia, 90: 434—458.),

a fairly recently introduced and fatal disease of P. radiata and other pines in

coastal northern California (McCain, A. H., C. S. Koehler & S. A. Tjosvold.

1987. Calif. Agric., 41: 22—23.). In Europe, E. mollis is documented to infest

cones of Douglas-fir, Pseudotsuga menziesii (Mirbel) Franco, and giant sequoia,

32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

Sequoiadendron giganteum (Lindley) J. Buchholz (Roques, A. 1983. Les insectes

ravageurs des c6nes et graines de coniféres en France, Institut National De La

Recherche Agronomique Publication.). A native species of Ernobius [punctulatus

(LeConte)] is known to infest cones of P. radiata (White, R. E. 1982. A Catalog

of the Coleoptera of America North of Mexico Family Anobiidae, USDA Agric.

Handbook No. 529-70, Furniss, R. L. & V. M. Carolin 1992. Western Forest

Insects, USDA For. Serv. Misc. Publ. No. 1339.) and to acquire the pitch canker

fungus through its association in the P. radiata cone with other cone-infesting

insects (Hoover, K., D. L. Wood J. W. Fox & W. E. Bros. 1995. Can. Entomol.,

127: 79—91.). Therefore, association of E. mollis with moribund P. radiata cone

tissue and the bark-xylem interface of branch and stem material appear to be a

likely means for this introduced insect to disseminate pitch canker disease in

northern Califormia. As both the insect and the fungal pathogen have wider host

ranges among conifers, E. mollis could play a role in transmission of the fungus

to new hosts and to new regions such as the montane forests of the Sierra Nevada

or the coastal forests of the Pacific Northwest.

Pinus radiata is frequently brought into coastal California homes as firewood,

and this example illustrates how easily larval and adult E. mollis might enter

homes. However, experiences from Europe, the southern hemisphere, and eastern

North America suggest that because of its requirement for bark-covered sapwood,

E. mollis will not be a pest in structures in northern Califormia unless it encounters

bark-covered timbers, or boards with unfinished edges or bark-occluded knots.

This example also illustrates that although an active infestation of E. mollis may

be present in firewood in a structure, if damage to other, unbarked wooden items

in the structure clearly occurred prior to manufacture, any relationship between

E. mollis and the damage can likely be discounted.

Finally, the original colony of E. mollis collected 22 Feb 1990 in Oakland was

maintained until Aug 1996 in a laboratory cage at ambient indoor temperature on

the original P. abies branch and stem material. On 12 Jul 1996, seven live male

and twelve live female adults were recovered from the cage. One pair of these

adults was observed in copulo and live larvae were also present. On 6 Aug 1996,

one live male, four live females, and two live larvae were also recovered from

the cage before the logs were frozen and disposed of. Thus, multiple, consecutive

generations of E. mollis successfully re-infested the same dry substrate for more

than 6.5 years.

Record.—USA. CALIFORNIA. ALAMEDA CO.: 1.5 km SE Albany Hill, Tal-

bot Street, Albany, 20 Sep 1993, S. J. Seybold, Pinus radiata.

Acknowled gment—I thank R. Kohl for bringing this occurrence of E. mollis

to my attention; A. J. Blomquist (University of Nevada, Reno) for helping to

process the specimens; D. L. Wood (University of California at Berkeley) and B.

J. Cabrera (University of Minnesota) for critical reviews of earlier drafts of this

report; and N. G. Rappaport (USDA Forest Service, Albany, California) for trans-

lation of French literature. This work was supported by a cooperative research

agreement (PSW-92-0014CA) between DLW and the Chemical Ecology of Forest

Insects Project, Pacific Southwest Research Station, U.S. Department of Agricul-

ture, Forest Service, Albany, California. Specimens of E. mollis recovered from

P. radiata were deposited in the California Academy of Sciences Entomology

Collection, Golden Gate Park, San Francisco, Califomia 94118.

2001 SCIENTIFIC NOTE 53

Steven J. Seybold, Departments of Entomology and Forest Resources, 219

Hodson Hall, 1980 Folwell Avenue, University of Minnesota, St. Paul, Minnesota

55108-6125.

Received April 9, 2000; Accepted July 6 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 54—56, (2001)

Scientific Note

RANGE EXTENSION OF PSENEO PUNCTATUS FOX AND

NOTES ON PREDATION OF AN INTRODUCED

SHARPSHOOTER, HOMALODISCA COAGULATA (SAY)

The tribe Psenini Bohart & Menke (Hymenoptera: Sphecidae: Pemphredoninae)

was described in 1976 (Bohart, R. M. & S. Menke 1976. Sphecid wasps of the

world, a generic revision, University of California Press, Berkeley, California,

USA). It is a poorly understood group, and basic biologies are unknown. Psenini

are small, slender, delicately constructed wasps that most commonly occur in

association with damp shady situations. Some genera and species within the tribe

are located in southern Califormmia (R. M. Bohart, personal communication). How-

ever, only one species of Pseneo has been recorded in California, Pseneo longiv-

entris (Cameron) (van Lith, J. P. 1975. Neotropical species of Psen and Pseneo

[Hymenoptera: Sphecidae: Psenini]. Tijdeschrift voor Entomologie. 118: 1—41).

We became aware of a large aggregate of digging wasps on the grounds of the

University of California Riverside in August 1998. Many of the nests seemed to

be completed, closed, nests, but the majority had entrances open with provisioning

occurring. Many nests were so close together that the mounds at the entrances

were overlapping. The wasps appeared to have little difficulty finding their own

nests, and no aggressiveness was observed between neighboring females. We

counted 181 nesting sites within 30 plant propagation trays (45 x 45 x 10 cm)

filled with U. C. mix #3 (Matkin, O. A. & P. A. Chandler. 1957. The UC soil

type mixes. U. C. Berkeley, California Agriculture Experiment Station, Manual

23) to a depth of 6 cm. All trays were located within a 10 m? area in an open

lathhouse on raised benches. The soil-filled trays were being temporarily stored

prior to being used for seedling establishment in unrelated experiments. Nesting

wasps were observed wherever the soil was exposed in any of the trays. We

observed wasps active in 53 (29%) of the observed 181 nesting sites.

Specimens were collected and identified as P. punctatus Fox, a species found

from North Dakota to southern Mexico. Features of the male, including the spe-

cies-specific details of the antenna and genital capsule, clearly match those of

typical P. punctatus. However, the Riverside wasps have extensive reddish col-

oration on the legs in both sexes; in van Lith’s (1975) key they run to P. carolina,

a southeastern U.S. species described exclusively from females (also originally

described as a subspecies of P. punctatus). Specimens of putative male P. carolina

from Texas raise questions as to the validity of the taxon, as they are nearly

identical to the California specimens of punctatus (A. Finnamore, personal com-

munication), and the resultant distribution of “‘carolina’’ would apparently bisect

the known distribution of punctatus. Our suspicion is that P. punctatus is a species

with a large distribution and substantial geographic variation, which has not yet

been sampled adequately to resolve the limits of the various forms, including P.

carolina, which was probably inappropriately elevated to species status by van

Lith. Pseneo punctatus represents the second Pseneo species collected in Cali-

fornia (Bohart, personal communication; Krombein, K. V. 1979. Superfamily

2001 SCIENTIFIC NOTE 55

Sphecoidea. pp. 1573-1740. Jn Krombein, K. V., P. D. Hurd, Jr., D. R. Smith &

B. D. Burks [eds.]. Catalogue of Hymenoptera of America North of Mexico. 2:

1119-2209. Smithsonian Institution Press, Washington, D.C., USA). Voucher

specimens are located at the University of California Riverside, Entomology Re-

search Museum.

During our observations of P. punctatus over a period of several days we noted

the following behaviors. Nest excavation was observed only in the afternoon.

Nests were excavated with the mandibles and front legs. Soil was loosened by

the mandibles, formed into a small uneven clump, and pushed backward beneath

the body with the front legs. Usually the clumps of soil are thrown clear of the

body by the initial thrust of the front legs. If not, the soil clumps are thrown clear

of the body with thé assistance of the middle and hind legs. The female typically

digs the burrow vertically straight down. As the wasp digs deeper, small clumps

of soil clog the nest entrance and hide the digging female. Occasionally, as soil

accumulates near the nest entrance, the wasp would back up and clear the entrance

hole. During mound building, most wasps would occasionally tamp the soil down

with their abdomens, presumably to prevent the nest entrance from becoming

obstructed with excavated soil. Nest entrances were 6.50 + 2.35 mm (mean +

SD, n = 16, range 5-12 mm) in diameter, and were sometimes hidden under

fallen leaves or other debris.

Over a period of hours to days, females repeatedly provisioned their nests with

large Homopteran sharpshooters (Cicadellidae). When the nest was fully provi-

sioned, the female would repeatedly emerge and re-enter the nest entrance drag-

ging clumps of soil with her forelegs back into the hole each time she entered.

When the upper portion of the burrow was filled with soil, the female would walk

over the top of the mound repeatedly scraping the surface with her forelegs and

tapping it with her abdomen. At this point, it was no longer possible to discern

that a burrow was present.

What was of particular interest to us was that P. punctatus was using both the

native smoketree sharpshooter, Homalodisca lacerta (Fowler) (Homoptera: Ci-

cadellidae), and the introduced glassywinged sharpshooter, Homalodisca coagu-

lata (Say), as prey. Homalodisca coagulata became established in California dur-

ing 1989-1990 (Sorenson, J. T. & R. J. Gill. 1990. Pan-Pacific Entomol., 72:

160-161). Ithas now been established in 11 counties and on numerous host plants

including citrus, grapes, oleander and a wide variety of ornamental landscape and

native plants. Currently, H. coagulata is the principle vector of Pierce’s disease

of grapevines in the southern portion of California (Costa, H. S., M. J. Blua, J.

A. Bethke, & R. A. Redak. 2000. [In press]. HortScience University of California,

Office of the President. 2000. Report of the Pierce’s Disease Research and Emer-

gency Task Force). The vast majority of prey we observed were H. coagulata

adults. Pseneo punctatus’ natural prey most likely is H. lacerta, and we believe

it has expanded its usable hosts to include the recently introduced sharpshooter.

When several of the nests were excavated, we usually observed three cells per

nest, but it was difficult at times to accurately determine how many cells each

nest contained. Many cells contained sharpshooters that were old, decayed and

untouched. We could not determine if they were abandoned or if they were part

of the normal provisioning of the nest and simply unused. Although the number

of sharpshooters per cell varied, they commonly contained about 4 adults. In some

56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

cells, we observed eggs deposited between the front pair of coxae on the venter

of a single sharpshooter. They were oblong, white, and slender, and on average

they were 2.29 + 0.35 mm long (mean + SD, n = 5, range 3.0—4.6).

Wasp cocoons were also observed. On average cocoons were ellipsoidal, 14.50

+ 1.34 mm in length, and 5.79 + 0.70 mm in width (mean + SD, n = 14, range

13-17, and 5—7 respectively). Cocoons were covered with small pieces of plant

material and sharpshooter fragments (mostly head capsules and wings).

Occasionally, we observed wasps emerging from the soil as adults. Emergence

holes were a mean of 3.64 + 0.50 mm in diameter (mean + SD, n = 11, range

3—4) and unlike active nest entrances, they were not marked by any debris or

mounds of dirt.

Specimens Examined CALIFORNIA. RIVERSIDE Co.: Riverside, University of California cam-

pus, lath house, 21 Aug 1998, Kathleen A. Campbell, swept, 2 females, deposited: University of

California, Riverside, Entomology Research Museum; same loc., 25 Aug 1998, reared from larva, 1

female, reared from pupae, (6 males, 2 females), deposited: University of California, Riverside, En-

tomology Research Museum; same loc., 27 Aug 1998, reared from larva, 1 female, deposited: Uni-

versity of California, Riverside, Entomology Research Museum; Riverside, University of California

campus, undeveloped area 2 meters west of the lath house, 11 Sept 1998, James A. Bethke, swept

from eucalyptus, (1 male, 3 females), deposited: University of California, Riverside, Entomology

Research Museum; Riverside, University of California campus, solid bin adjacent to Entomology

Annex I, swept, 1 female, deposited: University of California, Riverside, Entomology Research Mu-

seum.

Acknowled gment.—Authors thank R. Bohart for review of this manuscript, L.

Petro for helpful observations, and R. Bohart and A. Finnamore for helpful com-

ments.

James A. Bethke, Kathleen A. Campbell, Matthew J. Blua, Richard A. Redak,

and Douglas A. Yanega, Department of Entomology, University of California,

Riverside, California 92521.

Received 11 Mar 2000; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(1): 57-60, (2001)

Scientific Note

MONODONTOMERUS ARGENTINUS BRETHES

(HYMENOPTERA: TORYMIDAE): A PARASITOID OF

EUGLOSSA NIGROPILOSA MOURE (HYMENOPTERA:

APIDAE: EUGLOSSINAE)

Euglossine bees are the only group in the subfamily Apidoideae that do not

have eusocial behavior. Euglossines display a broad spectrum of social interac-

tions, from solitary to presocial, and are thus an important group for studying the

evolution of eusociability in bees (Gardéfalo, C. A. 1985. Entomol. Gener., 11:

77-83). An important factor involved in the origin of social behavior is the effect

of parasites and parasitoids on survival and reproduction (Roubik, D. W. 1989.

Ecology and natural history of tropical bees [1st ed.]. Cambridge University Press.

New York).

Although it is known that microhymenoptera are parasitoids of Euglossa (Zuc-

chi, R., S. FE Sakagami & J. M. EK de Camargo. 1969. J. Fac. Sci. Hokkaido Univ.,

Series VI, Zool., 17: 271-380), little is known about their relationship with their

hosts (Dressler, R. L. 1982. Ann. Rev. Ecol. Syst., 13: 373-394). Here I present

data on the parasitic behavior of the microhymenopteran Monodontomerus ar-

gentinus Brethes (Torymidae) and discuss its effect on the social structure of E.

nigropilosa. 1 compared the percentage parasitism of M. argentinus on both an

attended and an unattended nest of E. nigropilosa. This study was carried out

between August 1994 and April 1995.

Euglossa nigropilosa is a communal bee of NW South America, the colonies

of which have active bees all year. It is distributed between 700 and 1400 m in

the Andes of Colombia and Ecuador. This species builds nests with an envelope,

a resinous cover that protects the cells. Cells are of 10 X 6 mm, with soft resinous

walls. I found several nests of E. nigropilosa in wood cavities of timber buildings

at the Reserva Natural La Planada (RNLP), Colombia (77°24’ W, 1°5’ N). The

nests contained colonies of up to 22 females (Otero, J. T. 1996. Bol. Mus. Ent.

Univ. Valle. 4: 1-19).

Nests of E. nigropilosa were parasitized by M. argentinus, an ectoparasitic

wasp that is known to attack the solitary bee Eufriesea nigrescens Friese, another

euglossine, in the eastern part of the Andes in Colombia (Sakagami, S. FE & Strum.

1965. Insecta Matsumurana, 28: 83—97). This is the first record of a different host

for M. argentinus. Samples of both species were deposited in the Museo de En-

tomologia of Departamento de Biologia de la Universidad del Valle (Cali, Co-

lombia).

With the objective to study bee behavior inside the nest, on 20 Sept. 1994 I

placed a nest with four adult bees and 18 cells of E. nigropilosa in a wooden box

30 X 15 X 10 cm, with a glass plate covering the upper surface and a wood

cover which remained in place during non-observation hours. The bees had free

access through a one cm diameter hole. However, the adult bees abandoned the

box nest. This undefended nest was compared to a nearby undisturbed nest, in

which adult bees were still present, for levels of cell parasitism by M. argentinus.

58 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

This second nest provided a control for the degree to which adult bees can protect

their nest from parasitism. On 11 Nov. 1994 I opened all the cells of the aban-

doned box nest to check the level of parasitism of M. argentinus. For comparison,

I opened 20 randomly chosen cells from the undisturbed nest to determine whether

or not they were parasitized.

Comparison between the nests with and without females revealed that parasit-

ism dropped significantly (x? = 12.73, P = 0.0004) with the presence of adults

in the nest. The percentage of parasitism on the 18 cells in the box nest with

female bees absent was 72%. However, the percentage for the control nest, which

had between eight and 14 female bees, was only 15%, based on the sampled cells.

In the box nest cells there was an average of 13.69 (+ 9.04 SD, n = 13) wasp

offspring per cell. Individual wasps were found at all developmental stages: young

larvae, mature larvae, prepupae (pupae without pigmentation); and pupae (black

pupae).

I observed the behavior of 38 M. argentinus wasps inside the defended E.

nigropilosa nest. Monodontomerus argentinus arrived at the nest entrance after a

zigzag flight. The wasp flew around the entrance for up to five min, before landing

at the entrance hole. Parasitoid wasps entered the nest slowly, using the same

entrance hole that the bees used. Upon landing, the wasps drummed on the surface

of the nest with their antennae, moving them fast and harmoniously. The wasps

alternated each antenna in this process. The wasps advanced slowly and continued

drumming until they arrived at the entrance hole. Once inside the nest, the wasps

drummed with their antennae on the surface of a cell for up to five min. without

moving any other part of the body. When a wasp found an appropriate cell, it

continued drumming for several minutes before ovipositing. Oviposition lasted

for up to two min. For this process the wasp raised her abdomen, exerted her

Ovipositor and put it in contact with the resin wall of the cell. Occasionally the

parasitoids withdrew their abdomen and continued drumming on the next cell.

Euglossa nigropilosa appeared to recognize M. argentinus as a natural enemy.

These interactions were very strong. For example, a parasitoid, recently killed in

a cyanide bottle, was left exposed in the nest. This body was torn apart by a

resident bee. On first detecting the dead wasp, the bee became excited and circled

around it, touching it with her antennae. It then bit the wasp in the abdomen and

pushed it five cm away from the cell. The bee again found the dead wasp on the

ground and bit it repeatedly for more than a minute. Following this attack the

parasitoid lost two legs and had its wing and crushed abdomen nearly destroyed.

This strong reaction may, however, have been a response to the odor of the

cyanide, and not the presence of the parasitoid body. During observations of

attempts to parasitize a cell, I never observed the bees to perceive the wasp’s

presence. The wasps proved very adept at escaping from the bees notice in the

nest. When a parasitoid was approached by a bee, the wasp jumped to the ground,

so evading detection, and remained there for several minutes before attempting

to oviposit again.

Sakagami and Strum (1965) found an aggregation of nests of the solitary bee

Euplusia longipennis in Colombia in 1956 (Eufriesea nigrescens sensu Kimsey,

L. S. 1982. Systematics of bees of the genus Eufriesea. University of California

press). These nests had 81 highly elaborate resinous cells of approximately 19 X

9 mm, two of which were parasitized by M. argentinus (2.5% parasitism). Each

Table 1. Length of bees and cells, and number of broods per cell of the parasitic wasp Monodontomerus argentinus in two different euglossine bees in Colombia.

Euglossa nigropilosa Eufriesea nigrescens

Bee length (mm) 12 16

Cell length (mm) 10 19

Cell diameter (mm) 6 8

Number of broods by cell 13.7 (SD = 9.4; n = 13) 33.5 (SD = 2.1; n = 2)

AHLON OWILNAIDS 1007

6S

60 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(1)

parasitized cell had an average of 33.5 (SD + 2.1, n = 2) parasites. Eufriesea

nigrescens and Euglossa nigropilosa were attacked by the same parasitoid, M.

argentinus, but the two host species have different defense strategies. The solitary

species, E. nigrescens, provides cells with thick protective walls. In contrast E.

nigropilosa build cells with a thin, vulnerable wall. However, adult bees of E.

nigropilosa may protect the nest from M. argentinus. In addition, the presence of

a resinous envelope enclosing the nest may restrict the wasp’s entrance. These

defenses of E. nigropilosa, however, appear to provide less effective protection

from the parasitic wasps than the thick cell walls of E. nigrescens. Parasitism

incidence was higher in E. nigropilosa, between 72% in box nest and 15% in the

control nest, than in E. nigrescens (2.5%). There appears to be less parasitism

when nests are protected by many active females. In this case, the parasitoids

probably spent more time escaping from resident bees and were not able to ovi-

posit in cells.

From these observations I suggest that the presence of active bees in the nest

may affect the rate of M. argentinus parasitism of E. nigropilosa. Despite having

vulnerable cell walls, E. nigropilosa has active bees throughout the year and the

large colony size makes the presence of a female in the nest more likely, decreas-

ing the risk of M. argentinus attack. In contrast, E. nigrescens is a seasonal bee

with oviposition limited to only two months of the year. Females of Eufriesea

nigrescens die after oviposition and they can not protect the cells. Thus an in-

trinsic protective mechanism is needed, in this case a thick resinous cell wall that

is difficult for the parasitoid to penetrate.

Acknowled gment.—I thank Dr. Philip Silverstone-Sopkin and Dra. Patricia Cha-

c6n from Universidad del Valle and Dr. R. Ospina and G. Nates from Universidad

Nacional de Colombia for their support; Dr. E. E. Grissell from USDA for iden-

tification of wasps; Dr. J. Ackerman, Dr. W. O. McMillan, Dra. N. Flanagan and

Dr. P. Bayman for comments in the manuscript; FES foundation for logistic sup-

port in the Reserva Natural La Planada and the MacArthur Foundation for eco-

nomic support.

J. Tupac Otero, Department of Biology, University of Puerto Rico, San Juan,

Puerto Rico 00931-3360.

Received I Mar 2000; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

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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

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Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

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THE PAN-PACIFIC ENTOMOLOGIST

Volume 77 January 2001 Number 1

Contents

PUBLICATIONS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY ____-------------------

BRAILOVSKY, H. & E. BARRERA—The genus Notobitopsis Blote with the description of

two new species (Hemiptera: Heteroptea: Coreidae: Coreinae: Mictini)

SCHEFTER, P. W. & K. A. JOHANSON—Three new species of Helicopsyche from Vietnam

(Trichoptera: Helicopsychidae)

MILICZKY, E. R. & C. O. CALKINS—Prey of the spider, Dictyna coloradensis, on apple,

pear, and weeds in central Washington (Araneae: Dictynidae)

BERNAL, J. S., D. GONZALEZ & E. DAVID-DIMARINO—Overwintering potential in

California of two Russian wheat aphid parasitoids (Hymenoptera: Aphelinidae et Aphid-

iidae) imported from Central Asia

ANDREWS, F. G.—A new species of Corticarina from Arizona (Latridiidae: Corticariini)

SCHWARTZ, M. D. & M. A. WALL—Melanotrichus boydi, a new species of plant bug

(Heteroptera: Miridae: Orthotylini) restricted to the nickel hyperaccumulator

Streptanthus polygaloides (Brassicaceae)

ZACK, R. S., C. N. LOONEY, M. E. HITCHCOX & J. P. STRANGE—First records of

Leptopodidae in Washington State (Hemiptera: Heteroptera) with notes on habitat

SCIENTIFIC NOTES

FIERROS-LOPEZ, H. E. & J. L. NAVARRETE-HEREDIA—Altitudinal distribution and phe-

nology of three species of carrion beetles (Coleoptera: Silphidae) from Nevado de Coli-

ma, Jalisco, México

SEYBOLD, S. J.—Ernobius mollis (L.) (Coleoptera: Anobiidae): an exotic beetle colonizes

Monterey pine, Pinus radiata D. Don, in Northern California

BETHKE, J. A., K. A. CAMPBELL, M. J. BLUA, R. A. REDAK, AND D. A. YANEGA—Range

extension of Pseneo punctutus Fox and notes on predation of an introduced sharpshoot-

er, Homalodisca coagulata (Say)

OTERO, J. T—Monodontomerus argentinus Brethes (Hymenoptera: Torymidae): a parasitoid

of Euglossa nigropilosa Moure (Hymenoptera: Apidae: Euglossinae)

19

28

37

ao

47

45

51

54

a7

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 77 April 2001 Number 2

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

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71(2): 61-67, (2001)

INVERSE DENSITY-DEPENDENT PARASITISM OF

OPSIUS STACTOGALUS FIEBER (HOMOPTERA:

CICADELLIDAE) BY GONATOPUS SP.

(HYMENOPTERA: DRYINIDAE)

W. D. WIESENBORN

1410 Stacey Lane, Boulder City, Nevada 89005

Abstract—The relationship between Opsius stactogalus Fieber leafhopper density (abundance

per plant-sample mass) and parasitism by the dryinid wasp Gonatopus sp. was examined. Branch-

es cut from Tamarix ramosissima Ledebour shrubs, sustained by treated wastewater, were

weighed and sampled for parasitized and unparasitized leafhoppers. Leafhopper abundance in-

creased linearly as branch mass increased exponentially (mass®-®), in agreement with isometric

scaling laws relating leaf abundance to branch mass. The proportion of leafhopper nymphs

parasitized (18%) was greater than the proportion of adults parasitized (2.9%). Branches were

more likely to contain one or more parasitized nymphs, signifying exploitation of the branch by

Gonatopus, as nymph density increased. On branches containing at least one parasitized nymph,

increasing nymph density was related to an increase in the number of parasitized nymphs but

to a decrease in parasitism rate. Parasitism of O. stactogalus nymphs on branches exploited by

Gonatopus was inverse density dependent. Gonatopus appears not to regulate populations of O.

stactogalus or limit the leafhopper’s damage to T. ramosissima.

Key Words.—Insecta, Cicadellidae, Opsius stactogalus, Dryinidae, Gonatopus, Tamarix, para-

sitism.

The tamarix leafhopper, Opsius stactogalus Fieber, is a small, cryptic insect

primarily found on tamarisk, Tamarix spp. (Tamaricaceae) (Harding 1930, Liesner

1971). Tamarisk is a halophytic shrub or tree facultatively-dependent on shallow

groundwater (Brock 1994, Di Tomaso 1998). Species of Tamarix are native to

the Old World, occurring from the Mediterranean across southern Russia to east-

ern Asia, and were imported to the USA in the early 1800s for use as ornamentals

(Robinson 1965). Tamarisk since has spread beyond cultivation with the most-

recent survey estimating the plant’s extent to exceed 350,000 ha in the western

USA (Robinson 1965). Tamarisk’s invasiveness, water uptake, and low diversity

of dependent wildlife has made it the target of classical biological control (For-

nasari 1997). The taxonomy of New World Tamarix is unclear (Brock 1994, Di

Tomaso 1998); eight species occur in North America (Baum 1967) with five

species, four deciduous and one evergreen, currently recognized in California

(Wilken 1993).

Opsius stactogalus is the most abundant insect on tamarisk (Harding 1930,

Hopkins & Carruth 1954, Liesner 1971). The insect’s green color and size (length

0.81 mm for first-instar nymphs, 4.5 mm for adult females, Harding 1930) make

it difficult to distinguish from tamarisk’s sessile, scalelike leaves (1.5 to 3.5 mm-

long in Tamarix ramosissima Ledebour, Wilken 1993). The life cycle of O. stac-

togalus is comprised of five nymphal instars, adults, and eggs inserted within

stems (Harding 1930). Generations per year number three in Kansas (Harding

1930) and four in New Mexico (Liesner 1971). Like many other Cicadellidae, O.

stactogalus is a fluid-feeder, and the aggregate feeding imposed by populations

of the leafhopper can reduce tamarisk’s growth (Liesner 1971). Although the

62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

leafhopper was described as a new species in 1907 from collections in Texas, a

later comparison of Meditteranean and North American specimens and review of

the systematic literature revealed the 1907 species description as synonymous with

an original description in Europe during 1847 (Harding 1930). Opsius stactogalus,

like its host tamarisk, is native to the Old World.

Opsius stactogalus nymphs and adults are parasitized by the dryinid wasp Gon-

atopus spp. (Harding 1930, Liesner 1971). Dryinidae occur worldwide and par-

asitize only the homopteran suborder Auchenorrhyncha (Olmi 1984). Dryinid lar-

vae are individually enclosed in a sac of exuviae that protrudes between their

host’s thoracic or abdominal sclerites. Adult females of many Dryinidae species

also are predaceous on their hosts (Olmi 1984). The only published records of

Gonato pus parasitizing O. stactogalus in North America are of Gonatopus caroli

Olmi and Gonatopus curriei Krombein (Guglielmino & Olmi 1997). One adult

female Gonatopus reared from a parasitized O. stactogalus | collected at the

present study’s locality was identified as G. caroli (M. Olmi, personal commu-

nication).

The effect of parasitism by Gonatopus on the abundance of O. stactogalus is

uncertain. Harding (1930) described parasitism rate by Gonatopus as low and

unable to significantly affect rapid population increases of O. stactogalus. In con-

trast, Liesner (1971) described parasitism rates (30—44% of leafhoppers parasit-

ized) high enough to hinder his collecting leafhoppers for experimental trials.

Increasing parasitism rate with increasing O. stactogalus density (density depen-

dence) may regulate leafhopper populations (Turchin 1995) and decrease their

detrimental effect on tamarisk. Parasitism rate decreasing with increasing leaf-

hopper density (inverse density dependence) or unrelated to leafhopper density

(density independence) would not regulate populations of O. stactogalus and limit

the insect’s damage to tamarisk. A synopsis (Stiling 1987) of the relationships

between insect-host density and parasitism rate (171 published accounts) found

density independence most common (52%) followed by density dependence

(25%) and inverse density dependence (23%). This study examines the relation-

ship between density of O. stactogalus and parasitism rate by Gonatopus.

MATERIALS AND METHODS

The study site was located at the effluent discharge of the City of Boulder City,

Clark County, Nevada, wastewater treatment plant. The treatment plant discharges

approximately 2000 m? of secondary-treated municipal effluent per day equally

into two parallel unlined southward-flowing channels approximately 0.1 km apart.

Effluent discharged to each channel maintains a 2-m wide surface flow extending

approximately 3 km downstream. Sustained by the effluent, and bordering both

channels, are dense, linear stands of the deciduous tamarisk T. ramosissima al-

ternating with clumps of cat-tail, Typhus latifolia L. (Typhaceae). The site lies at

an elevation of 610 m within the Mojave Desert; creosote bush, Larrea tridentata

(Sessé & Mocino ex Candolle) Coville (Zygophyllaceae), and bursage, Ambrosia

dumosa (Gray) Payne (Asteraceae), are the dominant plant species surrounding

the effluent-supported vegetation.

Leafhoppers on tamarisk were sampled at 12 points along the west edge of the

east channel. Sampling points were located every 80 m along the channel begin-

ning 80 m downstream of the effluent discharge. Three samples were taken at

2001 WIESENBORN: GONATOPUS PARASITISM OF OPSIUS 63

each point during the morning on 17-18 Jul, 16—17 Aug, 16—17 Sep, and 20-21

Oct 1999. Each leafhopper sample was taken by bagging a 1-m long tamarisk

branch within a plastic trash bag (159 liter, 84 cm X 1.2 m, 76 pm thick) held

open with a 52-cm diameter metal hoop. The bag was quickly swept over the

branch and slipped off the hoop to constrict the bag around the proximal-end of

the branch. The branch then was cut at the top of the bag, pushed into the bag,

and the bag closed. Enclosed leafhoppers were killed with an aerosol insecticide

(Raid® Concentrated Deep Reach Fogger, S.C. Johnson & Son, Racine, Wiscon-

sin) containing 14 g of 1.7% cypermethrin. Approximately 10 min after treatment,

the bag was held upright and the branch shaken for 30 sec. Dead leafhoppers

shaken from the branch were collected into a 3-dram (15-mm diameter < 65-mm

long) patent-lip glass vial. Plant mass supporting the collected leafhoppers was

measured (+ 2 g) by removing the branch from the bag and weighing it with a

300-g capacity spring scale. Vial contents were examined with a microscope, and

collected leafhoppers were sorted from plant debris and stored in 70% ethanol.

Leafhoppers were examined at 30X and the number of parasitized and unpara-

sitized nymphs and adults counted. Parasitized and unparasitized O. stactogalus

in ethanol were deposited as vouchers at the University of California, Riverside,

Entomology Museum.

Opsius stactogalus density was estimated by first determining the relationship

between leafhopper abundance and plant mass. Leafhopper abundance (trans-

formed In[Y + 0.5]) was regressed (SPSS version 6.1, Chicago, Illinois) against

plant mass and life stage, nymph or adult coded as an indicator variable. Two

regressions were performed with plant mass in grams and with plant mass trans-

formed In(grams). Plant mass transformed In(grams) was selected, due to its great-

er partial 7?, as the better linear predictor of leafhopper abundance and used to

calculate leafhopper density. The interaction life stage < plant mass was added

to the regression to test if the trends in abundance across transformed plant mass

differed between nymphs and adults. For graphing, abundances of nymphs and

adults in each sample were summed, transformed In(Y + 0.5), and regressed

against transformed plant mass.

Parasitism of O. stactogalus was examined by first comparing (x? test) the

proportions of parasitized nymphs and adults. Parasitism of nymphs was further

examined at two spatial scales, the tamarisk branch and the individual leafhopper.

To evaluate exploitation of tamarisk branches by ovipositing Gonatopus, the in-

fluence of nymph density (dn[N + 0.5]/In[g]; NW = number of nymphs, g = grams

plant mass) on the probability of a leafhopper sample containing at least one

parasitized nymph was determined with logistic regression (Neter et al. 1996). A

predicted change and 95% confidence interval in the odds of a Gonatopus ex-

ploiting versus not exploiting a branch as nymph density increased was calculated

from the regression coefficient and its standard error.

To evaluate parasitism of individual leafhoppers on branches exploited by Gon-

atopus, the number of parasitized nymphs (G) in leafhopper samples with at least

one parasitized nymph was transformed In(G + 0.5) and regressed against nymph

density. One outlier leafhopper sample, collected on 17 Aug 1999, containing 35

unparasitized nymphs and 41 parasitized nymphs was excluded from the regres-

sion due to its extreme influence (standardized residual = 3.4) on the fitted func-

tion. Parasitism rate (R) on branches exploited by Gonatopus was quantified as

64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

«ep, «e@)

=~

Leafhopper Abundance, Y+0.5

ThY = 0.80 - 3.7X

Parasitism Rate, (G+ 0.5)/(N - G+ 0.5)

0.1 02 03 04 05 06 0.7 0.8

Plant Mass (g) Nymph Density, In(N + 0.5)/In(g)

Figures 1-2. Fig. 1 (left). Opsius stactogalus abundance (nymphs + adults) plotted on a logarith-

mic scale against Tamarix-branch mass. Curve was fit by linear regression with plant mass transformed

In(g). Fig. 2 (right). Parasitism rate of O. stactogalus nymphs versus nymph density (V = nymph

abundance) on exploited branches containing = 1 parasitized nymph. Parasitism rate is the ratio of

parasitized nymphs (G) to unparasitized nymphs plotted on a logarithmic scale. The size of each data

point represents its final weight determined from iteratively reweighted least squares regressions.

the ratio of the number of parasitized nymphs to the number of unparasitized

nymphs (N — G), transformed as an empirical logit (Agresti 1990), R = In[(G +

0.5)/(N — G + 0.5)] in each leafhopper sample. Iteratively reweighted least

squares regressions (see logistic regression with repeat observations, Neter et al.

1996) of parasitism rate against nymph density were performed to produce a

maximum likelihood estimate of the regression coefficient. The regression was

unweighted in the first iteration, and the resulting predicted parasitism rates (R)

were used to calculate predicted probabilities of parasitism, 7 = [exp(R)/[1 +

exp(R)]. A weighted regression then was performed with leafhopper-sample

weights calculated with N(m)(1 — tm). Predicted probabilities of parasitism from

the weighted regression were used to recalculate weights, and the process was

repeated (3 iterations) until the regression coefficient stabilized. Weights deter-

mined in the final iteration were adjusted downward (multiplied by 34/51) to

restore the unweighted error degrees of freedom. The leafhopper sample identified

above as an outlier again was excluded (standardized residual = 3.2) from the

analysis. A predicted change in the odds ratio of parasitized nymphs (+ 0.5) to

unparasitized nymphs (+ 0.5) as nymph density increased was calculated from

the regression coefficient.

RESULTS

Abundance of O. stactogalus was more linearly related to plant mass trans-

formed In(grams) (partial 7? = 0.092; F = 29.8; df = 1,285; P < 0.001) (Fig. 1)

than to plant mass in grams (partial 7 = 0.065; F = 20.5; df = 1,285; P <

0.001) and was greater (F = 9.1; df = 1,285; P = 0.003) in nymphs (634 leaf-

hoppers) than in adults (309 leafhoppers). The partial regression coefficient (0.64

+ 0.12 [SE]) for transformed plant mass therefore equaled the exponent of the

power function, leafhopper abundance = f (plant mass®?°*). Trends in abundance

2001 WIESENBORN: GONATOPUS PARASITISM OF OPSIUS 65

across transformed plant mass did not differ (F = 0.47; df = 1,284; P = 0.5)

between nymphs and adults.

The proportion of O. stactogalus nymphs parasitized (18.0%) by Gonatopus

was greater (x? = 41.6; df = 1; P < 0.001) than the proportion of adults para-

sitized (2.9%). Tamarisk branches supporting nymphs (99 of 144 total samples)

were more likely (x? = 41.4; df = 1; P < 0.001) to be exploited by Gonatopus

as nymph density increased. The odds of an ovipositing Gonatopus exploiting

versus not exploiting a tamarisk branch increased 2.3-fold (95% confidence in-

terval = 1.8 to 3.0) for a 2-fold increase, from 2 to 4 nymphs on a 100 g branch,

in nymph density. The increase in the odds of Gonatopus exploiting a branch

therefore was approximately proportional to the increase in nymph density.

On branches supporting at least one parasitized nymph (36 samples excluding

outlier), increasing nymph density was related to an increase in the number of

parasitized nymphs (F = 15.6; df = 1,34; P = 0.004) but to a decrease in the

parasitism rate of nymphs (F = 30.9; df = 1,34; P < 0.001) (Fig. 2). The odds

of a nymph being parasitized versus unparasitized decreased 38% with a 2-fold

increase, from 2 to 4 nymphs on a 100 g branch, in nymph density. Parasitism

of O. stactogalus nymphs on branches exploited by Gonatopus therefore was

inverse density dependent.

DISCUSSION

A diminishing rate of increase in O. stactogalus abundance, as the mass of the

tamarisk branch supporting the population increased, is predicted by the scaling

laws of isometric growth (Schmidt-Nielsen 1984). Tamarisk branches cut for leaf-

hopper samples were composed of a central, non-photosynthetic stem supporting

smaller, branching stems covered with sessile leaves. The central stem contained

vascular tissue transporting xylem and phloem to and from the leaves, the portion

of the branch supporting O. stactogalus. The capacity for this transport, limiting

the abundance of leaves (and leafhoppers) supported by the stem, would have

been proportional to the area of the stem’s cross-section. The mass of the stem,

however, would have been proportional to its volume. As the stem grew, its cross-

sectional area (A) would have increased more slowly than its mass (MV) according

to the equation A = M*? or A = M°°’. This function approximates the observed

relationship between O. stactogalus abundance and tamarisk-branch mass.

Greater parasitism of nymphs compared to adults, as observed in O. stacto-

galus, also has been reported in other leafhopper species. Dryinids parasitizing

eight species of leafhoppers inhabiting forage grass generally were found to attack

nymphs more than adults (Buntin 1989). In contrast, Liesner (1971) observed

higher rates of parasitism by Gonatopus in O. stactogalus adults (14—100%) than

in nymphs (5—54%). Greater parasitism of O. stactogalus nymphs compared to

adults is not due to different durations, and therefore exposure times, of the two

life stages; both first through fifth instar nymphs and adults live an average of 1

month (Harding 1930). Gonatopus parasitizing O. stactogalus preferred nymphs

over adults for hosts.

Increasing O. stactogalus nymph density was associated with increasing prob-

ability of a branch being exploited by Gonatopus and with increasing abundance

of parasitized nymphs. Gonatopus therefore appears to be attracted to, or arrested

by (Vinson 1984), populations of O. stactogalus. The increase in abundance of

66 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

parasitized nymphs, however, did not keep pace with the increase in nymph abun-

dance, resulting in a parasitism rate that decreased as nymph density increased.

Reproductive rates of parasitoids can lag behind those of their hosts due to lower

fecundities or long host-handling times (Stiling 1987).

Inverse density-dependent parasitism of O. stactogalus by Gonatopus is con-

trary to most insect host-parasite systems that have been studied (Stiling 1987).

Decreasing parasitism rate with increasing host density may be common in Dryi-

nidae, however, as it also has been observed in several dryinid species parasitizing

grassland leafhoppers (Waloff 1975). The only other insect known to parasitize

O. stactogalus in North America is the egg parasite Polynema saga (Girault)

(Hymenoptera: Mymaridae) (Lawson 1929). Harding (1930) found approximately

25% of O. stactogalus eggs in Kansas parasitized by P. saga, and Liesner (1971)

dissected P. (Barypolynema) saga from the leafhopper’s eggs in New Mexico.

Opsius stactogalus therefore supports two parasite species separately attacking

nymphs and adults or attacking eggs. In comparison, 34 of 50 cicadellid species

collected in Kansas grasslands supported two or more species of parasites infect-

ing nymphs and adults (Baldridge & Blocker 1980), and eggs of the leafhopper

Circulifer tenellus (Baker) in Iran were found to harbor seven species of parasites

(Walker et al. 1997). Although caged O. stactogalus were eaten by Chrysopidae

larvae (Harding 1930) and spiders (Liesner 1971), and leafhoppers may be eaten

by adult female Gonatopus, the contribution of predation to O. stactogalus mor-

tality in nature is unknown. Inverse density-dependent parasitism of O. stacto-

galus by Gonatopus may not only be due to intrinsic differences between the two

species, such as fecundity rate, but also to interactions across several trophic levels

(Hare 1992). The leafhopper’s abundant, predictable food source and low diversity

of natural enemies may combine to increase its reproductive rate beyond that of

its parasite.

Populations of O. stactogalus are not regulated (Turchin 1995) by the inverse

density-dependent parasitism imposed by populations of Gonatopus. Uncon-

strained population increase by O. stactogalus agrees with descriptions (Harding

1930, Hopkins & Carruth 1954, Liesner 1971) of the leafhopper occurring on

tamarisk in high densities. Liesner (1971) also suggested populations of O. stac-

togalus can be high enough to reduce tamarisk growth. Reduced growth expect-

edly would result from the photosynthate drain imposed on the plant by high

densities of the fluid-feeding leafhopper. Lack of population regulation by Gon-

atopus permits unhindered herbivory by O. stactogalus, considered beneficial

when viewed in the context of biological control against tamarisk. Populations of

O. stactogalus and herbivory by the leafhopper may be limited only by temper-

ature. Insect development and photosynthate extraction would slow during the fall

as temperatures decline and would cease when 7. ramosissima and other decid-

uous Tamarix drop their leaves and force O. stactogalus to overwinter in the egg

stage within stems.

ACKNOWLEDGMENT

I am grateful to M. Olmi of the Universita degli Studi della Tuscia, Viterbo,

Italy, for providing dryinid collection records and for identifying a specimen and

to J. D. Hare of the Department of Entomology at UC Riverside for critiquing

this manuscript. Appreciation is extended to the City of Boulder City, Nevada,

2001 WIESENBORN: GONATOPUS PARASITISM OF OPSIUS 67

for allowing the study on their lands and providing wastewater treatment plant

discharge quantities.

LITERATURE CITED

Agresti, A. 1990. Categorical data analysis. John Wiley & Sons, New York.

Baldridge, R. S. & H. D. Blocker. 1980. Parasites of leafhoppers (Homoptera: Cicadellidae) from

Kansas grasslands. J. Kansas Entomol. Soc., 53: 441-447.

Baum, B. R. 1967. Introduced and naturalized tamarisks in the United States and Canada (Tamari-

caceae). Baileya, 15: 19-25.

Brock, J. H. 1994. Tamarix spp. (salt cedar), an invasive exotic woody plant in arid and semi-arid

riparian habitats of western USA. Chapter 4. pp. 27-44. In deWaal, L. C., L. E. Child, P. M.

Wade, & J. H. Brock (eds.). 1994. Ecology and management of invasive riverside plants. John

Wiley & Sons, New York.

Buntin, G. D. 1989. Dryinid (Hymenoptera: Dryinidae) parasites of leafhoppers and planthoppers

(Homoptera) in forage-type bermudagrass. J. Kansas Entomol. Soc., 62: 602-606.

Di Tomaso, J. M. 1998. Impact, biology, and ecology of saltcedar (Tamarix spp.) in the southwestern

United States. Weed Technology, 12: 326-336.

Fornasari, L. 1997. Host specificity of Coniatus tamarisci (Coleoptera: Curculionidae) from France:

potential biological control agent of Tamarix spp. in the United States. Environ. Entomol., 26:

349-356.

Guglielmino, A. & M. Olmi. 1997. A host-parasite catalog of world Dryinidae (Hymenoptera: Chry-

sidoidea). Cont. Entomol. Int., 2: 165-298.

Harding, L. 1930. The biology of Opsius stactogalus Fieber (Homoptera, Cicadellidae). J. Kansas

Entomol. Soc., 3: 7-22.

Hare, J. D. 1992. Effects of plant variation on herbivore-natural enemy interactions. Chapter 12. pp.

278-298. In Fritz, R. S. & E. L. Simms (eds.). 1992. Plant resistance to herbivores and path-

ogens: ecology, evolution, and genetics. Univ. Chicago Press, Chicago, Illinois.

Hopkins, L. & L. A. Carruth. 1954. Insects associated with salt cedar in southern Arizona. J. Econ.

Entomol., 47: 1126-1129.

Lawson, P. B. 1929. A leafhopper parasite—Polynema saga (Girault) (Hymenoptera, Mymaridae).

Ann. Entomol. Soc. Am., 22: 130.

Liesner, D. R. 1971. Phytophagous insects of Tamarix spp. in New Mexico. M.S. Thesis, New Mexico

State University, Las Cruces.

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(4th ed.). McGraw-Hill, Boston, Massachusetts.

Olmi, M. 1984. A revision of the Dryinidae (Hymenoptera). Mem. Amer. Entomol. Inst., no. 37.

Robinson, T. W. 1965. Introduction, spread, and areal extent of saltcedar (Tamarix) in the western

States. U.S. Dept. Int., Geol. Surv. Professional Paper, 491-A.

Schmidt-Nielsen, K. 1984. Scaling: why is animal size so important? Cambridge University Press,

Cambridge.

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68: 844-856.

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Cappuccino, N. & P. W. Price (eds.). 1995. Population dynamics: new approaches and synthesis.

Academic Press, New York.

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(eds.). 1984. Chemical ecology of insects. Chapman & Hall, London.

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Homoptera) of acidic grassland. Trans. R. Entomol. Soc. London, 126: 637-686.

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Jepson manual: higher plants of California. Univ. Calif. Press, Berkeley.

Received 10 Mar 2000; Accepted 26 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(2): 68-70, (2001)

TAENIOGONALOS RAYMENTI CARMEAN & KIMSEY

(HYMENOPTERA: TRIGONALIDAE) REARED AS A

HYPERPARASITE OF STURMIA CONVERGENS

(WEIDEMANN) (DIPTERA: TACHINIDAE), A PRIMARY

PARASITE OF DANAUS PLEXIPPUS

(L.) LEPIDOPTERA: NYMPHALIDAE)

ANTHONY R. CLARKE! AND MyYRON P. ZALUCKI

Department of Zoology and Entomology, The University of Queensland,

Queensland, 4079, Australia

Abstract.—Taeniogonalos raymenti is confirmed as a hyperparasitoid of the tachinid Sturmia

convergens which parasitises larval Danaus plexippus. Trigonalids are indirect parasitoids and

in this case we have direct evidence that wasp eggs must have been laid on the caterpillar’s host

plant, Asclepias fruticosa, before the secondary host, but not necessarily before the primary

tachinid host, was present. Levels of hyperparasitism during our sampling period were very low

at less than two percent.

Key Words.—Insecta, Sturmia, Tachinidae, Asclepias.

Trigonalid wasps are unique among the parasitic Hymenoptera as they oviposit

onto foliage and the eggs are subsequently injested, along with the foliage, by

larval sawflies or herbivorous caterpillars. The host relationships of larval trigon-

alids are poorly known, but most appear to be either primary parasitoids of larval

sawflies, or hyperparasitoids of hymenopteran and tachinid parasitoids (Weinstein

& Austin 1991). The family has a cosmopolitan distribution, but is quite small

with only 100 known species. The biology, host relationships and systematics of

trigonalids are reviewed by Weinstein & Austin (1991) and Carmean & Kimsey

(1998). One of the major messages in both those reviews is that trigonalids remain

a very poorly known group, with little or no biological information for most

species. This note supplies a new host record for a trigonalid species from South-

east Queensland.

MATERIALS AND METHODS

Collection and Rearing.—As part of wider study, 222 wild, 5th instar Danaus

plexippus (L.) caterpillars were collected from four sites in South-east Queensland

from February to November 1998. Collections were made at Mt Crosby (27°32!

S, 152°49’ E), The University of Queensland Veterinary Research Farm Pinjarra

Hills (= Moggill Farm) (27°35' S, 152°53' E), Dunn’s Creek via Beenleigh

(27°43' S, 153°09' E) and Pine River via Boondal (27°17' S, 152°55’ E).

Caterpillars were collected from either Ascle pias fruticosa L. or A. curassavica

L., returned to the laboratory and reared individually on cut foliage of the host

species they were collected on until they pupated. A further 33 caterpillars were

obtained by placing D. plexippus eggs onto A. fruticosa plants, which did not

carry existing D. plexippus eggs or larvae, at Moggill Farm and then immediately

‘Current Address: Australian School of Environmental Studies, Griffith University Nathan Campus,

Qld, 4111.

2001 CLARKE & ZALUCKI: TAENJIOGONALOS REARED 69

Table 1. Fate of fifth instar Danaus plexippus (L.) larvae collected from sites in South-east Queens-

land. (P.1. = Paradrino laevicula Mesnil, S.c. = Sturmia convergens (Weidemann), W.s. = Winthemia

sumatrana (Townsend).

No. of adult No. of

No. of larvae No. of larvae tachinids emerging trigonalids

Collection site Date collected parasitised (P.1./S.c./W.s.) reared

Wild collections

Mt Crosby Feb 98 21 18 31 (25/1/5) 0

Dunn’s Ck Apr 98 31 18 25 (21/4/0) 0

Pine River May 98 2. 12 19 (16/3/0) 0

Pine River June 98 6 3 0 0

Dunn’s Ck June 98 22 19 36 (32/4/0) 0

Dunn’s Ck June 98 Pal 15 21 (21/0/0) 0

Dunn’s Ck June 98 40 20 24 (22/1/1) 0

Dunn’s Ck Nov 98 23 22 30 (17/13/0) 0

Moggill Farm Nov 98 21 19 40 (10/26/4) 1

Mt Crosby Nov 98 10 5 11 (6/5/0) 0

“Bag’’ reared

Moggill Farm Nov 98 33 33 48 (0/48/0) 6

bagging the plants with insect proof gauze. Caterpillars were harvested as 5th

instars and then reared as above.

Identification and Voucher Material.—Trigonalids were identified by J. C. Car-

dale, Australian National Insect Collection, CSIRO Entomology, Canberra.

Voucher material has been deposited with the University of Queensland Insect

Collection. Tachinids were identified by B. Cantrell, Queensland Department of

Primary Industries, Brisbane. Two species of tachinid were routinely reared from

monarch butterfly caterpillars during our research. We were able to determine

which tachinid was the trigonalid host from the remnant pupal cases, which are

noticeably different between the two tachinid species.

RESULTS

Danaus plexippus caterpillars were found to be heavily parasitised by tachinids,

most commonly Sturmia convergens (Weidemann) and Paradrino laevicula Mes-

nil. Both have been previously recorded at primary parasites of D. plexippus in

South-east Queensland (Zalucki 1981). As we will be reporting on primary par-

asitism elsewhere, we will not here discuss it further.

Seven trigonalid wasps emerged from seven tachinid pupal cases which had

emerged from seven individual caterpillars. These were identified as individuals

of Taeniogonalos raymenti Carmean & Kimsey (previously T. tricolor Rayment,

see Carmean & Kimsey 1998). All but one of these specimens were reared from

a single collection of caterpillars made from plants which had been bagged im-

mediately after placing butterfly eggs upon them (Table 1). In all cases the tachinid

host, based on pupal case identification, was S. convergens. Because plants had

been bagged immediately following artificial butterfly egg placement, it confirms

that both S. convergens and T. tricolor placed their eggs on the host plant (A.

fmtticosa) prior to the presence of the primary host insect. The identity of the

tachinid from which the seventh wasp was reared was not recorded, but the host

plant was again A. fruticosa.

70 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Overall hyperparasitism rate was only 1.8% (n = 397 tachinids reared), but for

the case where 6 wasps were reared from a single collection, the hyperparasitism

rate was 13% (n = 46 tachinids reared).

DISCUSSION

Taeniogonalos is the most widely distributed genus of the Trigonalidae and

Species in the genus have been reared previously as primary parasitoids of saw-

flies, or a hyperparasitoids of tachinid and ichneumonid parasitoids of lepidoptera

(Carmean & Kimsey 1998). This is the first rearing and oviposition substrate

record for JT. raymenti, but it is not the first time the genus Sturmia or the sub-

family Danainae have been associated as primary and secondary hosts of this

genus. Hirai & Ishi (1995) recorded T. fasciata (Strand) (previously Poecilogon-

alos fasciata) as hyperparasitising Sturmia bella (Meigen), a parasite of Parantia

sita (Moore) (the tiger chestnut butterfly), in Japan.

ACKNOWLEDGMENT

This work was undertaken while Anthony Clarke was in receipt of a University

of Queensland New Staff Grant and an Australian Research Council Large Grant.

Dr. Jan Green and Ms. Michelle Tung assisted in collecting and rearing of the

specimens. We thank Ms. Jo Cardale and Dr. Bryan Cantrell for the identifications.

LITERATURE CITED

Carmean, D. & L. Kimsey 1998. Phylogenetic revision of the parasitoid wasp family Trigonalidae

(Hymenoptera). System. Entomol., 23: 35-76.

Hirai, N. & M. Ishi 1995. Host relationships of the tachinid fly, Sturmia bella (Diptera: Tachinidae),

and the trigonalid wasp, Poecilogonalos fasciata (Hymenoptera: Trigonalyidae), parasitizing the

Chestnut Tiger butterfly, Parantica sita (Lepidoptera: Danaidae). Appl. Entomol. Zool., 30:

241-244.

Weinstein, P. & A. D. Austin 1991. The host relationships of trigonalyid wasps (Hymenoptera: Tri-

gonalyidae), with a review of their biology and catalogue to world species. J. Nat. Hist., 25:

399-433.

Zalucki, M. P. 1981. Temporal and spatial variation of mortality in Danaus plexippus L. Aust. Entomol.

Mag., 8: 3-8.

Received 20 Mar 2000; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(2): 71-78, (2001)

NEW SPECIES OF CERACLEA

(TRICHOPTERA: LEPTOCERIDAE: ATHRIPSODINI) AND

A FIRST RECORD OF ADICELLA

(TRICHOPTERA: LEPTOCERIDAE: TRIAENODINI)

FROM FAR EASTERN RUSSIA

TATYANA I. AREFINA! AND JOHN C. MORSE?

‘Institute of Biology and Soil Sciences, Vladivostok, Russia

Department of Entomology, Clemson University,

Clemson, South Carolina 29634-0365, U.S.A.

Abstract.—Three new species of the genus Ceraclea, belonging to the subgenus Athripsodina:

C. (A.) affinis, C. (A.) bilobulata and C. (A.) breviramosa from the southern Far East of Russia

are described and illustrated. The genus Adicella, represented by a new species, is newly recorded

for the Russian fauna.

Key Words.—Trichoptera, Leptoceridae, Ceraclea, Adicella, new species, new record, Far East-

ern Russia.

Until now, 9 genera and 59 species of the family Leptoceridae are known from

the Russian Far East (Arefina 1997; Vshivkova et al. 1997; J. C. Morse, L. Yang,

and I. M. Levanidova, unpublished data). During a survey of the caddisflies (Tri-

choptera) of the Ussuri River Basin, three Ceraclea species were collected that

are new to science. Furthermore, a representative of the genus Adicella, which

has not been previously known in Russia, was found.

The types of the new species are preserved in alcohol and held in the collections

of the Institute of Biology and Soil Sciences, Russian Academy of Sciences,

Vladivostok. In the present work, the terminology generally follows that in the

revision by Yang & Morse (1988).

CERACLEA (ATHRIPSODINA) DILUTA Group

Ceraclea (Athripsodina) affinis, NEW SPECIES

(Figs. 1-7)

Types.—Holotype male: KHABAROVSK TERRITORY (RUSSIA), Birushka

River, Ussuri River Basin, 23 Jul 1996, T. Arefina. Paratype: 1 female, same data

as holotype.

Description—Length of forewing: male—7.9 mm, female—8.1 mm. Head and body with white

and brown hairs mixed. Vertex of head, thorax and coxae brown, abdomen and legs lighter. Wings

yellowish brown with setae darker.

Male Genitalia (Figs. 1-4).—Tergum IX (IX) with pair of small papillae near poster meson. Preanal

appendages (pre. app) subtriangular, fused basally, each with acute apex. Tergum X (X) composed of

triangular median lobe and pair of lateral lobes, each with blunt apex in dorsal and lateral views,

lateral lobes extending slightly beyond median lobe. Main body of each inferior appendage (inf. app)

Straight in lateral view with subapicodorsal lobe (sap. do) bent caudad; basoventral lobe (bv) of each

inferior appendage nearly half as long as main body, obtusely angled about 110° from main body,

stout at basal half then abruptly tapering to digitate apex; harpago (har) tiny; mesal ridge of inferior

appendage (mes.rdg) produced in small, nearly trapezoidal process located slightly lower than harpago.

Phallus strongly curved ventrad and constricted ventrally two-thirds distance from base; both paramere

spines (par) seta-like with right spine positioned near apex, left spine almost straight and situated in

anterior half of phallus.

72 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

2

Figures 1-4. Male genitalia of Ceraclea (A.) affinis NEW SPECIES in lateral (1) and dorsal (2)

views; left inferior appendage (3) in caudal view; phallus (4) in lateral view. Abbreviations: bv =

basoventral lobe of an inferior appendage; har = harpago; infapp = inferior appendage; mes.rdg =

mesal ridge of an inferior appendage; par = paramere spines; pre.app = preanal appendage; sap.do

= subapicodorsal lobe of an inferior appendage; IX, X = abdominal segments IX and X.

Female Genitalia (Figs. 5—7).—Tergum IX rounded apically, with pair of small subdorsal processes.

Preanal appendages (pre.app) short, as long as broad, covered with short setae. Lamellae (lam) parallel-

sided in dorsal and ventral views, semicircular in lateral view, setose ventrally. Each gonopod plate

(go.pl; e.gon.IX of Nielsen 1980) with triangular caudal projection, plates approximate for most of

their length, with triangular excisions between their caudal projections; gonopod plates slightly con-

cave in middle with prominent mesal and lateral edges. Spermathecal sclerite (sp.sc) long, V-shaped,

with lateral arms curved mesad near caudal end. Pair of broad sclerotized plates (sc.pl) suspended

above spermathecal sclerite and extending cephalad nearly to two-thirds length of segment VUI.

Immature Stages.—Unknown.

Diagnosis.—Ceraclea affinis belongs to the C. (A.) diluta Group, which pres-

ently includes C. diluta (Hagen), C. perplexa (McLachlan) and C. trilobulata

Morse, Yang, and Levanidova. Male genitalia of the new species most closely

resemble those of C. trilobulata in the trilobate tergum X, in the shape of its

inferior appendages, and in the presence of a ventral constriction of the phallus.

It differs from C. trilobulata in the much longer base of the phallus, with the

position of the ventral constriction nearly two-thirds of the distance from the based

of the phallus; in the apparently longer middle lobe of tergum X; and in the

thicker base of each inferior appendage. Because the females of C. perplexa and

C. trilobulata are unknown, it is not possible to separate them from the female

2001 AREFINA & MORSE: NEW RUSSIAN TRICHOPTERA 73

pre.ap

7

(ox

Figures 5—7. Female genitalia of Ceraclea (A.) affinis NEW SPECIES (holotype) in lateral (5),

dorsal (6) and ventral (7) views. Abbreviations: go.p! = gonopod plate of gonopod IX; lam = lamella;

pre.app = preanal appendage; sc.pl = sclerotized plate of spermathecal sclerite; sp.sc = spermathecal

sclerite.

of the new species; however, the apicolateral lamellae of C. affinis are shorter

than those of C. diluta.

Distribution.—Known only from the type locality in Khabarovsk Territory

(Russia).

Etymology.—Latin, “‘similar,’’ with reference to the resemblance of the new

species with C. trilobulata.

CERACLEA (ATHRIPSODINA) ANNULICORNIS Group

Ceraclea (Athripsodina) bilobulata, NEW SPECIES

(Figs. 8-15)

Types.—Holotype male) KHABAROVSK TERRITORY (RUSSIA). Ussuri

River near Zabaikalskoe Village, 23 Jul 1996, T. Arefina. Paratypes: 7 males,

same data as holotype.

Description—Length of forewing: male—5.5—6.2 mm, female—5.3-5.9 mm. Head with mostly

74 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Figures 8-11. Male genitalia of Ceraclea (A.) bilobulata NEW SPECIES (paratype) in lateral (8)

and dorsal (9) views; left inferior appendage (10) in caudal view; phallus (11) in lateral view.

white hairs. Vertex of head and thorax yellow-brown, abdomen lighter, nearly white ventrally. Wings

straw yellow with yellow-brown setae.

Male Genitalia (Figs. 8—15)—Tergum IX protruding at apical center. Preanal appendages very short

and broad in dorsal view, separated basally, slightly shorter then tergum IX to slightly longer. Tergum

X extending far beyond preanal appendages, broad basally, divided into two lobes nearly to base, each

lobe with rounded apex; dorsal edge of tergum straight in lateral view, ventral edge of tergum upturned

from middle, apex rounded. Main body of each inferior appendage slender, both harpago and subap-

icodorsal lobe short; harpago nearly as long as semimembranous subapicodorsal lobe to slightly longer;

basoventral lobe of each inferior appendage vestigial, very short, with single spine directed somewhat

mesad; mesal ridge without project and bearing only one normal seta; main body with variable cau-

dolateral lobe bearing two large setae. Phallus curved ventrad, with sclerotization extended full length

ventrally; anterior portion shorter and broader than posterior portion; two paramere spines present, left

paramere spine about same size as right, but retracted somewhat more cephalad.

Remarks.—Different individuals of the species vary in the shape of the preanal

and inferior appendages and in size and location of the paramere spines of the

phallus. In spite of the variability of these characters, the available specimens

generally agree with the foregoing description.

Female and Immature Stages.—Unknown.

Diagnosis.—The male of this species resembles those of C. sibirica (Ulmer)

and C. hastata (Botosaneanu) in the presence of a spine at the basoventral position

of the inferior appendages; in the mesal ridge of each inferior appendage without

2001 AREFINA & MORSE: NEW RUSSIAN TRICHOPTERA 75

14 15

Figures 12-15. Male genitalia of Ceraclea (A.) bilobulata NEW SPECIES (paratype) in lateral

(12) and dorsal (13) views; left inferior appendage (14) in caudal view; phallus (15) in lateral views.

any projection; in tergum X lacking lateral processes; and in the sclerotization

extending the full length of the phallus ventrally. However, tergum X is divided

into two lobes in the new species whereas C. sibirica and C. hastata have no

excision on tergum X. Ceraclea bilobulata differs from both C. sibirica and C.

hastata and other known males of the C. (A.) annulicornis Group in the short and

basally separated preanal appendages, in the very short basoventral lobe of each

inferior appendage, in the mesal ridge of the inferior appendages having only one

seta, and in possessing two phallic paramere spines nearly equal in length.

Distribution.—Known only from the type locality in Khabarovsk Territory

(Russia).

Etymology.—Latin, “‘two-lobed,”’ with reference to the shape of tergum X in

the male genitalia.

CERACLEA (ATHRIPSODINA) RIPARIA Group

Ceraclea (Athripsodina) breviramosa, NEW SPECIES

(Figs. 16—23)

Types.—Holotype male: KHABAROVSK TERRITORY (RUSSIA). Ussuri

River near Zabaikalskoe Village, 23 Jul 1996, T. Arefina. Paratypes: 24 females,

same data as holotype; Khabarovsk Territory (Russia), Kiya River, Ussuri River

Basin, Ekaterinoslavka Village vicinity, 26 Jul 1996, T. Arefina, 4 females; Pri-

76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Figures 16-20. Male genitalia of Ceraclea (A.) breviramosa NEW SPECIES (holotype) in lateral

(16) and dorsal (17) views; left inferior appendage (18) in caudal view; harpago and subapicodorsal

lobe of left inferior appendage (19) in caudal view; phallus (20) in lateral view.

morye Region (Russia), Kabarga River, Ussuri River Basin, 28 Jul 1998, T. Tiu-

nova, 2 females.

Description—Length of forewing: male—5.8 mm, female—5.5—6.0 mm. Head with white hairs on

middle of front and vertex, side hairs brownish. Head and thorax yellow-brown, abdomen whitish,

abdominal dorsum pale brown. Wings straw yellow with yellow-brown setae.

Male Genitalia (Figs. 16—20).—Tergum IX protruding at apical center. Preanal appendages nearly

as long as broad, separated basally, about as long as tergum IX, obliquely truncate apically. Tergum

X about 2.5X as long as preanal appendages, broad at base, tapering to blunt apex, with V-shaped

cleft apically one-eighth of its length; tergum X with pair of smaller papillae near middle of tergum.

Basoventral lobe of each inferior appendage shorter than main body of appendage, directed somewhat

mesad, bearing single spine subbasally; triangular projection above basoventral lobe, nearly as long

as spine in lateral view. Main body of appendage slender; harpago slightly longer than subapicodorsal

lobe, each harpago setose, with two small spines at apex; each subapicodorsal lobe with several long

setae along dorsal surface and membranous apex bearing two setae. Mesal ridge of each inferior

appendage with two normal setae. Phallus strongly curved ventrad about half distance from base, with

sclerotization extended full length ventrally; anterior portion of phallobase slightly broader and longer

than posterior portion in lateral view. Paramere spines aligned, with apex of left spine slightly inserted

in elliptical opening of base of right spine, as typical for Group; left spine with short and straight

2001 AREFINA & MORSE: NEW RUSSIAN TRICHOPTERA 77

i

eosSree§

eye

eee SS

-as

Figures 21-23. Female genitalia of Ceraclea (A.) breviramosa NEW SPECIES (paratype from

Ussuri River near Zabaikalskoe Village) in lateral (21), dorsal (22) and ventral (23) views. Abbrevi-

ation: d.rdg = diagonal ridge of a gonopod plate.

dorsomedial projection; apical half of left spine sinuous, with upturned tip; right spine straight, curved

ventrad at three-quarters distance from base, with tip turned slightly outwards, to right.

Female Genitalia (Figs. 21—23).—Preanal appendages broad but very short, as typical for Group.

Lamellae setose ventrally, each with broadly rounded apex in lateral view. Gonopod plates widely

separated anteriorly, not concave, each with well-sclerotized diagonal ridge (d. rdg); each plate with

two caudal projections: mesal projection narrow and elongate, lateral projection rather large, oval in

lateral and ventral views. Anterior part of the spermathecal sclerite narrow, broadens in middle, convex

laterally in posterior part.

Immature Stages.—Unknown.

Diagnosis.—The male genitalia of this species resemble those of most species

of the C. (A.) riparia Group in possessing two phallic paramere spines which are

aligned, with the apex of the left spine inserted in an opening of the base of the

right spine, but in this new species the left spine has a dorsomedial projection

and an upturned tip. This is the only species with the C. (A.) riparia Group whose

male: (1) lacks a pair slender lateral processes of tergum X; (2) has the inferior

appendages each with such a short basoventral lobe; and (3) has the mesal ridge

78 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

of each inferior appendage bearing only two setae. The female of this species

resembles those of C. riparia (Albarda), C. yangi (Mosely) and C. nankingensis

(Hwang) in the shape of the caudal projections of the gonopod plates and of the

spermathecal sclerite, but it differs from all known species of the C. (A.) riparia

Group in possessing a diagonal ridge of each gonopod plate.

Distribution.—Known only from the type localities in Khabarovsk Territory

and the Primorye Region (Russia).

Etymology.—Latin, “‘short-branched,”’ with reference to the short basoventral

lobe of each inferior appendage in the male genitalia.

NEw RECORD

Adicella sp. n. (Yang and Morse, unpublished data)

Material Examined. —KHABAROVSK TERRITORY (RUSSIA). Kiya River, Ussuri River Basin,

Yekaterinoslavka Village vicinity, 26 Jul 1996, T. Arefina, 2 males.

Distribution.—Khabarovsk Territory (Russia), China.

ACKNOWLEDGMENT

The work was supported by the Russian Fund of Fundamental Investigations,

grant 96-04-50388.

LITERATURE CITED

Arefina, T. I. 1997. Three new to the Russia species of the caddisflies (Trichoptera: Leptoceridae). Far

Eastern Entomol., 47: 18.

Nielsen, A. 1980. A comparative study of the genital segments and genital chamber in the female

Trichoptera. Det Kongelige Dansk Videnskabernes Selskab Biologiske Skrifter, 23: 1-200.

Vshivkova, T. S., J. C. Morse & L. Yang. 1997. Fam. Leptoceridae. pp. 154—202. In Lehr, P. A. (ed.).

Key to the insects of Russian Far East V. Trichoptera and Lepidoptera Pt. 1, Vladivostok.

Yang, L. & J. C. Morse. 1988. Ceraclea of the People’s Republic of China (Trichoptera: Leptoceridae).

Contr. Amer. Entomol. Inst., 23: 1-69.

Received 3 Aug 1999; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

71(2): 79-89, (2001)

A FURTHER CONTRIBUTION TO THE AUSTRALIAN

SYSTEMATICS OF THE TRIBE COLPURINI

(HEMIPTERA: HETEROPTERA: COREIDAE: COREINAE)

HARRY BRAILOVSKY

Departamento de Zoologia, Instituto de Biologia,

Universidad Nacional Aut6noma de México, Apdo Postal 70153,

México D.EF, 04510, México

Abstract.—One new genus (Weirhygia) and four new species (Weirhygia faceta, Grosshygia

formosa, Grosshygia lepida, and Grosshygia pisina) from Australia are described in the tribe

Colpurini (Coreidae). A revised key to all known genera and species is included. Dorsal habitus

illustrations and drawings of male genital capsule and female genital plates are provided.

Key Words.—Insecta, Hemiptera, Heteroptera, Coreidae, Colpurini, new genus, new species,

Australia.

The Australian species of the tribe Colpurini have recently been reviewed by

Brailovsky (1993), Brailovsky & Barrera (1996) and Steinbauer & Clarke (1996).

During a visit to Queensland Museum, Brisbane, Australia, I found, a series of

specimens of Colpurini that I first thought to be a previous known species of

Grosshygia Brailovsky and Grosshygioides Brailovsky. A preliminary check of

the head, pronotum, development of the hemelytra, general shape of the male

genital capsule and abdominal sternite VII of the female revealed a new genus,

and species and three new species of Grosshygia.

This additional information is made possible largely by the extensive collec-

tions of G. B. Monteith and his colleagues.

One of the most striking developments in australian colpurini systematics has

been the discovery of an extensive fauna living on the ground of tropical rainforest

at high and low elevations of northern and northeastern Queensland. They live

associated with piles of freshly dead leaves from recently fallen trees.

REVISED KEY TO AUSTRALIAN COLPURINI

1. Each side of head immediately in front of eye with long pointed spine

Ochs Ae ete een ne aoa Sint, nha A ss Ae eS NE Pachycolpura manca Breddin

1’. Sides of head in front of eyes unarmed

2. Tylus projecting as single, large, acute spine

2'< Tylus globose, truncated or ‘Difid’ 2.5 5 2c. sate pe wnecace ee eres, 25) wae eed 5

3. Femora unarmed; callar region of pronotum conspicuously convex; pron-

otal disc behind midline with transverse wrinkle; male genital capsule

with small median projection ........... Acanthotyla fasciata (Walker)

3’. Femora strongly armed with long, sharp ventral spines; callar region of

pronotum weakly convex to flat; pronotal disc without transverse wrin-

kle; posteroventral edge of male genital capsule without median pro-

ACCUIOM Ve YR, hee gen ad gt Eun eg Oene fore LD Snes 8 ee 4

4. Mandibular plate armed with short projection; posteroventral edge of male

genital capsule convex, obtusely rounded ... Agathyrna praecellens Stal

80

4’.

125

Pass

U3:

13%:

14.

14’.

$5:

share

16.

16’.

THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Mandibular plate unarmed; posteroventral edge of male genital capsule

elongate and bifurcate ..... Woodwardhysgia bifida Brailovsky (in part)

Antenniférous*tubercles armed” 5) 0520. h ake: oar needeees Sa peo ae ed 6

Antenniferous tubercles unarmed ................0 0c eee eee ees 14

Bucculae armed with obvious spine near middle third ............... 7

2 Bucculaeniitormiy-roundeg: (022s Oe ee vente ee ee 8

Rostrum reaching middle third of abdominal sternite V; body robust,

longer than 10.00 mm; frontal angles of pronotum conspicuously

prominent; scutellum as wide or wider than long .................

en rr ee ehh, aera ea a ore ra ie Acantholybas kirkaldyi Bergroth

. Rostrum reaching posterior margin of abdominal sternite IV; body shorter

than 9.00 mm; frontal angles of pronotum scarcely exposed; scutellum

clearly longer than wide ............. Acantholybas brunneus Breddin

Micropterous, hemelytral membrane reduced to small flap; ocelli incon-

SPLCM OUST aly ye ee cote ee Cra eee te ea ewer a pee ele oe BUN eee ne an 2

. Macropterous to submacropterous, hemelytral membrane well developed;

ocelli clearly developed ....... Pachycolpuroides monteithi Brailovsky

Head dorsally flat; abdominal sternite VII of female without plica or

fissura ... Weirhygia faceta Brailovsky NEW GENUS NEW SPECIES

. Head dorsally convex; abdominal sternite VII of female with plica and

TUSSUGE Ae nase = Eke p iri ONS | ACE Rhea de Se aligt leche Foye Wc eects cade nese ne Maa 10

Apex of scutellum globose ............... Grosshygia nigra Brailovsky

= Apex Of seutelim SUDSCULED 22.0.8 ned lisiicdes sere) cwceeede 2 arena ee ates 11

Antenniferous tubercles each with external lobe arcuate, recurved, con-

verging anteriorly, almost touching the basal joint of antennal segment

De ese etch ge ea ee te ete Grosshygia lepida Brailovsky NEW SPECIES

. Antenniferous tubercles each with external lobe obliquely projecting, di-

VETSTI Seal LOLI@TU Ys ala ah Te Be ek leer the 5 btn he iden Seether aes too HARE 12

Head dorsally with vertex uniformly convex, without conical lobes; he-

melytral-menm Grane a WSent, oes A... ches ts eveyeet tei sa fer eee Ae Sori

gc pect chap yt Nees c eeu RR Grosshygia pisina Brailovsky NEW SPECIES

Head dorsally with a transverse depression separated into two conical

elevations; hemelytral membrane reduced to small flaps ........... 13

Antenniferous tubercles each armed with long lobe; antennal segment II

longer than 1.96 mm ................ Grosshygia lobatula Brailovsky

Antenniferous tubercles each armed with short lobe; antennal segment II

shorter than 1.73 mm .............. Grosshygia monticeps Brailovsky

Head dorsally convex; apex of scutellum globose ...................

BUSINES ys tl A oe ee tre Grosshygia formosa Brailovsky NEW SPECIES

Head dorsally flat; apex of scutellum subacute ..................... 15

Abdominal sternite VII of female without plica or fissura; frontal angles

of pronotum rounded, blunt, not produced ....................... 16

Abdominal sternite VII of female with plica and fissura; frontal angles

of pronotum produced forward as conical teeth .................. 20

External edge of gonocoxa I in lateral view with upper half conspicu-

ously exposed and lower half projected in a medium-sized convex lobe

2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 81

17. Posteroventral edge of male genital capsule with small V-shaped con-

cavity, laterally enclosed by two shorter arms; gonocoxa I in caudal

WITS Wi EC LOSE tee pbsanete nit Anke cop cc epee as Sciophyroides sortita (Horvath)

17’. Posteroventral edge of male genital capsule with U-shaped concavity,

enclosed by two lateral medium-sized robust arms; gonocoxa I in cau-

dal-view-opened. <9 in ee.. 2 eee, Scio phyroides sulcicrus (Breddin)

18. Body length longer than 10.30 mm; posteroventral edge of male genital

capsule with large U-shaped concavity, laterally enclosed by two

strong divergent arms ............ Sciophyrella australica (Brailovsky)

18’. Body length shorter than 10.10 mm; posteroventral edge of male genital

CASH le SHOLsAS TAB OVER. sacra ee AOR ay RR I cal tk Seger 19

19. Posteroventral edge of male genital capsule produced into medium-sized

and robust lateral projections, enclosing deep U-shaped concavity;

gonocoxa I in lateral view narrow .... Sciophyrella diminuta (Horvath)

19’. Posteroventral edge of male genital capsule with small V-concavity en-

closed by two short globose arms; gonocoxa I in lateral view enlarged

oP at bp hl dere, oe A Sciophyrella minuscula Brailovsky & Barrera

20. Mandibular plate armed with large prominent tubercle; pronotal disc with

deep longitudinal furrow along midline; hemelytra truncate, membrane

PLO s| ee eRe, Bienes Sota Grosshygioides mandibularis Brailovsky

20'. Mandibular plate unarmed; pronotal disc flat, without midline furrow;

macropterous; hemelytral membrane well developed .............. Zz

21. Femora armed with two rows of ventral spines: tylus apically bifid

Rete ee Seon yAr cae ratte Oe Woodwardhygia bifida Brailovsky (in part)

21’. Femora unarmed; tylus apically globose ..................0000000e-

et ie Oa Lo rere ae Hygia (Australocolpura) sandaracine Brailovsky

WEIRHYGIA BRAILOVSKY, NEW GENUS

Type species.—Weirhygia faceta Brailovsky, NEW SPECIES.

Description—Head longer than wide (across eyes), pentagonal, nondeclivent, and dorsally flat; tylus

unarmed, apically globose, extending anteriorly to and laterally higher than juga; juga unarmed, thick-

ened, apically rounded, shorter than tylus; antenniferous tubercle armed with raised lobe, diverging

anteriorly and rounded to quadrate apically; sides of head in front of eyes unarmed; genae and man-

dibular plates unarmed; antennal segment I robust, thickest, slightly curved outward; segments II and

Ill cylindrical, and slender; segment IV fusiform; antennal segment II longest, I shortest and III longer

than IV; ocelli not raised; preocellar pit obliquely deep; eyes small, globular; postocular tubercle

protuberant, globose; buccula rectangular, short, elevated, not projecting beyond antenniferous tubercle,

without teeth; rostrum long, reaching posterior margin of abdominal sternite VI or anterior margin of

VII; rostral segment IV longest, I shortest, and II longer than II. Thorax. Pronotum: Wider than long,

trapeziform, non declivent, and bilobate; anterior lobe longer than posterior lobe, each with antero-

lateral borders barely convex; collar wide; frontal angles projecting forward as conical teeth; humeral

angles rounded, projected into rounded lobes, elevated, directed outward, higher than posterior pronotal

disc; posterolateral and posterior borders almost straight; callar region weakly convex, separated me-

dially by deep longitudinal furrow, which extends to posterior margin. Anterior lobe of metathoracic

peritreme elevated, reniform, posterior lobe sharp, small. Legs: Unarmed; tibiae cylindrical, with lon-

gitudinal sulcus indistinct. Scutellum: Triangular, flat, wider than long, with apex subacute. Hemelytra:

Coleopteroid; clavus and corium fused into coriaceous pad, wings meeting each other along midline;

hemelytral membrane reduced to small flap, reaching middle third of abdominal tergite III, thus leaving

the abdominal terga exposed. Abdomen: Connexival segments practically at same level as abdominal

segments; posterior angle of connexivum extending into short and robust spine; abdominal sterna with

82 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

medial furrow extending to posterior border of sternite VI. Integument: Body surface dull; ventral

face of head, posterior lobe of pronotum, scutellum, clavus, corium, propleura, mesopleura and me-

tapleura, acetabula and abdomen with scattered punctures, each puncture with short decumbent silvery

bristle-like hair, intermixed with long erect bristles on antennal segments, legs, and abdominal sterna;

dorsal surface of head, anterior lobe of pronotum, and connexival segments impunctate.

Male Genitalia—Genital capsule. Posteroventral edge simple, transversely barely arcuate, with

shallow notch at midline (Fig. 3).

Female Genitalia.—Abdominal sternite VII without plica and fissura; gonocoxae I enlarged antero-

posteriorly, in lateral view with external face obliquely straight, in caudal view open; paratergite VIII

short, almost square, with spiracle visible; paratergite LX larged than paratergite VIII (Fig. 4).

Diagnosis.—The reduction of wings, the prominent postocular tubercle, the

pronotal disk with deep midline furrow, and tylus apically globose might suggest

a relationship with Grosshygioides Brailovsky.

In Weirhygia, the antenniferous tubercle armed, the mandibular plate unarmed,

posterior angle of connexivum extending into short and robust spine, abdominal

sternite VII of female without plica and fissura, and frontal angles produced for-

ward into conical teeth. In Grosshygioides, the antenniferous tubercle is unarmed,

the mandibular plate armed, posterior angle of connexivum unarmed, abdominal

sternite VII of female with plica and fissura, and frontal angles projecting forward

as small rounded teeth.

Distribution.—Only known from Australia.

Etymology.—Named for Tom Weir, distinguished Australian entomologist.

WEIRHYGIA FACETA BRAILOVSKY, NEW SPECIES

(Figs. 3, 4, 7)

Types.—Holotype male: Australia. NE Queensland, Isley Hills, 1050 m, 17°03’

S 145°42' E, 30 Nov 1993, Cook, Monteith and Janetzki. Deposited in Queensland

Museum, Brisbane, Australia. Paratypes: 1 male, 1 female; data: same as holotype.

Deposited in the ‘‘Colecci6n Entomoldgica del Instituto de Biologia, UNAM,

México, and Queensland Museum, Brisbane, Australia. 1 male; data: Australia:

NE Queensland, Upper Isley Ck., 750 m, 17°03’ S 145°41’ E, 29 Nov 1993,

Monteith, and Janetzki. Deposited in Queensland Museum, Brisbane, Australia.

Description—Male (holotype). Dorsal coloration: chestnut orange with inner face of antenniferous

tubercle, dorsal aspect of postocular tubercle, frontal angles, short longitudinal stripe at middle third

of posterior lobe of pronotal disk, and posterior angles of connexival segments II to VII yellow to

orange yellow; antennal segments I to III chestnut orange, IV yellow with basal join chestnut; he-

melytral membrane yellow with inner angle black. Ventral coloration: Chestnut orange with rostral

segments I to IV, buccula, acetabula (punctures chestnut orange), anterior and posterior lobe of meta-

thoracic peritreme, evaporative area, and genital capsule yellow to orange yellow; rim of abdominal

spiracular peritreme black; punctures of abdominal sternite V to VII reddish orange; coxae yellow

with dark brown spots; trochanters yellow and apically dark brown; femora yellow, and dorsally

covered with dark brown to light chestnut brown spots; tibiae yellow with four to five red rings; tarsi

yellow with diffuse reddish to dark brown marks.

Female—Dorsal coloration: head chestnut orange, sprinkled with small reddish tubercles and with

following areas yellow: dorsal aspect of postocular tubercle and longitudinal stripe adjacent to eyes;

antennal segment I chestnut orange, segment II dark yellow with apical third reddish orange, segment

III reddish orange with basal joint yellow, and IV yellow with basal joint reddish orange; pronotum,

scutellum, corium, clavus and dorsal abdominal segments with punctures reddish brown to dark brown;

connexival segments reddish brown with posterior angle yellow; hemelytral membrane black to dark

brown with middle third yellow. Ventral coloration: yellow with reddish brown to dark brown punc-

tures and irregular spots scattered throughout the body; rostral segments I-II chestnut brown, HI-IV

yellow (apex of IV chestnut orange); anterior and posterior lobe of metathoracic peritreme creamy

2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 83

Figures 1-2. Male genital capsule of Grosshygia spp.

Figure 1. G. formosa Brailovsky, NEW SPECIES.

Figure 2. G. pisina Brailovsky, NEW SPECIES.

Figures 3-4. Weirhygiafaceta Brailovsky, NEW GENUS, NEW SPECIES. Figure 3. Male genital

capsule. Figure 4. Female genitalia.

yellow and evaporative area yellow; coxae yellow with diffuse dark brown spots; trochanters yellow

with apex dark brown; femora and tibiae yellow, with four to five reddish orange to black irregular

rings; tarsi yellow with diffuse reddish to dark brown marks; abdominal spiracular peritreme black.

Measurements.—Male (female). Head length: 1.88 mm (1.98 mm); width across eyes: 1.52 mm

(1.64 mm); interocular space: 1.00 mm (1.02 mm); interocellar space: 0.50 mm (0.46 mm); preocular

distance: 1.30 mm (1.36 mm); antennal segments lengths: I, 1.12 mm (1.12 mm); IJ, 1.72 mm (1.76

mm); III, 1.40 mm (1.40 mm); IV, 1.20 mm (1.16 mm). Pronotal length: 1.72 mm (1.80 mm); width

across frontal angles: 1.68 mm (1.84 mm); width across humeral angles: 3.32 mm (3.56 mm). Scutellar

length: 1.00 mm (1.04 mm); width: 1.28 mm (1.48 mm). Maximum width of abdomen: 5.04 mm

(5.84 mm). Total body length: 10.20 mm (11.63 mm).

Etymology.—From the Latin, facetus, meaning elegant, fine.

GROSSHYGIA FORMOSA BRAILOVSKY, NEW SPECIES

(Figs. 1, 5)

Types.—Holotype male: Australia. Queensland, Kroombit Tops, 65 km SW

Gladstone (Sieved litter. Q.M. Berlesate no. 383, in rainforest), 1100 m, 22—26

Feb 1982, G. Monteith and G. Thompson. Deposited in Queensland Museum,

Brisbane, Australia.

84 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Figure 5. Dorsal view of Grosshygia formosa Brailovsky, NEW SPECIES.

Description—Male (holotype). Coloration: dark reddish brown; antennal segment I dark chestnut

orange II, and III pale chestnut orange, IV yellow with basal joint pale chestnut orange; hemelytral

membrane yellow; inner margin of acetabula, coxae, and trochanters dirty chestnut orange; femora

chestnut orange with basal joint, and few subapical spots dirty yellow; tibiae chestnut orange with

three to four irregular yellow rings; tarsi chestnut orange with dirty yellow reflections; upper margin

2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 85

of connexival segments black. Structural characters—Head: longer than wide across eyes, pentagonal,

conspicuously convex dorsally; antenniferous tubercles unarmed; rostrum reaching anterior margin of

abdominal sternite VII; vertex uniformly convex, without conical tubercles. Pronotum: slightly tra-

peziform, bilobed, nondeclivent; anterolateral margins convexly rounded, moderately elevated, and

slightly reflexed; callar region transversely nodulose; anterior and posterior lobe along the middle line

with wide longitudinal depression. Legs: unarmed. Scutellum: apex globose. Hemelytra: micropterous,

reaching posterior margin of abdominal segment I; hemelytral membrane reduced to small pads, sep-

arated from each other, leaving the abdomen exposed mesally. Genital capsule: posteroventral edge

simple, with median triangular expansion (Fig. 1).

Female—Unknown.

Measurements.—Head length: 1.86 mm; width across eyes: 1.34 mm; interocular space: 1.06 mm;

preocular distance: 1.32 mm; antennal segments lengths: I, 1.12 mm; II, 1.44 mm; III, 1.38 mm; IV,

1.28 mm. Pronotal length: 1.28 mm; maximum width of anterior lobe: 2.28 mm; maximum width of

posterior lobe: 2.92 mm. Scutellar length: 0.72 mm; width: 1.18 mm. Maximum width of abdomen:

4.60 mm. Total body length: 10.30 mm.

Discussion.—Grosshygia nigra Brailovsky and G. formosa, are the only species

of the genus with the scutellar apex globose. In the other known species, it is

subacute. In G. formosa the head is longer than wide, the vertex is uniformly

convex without conical lobes, the antenniferous tubercles are unarmed, and the

posteroventral edge of male genital capsule has a median triangular expansion

(Fig. 1). In G. nigra the head is wider than long, the vertex has two conical

elevations, the antenniferous tubercles are armed with extremely long lobes, and

the posteroventral edge of male genital capsule is transversely straight and entire.

Etymology.—From the Latin, formosus, meaning beautifully formed.

GROSSHYGIA LEPIDA BRAILOVSKY, NEW SPECIES

(Fig. 6)

Types.—Holotype female: Australia. N Queensland, Mt. Barflie Frere, Swiftlet

Cave, 900 m, 8 Dec 1990, Monteith, Thompson, Cook and Sheridan. Deposited

in Queensland Museum, Brisbane, Australia.

Description.—Female. (holotype). Coloration: dark chestnut orange with following areas dark yel-

low: dorsal aspect of postocular tubercle, rostral segments I to IV, posterior edge of connexival seg-

ments and abdominal pleural sterna III to VI, diffuse reflections on abdominal sterna III to VII, and

genital plates; head and scutellum reddish brown; antennal segments I to III chestnut orange with

basal joint of III yellow; antennal segment IV yellow with basal joint chestnut orange; anterior and

posterior lobe of metathoracic peritreme bright chestnut orange; coxae reddish orange; trochanters and

tibiae chestnut orange fore and middle femora chestnut orange with diffuse yellow rings; hind femur

yellow with apical joint and two or three incomplete chestnut orange rings; tarsi yellow. Structural

characters. Head: longer than wide across eyes, pentagonal, conspicuously convex dorsally; antenni-

ferous tubercles armed, lobes raised, arcute, recurved, converging anteriorly, apically subacute, almost

touching basal joint of antenna segment J; ocelli absent; rostrum reaching middle third of abdominal

sternite VI; vertex globose with thin transverse depression, dividing it into two elevations, the anterior

one broader, conical and higher than posterior one which is slightly rounded. Pronotum: scarcely

quadrate, bilobed, nondeclivent; anterolateral margins convexly rounded, moderately elevated, and

slightly reflexed; callar region transversely nodulose; anterior and posterior lobe along the middle line

with wide longitudinal depression. Legs: unarmed. Scutellum: apex subacute. Hemelytra: micropterous,

reaching middle third of abdominal segment III; wings reduced to small pads, separated from each

other, leaving abdomen exposed mesally.

Male—Unknown.

Measurements.—Head length: 1.80 mm; width across eyes: 1.72 mm; interocular space: 1.20 mm;

preocular distance: 1.30 mm; antennal segments lengths: I, 1.00 mm; II, 1.60 mm; III, 1.04 mm; IV,

1.04 mm. Pronotal length: 1.60 mm; maximum width of anterior lobe: 2.20 mm; maximum width of

86 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Figure 6. Dorsal view of Grosshygia lepida Brailovsky, NEW SPECIES.

posterior lobe: 2.68 mm. Scutellar length: 0.92 mm; width: 1.16 mm. Maximum width of abdomen:

4.84 mm. Total body length: 9.55 mm.

Discussion.—Grosshygia lepida and its closely related species G. lobatula Brai-

lovsky and G. monticeps Brailovsky, has the head longer than wide, the scutellar

2001

Figure 7.

BRAILOVSKY: NEW AUSTRALIAN COLPURINI

Dorsal view of Weirhygia faceta Brailovsky, NEW GENUS, NEW SPECIES.

87

88 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

apex subacute, and the hemelytra reaching the medial third of abdominal segment

III. In G. lepida the external lobes of the antenniferous tubercles converget an-

teriorly, conspicuously arcuate, recurved, and almost touching the basal joint of

antennal segment I (Fig. 6), and the vertex has two conical elevations separated

along middle third by a deep longitudinal depression. In the other two species

the external lobes of antenniferous tubercles are obliquely projecting, diverging

anteriorly, and the two anterior conical elevations of vertex are almost fused.

Etymology.—From the Latin, lepidus, meaning pleasant, elegant.

GROSSHYGIA PISINA BRAILOVSKY, NEW SPECIES

(Fig. 2.)

Types.—Holotype male: Australia. NE Queensland, 20 km WNW of Cape Trib-

ulation (Site 2), 50 m, 23 Sep 1982, Monteith, Yeates and Thompson. Deposited

in Queensland Museum, Brisbane, Australia.

Description—Male (holotype). Dorsal coloration: head reddish brown with dorsal aspect of pos-

tocular tubercle yellow; antennal segments I to III chestnut orange with basal joint of III yellow;

antennal segment IV yellow with basal joint chestnut orange; anterior lobe of pronotum reddish brown,

posterior lobe chestnut orange, anterolateral margins yellow; scutellum dark chestnut orange; heme-

lytral membrane chestnut orange; connexival segments bright reddish with posterior margins of seg-

ments III to VI yellow; dorsal abdominal segments reddish brown. Ventral coloration: Head reddish

brown, rostral segments I to IV and small spot close to eyes yellow; thorax reddish brown with

acetabula dark reddish, pro, meso, and metapleura dirty yellow with punctures dark brown; coxae

bright reddish; trochanters bright chestnut yellow; fore and middle femora chestnut orange with basal

third, subapical third, and few scattered spots yellow; hind femur yellow with apical third and four

irregular rings chestnut brown; tibiae and tarsi yellow; abdominal sterna dark reddish brown, scattered

with yellow diffuse areas; pleural abdominal sterna bright reddish with posterior third of sterna III to

VI yellow; genital capsule reddish brown with posteroventral margin pale orange yellow. Structural

characters. Head: longer than wide across eyes, pentagonal, conspicuously convex dorsally; antenni-

ferous tubercles each armed with short and robust lobe, obliquely projecting, diverging anteriorly and

subacute apically; rostrum reaching anterior margin of abdominal sternite VII; vertex uniformly convex

without conical elevations. Pronotum: slightly trapezoidal, bilobed, and nondeclivent; anterolateral

margins convexly rounded, moderately elevated, and slightly reflexed; callar region transversely nod-

ulose; anterior and posterior lobe along medially with wide longitudinal depression. Legs: unarmed.

Scutellum: apex subacute. Hemelytra: micropterous, reaching posterior third of abdominal segment

III; hemelytral membrane absent. Genital capsule: posteroventral edge simple, transversely concave,

entire (Fig. 2).

Female-——Unknown.

Measurements——Head length: 1.82 mm; width across eyes: 1.48 mm; interocular space: 1.00 mm;

preocular distance: 1.24 mm; antennal segments lengths: I, 1.14 mm; II, 1.84 mm; III, 1.14 mm; IV,

1.12 mm. Pronotal length: 1.48 mm; maximum width of anterior lobe: 1.88 mm; maximum width of

posterior lobe: 2.28 mm. Scutellar length: 0.80 mm; width: 0.96 mm. Maximum width of abdomen:

3.68 mm. Total body length: 9.10 mm.

Discussion.—Like G. monticeps Brailovsky, the head is longer than wide, the

scutellar apex is subacute, the antenniferous tubercles each is armed with a short

lobe, and the total length of antennal segment II is shorter than 1.86 mm. In G.

pisina, the vertex is uniformly convex without conical elevations, the membrane

of the hemelytra is absent, and the posteroventral edge of male genital capsule

simple, transversely concave and entire (Fig. 2). In G. monticeps the vertex is

globose with a thin transverse depression, dividing it into two conical elevations,

the hemelytra membrane is reduced to small lobe, and the posteroventral edge of

male genital capsule is indented medially and excavated laterally.

2001 BRAILOVSKY: NEW AUSTRALIAN COLPURINI 89

Grosshygia lobatula Brailovsky has the head longer than wide, the scutellar

apex subacute, the posteroventral edge of male genital capsule simple, the anten-

niferous tubercles each is armed with a long lobe, the length of antennal segment

II is longer than 1.96 mm, the vertex has conical elevations, and the hemelytra

membrane is reduced to a small flap.

Etymology.—From the Latin, pisina, small.

ACKNOWLEDGMENT

I thank G. B. Monteith (Queensland Museum, Brisbane, Australia) for the loan

of specimens. Special thanks to Cristina Urbina and Ernesto Barrera for the prep-

aration of illustrations (each one at the Instituto de Biologia, Universidad Nacional

Autonoma de México).

LITERATURE CITED

Brailovsky, H. 1993. A revision of the Tribe Colpurini from Australia (Hemiptera-Heteroptera-Cor-

eidae). Memoirs of the Queensland Museum 34: 35-60.

Brailovsky, H. y E. Barrera. 1996. Revisi6n del Complejo Sciophyrus (Hemiptera: Coreidae: Colpu-

rini). Folia Entomol. Mex. 96: 15-106.

Steinbauer, M. J. & A. R. Clarke. 1996. Revision of the Genus Acantholybas Breddin (Hemiptera:

Coreidae). Ann. Entomol. Soc. Am. 89: 519-525.

Received 14 Sep 1999; Accepted 7 Feb 2000.

PAN-PACIFIC ENTOMOLOGIST

71(2): 90-122, (2001)

SPIDERS FROM RIVERSIDIAN COASTAL SAGE SCRUB

WITH COMPARISONS TO DIEGAN SCRUB FAUNA

(ARACHNIDA: ARANEAE)

THOMAS R. PRENTICE!, JUTTA C. BURGER!, WENDELL R. ICENOGLE?,

AND RICHARD A. REDAK?!3

‘Department of Entomology, University of California, Riverside,

California, 92521

232631 Grand Avenue, P.O. Box 277, Winchester, California 92596

3Center for Conservation Biology, University of California, Riverside,

California 92521

Abstract.—We provide a list of 166 spider species collected from Riversidian coastal sage scrub

(CSS) located within the Southwestern Riverside County Multispecies Reserve in southern Cal-

ifornia. Spiders were collected over a two-year period from pitfall traps, vacuum samples, and

malaise traps; additional species were collected by hand or in yellow pan traps. One hundred

and twenty-seven species, representing 30 araneomorph families and one mygalomorph family,

were separated from the combined pitfall, vacuum, and malaise samples from 24 undisturbed

Riversidian CSS plots. Three species occurred only in malaise samples, 58 species occurred only

in pitfall samples, and 37 species occurred only in vacuum samples. An additional 39 species

were collected by hand or in yellow pan traps (two species), among which were representatives

of two mygalomorph families not represented in pitfall catches. Over 33% of the Riversidian

species were unrecorded from our recent Diegan CSS study bringing the combined total to 255

species. Eighteen species are new records for Riverside County. At least eleven species are

believed to be undescribed. Six non-native species, Holocnemus pluchei (Scopoli), Dysdera

crocata C. L. Koch, Zelotes nilicola O. P.-Cambridge, Oecobius annulipes Lucas, Steatoda trian-

gulosa (Walckenaer), and Menemerus bivittatus (Dufour), were among the species collected.

Phenology data are provided for the 12 most prevalent species. Natural history data are provided

for habitat preference of several species and on ant predation by three species. Non-native and

endemic spider taxa are discussed. Comparison of Riversidian fauna to Diegan fauna suggested

higher species diversity in Diegan CSS.

Key Words.—Arachnida, Araneae, spiders, species list, coastal sage scrub, endemic species.

We present here a significant addition to the knowledge of the spider fauna of

coastal sage scrub (CSS) from samples obtained from an ecologically threatened

southern California coastal sage scrub association, Riversidian CSS (Westman

1983). Because CSS communities, in general, are distributed at lower elevations

in California along the coast or adjacently inland, they are subject to extensive

clearing as the demands of urbanization are met. This study provides an invaluable

database for potential use by conservationists, developers, and planners, involved

either directly in CSS ecosystem research or indirectly in the planning and de-

velopment of reserves to protect this fragile ecotype in southern California. It

also provides arachnologists and ecologists alike with data pertaining to species

composition and richness, relative abundance, endemism, and the apparent impact

of non-native species in CSS. To date, this study and our previous Diegan CSS

study (Prentice et al. 1998) remain the only examinations of CSS spider fauna

and, together, create one of the most comprehensive lists of Araneae assembled

for any specific ecosystem type in the United States.

Our study sites were located within the Southwestern Riverside County Mul-

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 91

tispecies Reserve (SRCMR), a reserve owned by Metropolitan Water District

(MWD) and jointly managed by federal, state, county, and MWD personnel. This

area includes undisturbed coastal sage scrub surrounding Lake Skinner and ex-

tending 1 km south, 4 km east to the Tucalota Hills, and 9.5 km to the north,

climbing in elevation up through Rawson Canyon and then descending to (what

is now) Diamond Valley Lake. Chaparral communities, oak groves, and riparian

areas are associated with CSS in several regions within the reserve. The 1993

Winchester Fire burned large portions of SRCMR, leaving a mosaic of unburned

areas on the north side of the lake (Fig. 1, regions C (northern third), D, E). The

areas to the west and south of Lake Skinner were missed by the fire (Fig. 1,

regions A, B, C (southern two thirds)), but a large portion of the CSS and chap-

arral through Rawson Canyon to the northern boundary was destroyed (Fig. 1,

regions G, FP).

There are four major California CSS associations. Three have geographical

limits along the Pacific coast; one occurs further inland (Westman 1983). These

floristic associations are defined mainly by a geographic/climatic gradient of eva-

potranspiative stress, which increases from north to south and from coastal to

inland (Axelrod 1978; Desimone & Burk 1992; Westman 1981, 1983). Soil ni-

trogen levels and air pollution are also believed to be critical factors that influence

floristic associations (Westman 1981). Riversidian CSS extends from inland Los

Angeles County through the southwestern corner of San Bernardino County, west-

ern Riverside County, and into inland San Diego County and has a somewhat

different floristic composition than the more coastally distributed Diegan CSS

(Westman 1983). Although the percentage of annual forb and grass cover gen-

erally increases in CSS from north to south, the Riversidian region has a signif-

icantly higher percentage of annual cover (primarily introduced species) and a

lower percentage of shrub cover than the Diegan region (Westman 1983). The

relatively sparse shrub cover and dense annual cover in Riversidian CSS are

believed to be mainly the result of evapotranspiative stress (due to higher sum-

mertime temperatures) and increased ozone levels (Westman 1983). Vegetation

surveys (unpublished data) conducted within both our Diegan and Riversidian

sites showed that unburned sites at SRCMR had both relatively sparse shrub cover

and lower shrub stature than sites at MCBCP/MNAS. California Buckwheat (Er-

logonum fasciculatum), Yerba Santa (Eriodictyon crassifolium), Brittlebush (En-

celia farinosa), Black Sage (Salvia mellifera), and Bush Mallow (Malacothamnus

fasciculatus) were more common in unburned sites at SRCMR while California

Sage (Artemesia californica), White Sage (Salvia apiana), and Laural Sumac

(Malosma laurina) were more common in sites at MCBCP/MNAS. Bush Mon-

keyflower (Mimulus aurantiacus) and both Broom Baccharis (Baccharis saroth-

roides) and Coyote Brush (Baccharis pilularis) were common at MCBCP/MNAS

sites but absent from sites at SRCMR. In contrast, Bush Penstemon (Kekiella sp.)

was common at SRCMR but absent from sites at MCBCP/MNAS.

Our objectives are to: (1) develop an inventory of the spider fauna collected

in Riversidian CSS which includes the number of adults of each sex per species

per collecting period, (2) compare spider assemblages of Riversidian and Diegan

CSS, (3) develop phenology data for the most abundantly sampled Riversidian

Species with comparisons to the same taxa sampled in Diegan scrub (Prentice et

al. 1998), (4) propose a subset of probable endemic species to Riversidian and/

92 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

1 : Kilometers

Figure 1. Map of SRCMR in SW Riverside County, CA showing geographic regions of collections.

Regions ‘A—F’ encompass pitfall, vacuum, and malaise sampling sites. Regions ‘G’, ‘e-o-b’ (entrance

station outbuilding area), and the smaller encircled area within region ‘F (oak grove), were localities of

hand collection only. The open circles (CO) within regions ‘A—F’ represent unburned sites and the solid

circles (@) represent sites burned by the Winchester Fire of 1993. Regions ‘A’, ‘B’, and ‘e-o-b’ were

not burned; the understory of the oak groves (regions F and G) was burned but the oaks survived.

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 93

or Diegan CSS (area endemics), (5) discuss the apparent status of non-native

species in CSS, and (6) provide natural history data on habitat preference and ant

predation by two subsets of species, respectively.

MATERIALS AND METHODS

Study Sites.—We selected 24 circular sites representative of undisturbed Riv-

ersidian CSS scrub, each 100 m in diameter. Half (12) of these sites were located

in areas recovering from the Winchester Fire of 1993 and the other half (12) were

within areas that remained unburned (Fig. 1, regions A—F). Plots in the burned

areas had substantially reduced shrub canopy cover and greater annual grass and

forb cover than those plots in unburned areas which generally had dense canopies

and little forb and grass understory, except occasionally around the trunks of the

shrubs and under the overhanging branches. Unburned sites were chosen in a

manner that maximized site distribution over SRCMR without encroaching on

chaparral, oak, or riparian communities and that encompassed, within the com-

bined twelve sites, as much of the variation in CSS composition as possible.

Burned sites were selected, when possible, as pairs, adjacent to their unburned

counterparts, but were otherwise selected to maximize site distribution.

Selection of Geographical Regions.—The grouping of sites into regions (Fig.

1) was based on three criteria, provision of locality data, geographical relationship,

and practicality. Foremost, was to provide locality data for future workers wishing

to sample any of the taxa that we collected at SRCMR. Although we have more

precise locality data for each specimen sampled, inclusion here would prove both

cumbersome and unnecessary. Therefore, the only logical method of reporting

these data, was to assign sites to regions based on both proximity of the sites and

geographical distinction of the encompassed regions (Fig. 1). Sites in regions A

and B are on the south side of the lake or an arm of the lake (region B site is

within 50 m of the shoreline) in an area that was not burned by the Winchester

Fire. Sites in region C are the only sites that are relatively close to riparian areas,

although the northern site was burned in the 1993 fire. Sites in region D are

closely adjacent to the lake’s north shore and more exposed to the predominant

SW winds than other sites near the lake. Region E encompasses sites that are in

close proximity to one another (burned and unburned). Sites in region F are

isolated from other sites both by distance and elevation (640 m vs. 460 m). Region

G encircles an oak grove (closely adjacent to CSS) where various hand-collected

Species were taken; it is intermediate in elevation (550 m).

Sampling Periods.—Sampling was performed by pitfall trap, vacuum, and mal-

aise trap methods in September and December of 1996, in March, June, Septem-

ber, and December of 1997, and in March and June of 1998. Collections spanned

the first two weeks of each sampling month. Although sampling was also per-

formed in March and June 1996, pitfall and malaise traps were not set in June

1996 and only five pitfall traps were set (rather than seven, thereafter) in March

1996. As a result, specimens collected during these periods are excluded from the

data presented in this study, with the exception of representatives of thirteen

species, seven of which were collected only during these samplings (see Results).

These 17 specimens are listed because their inclusion contributes to the knowledge

of Riversidian spider fauna (Table 1).

Malaise Trap Sampling.—A single malaise trap (Bioquip® design, model

Table 1 A list of 166 species of Araneae from Riversidian CSS at Southwestern Riverside County Multispecies Reserve (SRCMR) in California. Of these, 127

species were collected from pitfall traps, vacuum samples, and malaise traps in September and December of 1996, March, June, September, and December of

1997, and March and June of 1998 (all specimens of 7 of these species were collected only in March or June 1996 samples and are reflected in the various totals).

Specimens of the opposite sex of 11 of the 127 species and all specimens of the additional 39 species listed were collected only by hand (hand, Yellow Pan Trap

(YPT), or sweepnet); hand-collected specimens are not included in the totals. Geographic regions are shown in Figure 1.

Species : Geographic Regions Coll. 3 Collection Periods Specim.

Burned Unbumed Method Mar Jun Sep Dec Total

MYGALOMORPHAE

Ctenizidae

Bothriocyrtum californicum (O. P.-Cambridge), 1874: 260 (E) (h) (1)

Cyrtaucheniidae

Aptostichus atomarius Simon, 1891:317 D A,(A,E) P (h) 43 4, (22)

Aptostichus sp. #1 (#118) * E R Te) l

Theraphosidae

Aphono pelma steindachneri (Ausserer), 1875: 199 (TRP coll.) (E,F) (h) (63 19)

Aphono pelma sp. #) (‘eutylenum' type’): Prentice, 1997 (TRP coll.) (D,E) (AE) (h) (93 49)

ARANEOMORPHAE

Agelenidae

Ageleno psis aperta Gertsch, 1934: 25 E P 19 1

Calilena stylophora Chamberlin & Ivie, 1941: 610 C,E,F E,F P 39 6312 4319 139 28

Rualena balboae (Schenkel), 1950: 82 F P 19 1

Amaurobiidae

species #1: undescribed species, new genus (CAS; TRP coll.) C,E B,E P 22 1312 4

Anyphaenidae

Anyphaena californica (Banks), 1904: 339 [E] [M] [1d] 1

Araneidae

Ar gio pe argentata (Fabricius), 1775: 443 () (h) (12)

Ar gio pe trifasciata (Forskal), 1775: 86 E A Vv 26 2

Cyclosa turbinata (Walckenaer), 1841: 140 Cc Vv 19 1

Eustala californiensis (Keyserling), 1885: 525 E D,E VM 1d 19 1é 3

Eustala conchlea (McCook), 1888: 199 (D) (h) (12)

Eustala rosae Chamberlin & Ivie, 1935: 22 (D) (h) (1)

Hypsosinga funebris (Keyserling), 1893: 37 [E] [V] [1d 12] 2

Larinia directa (Hentz), 1847: 478 D PY 19 1

Mastophora cornigera (Hentz), 1850: 20 [E] [V] [1d] 1

Metepeira crassi pes Chamberlin & Ivie, 1942: 66 D CE Vv 43 29 6

\O

£

LSIDO'IOWO.LNY OIWIOVd-NVd AHL

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Table 1. Continued.

Metepeira foxi Gertsch & Ivie, 1936: 66

Mete peira grandiosa grandiosa Chamberlin & Ivie, 1941: 17

Neoscona crucifera (Lucas), 1839: 42

Neoscona oaxacensis (Keyserling), 1863: 97

Caponiidae

Orthonops zebra Platnick, 1995: 15

Corinnidae

Castianeira athena Reiskind, 1969: 228

Castianeira occidens Reiskind, 1969: 211

Dictynidae

RCBlabomma sanctum Chamberlin & Ivie, 1937: 221

Blabomma sp. #4 (#436)

Blabomma sp. #5 (#446)

Blabomma sp. #6 (#445)

Cicurina utahana Chamberlin, 1919: 257

Dictyna abundans Chamberlin & Ivie, 1941: 6

Dictyna agressa Ivie, 1947: 2

Dictyna calcarata Banks, 1904: 342

Dictyna cholla Gertsch & Davis, 1942: 12

®CDictyna sierra Chamberlin, 1948: 8

®CE-mblyna consulta (Gertsch & Ivie), 1936: 6

Emblyna hoya (Chamberlin & Ivie), 1941: 7

Emblyna linda (Chamberlin & Gertsch), 1958: 148

Emblyna reticulata (Gertsch & Ivie), 1936: 7

RCvfallos pearcei Chamberlin & Gertsch, 1958: 45

Tivyna moaba (Ivie), 1947: 2

Tricholathys jacinto Chamberlin & Gertsch, 1958: 20

Tricholathys monterea Chamberlin & Gertsch, 1958: 22

Yorima angelica Roth, 1956: 4

Diguetidae

Diguetia canites (McCook), 1890, in 1889-1894: 135

Dysderidae

NNDysdera crocata C. L. Koch, 1839: 81

Filistatidae

Filistatinella sp. #3 (#522)

Gnaphosidae

Callile pis gosoga Chamberlin & Gertsch, 1941: 10

D,E,F

D,E

(F, oak grove)

F

(G)

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E

(F, oak grove)

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Table 1. Continued.

a een essa ea ee See

Cesonia classica Chamberlin, 1924: 619 A.E,F P 53 19 If 7

Drassyllus fractus Chamberlin, 1936: 14 E B,E P 8d 3¢ 11

Drassyllus insularis (Banks), 1900: 97 C-F A-F P 14812 268302 808249 103

®Drass yllus lamprus (Chamberlin), 1920: 193 E P 1d 1

Gnaphosa californica Banks, 1904a: 335 DE D,F P 4s 1d 29 7

Haplodrassus maculatus (Banks), 1904: 336 D,E,F A-F P 41d 262 12 43 72

Herpyllus hes perolus Chamberlin, 1928, in Cham. & Gert. 1928: 176 E M 1d 1

Her pyllus propinquus (Keyserling), 1887: 430 Cc P 1d 1

Micaria deserticola Gertsch, 1933: 2 EF] P[M] | [1¢]19 2

Micaria gosiuta Gertsch, 1942: 1 E P 12 1

Micaria icenoglei Platnick & Shadab, 1988: 57 F P ld 1

Micaria jeanae Gertsch, 1942: 4 DE A,B,E PVM ld 183° 19 6

R€Nodocion utus (Chamberlin), 1936a: 7 F P 19 1

Scopoides catharius Chamberlin, 1922: 156 (E) E P (h) 19 1,(1¢)

Zelotes gynethus Chamberlin, 1919: 7 D,E,F A,B,E,F P 26 152 29 1d 73 29 29

Zelotes monachus Chamberlin, 1924: 621 E A,D,E,F P 63 52 11

NNZelotes nilicola (O. P.-Cambridge), 1874: 380 AE P 161° 2

Zelotes skinnerensis Platnick & Prentice, 1999: 672 E P 2612 3

Hahniidae

Neoantistea santana Chamberlin & Ivie, 1942: 29 E P 19 19 2

Heteropodidae

Olios schistus Chamberlin, 1919: 10 (E) (h) (1d)

Homalonychidae

Homalonychus theologus Chamberlin, 1924: 631 D D,F P 3d 3

Linyphiidae

Ceraticelus sp. #1 (#7) [C] E Vv [1d] 2¢ 3

Erigone autumnalis Emerton, 1882: 58 E F 19 1

Erigone dentosa O. P.-Cambridge, 1894, in 1890-1902: 128 D,E,F B,D,E,F v*P 28d 192 26 49

Frontinella pyramitela (Walckenaer), 1841: 261 E Vv 1d 1

RCT inyphantes aliso Chamberlin & Ivie, 1942: 53 E Vv 12 1

RCT inyphantes micro ps Chamberlin & Ivie, 1942: 53 E,F Cc P 262° +

Linyphantes sp. #5 (E) (h) (1?)

Meioneta sp. #2 (#53) D,E,F B,D,E,F P 5d 5? 238 49 731° 24

Meioneta sp. #3 (#460P) E A,E VP 38 29 19 19 7

Microlinyphia mandibulata punctata Chamberlin & Ivie, 1943: 24 E E vp ld 1622 #4

Spirembolus erratus Millidge, 1980: 151 (B) (h) (1d)

Spirembolus phylax Chamberlin & Ivie, 1935: 19 C,E A vP 19 19 i) 3

NNT ee eee || ———————————————————————

LSIDO' TOWO.LNA OIWIOVd-NVd AHL

(Z)LL TOA

Table 1. Continued.

ee LL

Spirembolus pusillus Millidge, 1980: 128

Spirembolus tortuosus (Crosby), in Chamberlin, 1925: 116

Spirembolus redondo (Chamberlin & Ivie), 1945: 225

Tennesseellum formicum (Emerton), 1882: 71

®CWubana drassoides (Emerton), 1882: 72

Liocranidae

Drassinella gertschi Platnick & Ubick, 1989: 5

Phrurotimpus mateonus (Chamberlin & Gertsch), 1930: 141

CS cotinella kastoni (Schenkel), 1950: 73

Lycosidae

Alopecosa kochi (Keyserling), 1876: 636

Pardosa californica Keyserling, 1887: 483

Pardosa ramulosa (McCook), 1894, in 1889-1894, pl. 30

Schizocosa maxima Dondale & Redner, 1978: 165

Schizocosa mccooki (Montgomery), 1904: 283

Mimetidae

Mimetus hes perus Chamberlin, 1923: 5

Miturgidae

Cheiracanthium inclusum (Hentz), 1847: 451

Sys pira synthetica Chamberlin, 1924: 665

Oecobiidae

NNOecobius annuli pes Lucas, 1849: 102

Oonopidae

Oonops sp. #1 (TRP coll.)

Scaphiella hes pera Chamberlin, 1924: 594

Oxyopidae

Oxyopes salticus Hentz, 1845: 196

Oxyopes scalaris Hentz, 1845: 196

Oxyopes tridens Brady, 1964: 472

Peucetia longipal pis F. P.-Cambridge, 1902, in 1897-1905: 340

Peucetia viridens (Hentz), 1832: 105

Philodromidae

Apollophanes texanus Banks, 1904: 113

RCE bo albocaudatus Schick, 1965: 77

RC bo andreaannae Schick, 1965: 82

Ebo californicus (Gertsch), 1933: 14

RCE bo evansae Sauer & Platnick, 1972: 41

(B)

(A)

AB,DE,F

12

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3°

13842

2?

183 62

[lo]19

42

2612

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12

25d 2?

431?

2?

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[1¢]19

1619

2d 29

1419

1819

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106 1?

12 1?

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12

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90

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(1g 19)

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Table 1. Continued.

a,

Ebo mexicanus Banks, 1898: 256

Philodromus californicus Keyserling, 1884: 676

Philodromus chamisis Schick, 1965: 50

®¢T hanatus formicinus (Clerck), 1757: 134

Tibellus chamberlini Gertsch, 1933: 10

Pholcidae

Holocnemus pluchei (Scopoli), 1763: 404

Physocyclus californicus Chamberlin & Gertsch, 1929: 102

Psilochorus sp. #1 (#103)

Plectreuridae

Plectreurys conifera Gertsch, 1958: 14

Salticidae

Habronattus californicus (Banks), 1904: 117

Habronattus hirsutus (Peckham & Peckham), 1888: 64

Habronattus pyrrithrix (Chamberlin), 1924: 693

Habronattus schlingeri (Griswold), 1979: 135

Habronattus tranquillus Peckham & Peckham, 1901: 201

Menemerus bivittatus (Dufour), 1831; 369

Metaphidippus chera (Chamberlin), 1924: 683

Metaphidi ppus mannii (Peckham & Peckham), 1901: 326

Metaphidi ppus vitis (Cockerell), 1894: 207

Pelegrina aeneola (Curtis), 1892: 332

Pellenes limatus Peckham & Peckham, 1901: 217

Phidippus ardens Peckham & Peckham, 1901: 288

Phidippus californicus Peckham & Peckham, 1901: 289

Phidippus johnsoni Peckham & Peckham, 1883: 22

Phidippus nikites Chamberlin & Ivie, 1935: 41

Phidippus octopunctatus (Peckham & Peckham), 1883: 6

Salticus pal palis (Banks), 1904: 360

Sarinda cutleri (Richman), 1965: 133

Sitticus dorsatus (Banks), 1895: 97

Synageles occidentalis Cutler, 1987: 343

Tengellidae

RC Anachemmis dolicho pus Chamberlin, 1919: 139

Tetragnathidae

Tetragnatha guatemalensis O. P.-Cambridge, 1889, in 1889-1902: 8

®¢Tetragnatha pallescens F. P.-Cambridge, 1903, in 1897-1905: 436

D,E

C,F,(D,E)

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1é

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3d 3°

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1?

1d 4°

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2562

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38219 40

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26 1d 2? 17

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LSIDOTOWOLNA OIWIOVd-NVd FHL

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Table 1. Continued.

Tetragnatha versicolor Walckenaer, 1841: 215 D VM 2619 3

Theridiidae

Achaearanea tepidariorum (C. L. Koch), 1841: 75 (nr. e-o-b) (h) (12)

Di poena abdita Gertsch & Mulaik, 1936: 6 Cc F 19 1

CF noplognatha selma Chamberlin & Ivie, 1946: 3 (E) E,(B) P (h) 1d 1, (49)

Euryo pis californica Banks, 1904: 345 D Vv 19 1

Latrodectus hes perus Chamberlin & Ivie, 1935: 15 (E) (h) (12)

NNSteatoda triangulosa (Walckenaer), 1802: 207 (culvert nr. e-o-b) (h) (1d 29)

Steatoda washona Gertsch, 1960: 21 (E) (h) (1d)

Theridion dilutum Levi, 1957: 37 E A Vv ld 29? 3

®¢T heridion llano Levi, 1957: 28 [E] [A,D],(B) [VP] (h) [12] [29] 3, (16)

Theridion melanurum Hahn, 1931: (pl. 3, fig. a.) (culvert nr. e-o-b) (h) (1)

Theridion rabuni Chamberlin & Ivie, 1944: 53 B[D]E Vv [16]3? 4

Tidarren haemorrhoidale (Bertkau), 1880: 78 (F) (h) (12)

Thomisidae

Misumeno ps aikoae Schick, 1965: 131 vac shaff Vv ‘oe 1

Misumenops californicus (Banks), 1896: 91 [E] [V] [12] 1

Misemenops deserti Schick, 1965: 124 D,E Vv 1629 3

Misumeno ps importunus belkini Schick, 1965: 131 E B,C Vv 36 19 1d ld 6

Misumeno ps lepidus (Thorell), 1877: 498 C,E,F A,C Vv 8d 162° ll

Misumenops rothi Schick, 1965: 117 E D,E,F Vv 43 19 5

Xysticus californicus Keyserling, 1880: 37 C-F p*V 27d 32 39 33

Xysticus gertschi Schick, 1965: 159 E Vv 19 1

Xysticus montanensis Keyserling, 1887: 479 (F) (B) (h) (2?)

Total 354 §11 147 211 1223

' Geographic Regions A-—G; letters in Burned and Unburned columns refer to condition of sites within the various regions; e-o-b = entrance station outbuilding

(see Fig. 1).

2P = pitfall trap, V = vacuum sample, M = malaise trap, (h) = hand-collected. Collection methods are listed in order of greatest to least number of specimens

collected by the particular method; second letter preceded by an asterisk (*) indicates that very few specimens were collected by that particular method.

3 Number of specimens collected are recorded under each period.

4 Number following an undetermined species number is the OTU reference number assigned to the particular species in our collection (1.e., Aptostichus sp. #1

(#118)).

NN:RC (superscript to the left of listed species): NN = non-native species; RC = new species record for Riverside County, California.

() Hand collected specimens; parentheses around particulars in all except Species and Collection Period columns (dates of collection are provided in Table 2).

[ ] Specimens collected only in March or June 1996 by pitfall, vacuum, or malaise; brackets around particulars in all columns except Species and Total columns.

SYAdIdS €NaOs AOVS “TV LA AOLLNAYd 100¢c

66

100 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

#2875 AG) was erected. near the center of each site and left for three days during

each sampling period. Arthropods collected in the kill heads (containing KCN or

80% EtOH) were then transferred to plastic bottles containing 80% EtOH. A total

of 192 malaise samples (576 trap/days) were collected.

Vacuum Sampling.—Five one-minute vacuum samples were collected at each

site during each sampling period (Buffington & Redak 1998). Vegetation was

vacuumed along portions of five 25 m line transects established from a center

pole by randomly chosen degree points from North. Each sample consisted of

vacuuming a 10 X 1 m (10 m?’) band along the transect for one minute beginning

at either the 5 or 15 m mark (randomly selected). Live arthropods were extracted

from the collected chaff using one-gallon cardboard extraction drums. The drums

(ice-cream containers) were painted black inside and each lid fitted with the fun-

nel-shaped neck portion (glued in place, narrow end out) of one-liter plastic soda

bottle. Each funnel was roughened on the inside (for traction) and painted black.

A plastic vial cap was bored to tightly fit over the mouth of each funnel in a

manner that would allow a 40-dram collecting vial to be attached to the lid.

Contents of the vacuum bags were placed into the drums. Drums were turned

sideways with the plastic vial portion under blacklights for a period of three hours.

Arthropods that had collected in the vials were then anesthetized with carbon

dioxide and stored in 80% ethanol. Vacuum chaff was examined for additional

specimens for a period of one minute. A total of 960 vacuum samples were

collected and extracted.

Pitfall Trap Sampling.—Seven pitfall traps were placed at each site in a strat-

ified random manner and left in place for the entire course of the study. Each

trap consisted of a 473 ml (16 oz) plastic cup (9 cm, inside diameter) equipped

with a funnel that narrowed to 3.2 cm and a fluid filled collection cup (fluid

composition: 2.0 tablespoons odorless detergent and 1.0 tablespoon salt per gallon

water). Pitfall cups were placed so that the rims were flush with the substrate

surface. A square masonite board, supported by three 20-penny galvanized nails

at approximately 2 cm above each trap, was used to protect the trap from des-

iccation and/or flooding and provided a dark refuge for arthropods. Traps were

opened for a period of three days at each site during each sampling period. Pitfall

contents were strained and transferred to 80% ethanol within seven hours of

collection. A total of 1344 pitfall samples (4032 trap/days) were collected.

Vegetation Sampling.—Four 48 m transects were established at each plot. A

pole marked in decimeters was dropped at a random location (point) within every

2 m section along each transect (n = 96 points per plot). Canopy species, height,

and structure (by number of contacts within each decimeter), as well as ground

cover, were then recorded at each point. Along three of the four transects at each

plot, ground and canopy cover, per decimeter, were recorded. We were able to

estimate relative cover of major shrubs, shrub structure, ground cover and rock-

iness at each site with these data.

Statistical Analysis.—We used species accumulation curves to compare spider

diversity in Diegan and Riversidian CSS. This method allowed us to correct for

the disparity in total number of specimens collected from each location. Curves

and error bars were constructed with EcoSim, Version 3 (Gotelli & Entsminger

1999), by resampling using Monte Carlo randomizations of species abundance

data. All adults and any immatures, identifiable as distinct species, were included.

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 101

Specimens from malaise samples were not considered because malaise traps were

not implemented in Diegan sampling. We intend to include a more involved sta-

tistical comparison of the Arthropoda collected at both locations in a community-

level study (in preparation).

Manuscript Organization and Terms.—The organization of the manuscript fol-

lows, for the most part, that of our previous Diegan CSS study (Prentice et al.

1998) to facilitate comparison of Riversidian and Diegan spider fauna. The use

of the terms “Diegan’ and ‘Riversidian’ is in reference to our Diegan CSS study

(and to Diegan CSS) and our present Riversidian CSS study (and to Riversidian

CSS), respectively.

Specimens Examined and Species List.—A total of 5676 specimens from pitfall,

vacuum, and malaise samples, 108 hand collected specimens, 2 specimens from

yellow pan traps, and one specimen collected by sweepnet were examined. Only

adults were considered in the species list (Table 1). Specimens that were collected

by hand are not added in the various totals in Table 1; collection data for these

specimens are in Table 2. Taxonomy follows Platnick (1989, 1993, 1997). Un-

determined species are designated as sp. #1, sp. #2, etc.; listings starting with sp.

#2, #3, etc. indicate that the species are considered to be different congeneric

species than those designated as sp. #1, #2, etc. in Prentice et al. (1998). Gender

and number of specimens are provided for each species for each season collected;

collection method(s) and geographical region of collection are also provided for

each species (Table 1; Fig. 1). Table 1 is the central component of the manuscript

and should be referred to throughout the course of the text. Voucher specimens

of listed species (Table 1) are deposited in the University of California, Riverside,

Entomological Research Museum.

Abbreviations.—CAS, California Academy of Sciences; MCBCP, Marine Corps

Base Camp Pendleton (San Diego County); MNAS, Miramar Naval Air Station

(now Marine Corps Air Station Miramar) (San Diego County); SRCMR, South-

western Riverside County Multispecies Reserve.

Sampling Biases.—The trapping materials and methods used during this study

(excepting malaise traps) were essentially the same as those used during our

Diegan study. Therefore, the sampling biases discussed in Prentice et al. (1998)

also pertain to the present study. We provide here a brief summary of these biases.

The small size of our pitfall traps precluded the collection of specimens of large

size. Burrowing species were not accounted for unless collected by hand or an

occasional wandering male fell into the trap. Web spinners that used subterranean

retreats, had strong webs, and/or were nocturnal were minimally, if at all, collected

by the vacuum method. Specimens of many web spinners and arboreal hunters

and ambushers, that habitually drop or jump to the ground under conditions of

unusually strong vibration, were minimally collected by the vacuum method.

RESULTS AND DISCUSSION

Spider Fauna of Riversidian CSS.—In addition to the previously published

floristic distinctions between Riversidian and Diegan CSS types (Kirkpatrick &

Hutchinson 1977; Westman 1981, 1983; DeSimone 1995), we found accompa-

nying distinctions between the respective spider faunas. It appears that factors

influencing the floristic structures of Riversidian and Diegan CSS ecosystems,

such as seasonal rainfall patterns (including effective summer and winter mois-

102 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Table 2. List of species collected by hand (hand, YPT, or sweepnet). An asterisk * preceding a

binomial indicates that only specimens of the opposite sex, of the respective species collected by

pitfall, vacuum, or malaise methods, were hand-collected; all specimens of all other species were

collected only by hand. Dates of collection and data pertaining to particular habitats are provided.

Unless otherwise indicated, specimens were collected in CSS; other collections were in areas adjacent

to CSS communities. Collection of specimens other than by one of authors is denoted in the Collection

Date column.

Species Collection Date Habitat

MYGALOMORPHAE

Cyrtaucheniidae

*Aptostichus atomarius 13 Dec 1997, 2 9: Coll: J. Bond banks of ravine cuts

Theraphosidae

Aphono pelma steindachneri

Aphono pelma sp. #1

Ctenizidae

Bothriocyrtum californicum

ARANEOMORPHAE

Araneidae

Argiope argentata

Eustala conchlea

Eustala rosae

Metepeira foxi

Neoscona crucifera

Corinnidae

Castianeira athena

Dictynidae

Dictyna agressa

Dictyna calcarata

*Dictyna cholla

Emblyna consulta

Mallos pearcei

*Tivyna moaba

Tricholathys jacinto

Gnaphosidae

*Scopoides catharius

Heteropodidae

Olios schistus

Linyphiidae

Linyphantes sp #1

Spirembolus erratus

Spirembolus redondo

*Tennesseellum formicum

Liocranidae

Drassinella gertschi

17 Jul 1997, 2h; 21Jul 1997, 1d; 4 Aug 1997,

3d; 28 Sep 1997, 19

mid-May 1996, 12; 3 Sep 1997, 2¢; 11 Sep

1997, 2h; 28 Sep 1997, 3d; 29 Oct 1997,

16,12; 26 Nov 1997, 16; 7 May 1998, 12

early Jul 1997, 19

9 Sep 1997, 12

16 Mar 1997, 12

21 Mar 1997, 12

10 Feb 1998, 1¢

mid-Oct 1997, 12

7 Jun 1999, 1 2: Coll: C. Dunning

25 Mar 1997, 2 2

21 Mar 1998, 1¢

20 Dec 1998, 1¢

4 Jun 1998, 1¢

21 May 1999, 1 2: Coll: C. Dunning

6 Dec 1998, 39

26 May 1999, 1 2: Coll: C. Dunning

10 Mar 1998, 1¢ (pen.; def. molt 16 Mar

1998)

17 May 1998, 16

22 Nov 1997, 1?

1 Dec 1998, 16; 29 Dec 1998, 16; 31 Dec

1998,12

3-5 Mar 1998, 1 2: Coll: M. Buffington

11-13 Mar 1998, 16: Coll: M Buffington

31 Dec 1998, 10,12

(co) on dirt rds., (2) in

burrow

(od) on dirt rds., (2) in

burrows

in burrow

in web betw. shrubs

against stem of shrub

against mustard stem

on drag line betw. shrubs

riparian

in fallen acom, oak grove

in webs, entr. stn. outbldg.

in web, entr. stn. outbldg.

grass/leaf litter

in web, Calif. Buckwheat,

branch tip

on oak leaf, oak grove

grass/leaf litter

in grass litter, oak grove

under rock

under dead stump

on wild oat stem

grass litter

riparian, YPT

in shallow drainage, YPT

under dead root, chaparral

2001

Table 2. Continued.

Lycosidae

Pardosa californica

Schizocosa maxima

Oonopidae

Oonops sp. #1

Philodromidae

Ebo andreaanne

Ebo evansae

Philodromus californicus

Thanatus formicinus

Pholcidae

Holocnemus pluchei

Physocyclus californicus

Salticidae

Habronattus pyrrithrix

Habronattus schlingeri

Menemerus bivittatus

Pellenes limatus

*Phidi ppus ardens

*Phidip pus johnsoni

*Phidippus nikites

*Phidi ppus octopunctatus

Theridiidae

Achaearanea tepidariorum

*Fnoplognatha selma

Latrodectus hes perus

Steatoda triangulosa

Steatoda washona

*T heridion llano

Theridion melanurum

Tidarren haemorrhoidale

Thomisidae

Xysticus montanensis

PRENTICE ET AL.: SAGE SCRUB SPIDERS

9 Mar 1998, 1&: Coll: M. Buffington

16 Mar 1997, 1& (pen.; def. molt, late May-

early June 1997)

1 Dec 1998, 16,2 2; 20 Dec 1998, 86,72

19 Jun 1997, 1h (pen. def. molt, 6-7 Dec

1997)

17 Jul 1997, 12

27 Jan 1997, 1h; 26 Feb 1998, 19

21 Mar 1997,12

31 Aug 1998,12

16 Jan 1997, 12; 25 Jun 1997, 1; early Jul

1997, 136, 12; mid-Sep 1997, 1

12 Mar 1998, 12

6 Dec 1998, 1¢ (pen. def. molt, 4 Apr 1999)

11 Sep 1997, 1& (pen.), 19; (def. molt (¢),

21-25 Nov 1997)

1 Dec 1997, 1h (pen.; def. molt, early June

1998); 12 Mar 1998, 19

16 Jun 1997, 16; 25 Jun 1997, 1é

7 Mar 1997, 12; 16 Mar 1997, 12 (pen.;

w/&; molt, 20-24 Mar 1997); 20 Mar 1997,

1¢2;30 Mar 1997, 19; 6 Mar 1998, 12

30 Jul-3 Aug & mid-Jul 1998 (def. molts), 2¢

(both (&) reared from (¢ ) collected 10 Sep

1997)

4-7 Jun & 20-21 Jun 1998 (def: molts), 2¢

(both (¢) reared from (¢ ) collected 11 Sep

1997)

6 Feb 1997, 12

9-31 Dec 1998, 4 (pen.; molts 30 Dec 1998-

25 Jan 1999)

5 Mar 1997,12

27 Jan 1997, 12; 14 Mar 1997, 19; 7 Jun

1998, 1¢

4 Jun 1998, 1¢

29 Dec 1998, 1& (pen.; def. molt, 10-11 Jan

1998)

6 Feb 1997, 19

13 Mar 1997, 19

10 Mar 1997, 12; 1 Dec 1998, 12

es

in shallow drainage, YPT

under board

in fine, fluffed dead grass

on shrub

on shrub

entr. stn outbldg

on ground

in web, entr. stn. outbldg.

in web, entr. stn. outbldg.

on dirt road

sparse grass litter

on wall, entr. stn. outbldg.

on ground; field with low

grass

under masonite pitfall lid;

on Calif. Buck wheat

on ground.; under wood; in

PVC pipe centerpole

(2) under masonite over

pitfall trap

(2) on Calif. Buckwheat

in web on low shrub

grass/leaf litter

in cavity under rock

in web, inside culvert; in

webs, entr. stn. outbldg.

on ground

in web on shrub

in web, entr. stn. outbldg.

in web, cavity of oak

tree, oak grove

grass litter

rs

103

104 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

ture), variability in weather conditions and temperature, elevation gradients, gen-

eral topography, and soil types, also affect the associated spider faunas. Not only

did sampling comparisons between the Diegan and Riversidian CSS spider fauna

reveal differences in species composition and relative abundance, but suggested

a much higher species diversity in CSS (in general) than originally indicated in

our Diegan study.

A total of 166 species (Table 1), representing 96 genera in 30 araneomorph

and 3 mygalomorph families, were collected from Riversidian CSS communities

at SRCMR compared with 200 species (112 genera in 31 araneomorph and 4

mygalomorph families) collected from Diegan CSS (Prentice et al. 1998). Of those

collected at SRCMR, 127 species were collected by traditional trap or vacuum

methods while 39 species were collected by hand. Three species occurred only

in malaise samples, 58 species occurred only in pitfall samples, and 37 species

occurred only in vacuum samples. In samples from the traditional methods (mal-

aise, vacuum, and/or pitfall), Gnaphosidae were represented by the greatest num-

ber of species (19), followed by Salticidae (16), Dictynidae (14), Linyphiidae

(13), Araneidae (9), Thomisidae (8), Theridiidae and Philodromidae (each with

6), Oxyopidae (5), Agelenidae, Liocranidae, Lycosidae, and Tetragnathidae (each

with 3) (Table 3). The remaining families were represented by only one or two

species. In our Diegan study, the eight most speciose families (excluding hand-

collected species) in descending order were Gnaphosidae, Salticidae, Theridiidae,

Linyphiidae, Dictynidae, Araneidae, Thomisidae, and Philodromidae (Table 3).

Of the total number of species from SRCMR (including hand collected species),

over 33% (55 species) were absent in Diegan samples, although 14 of these

species were probably represented by juveniles (only) in Diegan samples Guve-

niles not included on species lists of either study).

Diversity within most taxa was generally greater in Diegan than in Riversidian

CSS (Table 3). This pattern remained after we corrected for differences in spec-

imen numbers across the respective CSS types (Fig. 2). For Riversidian CSS, the

maximum species richness for specimens collected from pitfall traps and vacuum

samples was 136. Given the species accumulation curves in Fig. 2, we estimated

spider species richness in Diegan CSS to be 18% greater than in Riversidian CSS

at the same specimen abundance (3302 specimens).

Mecicobothriidae (Mygalomorphae), Cybaeidae, and Nesticidae were repre-

sented only in Diegan samples but Caponiidae was represented only in Riversidian

samples (Table 3). However, various species of Mecicobothriidae and Caponiidae

have been collected from CSS/chaparral transition zones in both San Diego and

Riverside Counties. One species of Nesticidae is known from both counties, and,

at least, one species of Cybaeidae (most California species are distributed in

central and northern counties) has been collected in San Diego County.

At the genus level, diversity was greater in Diegan CSS within ten shared

families, most notably the Salticidae, but in Riversidian CSS diversity was greater

within only four shared families, most notably the Pholcidae (Table 3). Although

Species composition was, in general, richer in Diegan than in Riversidian CSS,

the reverse was true within six shared families, Cyrtaucheniidae, Araneidae, Dic-

tynidae, Oxyopidae, Philodromidae, and Pholcidae (most notably, Dictynidae,

Philodromidae, and Pholcidae), and seven shared genera (Aptostichus, Metepeira,

Oxyopes, Peucetia, Ebo, Steatoda, and Misumenops). Diegan fauna was substan-

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 105

tially more speciose within the Theridiidae and Gnaphosidae and, to a lesser

extent, within the Salticidae and Linyphiidae (Table 3).

March and June 1996 Sampling Additions—Specimens (11) of species that

were collected only in March and/or June 1996 samples are included on the

species list and counted in the various totals. The species are as follows: Hypo-

singa funebris (Keyserling), Mastophora cornigera (Hentz), Anyphaena califor-

nica (Banks), Phidippus californicus Peckham & Peckham, Theridion llano Levi,

Tetragnatha pallescens F P.-Cambridge, and Misumenops californicus (Banks).

The males of Ceraticelus sp. #1, Micaria deserticola Gertsch, Pardosa ramulosa

(McCook), Oxyopes salticus Hentz, and Theridion rabuni Chamberlin & Ivie, and

the female of Liocranoides dolichopus (Chamberlin) were also collected during

these sampling periods and are included on the species list and counted in the

various totals (6 specimens).

Hand-Collected Species.—TYable 2 provides additional collection and habitat

data, not provided in Table 1, for specimens of 50 species collected by hand. All

specimens of 39 of these species were collected only by hand (hand, Yellow Pan

Trap, or sweepnet). For each of the remaining 11 species, hand-collected speci-

mens of only one sex (either male or female) are recorded in Table 2; specimens

of the opposite sex of each of these species, respectively, were collected in pitfall,

vacuum, and/or malaise samples.

New Riverside County Records—Eighteen species are new records for River-

side County (indicated in Table 1 by the superscript “RC’). County listings are

provided here for three species reported only in counties north of and including

Santa Barbara County: Nodocion utus (Chamberlin): Ventura County (Platnick &

Shadab 1980); Thanatus formicinus (Clerck): Santa Barbara County, Monterey

County, Yosemite National Park (Mariposa and/or Tuolumne Counties) (Dondale,

Turnbull, & Redner 1964); Dictyna sierra Chamberlin: Sierra County (Chamberlin

& Gertsch 1958). Boe (1986—unpublished; available at UC Riverside Science

Library) reported D. sierra from Riverside County but we were unable to au-

thenticate this record.

Females, determined to be D. sierra: (Chamberlin & Gertsch 1958), were pre-

viously known only from SE Oregon. The female epigynum as illustrated in

Chamberlin & Gertsch (1958) is, decidedly, not the epigynum of the female that

we collected at SRCMR. However, general coloration and morphological simi-

larities between our male and female, as well as the close proximity of the sites

from which they were collected, suggest that our specimens are conspecific.

Undescribed Species.—Approximately 6.5% and 9.5% of the listed Riversidian

and Diegan species, respectively, are believed to be undescribed taxa. Taken to-

gether, the Riversidian and Diegan studies provided 26 undescribed species or

approximately 10% of the combined totals of 255 species. These percentages, as

well as the 18 new species records (above) for Riverside County and the 35 new

records for San Diego County (Diegan study), are indicators of just how little we

know of the arthropod community structure within the CSS system, in general.

The following 11—12 species (ten or eleven araneomorph and one mygalomorph

species) collected at SRCMR, are believed to be undescribed taxa: Blabomma sp.

#4, #5, #6 (Dictynidae); sp. #4 is probably the female of sp. #5 (undescribed

status confirmed by D. Ubick (CAS)); species #1 (m, f), undescribed genus

(Amaurobiidae) (family status suggested by D. Ubick, (CAS)); Filistatinella sp.

Table 3. Differences in the generic composition and the number of species within spider families represented in both Riversidian CSS samples (present study)

and Diegan CSS samples (Prentice et al. 1998). Unlisted families were either represented in only one of the studies or were represented in both studies by the

same species. Juveniles were considered in the elimination of unique genera. Representatives (Riversidian species) of a probable new genus/new species (n. g./n.

Ss.) are tentatively placed in the Amaurobiidae. Riversid. = Riversidian.

Family

Agelenidae

Amaurobiidae

Anyphaenidae

Araneidae

Corinnidae

Cyrtauchentidae

Dictynidae

Filistatidae

Gnaphosidae

Hahniidae

Linyphiidae

Liocranidae

Lycosidae

Mimetidae

Oonopidae

Oxyopidae

Philodromidae

Pholcidae

Plectreuridae

Salticidae

Riverside.

# gen.

3

1

Diegan

# gen.

4

2

1

OW

Riversid.

unique gen.

n. g./n. s.

?Anyphaena

Mastophora

Mallos

Wubana

Oonops

Thanatus

Holocnemus

Physocyclus

Diegan

unique gen.

?Hololena

Meraltella

Zanomys

Teudis

Corinna

Meriola

Urozelotes

Calymmaria

Idonella

Ostearius

Agrocea

Opopaca

Orchestina

Kibramoa

Metacyrba

Pckhamia

Phanias

Terralonus

Thiodina

Riverside.

sp./fam.

Diegan

sp./fam.

—

io)

LSIDO'IONOLINA OMIOVd-NVd HHL

(LL TOA

Table 3. Continued.

Family

Tengellidae

Theridiidae

Thomisidae

Riverside.

# gen.

1

8

Diegan Riversid.

# gen. unique gen.

1

10

3

Diegan

unique gen.

Argyrodes

Crustulina

?Thymoites

Coriarachne

Riverside.

sp./fam.

1

17

Diegan

sp./fam.

2

24

SYACIdS €NAOS AOVS *"IV LA AOLLNAYd 100d

LOI

108 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

250

200

150 iat

oO Diegan CSS

100 @_ = Riversidian CSS

Richness

0 2000 4000 6000 8000 10000 12000 14000

Abundance

Figure 2. Species diversity of Araneae in Diegan and Riversidian CSS. Species accumulation

curves were constructed by resampling species abundance data. Only adults and immature specimens,

distinct enough not to be confused with other species, were included in calculations. Error bars rep-

resent 95% confidence intervals from Monte Carlo randomizations.

#3 (Filistatidae); Ceraticelus sp. #1, Linyphantes sp. #5, Meioneta sp. #2, #3

(Linyphiidae); Oonops sp. #1 (Oonopidae); Psilochorus sp. #1 (Pholcidae); Ap-

tostichus sp. #1 (Cyrtaucheniidae) (status determined by J. Bond (Field Museum,

Chicago)). Specimens of only four of these species, Ceraticelus sp. #1, Meioneta

sp. #2, Psilochorus sp. #1, and Aptostichus sp. #1, were found in Diegan samples.

Phenology.—Twelve species, eleven from pitfall traps and one from vacuum

samples, were collected in numbers large enough (n > 25) to provide an abstrac-

tion of their life cycles or, more precisely, their adult activity patterns. The phe-

nologies of Calilena stylophora Chamberlin & Ivie (n = 28) and six other species

(n = 16-24) are not discussed either because the species was not observed in the

field, revision of genus in question is in need, comparative natural history liter-

ature was lacking, and/or specimens were hand collected.

In the majority of spider species with discreet breeding periods, first male

maturation just precedes or roughly corresponds to the beginning of the respective

breeding season, with maximum male activity peaking some time later. Peak fe-

male activity, on the other hand, roughly corresponds to copulatory periods and

increased foraging or searching for oviposition sites prior to egg deposition

(DeKeer & Maelfait 1997, Draney 1997). Phenograms (Fig. 3) do not reflect

juvenile activity because many congeneric immatures could not be reliably sep-

arated.

Our two most prevalent species, Callilepis gosoga Chamberlin & Gertsch and

Drassyllus insularis (Banks) (Figs. 3A and 3B, respectively), are apparently most

active in Riversidian CSS between late spring to late summer and presumably

mate during this period. In Diegan CSS, both species likely remain active some-

what longer (C. gosoga with slightly decreasing activity and D. insularis with

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 109

increasing activity between June and August) (Prentice et al. 1998). Although

adult D. insularis did not occur in December samples, two of us (TRP & WRI)

have hand-collected adults from November—February. Platnick & Shadab (1982)

also reported that adults occur year-round. Adults of C. gosoga have been col-

lected only between mid-April and early September (Platnick 1975). The female

to male ratio of C. gosoga (~1:3) in samples (Fig. 3A) is decidedly lower than

in D. insularis (~1.1:1) (Fig. 3B) which suggests foraging dissimilarities between

females of the two species. We hypothesize that C. gosoga is an ant specialist

(see: Natural History subsection: Ant Predators below). Mature females, in prep-

aration for egg deposition, may restrict their hunting ranges to areas near harvester

ant nests, in which case, they may only occasionally fall into randomly placed

pitfall traps. Contrarily, males probably wander in search of females (rather than

in search of prey), which increases their chances of falling into pitfall traps. Adults

of D. insularis are probably active generalist predators with, perhaps, broad hunt-

ing ranges which may render females (and males) more subject to pitfall collec-

tion.

The remaining two common gnaphosids, Haplodrassus maculatus (Banks) and

Zelotes gynethus Chamberlin, show increasing activity from late fall to early

spring with peak activity in March (Figs. 3C, 3D, respectively). Zelotes gynethus

seems to be more prevalent than H. maculatus in December, which may indicate

an earlier and/or more prolonged mating period than that of the latter species.

Moreover, Z. gynethus adults occurred year-round but those of H. maculatus were

absent in September samples. Because only one H. maculatus female was col-

lected in June samples, it seems likely the majority of adults expire during May.

According to Platnick and Shadab (1975), adults had been previously taken from

early January through mid-May. The lack of March samples from our Diegan

study may have accounted for the apparent absence of H. maculatus; only a few

juveniles were collected, which may have been either H. maculatus or H. signifier.

Although California collection records indicate an inland distribution of H. ma-

culatus, specimens have been collected within approximately 3 km of the coast

in Baja, California (Platnick & Shadab 1975). In all probability, H. maculatus

inhabits southern California Diegan CSS although it may not be as common as

in Riversidian CSS.

In both our Diegan and Riversidian CSS studies, Alopecosa kochi (Keyserling)

and Schizocosa mccooki (Montgomery) were the two most frequently collected

wolf spiders; specimens of both species were collected together in many of the

same plots during both studies. Although A. kochi are generally smaller than S.

mccooki, there is substantial size overlap between the species which may result

in competition for like-size prey items. The partitioning of resources may be

accomplished both by differences in breeding seasons and daily activity patterns.

Alopecosa kochi displays increasing activity from December to March (Fig. 3)

and is diurnal (Hagstrum 1970) whereas S. mccooki shows peak activity in Riv-

ersidian CSS in June (Fig. 3G) and is nocturnal (Dondale & Redner 1978).

In Diegan CSS peak activity of S. mccooki occurred in August. The presence

of penultimates (only) in late May and early June Diegan samples and the col-

lection of two females in late August (26th and 29th) with partial broods still

clinging to their abdomens suggest that breeding probably occurs between late

June and late July in Diegan CSS. Had we sampled in July and early August

110 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

120 | A Callilepis gosoga oi B Drassyllus insularis

100 | 3 oe

25

80 |

20

60

15

40 ‘6

20 7 5

0 4 Z c 0 |

Mar Jun Sep Dec Mar Jun Sep Dec

50 16 ,

a C Haplodrassus maculatus D Q Zelotes gynethus

w 14 4 :

> 40

Oo 12 |

>

oO 30 f 10 7

_—

ro =

- 205 6

®

O 44

Ss 10

= | > |

Zz

0 - 04

20 30

a F Schizocosa mccooki

16 | =

14 | 20

12.4 |

10 | 15 ;

8

6 | 10 |

4 5 |

2

0 —o pe 0 4 de 1

Mar Jun Sep Dec Mar Jun Sep Dec

Figure 3. Phenograms of twelve spider species (adults only): A-D, Gnaphosidae; E-K Lycosidae;

G-H, Linyphiidae; I, Pholcidae; J, Salticidae; K, Dictynidae; L, Thomisidae. Collecting periods illus-

trated on the abscissa were during the first two weeks of each month listed. Pitfall traps (7 per plot;

24 plots) were left open for 72 h + 4 h. Five one-minute vacuum samples perxplot (24 plots) were

collected during each sampling period. Closed circles (@) connected by solid lines represent male

collections; open circles (O) connected by dashed lines represent female collections.

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 111

28 1G Erigone dentosa 100 H _ Spirembolus tortuosus

24 80 -

20 |

60 -

15 |

40 -

10 ;

5. 20 |

|

0 | 0 ts ag Le 2

Mar Jun Sep Dec

30 ; 25 5

| l Psilochorus sp.1 | J Habronattus californicus

wT 25 | °

© 20 | fi

<5 |

zz 20 4

eet] |

2 15 | ei

S fg

10 | 4 :

he 10 : ’

0) |

OQ 5; |

= | 5 |

0) 4 }

za |

T T T T 0

Mar Jun Sep Dec

35 ) K Tricholathys monterea 30 | L Xysticus californicus

30 25.

25 | |

| 20

20 - |

| 15 4

15 4 |

| |

10 |

10 5

5 | oa

0 | 0

Sep Dec

Figure 3. Continued.

during our Riversidian study we may have seen an increase in female activity

past the apparent June peak (Fig. 3G). However, the absence of males in early

September Riversidian samples may be an indication of a slightly earlier breeding

season than that of Diegan populations. The large size of S. mccooki and the

presence of two distinct juvenile size classes in samples suggest both a biennial

112 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

life cycle and alternating generations (Dondale 1977, Hagstrum 1970). However,

in populations from pinon-juniper forests in New Mexico, Stratton & Lowrie

(1984) found S. mccooki to have an annual life cycle.

Alo pecosa kochi, inhabiting pine forests in the San Bernardino National Forest,

is reported to have a similar two-year life-cycle with alternate generations (Hags-

trum 1970). In the mountain populations, penultimates mature in fall with adults

overwintering, followed by mating in early spring and egg deposition in early

May. In our studies, December and March pitfall catches, as well as hand collec-

tion of both males and females in early February from both Riversidian and

Diegan study sites, indicate continuous activity throughout the winter months.

Unlike pine forest inhabitants, CSS inhabitants presumably mate during the winter

months and produce eggs in early March; females with egg sacs were collected

as early as 4 March from SRCMR. The absence of adults in both June and

September Riversidian samples and in August Diegan samples suggests that adults

may expire prior to the hottest and driest part of the summer.

Spirembolus tortuosus (Crosby) and Erigone dentosa O. P.-Cambridge were the

two most prevalent linyphiid species. All female and all except two males of E.

dentosa were collected in June 1998 vacuum samples (Fig. 3G); only one male

was taken (in June) during the Diegan study. Yeargan & Dondale (1974) reported

the take of EF. dentosa by all methods used (vacuum, sweepnet, pitfall trap) in

their study of the spider fauna in alfalfa fields of northern California. Crosby &

Bishop (1928) indicate collection records of both sexes from May to September.

However, two of our males occurred in March 1996 samples (data not used here)

from SRCMR and males, together with females, were collected by hand in early

March at SRCMR and in early February at MCBCP. It appears as though adults

are present throughout the year but may likely be most active between early June

and middle or late August. Males of S. tortuosus (Crosby) were collected exclu-

sively in December samples from both Diegan plots and Lake Skinner (Fig. 3H);

the few Riversidian females taken occurred only in March samples (December in

Diegan samples) and, of these, two were obviously gravid when collected. Mil-

lidge (1980) reported male collections in October, November, and January and

female collections in September, October, and December through February or

March. During independent collecting efforts by one of us in 1998 (TRP), both

penultimates and adults were taken from low grasses and by litter sifting between

the last week of November and the first week in December. For the remainder of

December, only adults were found, suggesting that the peak breeding period oc-

curred in Lake Skinner populations after the first week of December. In agreement

with Millidge (1980), we believe that the species breeds during the winter months

prior to early March when only females are present. The small proportion of

females in samples is probably due to both habitat structure and female behavior.

Like many erigonine species, female and juvenile S. tortuosus may live in small

webs (captive adults spin small, flat webs) that are constructed within moist leaf

litter, whereas males actively search for females during the breeding season. Such

behavioral differences would account for the disproportionate catch of males in

both Diegan and Riversidian studies. |

Phenograms of the prevalent pholcid and salticid species, Psilochorus sp. #1

and Habronattus californicus (Banks), respectively, reflect similar activity patterns

(Figs. 31, 3J, respectively). Field collections of both male and female Psilochorus

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 113

sp. #1 in December and February at MCBCP and at SRCMR indicate, however,

that adults of the species are most likely present year-round. December field col-

lections at both locations yielded only female H. californicus; males were not

found in December at MCBCP nor between November and late January at

SRCMR. Both species are thought to have an annual life cycle. Maturation of

captive H. californicus spiderlings (egg sacs produced and young reared in lab)

occurred in approximately six to seven months. The presence of juveniles of

several size classes within both H. californicus and Psilochorus in a given June

or September sample suggests that the respective females may breed and produce

egg sacs during several months of the year. This stands to reason given that

Psilochorus males are found throughout the year, H. californicus males occur

during ten of the twelve months, and gravid H. californicus females were collected

in both June and September samples. The September peak of H. californicus (Fig.

3J) may be indicative of increased foraging activities, which may be necessary

due to the scarcity of prey. By September, most of the vegetation, that the spider’s

prey may depend upon for sustenance, has either long since senesced or become

summer dormant. Unlike that of H. californicus, the September peak of Psilo-

chorus (Fig. 31) may reflect shelter-seeking activity, which may be necessary to

avoid desiccation from the summer heat. The vast majority of the specimens

observed in the field were discovered beneath rocks or under other objects that

provided both a dark refuge and insulation from the heat. Although unset pitfall

cups were covered with both plastic lids and masonite boards (weighted with

rocks), Psilochorus adults and subadults were repeatedly discovered in small webs

within the cups, immediately before the traps were set in early September. When

initially opened, most pitfall cups had condensation droplets on the inside surfac-

es. Set traps likely provided a very similar environment, attracting Psilochorus

seeking shelter, and, in turn, accounting for the September take.

Phenograms of the dictynid, Tricholathys monterea (Chamberlin & Gertsch),

and the thomisid, Xysticus californicus Keyserling, indicate late winter activity of

males but little cursorial activity of females in either late winter or spring (Figs.

3K, 3L, respectively). In Diegan CSS, both males and females of T. monterea

were relatively common in mid- to late spring but the only specimens of X.

californicus collected (3 females) were taken in May/June samples. Chamberlin

& Gertsch (1958) report collection dates for adult T. monterea from late March

to late May with the majority of specimens examined collected in early to mid-

May. Taken together with our data, it appears that the species’ breeding season

peaks between early and late spring. Egg deposition may occur during the early

part of the summer. Schick (1965) reported common collection of X. californicus

adults from March to July throughout most of the species’ southern California

distribution. However, during several visits to both study areas in early February

of 1997 and 1998, we observed an abundance of penultimates and adults of both

sex, although the adult to penultimate ratio was apparently greater in males (>

0.5) than in females (< 0.5). Based on the apparent disappearance of males in

Riversidian CSS by early June (Fig. 3L) we can predict a mid-winter to mid-

spring breeding season within these populations.

Occurrence Patterns in Burned vs. Unburned Plots.—For characterizations of

burned and unburned plots refer to Study Sites under Methods. Few species were

collected in large enough numbers to discern a pattern of occurrence in burned

114 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

vs. unburned plots. Of the 12 most prevalent species discussed immediately above,

five species occurred in equal numbers (+ 4%) in burned and unburned plots.

One species, Alopecosa kochi, was collected more frequently in unburned plots

(60% of specimens), and six species, S. mccooki, S. tortuosus, E. dentosa, Psil-

ochorus sp. #1, X. californicus, T. monterea, occurred more frequently in burned

plots (59, 79, 76, 60, 100, and 66% of specimens, respectively). Ten of the 11

Zelotes monachus Chamberlin and all specimens of Habronattus hirsutus (Peck-

ham & Peckham) were taken from unburned plots, 5 of 6 Scaphiella hespera

Chamberlin and 10 of 13 Dictyna abundans Chamberlin & Ivie specimens came

from burned plots. Spirembolus tortuosus, X. californicus, and several other spe-

cies (not listed above) that exhibited patterns of occurrence are further discussed

below in the Natural History subsection, Habitat Preferences.

Probable Endemic Species.—Because we did not sample in chaparral com-

munities, nor in other communities which are often intimately associated with

CSS communities in southern California, we use the term endemic to mean “‘re-

stricted to a particular area’’ following Anderson (1994). Eight species are here

considered as probable endemics to southern California (Riverside, San Bernar-

dino, Los Angeles, Orange, and/or San Diego counties) exclusive of the mountain

and desert regions: Orthonops zebra Platnick, Micaria capistrano Platnick &

Shadab, M. icenoglei Platnick & Shadab, Zelotes skinnerensis Platnick & Prentice,

Linyphantes aliso Chamberlin & Ivie, L. microps Chamberlin & Ivie, L. obscurus

Chamberlin & Ivie, Scotinella kastoni (Schenkel). One species, Plectreurys tecate

Gertsch is believed to be endemic to SW San Diego County and adjacent Baja,

California. Refer to Prentice et al. (1998) for additional data on species collected

only during our Diegan study.

Orthonops zebra (Caponiidae): In his revision of Orthonops, Platnick (1995)

examined 41 specimens, most of which were collected by one of us (WRI) from

Riversidian CSS near Winchester (approximately 15 km N of SRCMR). Two

Specimens were collected in chaparral communities in the Santa Ana Mountains

(Orange and Riverside counties, respectively), west of Murietta. We collected two

additional males in June pitfall samples from Riversidian CSS at SRCMR (the

majority of the males that Platnick examined were collected in March). All known

specimens were taken within an 18 km radius of Murietta. Three of the other nine

known Orthonops species are presumed to have limited distributions in southern

California (Platnick 1995); one of these species, O. johnsoni Platnick, is known

only from Johnson Canyon (San Diego County), a habitat of chaparral and Diegan

CSS.

Micaria capistrano (Gnaphosidae): Seven of the nine known specimens (male

unknown) were collected from either Diegan CSS or chaparral communities (Baja

specimens) in either SW San Diego County (MCBCP and MNAS) (3 females)

or in Baja California Norte (4 females, see Platnick & Shadab 1988). These

localities are separated by approximately 265 km. Of the two remaining known

specimens, one was collected from the Box Springs Mountains, Riverside County

(Riversidian CSS), the other in the Santa Ana Mountains near the Riverside Coun-

ty border (probably from chaparral). The lack of specimens in Riversidian samples

during the present study and in independently collected CSS samples (collections

spanning nearly 30 years) near Winchester by one of us (WRI), suggests that the

Species is very rare, if it naturally occurs, this far north.

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 115

Micaria icenoglei (Gnaphosidae): A total of 23 specimens, collected exclu-

sively from or closely adjacent to CSS, are known. Eight of these were collected

from Riversidian CSS, seven near Winchester (two males were collected near

buildings adjacent to CSS) (see Platnick & Shadab 1988), and one male during

our present study. The remaining 15 specimens were collected during our Diegan

CSS study. If the distribution of M. icenoglei is continuous from Winchester to

San Diego County it seems likely that the species also inhabits chaparral com-

munities, at least the lower elevation communities.

Zelotes skinnerensis (Gnaphosidae): The three known specimens (Platnick &

Prentice 1999), two males and one female, were collected from two plots within

600 m of the NE lakeshore of Lake Skinner (SRCMR). Because the revision

(Platnick & Shadab 1983) of the genus was based on a large amount of material

and because one of us (WRI) has collected Araneae in southern California for

thirty years without encountering the species, Z. skinnerensis is here considered

either rare or very secretive.

Linyphantes aliso (Linyphiidae): This species was described from a female

holotype from Laguna Beach (also female paratypes); the male was then unknown

and as yet remains undescribed. Although degrees of latitude and longitude were

provided in the original description (Chamberlin & Ivie 1942), it is next to im-

possible to determine the exact habitat from which the specimens were taken in

light of the development within the Laguna Beach area since 1942. Both CSS

and chaparral are (presently) common in the undeveloped and undisturbed areas

just north of the town proper. An additional 30 specimens were collected from

Diegan CSS, including 10 males, several of which were present with females in

various plot samples. The species was absent from Riversidian samples although

one male and one female were collected in pitfall traps at Santa Rosa Plateau

Ecological Reserve in SW Riverside County. The most southerly-inhabited region

(within MNAS) is approximately 100 km S of the type locality; all known col-

lection localities are within approximately 40 km of the coast.

Linyphantes microps (Linyphiidae): This species was described from a single

male taken in Irvine, Orange County (latitude and longitude provided in original

description) (Chamberlin & Ivie 1942). To our knowledge, the male holotype is

the only published record of L. microps prior to our work here. During our present

study, we collected two males in March samples from Riversidian CSS along with

two females that we determined to be conspecifics.

Linyphantes obscurus (Linyphiidae): This species was described from a single

female collected in December; the type locality is the same as that of L. aliso.

We collected 17 males (Linyphantes sp. #3) in December samples during our

Diegan study but were unable to determine the species then because the male of

L. obscurus has not been described. A female of L. obscurus, along with 28 males

(determined to be conspecific to both the female and the 17 males from San Diego

County) were taken (Coll.: C. Dunning) in December by pitfall trap at Santa Rosa

Plateau Ecological Preserve in SW Riverside County. The plots from which these

Specimens came are composed primarily of Engelmann oak, coast live oak, and

native perennial grassland. Specimens were not discovered at SRCMR. To date,

the species is known only from San Diego, Orange, and Riverside counties.

Fourteen of the 19 described Linyphantes species occur in California (Cham-

berlin & Ivie 1942), 11 of which are known only from California. The type

116 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

species, L. aeronauticus (Petrunkevitch) is the only member of the genus known

to have an extensive distribution, occurring in Idaho, Wyoming, Oregon, Utah,

Nevada, California, Arizona, and probably several additional states. All other

congeners are known only from western coastal localities (Canada, Washington,

Oregon, California, or Mexico). At least three of our seven purported Linyphantes

species (from both studies) are believed to be undescribed. When generic revision

of Linyphantes is eventually undertaken, we may find that there are several Lin-

yphantes species that are endemic to our region.

Scotinella kastoni (Liocranidae): This species was described from a single fe-

male collected 28 Dec 1938 from Balboa Park, San Diego (Schenkel 1950); the

male has not been described. Because the hills in and around the zoo are of low

elevation and were likely to have supported CSS (rather than chaparral) prior to

urbanization, it is probable that the landscape from which the specimen was col-

lected was composed primarily of CSS. We have collected an additional 108

specimens, 25 from CSS (7 males, 14 females during our Diegan study and 4

females during our present study) and 83 from oak-savanna habitats at Santa Rosa

Plateau. To our knowledge, there are no additional published locality records of

S. kastoni. The apparent distributional limits of the species (assuming a continuous

distribution) extend from Lake Skinner to Balboa Park, a terrain composed of

chaparral and CSS, covering a distance of approximately 100 km.

Plectreurys tecate (Plectreuridae): The only three heretofore known specimens

(2 males, 1 female) were collected 16 May 1952, 8 mi. W of Tecate, Baja Cali-

fornia (Gertsch 1958). CSS is present in the general area although it appears to

be either restricted to relatively small discontinuous patches or intermixed with

Californian chaparral, which provides the major canopy cover in the region. Spec-

imens of P. tecate were not taken in Riversidian samples but two additional males

were collected from the same plot at MNAS (between 12-19 May 1996). The

canopy cover of this plot has both CSS and chaparral components, the latter

including Adenostoma fasciculatum Hooker & Amott (chamise) and Yucca whip-

plei Torrey. Throughout the approximate 40 km gap between the type locality and

MNAS both CSS and chaparral are the primary vegetation types. Data indicate

that P. tecate prefers relatively dry habitats with enough surface soil and moisture

to support the growth of the transitional (chaparral to CSS) chaparral elements.

In all likelihood, the distribution of this species is probably more extensive in

Baja, California than in California. The remaining species in the P. castanea group

(9 species) are known only from California. Seven of these species apparently

have restricted distributions in southwestern part of the state; the other two spe-

cies, P. monterea Gertsch and P. castanea Simon, are known only from Monterey

County and from San Mateo County to San Diego County, respectively (Gertsch

1958).

Non-native Species.—The following six known non-native species were col-

lected either in Riversidian CSS samples or by hand (Table 1; denoted by the

superscript ‘NN’): Oecobius annulipes Lucas, Zelotes nilicola (O.P.-Cambridge),

Dysdera crocata C. L. Koch, Holocnemus pluchei (Scopoli), Steatoda triangulosa

(Walckenaer), and Menemerus bivittatus (Dufour). Only one male and female of

both O. annulipes and Z. nilicola and one female of D. crocata were found in

Riversidian samples. Specimens of H. pluchei, S. triangulosa, M. bivittatus were

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 117

collected by hand at or near the entrance station and are discussed below. Refer

to Prentice et al. (1998) for data on O. annulipes, Z. nilicola, and D. crocata.

Holocnemus pluchei (Pholcidae): Of Mediterranean origin (Porter & Jakob

1990), H. pluchei appeared in California during the early 1970s. One of us (WRI)

collected the species in Sutter County, California in 1974, which may be the

earliest California record (Porter & Jakob 1990). We found no records that indi-

cated when the species was first reported within the United States. Boe (1986-

unpublished) reported collection records in Imperial, Los Angeles, Napa, River-

side, and Sonoma Counties and from the San Joaquin and Sacramento Valleys.

Porter & Jakob (1990) conducted their studies of H. pluchei from populations in

Riverside, Los Angeles, Kern, Fresno, and Yolo Counties. The species has since

been observed by one of us (TRP) in San Bernardino County (Loma Linda) and

in San Diego Co. (Spring Valley). Our one specimen (female) was collected in

1998 from its web in the angle of the soffet and walls of the entrance station

outbuilding. Two additional specimens appearing to be subadults were also present

in the web. In both 1996 and 1997, another pholcid, Physocyclus californicus

Chamberlin & Gertsch, had occupied the available spaces under these eaves but

in 1998 was absent, suggesting displacement by H. pluchei (not observed in

previous years). In another example of probable displacement by H. pluchei, we

refer to a third pholcid species, Artema atlanta Walckenaer. One of us (WRI) had

observed, for several years, a healthy population of this very large, primarily

neotropical species under the Hwy 115 bridge crossing the Alamo River just west

of Holtville, Imperial County, California. In October 1998, two of us (WRI &

TRP) revisited the site to find a dwindling A. atlanta population and, for the first

time, H. pluchei, the population of which outnumbered that of A. atlanta by an

estimated 2/1 ratio.

Steatoda triangulosa (Theridiidae): Quite possibly of European origin, S. trian-

gulosa may have been reported in the United States (Georgia) as early as 1850

(see Levi 1957). The majority of early collection records indicate that the species

was originally introduced in one of the eastern seaboard states prior to 1900. Levi

(1957) indicated records in California in only two counties, Alameda and Solano.

Boe reported its distribution from San Diego and Imperial counties north to Sis-

kiyou and Nevada counties. Two of our three specimens (male, female) were

collected from the same outbuilding on which the pholcids had resided. Notably,

both specimens were collected from small webs confined to the extreme corners

slightly above the expanded webs of the resident pholcids, the female above

Physocyclus californicus in 1997 and the male above Holocnemus pluchei in

1998. The third specimen (female) was collected from its web in a recess on the

upper inner surface of a road culvert (culvert approximately 1 km S of entrance

outbuilding).

Menemerus bivittatus (Salticidae): Probably originating in the Old World Trop-

ics (Edwards 1979), the species now occurs in Puerto Rico, Costa Rica, and

Mexico, north to the southern United States. In the U.S., there are records in most

of the southern states from Florida to California. In California the species is found

as far north as Sonoma County. Both specimens collected at SRCMR were found

on the walls of the entrance station outbuilding. Throughout the course of our

study, both juveniles and adults were commonly seen on the building when the

days were warm and sunny, but were never observed within the native vegetation.

118 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Non-native spider invasions appeared to be minimal in Riversidian CSS com-

pared to those in coastal Diegan CSS. Evidence from our Diegan study suggests

that the non-native spider species are more likely to occur in greater numbers in

CSS habitats that are less variable in temperature and have higher summertime

humidity than Riversidian CSS habitats. Secondly, human traffic promotes the

introduction and subsequent spread of non-native species inadvertently hitching

rides. Obviously, the human element has a substantially greater impact on the

introduction and spread of non-native species at MCBCP and MNAS (both heavi-

ly traveled and in very populated areas) than at SRCMR (most parts are closed

to human traffic and the reserve is in a relatively isolated area). Holocnemus

pluchei, S. triangulosa, and M. bivittatus were probably not encountered during

our Diegan study because we failed to search manmade structures at either base.

All three species have been collected by one of us (TRP) from two residences in

Spring Valley, San Diego County.

Natural History.—Habitat Preference: Only a few of the species collected in

either Riversidian or Diegan CSS were prevalent enough in samples to provide

any insight into their habitat preferences. Collection results suggest that two of

the common Riversidian species, Spirembolus tortuosus (Linyphiidae) and Xys-

ticus californicus (Thomisidae) prefer relatively open microhabitats with an un-

derstory of grassy areas and a significant amount of grass or leaf litter or small

prostrate forbs. Spirembolus tortuosus seems to prefer valleys or gentle south

facing hillslopes with a significant amount of ground litter. In our Diegan study,

S. tortuosus was distributed in 40 of the 60 plots. However, plots with sparse

shrub cover and ample ground litter provided nearly half of the specimens. Of

the specimens that occurred in plots with a dense shrub cover, most (70%) were

from valley plots with grassy areas and abundant leaf or grass litter. Over 80%

of the specimens collected during our present study occurred in plots recovering

from the 1993 Winchester burn (burned plots); all of these plots were character-

ized with a sparse shrub cover, ample annual grass and forb cover, and abundant

litter. The ground-dwelling thomisid, Xysticus californicus, also seems to prefer

open grassy areas with either grass and/or leaf litter or a patchy prostrate forb

layer. All of the Riversidian specimens occurred in samples from recovering

burned plots; the three specimens collected in Diegan samples came from plots

with similar percentages of shrub cover and understory composition. The largest

number of specimens (> 50) that we observed at any one time was on a gentle,

bare SW slope (MCBCP) carpeted with an Erodium sp. (storksbill).

Among the Dictynidae, collection results from both our Riversidian and Diegan

studies suggest that Blabomma spp. prefer CSS habitats that offer the most humid

microhabitat conditions (i.e., valleys, northern exposures, dense canopy layer, sig-

nificant leaf litter, and grassy areas). Blabomma sanctum Chamberlin & Ivie oc-

curred only in unburned plots with dense shrub canopies at SRCMR; nearly 80%

of the specimens came from north facing plots. Of these, close to 95% were taken

from one plot (Fig. 1, region E) characterized by a very dense shrub cover pri-

marily of California Sage (Artemesia californica) and secondarily of California

Buckwheat (Eriogonum fasciculatum) and Black Sage (Salvia mellifera) and an

essentially bare substrate with sparsely distributed Schismus grass. Leaf litter was

abundant but present only near the shrub bases. The two specimens occurring in

Diegan samples were from plots with patchy shrub cover but with a relatively

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 119

dense understory of annual grasses. All specimens of the undescribed (or unde-

termined) Blabomma species (sp. #1, #2, #3) from Diegan samples were taken

from plots with chaparral elements (primarily chamise) as either major or impor-

tant minor shrub components (the presence of chaparral elements is indicative of

more moist conditions than those supporting only CSS). Two of these plots were

in valleys and two were SW and SSE exposures, respectively, with dense shrub

covers and little bare ground between shrubs. All except one specimen (Blabomma

sp. #5) of the three undescribed (or undetermined) Riversidian Blabomma species

(sp. #4, #5, #6) were collected from plots that either had dense shrub canopies,

were located on north slopes, or had abundant grassy patches and ground litter.

Of the 41 total Blabomma specimens taken, only four males of Blabomma sp. #5

were in samples from burned plots.

Among the Tetragnathidae, the three species of Tetragnatha collected at

SRCMR occurred only in plots near Lake Skinner’s north shore. Both T. guate-

malensis O. P.-Cambridge and T. versicolor Walckeraer were taken within 250 m

of the shoreline (Fig. 1 region D), while T. pallescens E P.-Cambridge was found

at a maximum distance of 600 m from shoreline (Fig. 1, region E). Levi (1981)

reported T. versicolor as one of two Tetragnatha species occurring furthest away

from water. Given the hot and dry summer conditions of inland southern Cali-

fornia, it was not surprising to find this species so close to water (Fig. 1, region

D). The 7. pallescens female found furthest from water was collected from a

gentle north facing hillslope. During July and August, literally hundreds of T.

guatemalensis webs were spun every evening between the stems of dead mustard

plants along a service road, less than 100 m from the N shoreline (Fig. 1, region

D). Males and females, in copula, were observed at night in the female webs

throughout July and early August. By day, only traces of the webs were seen, the

inhabitants almost invisibly stretched out against the mustard stems. Larinia di-

recta (Hentz) and Neoscona (primarily juveniles) webs were commonly inter-

spersed among the tetragnathid webs during these months.

Although relatively few adults of the araneid species, Metepeira crassipes

Chamberlin & Ivie and M. grandiosa grandiosa Chamberlin & Ivie, were col-

lected in samples, both species are very common at SRCMR, particularly on the

north side of Lake Skinner (Fig. 1, regions D, E). Observational evidence suggests

that these congeners prefer to construct their webs in CSS in areas where the

canopy cover is patchy and in vegetation types that provide natural spaces within

the shrub itself for web construction. Eriogonum fasciculatum (California buck-

wheat) appeared to be, by far, the most commonly used CSS shrub (refer to Levi

1977), generally occupied by M. grandiosa grandiosa in the spring and by M.

crassipes in the fall. Secondary web sites observed include dead mustard (where

dense) and S. mellifera (black sage). Of the major CSS shrub components, A.

californica (California sage) appeared to be the least frequently used (contra nat-

ural history reported by Levi (1977)).

Ant Predators: On various occasions in the spring of 1999, two gnaphosid

species and one theridiid species were observed attacking and feeding on ants at

MCBCP. A penultimate Callilepis gosoga male was observed attacking a har-

vester ant worker (Pogonomyrmex subnitidus Emery) approximately 3 m from

the harvester mound. While still in its penultimate instar, the C. gosoga male was

provided a second species of harvester ant, Messor andrei (Mayr) which it at-

120 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

tacked and fed on. The European congeneric species, Callile pis nocturna (L.), is

a known ant specialist (Heller 1974, 1976; in Foelix 1982).

The male of the gnaphosid species, Gnaphosa californica Banks, was observed

attacking a worker formicine (Formica francoeuri Bolton) in a fashion similar to

that of a wolf spider attacking large prey. In such an attack, the prey is bitten and

simultaneously wrapped with all eight legs of the spider.

The theridiid species, Steatoda fulva (Keyserling) is known to feed on harvester

ants (Hdlldobler 1970). This species was not collected during the course of either

this or the Diegan study but was subsequently collected at MCBCP. A penultimate

male was observed in the process of wrapping a struggling harvester ant (Po-

gonomyrmex subnitidus) within its web in a depression on the ant mound.

CONCLUSIONS

Species diversity within the majority of taxonomic levels examined was greater

in Diegan CSS than in Riversidian CSS, especially within the Theridiidae, Gna-

phosidae, and Salticidae. This, we believe, is largely due to the coastal climate,

which produces both more summertime moisture and moderate temperatures than

the corresponding inland areas and promotes the growth of a wider variety of

shrub species. For example, several of the theridiid species sampled in coastal

Diegan CSS are known primarily from coastal localities and/or inland canyons.

Such species include Dipoena atopa (Chamberlin), D. prona (Menge), Thymoites

expulsus (Gertsch & Mulaik), Theridion punctipes Emerton, and Wamba crispulus

Simon (Theridion intervallatum Emerton, synonym). The relatively flat topogra-

phy of SRCMR and absence of deep canyons, as well as dry summertime con-

ditions, may preclude the establishment of the species. Similarly, gnaphosids such

as Callile pis eremella Chamberlin, Drassyllus conformans Chamberlin, D. saphes

Chamberlin, and Zelotes gabriel Platnick & Shadab are generally known from

more mesic localities than those found at SRCMR. These species were collected

in coastal Diegan CSS but were not found during this study.

Both the greater number of non-native species (especially cosmopolitan species)

and the relatively greater density, per non-native species, in Diegan versus Riv-

ersidian samples, suggest that the coastal conditions provide a more hospitable

environment than the inland lowlands do for species that desiccate easily. We also

found proportionately greater percentages of both new species records and un-

described species for San Diego County (17% and 9.5%, respectively) than for

Riverside County (11% and 6.5%, respectively). These records likely reflect a

combination of both species immigration and inadequate sampling in the past.

Assuming that these variables are approximately equal in both CSS systems, the

proportionally greater percentages for San Diego County suggest that the Diegan

climate is more conducive to habitation by less adaptable forms than the more

extreme Riversidian climate. Proportionately fewer species can tolerate the xeric

summertime conditions and seasonal temperature extremes of the inland River-

sidian system.

ACKNOWLEDGMENT

We thank the following people: N. I Platnick and G. B. Edwards for their

assistance in determination of various species; M. Buffington and K. Campbell

for their help in fieldwork; S. Kensinger, L. Truong, B. Haubruge, S. Morita, M.

2001 PRENTICE ET AL.: SAGE SCRUB SPIDERS 121

Escobar, N. Davis, and M. Siddiqui for their assistance in sorting samples and

computer entry; J. Bethke, C. Dunning, M. Blua, and M. A. Patten for reviewing

earlier drafts of this manuscript; M. Misenhelter for creating the original version

of the map figure; and two anonymous reviewers for their valuable suggestions

to improve the quality of the manuscript. This research was funded by a grant

from the Metropolitan Water District of Southern California to R. A. R. (Agree-

ment 4602, Task Order 1) for arthropod studies at SRCMR.

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C. L. Koch (Araneae: Thomisidae). Can. Entomol., 96: 636-656.

Draney, M. L. 1997. Ground-layer spiders (Araneae) of a Georgia piedmont floodplain agroecosystem:

species list, phenology and habitat selection. J. Arachnol., 25: 333-351.

Edwards, G. B. 1979. Pantropical jumping spiders occurring in Florida (Araneae: Salticidae). Florida

Department of Agriculture and Consumer Services, Entomology Circular No. 100.

Foelix, R. E 1982. Biology of spiders. Harvard University Press. 306 pp.

Gertsch, W. J. 1958. The spider family Plectreuridae. Am. Mus. Novit., 1920: 1-53.

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Hagstrum, D. W. 1970. Ecological energetics of the spider Tarentula kochi (Araneae: Lycosidae). Ann.

Entomol. Soc. Am., 63: 1297-1304.

Heller, G. 1974. Zur biologie der ameisenfressenden spinnen Callilepis nocturna L., 1758 (Aranea,

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Heller, G. 1976. Zum Beutefangverhalten der ameisenfressenden Spinne Callilepis nocturna (Arach-

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Kirkpatrick, J. B. & C. EK Hutchinson. 1977. The community composition of Californian coastal sage

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Mexico (Araneae: Araneidae). Bull. Mus. Comp. Zool., 148: 185-238.

p22 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

Levi, H. W. 1981. The American orb-weaver genera Dolichognatha and Tetragnatha north of Mexico

(Araneae: Araneidae, Tetragnathinae). Bull. Mus. Comp. Zool., 149: 271-318.

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(Araneae, Gnaphosidae). Bull. Am. Mus. Nat. Hist., 174: 97-192.

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(Araneae, Pholcidae) in California. J. Arachnol., 18: 313-319.

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(Araneae, Mygalomorphae, Theraphosidae). J. Arachnol., 25: 137-176.

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scrub (Arachnida: Araneae). Pan-Pac. Entomol., 74: 181-202.

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28-92.

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Ecology, 62: 439-455.

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and the community/continuum debate. Vegetatio, 52: 3-19.

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Ent. Soc. Am., 67: 861-862.

Received 7 Mar 2000; Accepted 6 Jul 2000.

PAN-PACIFIC ENTOMOLOGIST

77(2): 123-125, (2001)

Scientific Note

A NEW AND UNUSUAL HOST RECORD FOR

HEMICOELUS GIBBICOLLIS (LECONTE)

(COLEOPTERA: ANOBIIDAE)

Hemicoelus gibbicollis (LeConte), referred to as the California deathwatch bee-

tle (Linsley, E. G. 1943. Pests and Their Control, 11: 11-14, 23—26.; Ebeling W.

1975. Urban Entomology, pp. 128—216, Univ. Calif. Div. Agric. Sci., Los An-

geles, California; Kramer R. D. 1997. Wood-Boring Beetles. In: Mallis, A. Hand-

book of Pest Control, Mallis Handbook and Training Company [8th ed.]. pp. 357—

391.), is the most economically significant anobiid infesting structures along the

Pacific Coast of North America (Suomi, D. A. & R. D. Akre. 1992. J. Econ.

Entomol., 85: 1188—1193.). In nature, H. gibbicollis larvae feed on a wide range

of softwoods and hardwoods (Knutson, L. V. 1963. Proc. Entomol. Soc. Wash.,

65: 177-195; see attached Host List). In a recent survey of structures in the Pacific

Northwest, Suomi and Akre (1992) (J. Entomol. Soc. Brit. Columbia, 89: 63-70.)

found that the primary wood species infested by H. gibbicollis was Douglas-fir,

Pseudotsuga menziesii (Mirbel) Franco, followed by western redcedar, Thuja pli-

cata Donn ex D. Don, and western hemlock, Tsuga heterophylla (Rafinesque)

Sargent. The high frequency of occurrence in P. menziesii probably reflects this

wood’s extensive use in home construction.

On 21 Nov. 1990, I inspected a home in San Francisco where the homeowner

had reported a beetle infestation of the joists and subflooring. During my inspec-

tion I noticed that the basement was extremely damp with standing water in

peripheral floor gutters. There were numerous emergence holes in the wooden

substructure of the home with pelleted frass in the galleries. This type of damage

in the San Francisco Bay Area is characteristic of colonization by H. gibbicollis.

The homeowner explained that vernal springs occurred in this area of the city

near the Presidio and this water contributed to the consistently high moisture in

the basement area. As evidence of the high population density of the insect, the

homeowner showed me the heavily infested wooden housing of an old television

set that had been kept in the basement (wood species unknown). I also noticed a

stand of bamboo that shaded the basement entry and further exacerbated the high

moisture conditions in the basement. Examination of the bamboo culms (stalks)

revealed many circular emergence holes, some of which contained dead anobiid

adults. The culms contained numerous frass-packed galleries indicative of larval

feeding and development. The fecal pellets were characteristic of H. gibbicollis

and dead adults removed from the bamboo were identified as H. gibbicollis. Spe-

cies-level identification of bamboo is not possible from the culms, but anatomical

analysis of the infested bamboo revealed that it was a species in a leptomorphic

(Type I) genus such as Arundinaria, Phyllostachys, Fargesia, Sasa, or Pseudo-

sasa, all of which are temperate climate bamboos (Liese, W. L. 1995. European

Bamboo Soc. Journal, May 6th, pp. 5—12).

Bamboo is frequently noted as a host for true powderpost beetles (Lyctidae)

(because the bamboo has a large pore size required for oviposition or because it

124 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(2)

“mea

~~,

LAs SY Me

Figure 1. Feeding damage to bamboo by (A) Lyctidae and (B) Hemicoelus gibbicollis (Anobiidae).

Arrows indicate the finely grained or flour-like frass produced by the lyctid and the pelleted frass

produced by the anobiid.

has a high starch content) and for false powderpost beetles (Bostrichidae) (because

of their abundance in the tropics where many bamboo products are derived) (Ebel-

ing 1975, Kramer 1997). Compared with lyctid-infested bamboo, which has a

finely grained or flour-like frass (Fig. 1A), H. gibbicollis-infested bamboo has

galleries packed with pelleted frass (Fig. 1B). A review of the literature indicates

that bamboo has never been reported before as a host for H. gibbicollis. It is

likely, in this instance, that the population density of H. gibbicollis was so high

in and around the structure that the beetles attacked this uncommon host. Addi-

tionally, H. gibbicollis is known to develop optimally when wood moisture con-

tent is between 14% and 17% (Suomi, D. A. & R. D. Akre. 1993. Pan-Pac.

Entomol., 69: 221—235.), which may be typical for sub areas and basements in

coastal northern California. Presumably, the high level of moisture in the wood

in the basement and in wood in the surrounding area contributed to colonization

of the bamboo as well.

Host List for Hemicoelus gibbicollis

Alder [Alnus spp.]*

Red alder [Alnus rubra Bong.]’

Blueblossom [Ceanothus thyrsiflorus Eschsh.]}’

Cherry [Prunus spp.]*

2001 SCIENTIFIC NOTE 125

Bitter cherry [Prunus emarginata (Dougl.) Walp.]*°’

Wild prune [Prunus subcordata Benth.]!

California hazel [Corylus californica (A. DC.) Sharp]’

Douglas-fir [Pseudotsuga menziesii (Mirbel) Franco]?*’

Fir [Abies spp.]*

White fir [Abies concolor (Gordon & Glendinning) Hildebrand]’

Grand fir [Abies grandis (Dougl. ex D. Don) Lindl.]’

Hemlock, Western [Tsuga heterophylla (Raf.) Sarg.]*’

Maple [Acer spp.]!*

Bigleaf maple [Acer macrophyllum Pursh]?®’

Oak [Quercus spp.}*

Interior live oak [Quercus wislizensii A. DC.]’

Redcedar, Western [Thuja plicata Donn ex D. Don]?

Redwood [Sequoia sempervirens (D. Don) Endl.]*°

Spruce [Picea spp.]*°

Willow [Salix spp.]*

Pacific or black willow [Salix lasiandra Benth.]’

Yew, Pacific [Taxus brevifolia Nutt.]’

Record.—USA. CALIFORNIA. SAN FRANCISCO CO.: ~2 km E. Mountain

Lake (in Presidio), Baker Street, San Francisco, 21 Nov 1990, S. J. Seybold,

Bamboo.

Acknowled gment.—I thank P. Klausen (University of Minnesota) for helping to

process the specimens of H. gibbicollis; D. L. Wood (University of California at

Berkeley) and B. J. Cabrera (University of Minnesota) for critical reviews of

earlier drafts of this report; Prof. Dr. W. Liese (Universitat Hamburg) for inden-

tification of bamboo; and D. Blackford for assistance with photography. Manu-

script preparation was supported by a grant from the California Department of

Consumer Affairs, Structural Pest Control Board awarded to V. R. Lewis (Uni-

versity of California at Berkeley) and SJS. Specimens of H. gibbicollis recovered

from bamboo were confirmed by Dr. R. E. White (USNM, retired) and deposited

in the California Academy of Sciences Entomology Collection, Golden Gate Park,

San Francisco, California 94118.

Steven J. Seybold, Departments of Entomology and Forest Resources, 219

Hodson Hall, 1980 Folwell Avenue, University of Minnesota, St. Paul, Minnesota

55108-6125.

Received April 15, 2000; Accepted July 6, 2000.

‘Chamberlain, W. J. 1960. Insects Affecting Forest Products and Other Materials. Oregon State

College Coop. Assoc., Corvallis. 159 pp.

* Ebeling, W. 1975. Urban Entomology. Univ. Calif. Div. Agric. Sci., Los Angeles, California

3 Furniss, R. L. 1938. Proc. Entomol. Soc. Brit. Columbia, 35: 5-8.

4 Furniss, R. L. & Carolin, V. M. 1992. Western Forest Insects, USDA For. Serv. Misc. Publ. No.

1339.

> Hatch, M. H. 1946. J. Econ. Entomol., 39: 274.

6 Keen, F P. 1952. Insect Enemies of Western Forests, USDA Misc. Publ. No. 273

7 Knutson, L. V. 1963. Proc. Entomol. Soc. Wash., 65: 177-195.

8 Suomi, D. A. & Akre, R. D. 1992. J. Entomol. Soc. Brit. Columbia, 89: 63-70.

PAN-PACIFIC ENTOMOLOGIST

71(2): 126, (2001)

Scientific Note

MALFUNCTION OF ELECTRICAL EQUIPMENT CAUSED

BY SOLENOPSIS AUREA WHEELER

(HYMENOPTERA: FORMICIDAE)

Fire ants of the genus Solenopsis, especially S. invicta Buren and S. xyloni

McCook, attack and destroy the efficacy of electrical equipment (Mackay, W., S.

Majdi, S. B. Vinson & C. Messer. 1990. Effects of the fire ant, Solenopsis invicta,

on electrical equipment. pp. 496-503. In Applied Myrmecology, A World Per-

spective, R. Vandermeer, K. Jaffe & A. Cedeno (eds.). Westview Press; Mackay,

W., D. Sparks & S. B. Vinson. 1990. Pan-Pac. Entomol., 66: 174-175). Although

ants appear to be attracted to electrical fields generated by such equipment (Mack-

ay, W., S. Majdi, J. Irving, S. B. Vinson & C. Messer. 1992. J. Kans. Entomol.

Soc., 65: 39-43; Mackay, W., S. B. Vinson, J. Irving, S. Majdi & C. Messer.

1992. Environ. Entomol., 21: 866-870), this may not occur (Slowik, T., H. Thor-

vilson & B. Green. 1996. J. Econ. Entomol., 89: 347-352). Control of these pests

in electrical equipment can often be relatively easy (Mackay, W., S. B. Vinson,

J. Irving, S. Majdi & C. Messer. 1991. Sociobiology, 19: 309-322).

We here describe damage caused by a third species of fire ant, the native S.

aurea Wheeler. This species infests intrusion monitors (or vibration sensors) in

Big Bend National Park, Texas. The ants were able to penetrate the O-ring on the

periphery of the monitor box, and gain access to the electronic sensors. Once in

the box, they become wedged in the electronic system and act as conductors of

false signals. The large numbers of ants in the systems cause malfunction of the

equipment and must be removed by technicians. The system, made by Spartan

Technology (Model No. 4411-PT) is powered by four 6-volt batteries and the

electronics are controlled by sets of dipswitches. The solution to this problem

appears to be the use of hard rubber O-rings. None of the older devices, with

hard rings, placed in the same general areas were infested. It is not clear what

attracts the ants, but it is suspected it is the electrical system of the unit. The unit

does not vibrate, it only senses the vibration.

Acknowled gment.—Our research was supported by Science and Resource Man-

agement, Big Bend National Park, by the Fire Ant Initiative of the Texas De-

partment of Agriculture, and by the Department of Biological Sciences of the

University of Texas.

W. P. Mackay,! A. E Van Pelt,” & I. Moreno,!!Fire Ant Laboratory, Department

of Biological Sciences, The University of Texas, El Paso, Texas 79968; 7203

Howell Place, Greensboro, North Carolina 27455.

Received 23 Aug 1999; Accepted 23 Dec 1999.

PAN-PACIFIC ENTOMOLOGIST

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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. intemational 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.

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THE PAN-PACIFIC ENTOMOLOGIST

Volume 77 April 2001 Number 2

Contents

WIESENBORN, W. D.—Inverse density-dependent parasitism of Opsius stactogalus Fieber

(Homoptera: Cicadellidae) by Gonatopus sp. (Hymenoptera: Dryinidae)

CLARKE, A. R. & M. P. ZALUCKI—Taeniogonalos raymenti Carmean & Kimsey (Hymenop-

tera: Trigonalidae) reared as a hyperparasite of Sturmia convergens (Weidemann) (Diptera:

Tachinidae), a primary parasite of Danaus plexippus (L.) (Lepidoptera: Nymphalidae)

AREFINA, T. I. & J. C. MORSE—New species of Ceraclea (Trichoptera: Leptoceridae:

Athripsodini) and a first record of Adicella (Trichoptera: Leptoceridae: Triaenodini) from

far eastern Russia

BRAILOVSKY, H.—A further contribution to the Australian systematics of the tribe Colpurini

(Hemiptera: Heteroptera: Coreidae: Coreinae)

PRENTICE, T. R., J.C. BURGER & W. R. ICENOGLE—Spiders from Riversidian coastal sage

scrub with comparisons to Diegan scrub fauna (Arachnida: Araneae)

SCIENTIFIC NOTES

SEYBOLD, S. J—A new and unusual host record for Hemicoelus gibbicollis (Leconte)

Coleoptera: Anobiidae

MAKAY, W. P,, A. F. VAN PELT & I. MORENO—Malfunction of electrical equipment caused

by Solenopsis aurea Wheeler (Hymenoptera: Formicidae)

61

68

71

79

90

123

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 77 July 2001 Number 3

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The Pan-Pacific Entomologist

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PAN-PACIFIC ENTOMOLOGIST

71(3): 127-143, (2001)

STUDIES IN THE EUPHORIINA OF THE AMERICAS

(COLEOPTERA: SCARABAEIDAE)

Il. STATUS OF NAMES IN EUPHORIA, TYPES AND

SYNONYMIES, WITH NOTES ON THE

SOUTH AMERICAN SPECIES!

ALAN R. HARDY

Insect Taxonomy Laboratory, Plant Pest Diagnostic Center,

California Department of Food and Agriculture,

3294 Meadowview Rd., Sacramento, California 95832

Abstract—The nomenclature of taxa used in the genus Euphoria is reviewed, and numerous

synonymies, based upon an examination of the available type material and approximately 20,000

specimens are proposed. Notes upon the types examined are presented, and lectotypes designated

where necessary. A listing of the 125 names and variant spellings is presented, along with their

current taxonomic status. Three entities are currently unrecognizable or unplacable.

Comments upon the six recorded South American species are presented with their synonymies.

Relevant literature is cited.

Key Words.—Coleoptera, Scarabaeidae, Cetoniinae, Euphoriina, Euphoria, nomenclature, types,

lectotypes.

Material used in this study was borrowed from numerous institutions and in-

dividuals (see Part I). Most of this has been recently returned, with labels reflect-

ing the taxonomic arrangement presented below. This paper is presented in order

to permit understanding of the changes proposed. In addition to the taxa listed

below, there are an additional half-dozen species yet to be described. The gemi-

nata species group is very confused and much more material is needed for a

satisfactory resolution of the numerous problems encountered. There are probably

a number of unrecognized, undescribed species included in the material examined

from this group, but the range of variability within all species couldn’t be deter-

mined with the material at hand. The conclusions presented below are based on

an evaluation of material as was considered adequate to indicate morphological

variability in the species involved.

The species groups proposed below are tentative, pending further analysis, but

serve to group species sharing morphological characters, although the variable

nature of the group makes definition difficult in some cases.

Literature citations given below are not comprehensive; only those dealing with

synonymies or nomenclatorial changes are cited with a few exceptions.

An attempt was made to locate and examine all primary types. Where types

were not located or examined, it has been so noted. No attempt was made to

examine types of infrasubspecific taxa.

‘Part I, Introduction and Generic Concepts, appeared in The Coleopterist’s Bulletin, 42(1): 1-9,

1988.

128 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

E{UPHORIA

FULGIDA GROUP

EUPHORIA FULGIDA FULGIDA

Cetonia fulgida Fabricius 1775: 48

Type locality “‘Carolina.”’ Type in British Museum [not seen].

Erirhipis fulgida; Burmeister 1842: 393.

Euphoria fulgida; Horn 1880: 405.

Scarabaeus turchesinus Voet 1778: 8. Type locality not given. Type lost ? NEW

SYNONYMY.

EUPHORIA FULGIDA LIMBALIS, NEW COMBINATION

Euphoria limbalis Fall 1905: 273. Type locality Enterprize, Florida. Type in MCZ

[seen].

EUPHORIA FULGIDA FUSCOCYANEA

Euphoria (Erirhipis) fuscocyanea Casey 1915: 303. Type locality ““Texas.”’ Type

in USNM [seen].

E. fulgida fuscocyanea; Brown 1930: 5.

EUPHORIA FULGIDA HOLOCHLORIS, NEW COMBINATION

Euphoria holochloris Fall 1905: 273. Type locality Fort Huachuca, Arizona. Type

in MCZ; Lectotype here designated: male, with the following labels: Ft. Hu-

achuca Ariz. [printed]/ holochloris TYPE [printed and handwritten]/ M.C.Z.

Type 24876 [printed and handwritten, red]/ Lectotype Euphoria holochloris Fall

1905 By A. R. Hardy 1977 [handwritten, red].

E. (Erirhipis) holochloris] Casey 1915: 302

EUPHORIA SEPULCRALIS

Cetonia sepulcralis Fabricius 1801: 156. Type locality “‘Carolina.”’ Type in Kiel

[not seen].

Euphoria sepulcralis; Burmeister 1842: 376.

Euphoria (Euphoria) sepulchralis [sic]; Casey 1915: 321.

Cetonia lurida Olivier 1789: 43 [nec Fabricius]; 1790: 422.

Euphoria lurida; Schaum 1849: 266 [placement of Olivier’s concept as sepul-

cralis].

Cetonia reich Gory & Percheron 1833: 58, 210. Type locality ‘“‘Mexique.’’ Type

female in Geneva [seen].

Cetonia reichii; Burmeister 1842: 376 [as synonym of sepulcralis]; Bates 1889:

364 [as synonym of leucographa]; Casey 1915: 321 [as synonym of sepul-

cralis |.

Euphoria (Euphoria) sepulchralis crinitula Casey 1915: 321. Type locality

‘Southern [linois at St. Louis, Missouri.’’ Type in USNM [seen]. NEW SYN-

ONYMY.

Euphoria (Euphoria) sepulchralis cuprascens Casey 1915: 322. Type locality

**Kansas (Medora).’’ Type in USNM [seen]. NEW SYNONYMY.

Euphoria (Euphoria) sepulchralis floridana Casey 1915: 321. Type locality ‘‘Flor-

ida (East coast, from Jacksonville to Palm Beach).’’ Type in USNM. Lectotype

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 129

here designated; male, with the following labels: 1) Jackson/ville Florida [hand-

written] 2) CASEY/bequest/1925 [printed] 3) TYPE USNM/48677 [Printed,

handwritten, red] 4) floridana/Csy [handwritten] 5) LECTOTY PE/Euphoria/se-

pulcralis/floridana Csy./By A. R. Hardy ’78 [handwritten, red]. 14 paralecto-

types. NEW SYNONYM Y.

Euphoria (Euphoria) sepulchralis kansana Casey 1915: 321. Type locality Man-

hattan and Muncie, Kansas. Type in USNM. Lectotype here designated; male,

with the following labels: 1) Manhattan/Ks [printed] 2) CASE Y/bequest/1925

[printed] 3) kansana-3/PARATYPE USNM/48675 [printed, handwritten, red] 4)

LECTOTYPE/Euphoria/sepulcralis/kansana Csy./By A. R. Hardy 1978 [hand-

written, red]. 15 paralectotypes. NEW SYNONYMY.

Euphoria (Euphoria) appalachia Casey 1915: 322. Type locality Maryland to

northern Alabama. Type in USNM. Lectotype here designated; male with the

following labels: 1) Southern Pines/V-NC/A. H. Manee 09 [handwritten, print-

ed] 2) CASEY/bequest/1925 [printed] 3) appalachia-2/PARATYPE USNM/

48679 [handwritten, printed, red] 4) LECTOTYPE/ Euphoria/ appalachia/ By

A. R. Hardy/ 1978 [handwritten, red]. 6 paralectotypes. NEW SYNONYMY.

Euphoria (Euphoria) scolopacea Casey 1915: 322. Type locality North Carolina

to Florida. Type in USNM. Lectotype here designated; male with the following

labels: 1) Black/ Mts. North Carolina/ [handwritten] 2) CASEY/ bequest/ 1925

[Printed] 3) TYPE USNM/ 48680 [printed, handwritten, red] 4) scolopacea/

Csy [handwritten] 5) LECTOTYPE/ Euphoria/ scolopacea/ CASEY=By/ A. R.

Hardy [handwritten, red]. 25 paralectotypes. NEW SYNONYMY.

Euphoria (Euphoria) oxysternum Casey 1915: 323. Type locality Indiana. Type

female in USNM. NEW SYNONYMY.

EUPHORIA SEPULCRALIS NITENS, NEW COMBINATION

Euphoria (Euphoria) nitens Casey 1915: 320. Type locality ““Texas.’’ Type in

USNM. Lectotype here designated: male with the following labels: 1) melan-

cholica/ Gory [handwritten, folded] 2) CASEY/ bequest/ 1925 [printed] 3)

TYPE USNM/ 48683 [printed, handwritten, red] 4) nitens/ Csy [handwritten]

5) Euphoria/ nitens/ Casey [handwritten] 6) LECTOTYPE/ Euphoria/ nitens

Csy./ By A. R. Hardy 1978 [handwritten, red]. (Casey identified three males

and seven females as being in the type series. There are currently 14 specimens

labelled as nitens in the Casey material at the USNM. One male specimen is

of the dark color form formerly known as E. kerni texana, and so is here

excluded from further consideration. Of the remaining 13, there are three males

and 10 females. The three males (specimens numbered 1, 11, and 13) will be

considered as part of the original cotypical series. The first specimen in the

series is without a data label, but bears the label “‘melancholica Gory’’, and

appears to be in Casey’s hand. The remaining 10 (females) are all indistinguish-

able as to which were a part of the original series, and all have the label ‘“Tex.”’

For this reason, only the two remaining males were labeled as paralectotypes.

EUPHORIA SEPULCRALIS LEUCOGRAPHA, NEW COMBINATION

Cetonia leucographa Gory & Percheron 1833: 58, 208. Type locality ‘“‘Mexico.”’

Type in Paris? [Dejean colln., Oberthur colln.? Not seen.]

Euphoria leucographa; Burmeister 1842: 375.

130 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

Euphoria (Erirhipis) leuacographa; Casey 1915: 305.

Euphoria limatula Janson 1881: 585. Type locality Guatemala. Type male in

Leiden [seen]. NEW SYNONYMY.

Euphoria yucateca Bates 1889: 365. Type locality ‘““Mexico, Temax in N. Yu-

catan.”’ Type in BMNH. Lectotype here designated: male with the following

labels: 1) SYN-/TYPE [white disc with blue edge, printed 2) Temax,/ N. Yu-

catan,/ Gaumer [printed] 3) B.C.A., Col., II(2)./ Euphoria/ yucateca [printed]

4) LECTOTYPE/ Euphoria/ yucateca/ Bates=By/ A. R. Hardy [handwritten,

red]. 8 paralectotypes. NEW SYNONYMY.

Euphoria submetallica Casey 1915: 319. Type locality “‘Panama (Nata).’’ Type

in USNM [seen]. NEW SYNONYMY.

EUPHORIA SEPULCRALIS RUFINA, NEW COMBINATION

Cetonia rufina Gory & Percheron 1833: 58, 206. Type locality “‘Mexico.”’ Type

in Dupont Collection (part of the Obertheur Collection), Paris Museum [seen].

Euphoria rufina; Burmeister 1842: 375.

Euphoria leucographa var. rufina; Bates 1889: 364.

Euphoria (Erirhipis) rufina; Casey 1915: 306.

Euphoria (Erirhipis) testacea Casey 1915: 305. Type locality ““New Mexico and

Arizona (Baboquivari, Santa Rita and Huachuca Mts.).’”’ Type in USNM. Lec-

totype here designated; male specimen with the following labels: 1) Huachuca

Mts./ July 1905 Ariz [printed]; 2) CASEY/ bequest/ 1925 [printed]; 3) testacea-

2/ PARATYPE USNM/ 48668 [printed, handwritten, red]; 4) Lectotype/ Eu-

phoria/ testacea Csy/ By A. R. Hardy 1978 [handwritten, red]. 7 paralectotypes.

NEW SYNONYMY.

Euphoria subtomentosa var. histrionella Schaeffer 1907: 74 [nomen nudum].

EUPHORIA FASCIFERA FASCIFERA

Euryomia fascifera Leconte 1861: 336. Type locality “‘Cape San Lucas.’ Type

male in MCZ [seen].

Euphoria fascifera; Horn 1880: 398, 404.

Euphoria (Euphoria) fascifera; Casey 1915: 324, 325.

EUPHORIA FASCIFERA TRAPEZIUM

Euphoria (Euphoria) fascifera trapezium Casey 1915: 324. Type locality “‘Ari-

zona (Cochise Co.).’? Type in USNM. Lectotype here designated; male with

the following labels: 1) San Bernardino/ Ranch 3750 ft. [printed]; 2) CASE Y/

bequest/ 1925 [printed]; 3) TYPE USNM/ 48684 [printed, handwritten, red];

4) trapezium/ Csy [handwritten]; 5) LECTOTYPE/ Euphoria/ trapezium/ Cas-

ey=By/ A. R. Hardy ’78 [handwritten, red]. 2 paralectotypes.

EUPHORIA CANDEZEI

Euphoria candezei Janson 1875: 375. Type locality ‘““Central America; Guatemala;

San Domingo, Chontales, Nicaragua.” Type male in Leiden [seen].

Euphoria belti Sharp 1877: 137. Type locality “‘Chontales, Nicaragua.’’ Type in

BMNH. Lectotype here designated: male specimen with the following labels:

1) SYN- / TYPE [blue edged disc, printed]; 2) Chontales/ Nicaragua/ T. Belt

[printed]; 3) Euphoria/ Belti/ Ind. typ./ D.S. [handwritten]; 4) B.C.A.,

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 131

Col.,II(2)./ Euphoria/ candezei [inverted, printed]; 5) LECTOTYPE/ Euphoria/

belti/ Sharp=By/ A. R. Hardy ’78 [handwritten, red]. 2 paralectotypes.

HERBACEA GROUP

EUPHORIA LESUEURI

Cetonia Lesueur Gory & Percheron 1833: 58, 212. Type locality “‘Mexique.”’

Type female[?] in Geneva [seen].

Euphoria Lesueuri; Burmeister 1842: 372.

Euphoria lesueuri; Schenkling 1921: 340.

Cetonia Latreille Gory & Percheron 1833: 58, 213. Type locality ““Mexique.”’

Type male in Geneva [seen].

Euphoria Latreillii; Burmeister 1842: 373.

Cetonia notulata Gory & Percheron 1833: 91, 244. Type locality ‘“‘Mexique.”’

Type male[?] in Leiden [seen].

EUPHORIA STEINHEILI

Euphoria Steinheili Janson 1878: 303. Type locality “‘Panama.’’ Type male in

Leiden [seen].

Euphoria steinheili; Bates 1889: 364.

EUPHORIA MONTICOLA

Euphoria monticola Bates 1889: 415; Schoch 1897: 474. Type locality ““Mexico,

Canelas in Sia. Madre de Durango.”’ Type female in Paris.

EUPHORIA WESTERMANNI

Cetonia Westermann Gory & Percheron 1833: 212. Type locality “‘Mexico.”’ Type

female[?] in Geneva [seen].

Euphoria Westermanni; Burmeister 1842: 373.

Euphoria westermanni; Bates 1889: 363.

EUPHORIA HERBACEA

Cetonia herbacea Olivier 1789: 35; Burmeister 1842: 393 [in the treatment of

Erirhipis| Type locality ’l?Amerique septentrionale.“* Type in Geneva? [not

seen].

Euphoria herbacea; Horn 1880: 399, 406.

Cetonia antennata Gory & Percheron 1833: 56, 177; Burmeister 1842: 391 [syn-

onymy with pubera]. Type locality ‘“‘Philadelphie.’’ Type female in Geneva

[seen].

Cetonia pubera Gyllenhal in Schoenherr 1817: 138, 53. Type locality “‘America

septentrionali.”’ Type in Uppsala [not seen].

Erirhipis pubera; Burmeister 1842: 391; Schaum 1844: 375 [synonymy with her-

bacea]; Horn 1880: 408 [synonym of herbacea].

Erirhipis (Euphoria) herbacea var. occidentalis Knaus 1916: 84. Type locality

‘““Crawford Co. and Cherokee Co., Kansas.’ Type in Lawrence Kansas [not

seen]. NEW SYNONYMY.

132 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

EUPHORIA SUBMACULOSA

Cetonia submaculosa Gory & Percheron 1833: 58, 211. Type locality ‘“‘Mexico.”’

Type female in Geneva [seen].

Erirhipis submaculosa; Burmeister 1842: 392.

Euphoria submaculosa; Bates 1889: 369; Schoch 1895: 125.

Euphoria (Erirhipis) submaculosa; Casey 1915: 304.

EUPHORIA HUMILIS

Euphoria humilis Blanchard 1850: 13. Type locality ‘‘Mexico.’’ Type female in

Paris [seen].

EUPHORIA IRIDESCENS

Cetonia iridescens Schaum 1841: 45. Type locality “‘Guatemala.’’ Type female

in Geneva [seen].

Erirhipis tridescens; Burmeister 1842: 388.

Euphoria iridescens; Bates 1889: 368.

EUPHORIA MYSTICA

Euphoria mystica Thomson 1878: 27. Type locality “‘Mexico.”’ Type male in Paris

[seen].

segmentata Guerin [in litt., ms. name?]

E{UPHORIA CHONTALENSIS

Euphoria chontalensis Bates 1889: 368. Type locality “Nicaragua, Chontales.”’

Type in BMNH. Lectotype here designated: specimen with the following labels:

1) SYN-/ TYPE [blue edged disc, press printed]; 2) Chontales/ Nicaragua/ T.

Belt. [press]; 3) B.C.A., Col., I1(2)./ Euphoria/ chontalensis, Bates [handwritten,

printed]; 4) LECTOTYPE/ Euphoria/ chontalensis/ Bates=By/ A.R. Hardy ’78

[handwritten, red]. 1 paralectotype [evidently 3 other specimens scattered].

EUPHORIA PRECARIA

Euphoria precaria Janson 1881: 583. Type locality “‘New Grenada’”’ [Colombia].

Type male in Leiden [seen].

Euphoria morosa Janson 1881: 582. Type locality “Quito, Ecuador.’’ Type male

in Leiden [seen]. NEW SYNONYMY. [Under the first revisor rule (Recom-

mendation 24A) page priority can be suspended in cases where improved sta-

bility would result. As first revisor, I select precaria as the valid name, since

it was used in the Biologia Centrali-Americana, and morosa has not been used

since it was originally described. ]

Euphoria (Erirhipis) comminuta Casey 1915: 307. Type locality “‘Chiriqui’’ [Pan-

ama]. Type female in USNM [seen].

INDA GROUP

E{UPHORIA INDA

Scarabaeus indus Linnaeus 1760: 352. Type locality “‘India.”’ Type in Uppsala

[fide Landin 1956: 12; not seen].

Trichius indus; Fabricius 1775: 40.

Cetonia inda; Pallas 1781: 25.

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 133

Erirhipis inda; Burmeister 1842: 389.

Euphoria inda; Horn 1880: 399, 407.

Euphoria Erirhipis inda; Casey 1915: 308.

Cetonia barbata Say 1823: 239; Burmeister 1842: 389 [under Erirhipis inda; new

synonymy?]. Type locality ‘““Council Bluffs on the Missouri and on the Arkan-

sas.’ Type probably destroyed.

Cetonia brunnea Gory & Percheron 1833: 63, 267; Burmeister 1842: 389 [under

Erirhipis inda; new synonymy?]. Type locality “‘l? Amerique boreale.’’ Type not

located. [There is a male specimen in Geneva which has been suggested as the

possible type. The label reads: Inda Linne var./ Amer. Bor./ W. Murray [hand-

written]. Horn & Kahle list a W. Murray Coleoptera and Lepidoptera collection

from an individual who died in 1885. Since Gory & Percheron was published

52 years before this date, it is conceivable, although not likely, that this spec-

imen was in europe prior to 1833. There is no evidence of correspondence with

J. LeConte, a contemporary, so it further seems unlikely that W. Murray was

very active. ]

Cetonia marilandica von Frolich 1792: 116; Burmeister 1842: 389 [under Erir-

hipis inda; new synonymy?]. Type locality unknown to me. Type in Munich?

[not seen].

Euphoria inda var. nigripennis Klages 1894: 198. Type locality “‘Pittsburgh, Pa.”

Type in Cornell [not seen]. NEW SYNONYMY.

Euphoria (Erirhipidia) inda nigripennis; Casey 1915: 309.

Euphoria (Erirhipidia) rufobrunnea Casey 1915: 309. Type locality ““New Mex-

ico (fort wingate), Arizona, Colorado (Boulder Co.) and Washington State.”

Type in USNM. Lectotype here designated: specimen with the following labels:

1) Ft. Wingate/ N. Mex. [printed]; 2) CASEY/ bequest/ 1925 [printed]; 3) ru-

fobrunnea 20/ Paratype USNM/ 48674 [handwritten, printed; red]; 4) Lectotype/

Euphoria/ rufobrunnea Csy/ By A.R./ Hardy 1978 [handwritten, red]. 19 par-

alectotypes. NEW SYNONYMY.

EUPHORIA HIRTIPES

Euphoria hirtipes Horn 1880: 398, 401. Type locality “‘Dodge co., Neb.’’ Type

in MCZ. Lectotype here designated: female[?] specimen with the following

labels: 1) Dodge Co. Nebr. [printed]; 2) Type no. 3708 Euphoria hirtipes G. H.

Horn [handwritten, printed; red]; 3) E. hirtipes Horn [handwritten, inverted;

appears to be Horn’s hand]; 4) Lectotype Euphoria hirtipes Horn 1880 by A.

R. Hardy 1977 [handwritten, red] [This specimen seems, on the basis of mea-

surements, to be the male as described by Horn]. 1 paralectotype.

Euphoriaspis hirtipes; Casey 1915: 298, 333.

EUPHORIA BASALIS

Cetonia basalis Gory & Percheron 1833: 64, 274. Type locality ‘““Mexique.”’ Type

male in Geneva [seen].

Euphoria basalis; Burmeister 1842: 380.

Euphoria (Euphoria) basalis; Casey 1915: 324, 325.

Euphoria (Euphoria) basalis crinicauda; Casey 1915: 324, 325. Type locality

““Mexico (Guerrero).’”’ Type in USNM. Lectotype here designated: male with

the following labels: 1) Guer. [printed]; 2) CASEY/ bequest/ 1925 [printed]; 3)

134 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

crinicauda-4/ PARATYPE USNM/ 48685 [printed, red]; 4) LECTOTYPE/ Eu-

phoria/ basalis/ crinicauda Csy./ By A. R. Hardy ’78 [handwritten, red].

HISTRIONICA GROUP

EUPHORIA HISTRIONICA

Euphoria histrionica Thomson 1878: 27. Type locality “‘Mexico.’’ Type male in

Paris [seen].

Euphoria (Euphorhipis) histrionica; Casey 1915: 315.

Euphoria histrionica var. sonorae Bates 1889: 374. Type locality ‘“‘Sonora.”’ Type

in BMNH [? not seen]. NEW SYNONYMY.

Euphoria sonorae; Casey 1915: 315.

Euphoria (Euphorhipis) scabiosa Casey 1915: 316. Type locality ‘Arizona

(Southern).’’ Type in USNM. Lectotype here designated: female with the fol-

lowing labels: 1) Ariz. [printed]; 2) CASEY/ bequest/ 1925 [printed]; 3) TYPE

USNM/ 48673 [handwritten, printed, red]; 4) scabiosa/ Csy [handwritten]; 5)

Euphoria/ scabiosa/ Casey [handwritten]; 6) LECTOTYPE/ Euphoria/ scabiosa

Csy./ By A. R. Hardy 1978 [handwritten, red]. 1 paralectotype. NEW SYN-

ONYMY.

EUPHORIA SUBTOMENTOSA

Cetonia tomentosa Steven [nomen nudum]. Mannerheim indicates that Cetonia

tomentosa Steven was never named by Steven, the name was given by Sturm,

which is how Dejean carries it (1837: 192). Because of homonomy with Cetonia

tomentosa Klug (= Anoplochilus tomentosus), Dejean cites the name as sub-

tomentosa Hope for replacement. This name evidently was never validated, and

subtomentosa should be credited to Dejean, not Mannerheim (see below).

Cetonia tomentosa Sturm [nomen nudum].

Cetonia tomentosa Gory & Percheron 1833: 64, 278 [preoccupied; nec Klug; see

Gory & Percheron 1833: 63, 139, 266]. Type locality “‘Mexique.’’ Type fe-

male[?] in Geneva [seen].

Cetonia subtomentosa Hope [nomen nudum?].

Cetonia subtomentosa Dejean 1837: 192 [replacement for tomentosa Sturm, =

tomentosa Gory & Percheron].

Erirhipis subtomentosa; Burmeister 1842: 390.

Euphoria subtomentosa; Bates 1889: 373, 416.

EUPHORIA DEVULSA

Euphoria devulsa Horn 1880: 398, 402. Type locality “‘San Antonio, Texas.”’

Type in MCZ. Lectotype here designated: male specimen with the following

labels: 1) Tex. [printed]; 2) Type No. 3709 Euphoria devulsa G. H. Horn [hand-

written, printed, red]; 3) E. devulsa Horn [inverted, handwritten; Horn’s hand?];

4) Lectotype Euphoria devulsa Horn 1880 By A. R. Hardy 1977 [handwritten,

red]. No paralectotypes seen.

Euphoria Usorhipina) devulsa; Casey 1915: 331, 332.

EUPHORIA LINEOLIGERA

Euphoria lineoligera Blanchard 1850: 13. Type locality “‘Mexico.”’ Type in Paris

[seen].

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 135

Euphoria Usorhipina) lineoligera “‘Bates’’; Casey 1915: 330.

Euphoria lineogera; Schoch 1895: 124 [error].

Euphoria xanthomelas Thomson 1878: 26. Type locality ‘““Mexico.”’ Type in Paris

[seen].

E{UPHORIA BIGUTTATA

Cetonia Bi-Guttata Gory & Percheron 1833: 64, 274. Type locality “‘Mexique.”’

Type female in Geneva [seen].

Euphoria biguttata; Burmeister 1842: 384.

Euphoria Usorhipina) biguttata; Casey 1915: 329.

Euphoria Usorhipina) biguttata binoculata Casey 1915: 330. Type locality “‘Mex-

ico (Guerrero).’’ Type in USNM. Lectotype here designated: male [?] with the

following labels: 1) Guer. [printed]; 2) CASEY/ bequest/ 1925 [printed]; 3)

TYPE USNM/ 48686 [handwritten, printed, red]; 4) binoculata/ Csy [hand-

written]; 5) LECTOTYPE/ Euphoria/ biguttata/ binoculata/ CASEY=by/ A.R.

Hardy ’78 [handwritten, red]. 2 paralectotypes. NEW SYNONYM Y.

Euphoria UIsorhipina) biguttata biplagiata Casey 1915: 330. Type locality “‘Gua-

temala.’’ Type in USNM. Lectotype here designated: male[?] with the following

labels: 1) G.T. [handwritten]; 2) CASEY/ bequest/ 1925 [printed]; 3) biplagiata-

2/ PARATYPA USNM/ 48687 [handwritten, printed, red]; 4) LECTOTYPE/

Euphoria/ biguttata/ biplagiata/ CASEY=By/ A.R. Hardy ’78 [handwritten,

red]. 2 paralectotypes. NEW SYNONYMY.

EUPHORIA CANESCENS

Cetonia canescens Gory & Percheron 1833: 64, 277. Type locality “‘Mexique.”’

Type female[?] in Geneva [seen].

Euphoria canescens; Burmeister 1842: 383.

Euphoria ([sorhipina) canescens; Casey 1915: 336.

Euphoria UIsorhipina) solidula Casey 1915: 329. Type locality “‘Guatemala”’ [la-

bel on type reads: Nebaj, Quiche, Guatemala]. Type male in USNM [seen].

NEW SYNONYMY.

E{UPHORIA LEUCOPYGE

Euphoria leucopyge Bates 1889: 366. Type locality ‘“‘Mexico, S.L.P; Alamos;

Ventanas in Dgo.”’ Type in BMNH. Lectotype here designated: specimen with

the following labels: 1) SYN-/ TYPE [printed, blue edged disc]; 2) Type [print-

ed, red edged disc, inverted]; 3) Hacienda de/ Bleados,/ S. Luis Potosi./ Dr.

Palmer. [printed]; 4) Euphoria/ leucopyge/ Bates [handwritten]; 5) B.C.A., Col.,

I(2)./ Euphoria [printed, inverted]; 6) LECTOTYPE/ Euphoria/ leucopyge/

Bates=By/ A.R. Hardy ’78 [handwritten, red]. 1 paralectotype.

Euphoria leucopyga; Schoch 1895: 125 [error].

EUPHORIA DIMIDIATA

Cetonia dimidiata Gory & Percheron 1833: 64, 275. Type locality piace

Type male in Geneva [seen].

Euphoria dimidiata; Burmeister 1842: 380.

Euphoria Usorhipina) dimidiata; Casey 1915: 331.

136 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

PULCHELLA GROUP

EUPHORIA PULCHELLA

Cetonia pulchella Gory & Percheron 1833: 64, 279. Type locality ““Mexique.”’

Type male in Geneva [seen].

Euphoria pulchella; Burmeister 1842: 381.

Euphoria pulchela; Schoch 1895: 123 [error].

Euphoria Usorhipina) pulchella; Casey 1915: 327.

Cetonia ferrugata Gory & Percheron 1833: 64, 276. Type locality “‘Mexique.”’

Type female[?] in Geneva [seen].

Cetonia montesuma Gory & Percheron 1833: 64, 277. Type locality ““Mexique.”’

Type from “‘Colln. Children”’’ not located, presumed lost.

Cetonia montezuma; Burmeister 1842: 382.

Cetonia children Gory & Percheron 1833: 64, 275. Type locality “‘Mexique.”’

Type from ‘“‘Colln. Children’? not located, presumed lost. NEW SYNONYMY.

Euphoria childrenii; Burmeister 1842: 382.

Euphoria childreni; Bates 1889: 416.

Euphoria Usorhipina) childreni; Casey 1915: 328, 331.

Cetonia thelasco Gory & Percheron 1833: 64, 279. Type locality “‘Mexique.”’

Type from “Colln. Children’’ not located, presumed lost. NEW SYNONYM Y.

Euphoria Usorhipina) thelasco; Casey 1915: 327.

FE{UPHORIA KERNI

Euphoria Cernii Haldeman 1852: 374. Type locality not given. Type male in

MCZ [seen] [unnecessary emendation of patronomic “‘Kern’’].

Euphoria kerni; Bates 1889: 374; Schoch 1895: 122.

Euphoria (Haplophoria) kerni; Casey 1915: 313.

Euphoria Clarkii LeConte 1853: 441. Type locality Texas. Type female in MCZ

[seen]. NEW SYNONYM Y.

Euphoria kerni var. Clarkii; Horn 1880: 402.

Euphoria texana Schaufuss 1863: 113. Type locality Texas [?]. Type location

unknown to me.

Euphoria (Haplophoria) texana; Casey 1915: 313.

Euphoria kerni var. texana; Schenkling 1921: 339.

Euphoria (Haplophoria) wichitana Casey 1915: 312. Type locality ‘“‘Kansas.”’

Type female in USNM [seen]. NEW SYNONYM Y.

Euphoria (Haplophoria) retusa Casey 1915: 311. Type locality ““Mexico.”’ Type

male in USNM [seen]. NEW SYNONYM Y.

Euphoria (Haplophoria) connivens Casey 1915: 312. Type locality ““Texas &

Arizona.”’ Type in USNM. Lectotype here designated: male specimen with the

following labels: 1) Tex [printed]; 2) CASEY/ bequest/ 1925 [printed]; 3) TYPE

USNM/ 48671 [handwritten, printed, red]; 4) connivens/ Csy [handwritten]; 5)

Euphoria/ connivens/ Casey [handwritten]; 6) LECTOT YPE/ Euphoria/ conni-

vens Csy/ By A. R. Hardy 1978 [handwritten, red]. NEW SYNONYMY.

EUPHORIA DISCICOLLIS, NEW COMBINATION

Stephanucha discicollis Dejean 1837: 192 [nomen nudum].

Stephanucha discicollis Thomson 1878: 28. Type locality “‘Amer. bor.’? Type

male[?] in Paris [seen].

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 137

Stephanucha atrata Dejean 1837: 192 [nomen nudum].

Stephanucha discicollis var. atrata Thomson 1878: 28. Type locality ‘“‘Amer. bor.”

Type in Paris[?] [not seen]. NEW SYNONYMY.

Euphoria aestuosa Horn 1880: 398, 400. Type locality ‘“‘Kansas.’’ Type female

in MCZ [seen]. NEW SYNONYMY.

Euphoria (Haplophoria) aestuosa; Casey 1915: 314.

Euphoriaspis aestuosa; Blackwelder 1939: 55.

GEMINATA GROUP

EUPHORIA GEMINATA

Cetonia geminata Chevrolat 1835: 107. Type locality ““Tampico.”’ Type male in

Leiden [seen]. [There are two examples in the Leiden Museum from the Chev-

rolat material, one labelled ‘‘Vera Cruz,” and the other ““Tampico.’’ The spec-

imen labelled ““Tampico”’ was considered to be the type.]

Erirhipis geminata; Burmeister 1842: 386.

Euphoria geminata; Bates 1889: 373.

Euphoria (Rhipiphoria) geminata; Casey 1915: 317.

Cetonia cheniaci Laporte; Burmeister 1842: 386. [Nomen nudum?]. Type locality

and location of type not known to me.

EUPHORIA LEPROSA

Euphoria leprosa Burmeister 1842: 379. Type locality ‘““Mexico.”’ Type location

unknown to me.

FE{UPHORIA EXIMA

Euphoria exima Bates 1889: 372. Type locality “‘Costa Rica.’’ Type in BMNH.

Lectotype here designated: male with the following labels: 1) SYN-/TYPE

[printed, blue edged disc]; 2) Type [printed, red edged disc, inverted]; 3) Costa

Rica [printed]; 4) Salle Coll [printed]; 5) 1340 [printed, green, inverted]; 6)

Euphoria/ exima/ Bates [handwritten]; 7) B.C.A., Col., I(2)./ Euphoria [printed,

inverted]; 8) LECTOTYPE/ Euphoria/ exima/ Bates=By/ A.R. Hardy ’78

[handwritten, red]. The other specimen from the cotypical series not located.

EUPHORIA SCHOTII

Cetonia vestita Gory & Percheron 1833: 63, 271 (nec Say 1825: 201). Type

locality “‘“Mexico’’ Dejean Collection, Oberthur collection, Paris [seen]. NEW

SYNONYMY.

Erirhipis vestita; Burmeister 1842: 387.

Euphoria vestita; Bates 1889: 371.

Cetonia irregularis Gory & Percheron 1833: 63, 272 (nec Olivier 1789: 6, 51,

62). Type locality ““Mexique.’’ Type in Hope collection, Oxford [not seen]?

Euphoria schotti LeConte 1853: 441. Type locality “‘Eagle Pass.’ Type in MCZ.

Lectotype here designated: male with the following labels 1) [red disc]; 2)

schotti 3 [handwritten]; 3) Lectotype E. schotti LeC. by A.R. Hardy 1977. 2

paralectotypes.

Euphoria (Erirhipidia) schotti; Casey 1915: 309, 318.

138 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

E{UPHORIA AVITA

Euphoria avita Janson 1881: 584. Type locality Guatemala. Type in Leiden Mu-

seum [seen].

Euphoria (Haplophoria) longula Casey 1915: 314. Type locality “‘Honduras (San

Pedro Sula).’’ Type in USNM [seen]. NEW SYNONYMY.

EUPHORIA FULVEOLA

Euphoria fulveola Bates 1189: 370. Type locality “‘Mexico, Zacualtipan in Hi-

dalgo, Oaxaca.”’ Type in BMNH. Lectotype here designated, specimen with the

following labels: 1) SYN-/TYPE [blue edged disc, press printed]; 2) Oaxaca,/

Mexico./ Hoege. [press printed]; 3) Euphoria/ fulveola/ Bates [handwritten]; 4)

B.C.A., Col., II(2)./ Euphoria [press printed, inverted]; 5) LECTOTYPE/ Eu-

phoria/fulveola/Bates=By/A.R. Hardy ’78 [red, handwritten]. 2 paralectotypes.

Euphoria nigriventris Bates 1889: 371. Type locality ‘“‘Mexico, Peras, Etla, Ja-

lapa.”’ Type in BMNH. Lectotype here designated: male with the following

labels; 1) SYN /TYPE [blue edged disc, press printed]; 2) Jalapa,/ Mexico./

Hoege [press printed]. 3) B.C.A., Col., If(2)./ Euphoria/nigriventris/Bates [press

printed, handwritten]; 4) LECTOTYPE/ Euphoria/nigriventris/Bates=By/A.R.

Hardy ’78 [red, handwritten]. 3 paralectotypes (4 missing). NEW SYNONY-

MY.

EUPHORIA QUADRICOLLIS

Euphoria quadricollis Bates 1889: 371. Type locality ““Durango City.’’ Type in

BMNH. Lectotype here designated, male with the following labels: 1) SYN-/

TYPE [blue edged disc, press printed]; 2) Type [red-edged disc, press printed,

inverted]; 3) Durango City/Mexico./Hoge. [press printed]; 4) Euphoria/ quad-

ricollis/ Bates 6 [handwritten]; 5) B.C.A., Col., II(2)./ Euphoria [press printed,

inverted]; 6) LECTOTYPE/ Euphoria/quadricollis/Bates=By/A.R. Hardy ’78

[red, handwritten]. Second specimen may be lost.

Euphoria arizonica Schaeffer 1907: 15. Type locality “‘Huachuca Mts., Az.’ Type

in USNM. Lectotype here designated, male with the following labels: 1) Type

[red, press printed, genitalia glued on]; 2) Palmerly/Cochise Co./July. Ariz.

[handwritten and press printed] 3) BROOKLYN/MUSEUM/COLL, 1929 [press

printed]. 4) Catal. No. /1351 [handwritten and press printed]; 5) Cotype/No.

42588/ U.S.N.M. [red, press printed and handwritten]; 6) Euphoria/arizonica/

Schaef. [handwritten]; 7) LECTOTYPE/ Euphoria/arizonica Schf./By A.R. Har-

dy 1979 [red, handwritten]. NEW SYNONYMY.

FE{UPHORIA ATRA

Euphoria atra Bates 1889: 371. Type locality ‘““Mexico, Peras, Oaxaca.”’ Type in

BMNH. Lectotype here designated, male with the following labels: 1) SYN-/

TYPE [blue edged disc, press printed]; 2) Oaxaca,/ Mexico/ Hoege. [press print-

ed]; 3) B.C.A., Col., If(2)./Euphoria/atra, Bates [press printed and handwritten];

6) LECTOTYPE/ Euphoria/atra/Bates=By/A.R. Hardy ’78 [red, handwritten].

1 paralectotype, one other BMNH specimen, believed to not be a part of the

co-typical series, labelled ‘“‘Oajaca’’ Mexico.

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 139

EUPHORIA CASSELBERRYI

Euphoria casselberryi Robinson 1937: 163. Type locality ‘“‘Davis Mountains Tex-

as.”” Type in USNM.

INCERTAE SEDIS

subguttata Burmeister

melancholica (Gory & Percheron) [= lurida or nitens?]

segmentata Guerin [ms. name’?]

SOUTH AMERICAN SPECIES

Of the six species of Euphoria (with recognized synonyms) recorded as re-

stricted to South America, one was not present in the material examined (E.

trivittata Nonfried), and is apparently known only from the type material. I felt

that a comprehensive review of the South American fauna should await the ac-

cumulation of more material. Several of the species examined (boliviensis Blan-

chard; lurida Fabricius; punicea Janson), have not been verified by the exami-

nation of type material. Some species show close afinities to species treated here.

E. punicea, as mentioned by Janson (1881: 584), is close to steinheili, and I

believe may be a synonym of that species. Euphoria boliviensis is related to

subtomentosa, and lurida to sepulchralis. Euphoria acerba Janson (type exam-

ined) is a valid species, resembling, as stated by the author, FE. leseuri Gory &

Percheron.

The South American species:

E. abreona Janson 1878: 304; Colombia

E. acerba Janson 1881: 582; Ecuador

E. boliviensis Blanchard 1850: 13; Bolivia

andina Schein 1950: 273

E. lurida (Fabricius) 1775: 49; Brazil, Argentina

adspersa (Weber 1801: 71)

bivittata (Gory & Percheron 1833: 58, 207)

fasciolata Eschscholtz 1822: 25

rufescens (Gory & Percheron 1833: 68)

sordens (Linnaeus 1790: 1573)

E. punicea Janson 1881: 584; Ecuador

E. trivittata Nonfried 1894: 130; Ecuador

STATUS OF NAMES IN EUPHORIINA

In the history of Euphoriina, there have been many names used in various

genera. The following list is presented as an aid to placement of these names.

abronea Janson valid- S. America

acerba Janson valid- S. America

adspersa Weber =lurida (in Blackwelder)

aestuosa Horn =discicollis Thomson

andina Schein =boliviensis (in Martinez 1957)

antennata Gory & Percheron =herbacea (Leng)

appalachia Casey =sepulchralis sepulchralis

areata Fabricius Stephanucha

arizonica Schaeffer =quadricillis

140 THE PAN-PACIFIC ENTOMOLOGIST

atra Bates

avita Janson

barbata Say

basalis Gory & Percheron

belti Sharp

biguttata Gory & Percheron

binoculata Casey

biplagiata Casey

bis pinis Bates

bivittata Gory & Percheron

boliviensis Blanchard

brunnea Gory & Percheron

californica LeConte

candezei Janson

canescens Gory & Percheron

casselberryi Robinson

cerni Haldeman

chapini Cartwright

cheniaci Laporte

childern Gory & Percheron

childreni Gory & Percheron

chontalensis Bates

clarki LeConte

comminuta Casey

connivens Casey

crinicauda Casey

crinitula Casey

cuprascens Casey

devulsa Horn

dimidiata Gory & Percheron

discicollis Thomson

exima Bates

fascifera LeConte

fasciolata Escholtz

ferrugata Gory & Percheron

flavoviridis Thomson

floridana Casey

fulgida Fabricius

fulgida var. schwarzi AUTHOR

fulveola Bates

fuscocyanea Casey

geminata Chevrolat

hera Burmeister

herbacea Olivier

hirta Fabricius

hirtella Linnaeus

hirtipes Horn

histrionella Schaeffer

valid

valid

=inda (in Leng)

valid

=candezei

valid

=biguttata

=biguttata

Stephanucha

=lurida

valid- S. America

=inda

= Oxycetonia jucunda

valid

valid

valid

=kerni

Chlorixanthe

= geminata (in Casey)

=pulchella

=pulchella

valid

=kerni

=precaria

=kerni

=basalis

=sepulchralis sepulchralis

=sepulchralis sepulchralis

valid

valid

valid

valid

valid

=lurida

=pulchella

Chlorixanthe

=sepulchralis sepulchralis

valid

=fulgida limbalis

valid

valid- ssp. fulgida

valid

Euphoriopsis

valid

=Tropinota

=Tropinota

valid

=sepulchralis rufina

2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES

histrionica Thomson

holochloris Fall

humilis Blanchard

inda Linnaeus

insignis Casey

iridescens Schaum

irregularis Gory & Percheron

kansana Casey

kerni Haldeman

latreillei Gory & Percheron

leprosa Burmeister

leseuri Gory & Percheron

leucographa Gory & Percheron

leucopyge Bates

limatula Janson

limbalis Fall

lineoligera Blanchard

longula Casey

lurida Fabricius

lurida Olivier

marilandica von Frolich

melancholica Gory & Percheron

montesuma Gory & Percheron

monticola Bates

morosa Janson

mystica Thomson

nigripennis Klages

nigriventris Bates

nitens Casey

notulata Gory & Percheron

occidentalis Knaus

oxysternum Casey

pilipennis Kraatz

precaria Janson

pubera Gyllenhal

pulchella Gory & Percheron

punicea Janson

quadricollis Bates

reichi Gory & Percheron

retusa Casey

refescens Gory & Percheron

rufina Gory & Percheron

rufobrunnea Casey

scabiosa Casey

schotti LeConte

scolopacea Casey

sepulchralis Fabricius

solidula Casey

valid

valid- ssp. fulgida

valid

valid

= Oxythyrea funesta

valid

= schotti

=sepulchralis sepulchralis

valid

=leseuri

valid

valid

valid- ssp sepulchralis

valid

=sepulchralis leucographa

valid- ssp. fulgida

valid

=avita

valid

=sepulchralis sepulchralis

=inda

=lurida or nitens

=pulchella

valid

=precaria

valid

=inda

=fulveola

valid- ssp. sepulchralis

=leseuri

=herbacea

=sepulchralis sepulchralis

Stephanucha

valid

=herbacea

valid

valid- S. America

valid

=sepulchralis sepulchralis

=kerni

=lurida

valid- ssp. sepulchralis

=inda

=histrionica

valid

=sepulchralis sepulchralis

valid

=canescens

141

142 THE PAN-PACIFIC ENTOMOLOGIST

sonorae Bates

sordens Linnaeus

steinheili Janson

subguttata Burmeister

submaculosa Gory & Percheron

submetallica Casey

subtomentosa Dejean

testacea Casey

texana Schauf [COMPLETE NAME]

thelasco Gory & Percheron

thoracica Casey

tomentosa mannerheim

trapezium Casey

trivittata Nonfried

verticalis Horn

vestita Gory & Percheron

vestita Say

westermanni Gory & Percheron

wichitana Casey

xanthomelas Thomson

yucateca Bates

=histrionica

=lurida

valid

Incertae sedis

valid

=sepulchralis leuacographa

valid

=sepulchralis rufina

=kerni

=pulchella

Stephanucha

= subtomentosa

valid- ssp. fascifera

valid- S. America

Stephanucha

= schotti

=Tropinota hirta

valid

=kerni

=lineoligera

=sepulchralis leuacographa

Vol. 77(3)

LITERATURE CITED

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West Indies, and South America. Part 2. Bull. Smith. Inst., 185: 189-341.

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von Frolich, J. A. 1792. Bermerkungen uber einige seltene Kafer aus der Insecten-Sammlung des Hr.

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familles naturelles de Latreille, la division des scarabées mélitophiles. 4 pp., 77 pls. Paris.

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2001 HARDY: EUPHORIA NAMES, TYPES AND SYNONYMIES 143

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397-408.

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(1758). (Col.). Entomol. Tidskr., 77: 1-17.

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Nonfried, A. E 1894. Beschreibungen neuer Lamellicorner, Buprestiden und Cerambyciden aus Cen-

tral- und Sud-Amerika. Entomol. Nachrichten, 20: 129-142.

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Received 10 Apr 2000; Accepted 4 Oct 2000.

PAN-PACIFIC ENTOMOLOGIST

71(3): 144-155, (2001)

A NEW SPECIES OF NOSERUS

(COLEOPTERA: ZOPHERIDAE) FROM MEXICO

MARIO GARCIA-PARIS!?, MILAGRO Coca-ABIA!, & GABRIELA PARRA-OLEA?

‘Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones

Cientificas, José Gutiérrez Abascal, 2. 28006 Madrid, Spain

*Museum of Vertebrate Zoology, 3101 Valley Life Sciences,

University of California, Berkeley, California 94720 USA

Abstract—A new species of the family Zopheridae, Noserus doyeni Garcia-Paris, Coca-Abia,

and Parra-Olea, NEW SPECIES, is described from the State of Nuevo Leén in México. The

species is characterized by the presence of a dense velvety cover of scaliform setae and by its

smooth non-prominent elytral tuberculation. The range of the genus Noserus extends along two

disjunct geographic areas. The first, which includes a group of poorly defined forms allied to N.

plicatus, is restricted to the Pacific regions of California. The second group includes two species,

N. doyeni, so far restricted to Cerro Potosi in the uplands of Nuevo Leon, and N. emarginatus

from Texas. The external appearance of the three species of Noserus is strikingly diverse, and

they may represent at least two independent phyletic lineages not necessarily sister to each other.

Key Words.—Insecta, Taxonomy, Coleoptera, Zopheridae, Noserus doyeni n. sp., North America,

México.

Resumen.—Se describe una especie nueva de la familia Zopheridae, Noserus doyeni Garcia-

Paris, Coca-Abia, and Parra-Olea, SPECIES NOVA, del Estado de Nuevo Leén en México. La

especie se caracteriza por el denso recubrimiento aterciopelado de sedas escamosas y por la poca

prominencia de los tubérculos elitrales, que sobresalen poco o nada de la superficie elitral. Las

especies del género Noserus se incluyen en dos grupos con distribuci6n geografica disjunta. El

primer grupo incluye un conjunto de razas poco definidas relacionadas con N. plicatus, y se

restringe a las regiones del Pacifico de California. En el segundo grupo se incluyen dos especies,

N. doyeni hasta el momento exclusiva del Cerro Potosi en las montafias de Nuevo Leé6n, y N.

emarginatus de Texas. El] aspecto externo de las tres especies de Noserus es sorprendentemente

diverso, de forma que el género podria incluir al menos dos linajes independientes, no necesa-

riamente hermanos.

The family Zopheridae, first proposed by Boving & Craighead (1931) on the

basis of larval characters, is currently integrated by two subfamilies, Usechinae

and Zopherinae and a few Australian genera, Docalis Pascoe, Cotulades Pascoe,

and Latometus Erichson, of difficult placement within either zopherid subfamily

(Doyen & Lawrence 1979, Lawrence 1994, Lawrence & Newton 1995). The

content and diagnosis of the zopherid subfamilies Usechinae and Zopherinae have

suffered relatively few modifications since Gebien’s World catalogues of Tene-

brionidae (1910, 1937). Only the addition of new species, the transfer of Exeniotis

Pascoe to the Tenebrionidae s. str., or the placement of some of Casey’s (1907a,

b) genera as junior synonyms of Zopherus Gray and Nosoderma Solier (Triplehorn

1972, Doyen & Lawrence 1979) are significant. The subfamily Zopherinae, a

relatively homogeneous group on external appearance, includes 65 species dis-

tributed in 8 genera. Keys to genera of New World Zopheridae, and generic

synonyms, have been provided by Doyen & Lawrence (1979), and the genus

Zopherus has been comprehensively revised by Triplehorn (1972), but no other

genera of Zopherinae have been critically revised.

The genus Noserus LeConte (1862), is characterized within the subfamily Zo-

pherinae by having the two apical segments of the antennae fused, the inner

2001 GARCIA-PARIS ET AL.: NEW SPECIES OF NOSERUS 145

surface of the femora of males with tuberosities, and by the presence of a shallow,

but marked short hypomeral groove that does not completely contain the antennae

while in repose (Doyen & Lawrence 1979). Until present, the genus included a

group of poorly defined species closely related to Noserus plicatus (LeConte),

1859, restricted to California (Casey 1907a, b), and Noserus emarginatus Horn

(1878), a well defined species known from Texas (USA) and Mexico (Garcia-

Paris et al. 2000).

Based on comparisons among specimens of the North American genera Mer-

alius, Noserus, Nosoderma, Phellopsis, Phloeodes, and Zopherus, and a series of

zopherine beetles collected at Cerro Potosi (Nuevo Leén, México), we believe

the latter share more traits with Noserus than with any other genera of Zopheridae,

including: (1) the presence of a short, shallow hypomeral groove that does not

completely contain the antennae while in repose, (2) the two apical segments of

the antennae fused, and (3) the presence of a narrow longitudinal area not covered

by setae in the ventral mid-line of each tarsal segment, which appear to be

grooved. These specimens do not match any known species of zopherine beetle

and therefore we include their description in the following paragraphs.

Depository Abbreviations—The following abbreviations are used to indicate

deposition of specimens: CAS (California Academy of Sciences, San Francisco,

USA), EM-UCB (Essig Museum of Entomology, University of California at

Berkeley, USA), IB-UNAM (Instituto de Biologia, Universidad Nacional Autén-

oma de México, México), MCZ (Museum of Comparative Zoology, Harvard Uni-

versity, Cambridge, USA), and MNCN (Museo Nacional de Ciencias Naturales,

CSIC, Madrid, Spain).

Methodology.—Dry mounted specimens were used for descriptions and draw-

ings. External sculpture and pubescence and dissected mouth parts were studied

with Scanning Electron Microscopy (SEM). Preparations of male and female gen-

italia were obtained from alcohol preserved specimens. Aedeagus and median

lobe were removed and studied at the dissecting microscope immersed in distilled

water, and with SEM. Female genitalia and genital segment were cleared with a

hot 5% KOH solution, progressively dehydrated in 70%, 90%, 95% and 100%

ethanol, and mounted in Euparal for examination at the dissecting microscope.

Drawings of the female genitalia were performed using camera-lucida.

NOSERUS DOYENI GARCiA-PARiS, COCA-ABIA & PARRA-OLEA NEW SPECIES

(Figs. 1—30)

Types.—Holotype: MEXICO: NUEVO LEON: Cerro Potosi, 2600—3200 m, 23

May 1996, M. Garcia-Paris and G. Parra-Olea: IB-UNAM. Paratypes (36 speci-

mens): 1—3: same data as holotype; 4: same data as holotype, except: MNCN; 5—

33: same data as holotype, except: 10,300 ft., 25 Sep 1975, J. Powell, J. Chemsak,

and T: Freilander: EM-UCB; 34—36: same data as holotype, except: 11,000 ft,

Galeana, 5 Aug 1938, H. Hoogstraal: CAS.

Diagnosis.—Zopherine beetle distinguished by its smooth, non-prominent dor-

sal and lateral tuberculation, completely covered by dense scaliform setae. Pos-

terior declivity of the elytra smooth, without tubercles. The two apical segments

of the antennae are fused. A shallow but marked short hypomeral groove that

does not completely contain the antennae while in repose is present on the ventro-

THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

146

Habitus of Noserus doyeni sp. nov. from Cerro Potosi (Nuevo Leon, México).

Figure 1.

2001 GARCIA-PARIS ET AL.: NEW SPECIES OF NOSERUS 147

the labial palp insertion region. Note the juxtaposed or slightly imbricate tegulae. Figure 3. Ventral

view of the mouth region showing the gular v-shaped incisions which accommodate the maxillary

basistipe. Figure 4. Ventral view of the mentum and labial palpi. Figure 5. Dense pubescence at

the mental margins. Figure 6. Sensory organs at terminal opening of the labial palp. Figure 7. Dor-

sal pubescence at the insertions of the labial palp.

lateral surface of the pronotum. Posterolateral corners of the pronotum smoothly

rounded, not angular.

Description—Habitus as in Fig. 1. Coloration uniformly castaneous, with most of the body closely

covered with squamiform, curved, golden to brown setae, with scattered hooked black to brown setae

particularly evident at the margin of the prothorax. Live specimens present a velvety, almost furry,

aspect. Tegument formed by irregularly shaped tegulae, juxtaposed or partially imbricated (Fig. 2).

Head inserted into the prothorax as far as the antennal insertions while in repose; epistoma medially

148 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

sani x

Figures 8-12. Buccal morphology of N. doyeni. Figure 8. Ventral view of the maxilla. Note the

pubescence of the lacinia and galea. Figure 9. Terminal segment of the maxillary palp. Figure

10. Terminal ovate fossa of the maxillary palp with internal sensory papillae. Figure 11. Ventral

view of the mandible. Note the shape of the incissor and molar lobes. Figure 12. Lateral view of

the mandible.

depressed, laterally inflated, densely covered by squamiform, golden setae. Gular suture arcuate; deep

gular v-shaped incisions, broadly opened, which accommodate the maxillary basistipe (Fig. 3). Men-

tum (prementum) subrectangular with arcuate posterior margin (labial suture), and smoothly emargin-

ate anterior edge (Fig. 4); ventral surface of the mentum setose with shallow broad fossae bearing

laterally a single long curved hair; mental margins densely setose (Fig. 5); labial palpi robust with a

broad terminal opening, showing the internal, non-protruding, sensory organs (Fig. 6); insertions of

the labial palpi with tuffs of dense long setae directed inwards towards the oral cavity (Fig. 7). Maxillae

with lacinia and galea densely setose (Fig. 8); maxillary palpi with non-terminal segments short and

robust, covered by setae on most of their dorsal surface; terminal segment of the palpi (Fig. 9) elongate,

2001 GARCIA-PARIS ET AL.: NEW SPECIES OF NOSERUS 149

Figures 13-17. Buccal morphology and antennae of N. doyeni. Figure 13. Dorsal view of the

labrum. Figure 14. Detail of the setiferous fossae on the dorsal surface of the labrum. Figure

15. Anterior edge of the labrum in ventral view. Figure 16. Antennal segments 7-11. Note the

fusion of the terminal segments and the protruding denuded area in segment 9. Figure 17. Detail of

the protruding smooth area in the 9th segment of the antenna.

fusiform, with scattered long setae and a terminal ovate fossa with internal sensory papillae (Fig. 10);

basistipe well developed, with a marked hump. Mandibles robust and strong, joined to the head by a

single point of articulation and several muscles; each mandible has a triangular base, and its mesal

surface is differentiated into a very sharp, distal toothed incisor lobe (Fig. 11) and a proximal molar

lobe flat and smooth, with poorly defined transversal grooves (Fig. 12). Labrum relatively small (Fig.

13), strongly punctured ventrally, with deep dorsal setiferous fossae (Fig. 14), and small setae at the

anterior edge (Fig. 15). Antennae with segments 2 and 4—8 subquadrate, wider than long, densely

covered by strong acute setae; segment 3 longer than wide; segment 9 conical, wider than the other

150 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

segments at its distal portion (Fig. 16), with a well marked smooth denuded area slightly protruding

anteriorly (Fig. 17); articles 10 and 11 fused forming a sub-sphaeric terminal segment (Fig. 16).

Prothorax subrectangular in dorsal outline, longer than wide, slightly wider at its first third, with

prominent anterior lobed lateral angles that cover genae and eyes; posterolateral corners smoothly

rounded, not prominent (Fig. 1); base of thorax arquate; dorsal surface completely covered with

squamiform, curved golden-brown setae; ventral surface of the prothorax densely covered by setae,

with occasional deep fossae, and a very short but marked hypomeral groove in the anterior portion

(Fig. 18). Scutellum hidden under the elytra, small and densely covered by postrate setae, each orig-

inating from basal granuli (Fig. 19).

Legs completely pilose except a smooth patch on the internal surface of the femora of males (Fig.

20); the surface of these black ovate tuberosities is smooth with fine pores unevenly dispersed (Fig.

21); ventral surface of tarsus with two rows of divergent setae that give a grooved appearance to the

tarsus (Fig. 22); terminal tarsal segment with strong claws (Fig. 23).

Elytra 1.6 to 1.8X longer than wide, with shallow humeral impressions, and faded posterior tuber-

culation, densely covered by squamiform golden brown setae (Fig. 24); dispersed small setiferous

fossae are almost completely obliterated by pilosity (Fig. 25); posterior elytral depressions absent;

sutural region depressed; apices rounded, not touching each other (Fig. 1).

Male genitalia including a well developed genital segment which corresponds to the eighth and

ninth abdominal sternites. Aedeagus with parameres dorsally fused, their apices pubescent and less

sclerotized than the rest (Fig. 26); phallobase ventrally opened and shorter than the parameres (P/Ph

ratio about 4). Median lobe long and thin (Fig. 27), its apex obliquely truncated and distally opened,

with the terminal portion narrowly rounded and with scattered deep fossae (Fig. 29). The endophallus

is a membranous sac that seems empty at low magnification.

The female genitalia (Fig. 30) consist of two ovaries, two lateral oviducts converging posteriorly

from the ovaries, a median oviduct (oviductus communis) receiving the lateral ducts anteriorly, the

gonopore opened to the exterior on the eighth abdominal segment, a spermatheca (receptaculum sem-

inis), two accessory glands, and the vagina, a tubular exit passage from the median oviduct to the

genital chamber. The eighth abdominal segment is well developed and it is associated with the genital

chamber which is modified in an organ formed of appendicular parts, serving for the deposition of

the eggs; this organ is not a real ovipositor but it can be protracted like a tube, having the opening

of the egg passage near its distal end; the functional ovipositor is composed of a pair of dorsal and

ventral elongate processes or threads, which give stiffness to this part of the female genitalia; the

ventral wall is longer than the dorsal and it has a sclerotized median rod in the middle and two distal

genital palpi with sensillae.

Length (elytra plus prothorax), 14.5-17.7 mm.

Variability.—Paratypes 5—33 are covered by a dark exudate that obliterates most of the scalation,

tuberculation, and fossae. Coloration of these specimens is brown-black with dorsal shiny reflections

from the exudate. Some specimens present small dorsal tubercles along the striae, completely covered

by golden setae.

Secondary sexual characters.—Males have a large oval black tuberosity com-

pletely free of setae on the interior surface of the femora.

Etymology.—This species is named to honor John T: Doyen whose contribu-

tions to the systematics of Neotropical and Neartic tenebrionoids have been sub-

stantial to present and future workers on the group, and who kindly provided us

with his material for the description of the new species.

Geographic Range.—Known only from the type locality, the Cerro Potosi in

Nuevo Leén, México.

Ecological Remarks.—The species was found under loose bark of dead pine

stumps, often in company of the plethodontid salamander Chiropterotriton priscus

Rabb 1956, at elevations ranging from 2600 to 3200 m.

Material Examined.—37 specimens of Noserus doyeni: The type series indicated above plus one

additional specimen used for the SEM study, MEXICO: NUEVO LEON: Cerro Potosi, 2600-3200

m, 23 May 1996, M. Garcfa-Paris and G. Parra-Olea: 1 specimen (MNCN).

Additional specimens of N. plicatus from California (n = 11; MNCN), N. emarginatus from Texas,

2001 GARCIA-PARIS ET AL.: NEW SPECIES OF NOSERUS 151

Figures 18-23. Taxonomic characters of N. doyeni. Figure 18. Ventral view of the posterior

portion of the head and the lateral area of the prothorax. Note the short hypomeral groove and the

ventral fossae. Figure 19. Detail of the pubescence on the scutellum. Figure 20. Oval tuberosity

on the internal surface of the femora of males. Figure 21. Detail of the surface of the femoral

tuberosity, showing the pores and the lack of pubescence. Figure 22. Ventral view of the tarsus.

Figure 23. Terminal claw of the tarsus.

including the holotype (n = 3; MCZ), and representatives of all the North American genera of Zo-

pherinae: Phloeodes, Nosoderma, Meralius, Phellopsis and Zopherus (CAS, EM, MNCN) were also

examined.

KEYS TO SPECIES OF ADULT NOSERUS

In the following key all the geographic forms restricted to California (Casey

1907a, b) and closely related to Noserus plicatus are included under that name.

152 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

Figures 24-29. Taxonomic characters of N. doyeni. Figure 24. Dorsal view of the elytral pubes-

cence at the disc region. Figure 25. Detail of the elytral pubescence. Note the density of the squam-

iform setae and the isolated long hair corresponding to the hidden dorsal setiferous foveae. Figure

26. Dorsal view of the parameri of the male genitalia. Note the pubescence on the lateral regions.

Figure 27. Lateral view of the median lobe of the male. Note the obliquely truncate apex. Figure

28. Distal opening of the median lobe. Figure 29. Detail of the deep fossae of the distal region of

the median lobe.

1. Elytral tuberculation reduced, not prominent to almost obliterate, and

completely hidden by dense pilosity. Posterior declivity of elytra

SII@OEl EM ery eo) os eater So et MM eg N. doyeni

*. Elytral tuberculation strong, especially prominent at the posterior half.

2001 GARCIA-PARIS ET AL.: NEW SPECIES OF NOSERUS 153

Figure 30. Female genitalia. Broken lines indicate the region is longer than shown.

Dorsal pilosity does not hide the posterior tubercules. Posterior decliv-

Wy (Ol Sly era Daal poe odes Se laa Ate ee Cees Av ek eee 2

2. Apex of the elytra deeply emarginate. Undersurface with obvious fossae

and without prominent tubercles, covered by dense squamiform pilosity

et Be MD, Se OIA «Nya eee Me, es are ER wn, 3, See N. emarginatus

Apex of the elytra not emarginate. Undersurface without large fossae,

with dispersed long setae placed at the base of rounded black tubercles

Ne eee ee aT es RO ee we ee ee ore eens ee N. plicatus

*

Discussion.—The range of the genus Noserus includes two disjunct geographic

areas. The first group is restricted to the Pacific regions of California. It includes

a group of poorly defined forms allied to N. plicatus, which do not deserve a

specific taxonomic status because most of their diagnostic characters are variable

features widespread along the geographic range of N. plicatus. The second unit

is distributed along the Atlantic regions of Texas (USA) and the State of Nuevo

Leén (México) (Garcfa-Paris et al. 2000). This group includes two species, N.

doyeni, so far restricted to a single mountain, the Cerro Potosi in the uplands of

Nuevo Le6n, and N. emarginatus from Texas. The Atlantic (Gulf of México)

species share a dense dorsal velvety vestiture and the presence of deep setiferous

fossae on the undersurface, not present in the Pacific N. plicatus.

Noserus doyeni with its furry vestiture and lack of dorsal and posterior elytral

tuberculation, is one of the most easily recognized species of the Zopherinae. The

single known population of this species may represent a relict of a wider distri-

bution, along the mountains of the Sierra Madre Oriental, that was progresively

restricted as a consequence of the Plio-Pleistocene aridification of the uplands of

Nuevo Leon (Braun 1955). The range of N. emarginatus covers part of the humid

lowlands of southern Texas, and montane forests in northeastern Mexico, but its

frequency of capture is very low as inferred from the scarcity of material available

in museum collections (Garcia-Paris et al. 2000).

Casey (1907b) divided the Zopherinae into two tribes, the Zopherini and the

Nosodermini. Casey’s (1907b) Zopherini included only Zopherus sensu Triplehorn

(1974), and Nosodermini included a highly heterogeneous assemblage of the re-

maining genera including Noserus. We follow Doyen & Lawrence (1979) and do

not consider justifiable to subdivide the Zopherinae at this point, since any pos-

sible arrangement probably requires the recognition of many monotypic tribes.

Noserus is morphologically close to the American genera Nosoderma, Meralius

154 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

and Phloeodes. These taxa share the fusion of the two apical segments of the

antennae and the presence of tuberosities on the internal surface of the femora of

males. The only apparently constant discriminating character among them is the

presence of a deep hypomeral groove that completely contains the antennae when

in repose in Phloeoedes, the complete absence of the hypomeral groove in No-

soderma and Meralius, and the presence of a shallow, but marked short groove

that does not completely contain the antennae in Noserus (Doyen & Lawrence

1979). The hypomeral groove in Atlantic Noserus is less developed than in the

Pacific species, and altogether, Noserus seems to be a transitional taxon between

Nosoderma and Phloeodes. The relevance of the hypomeral groove as a discrim-

inant character within Zopherinae is undisputable, however its phylogenetic rel-

evance is less clear because it is likely subjected to adaptive pressures, and it

seems to have evolved two or three times within the family, because it also occurs

in the American Zopherus Gray, and the Australian Zopherosis White, genera

apparently not closely related to each other nor to Phloeodes or Noserus (Doyen

& Lawrence 1979).

The external appearance of the three known species of Noserus is strikingly

diverse, and they may represent two independent phyletic lineages, the Pacific

and the Atlantic groups, rendering the genus Noserus possibly paraphyletic. The

presence of dense dorsal velvety vestiture, deep setiferous fossae on the under-

surface, and poorly defined tarsal grooves in Atlantic Noserus is shared by species

of Nosoderma. However Nosoderma is not a morphologically homogeneous taxon

(Casey 1907b, Doyen & Lawrence 1979), and any re-evaluation of the status of

Noserus must wait for the revision, currently in progress, of this diverse genus.

ACKNOWLEDGMENT

We are grateful to J. T. Doyen for kindly allowing the examination of his

material deposited at the Essig Museum, and to C. B. Barr (Essig Museum), D.

H. Kavanaugh and R. Brett (California Academy of Sciences), C. Martin (Museo

Nacional de Ciencias Naturales) and P. Perkins (Museum of Comparative Zool-

ogy) who helped us during the revision or provided invaluable material for study;

C. A. Triplehorn who also had recognized this species for his comments and

suggestions, EF Flores (México) who kindly helped us during field work, J. Bedoya

and E. Soteras (Museo Nacional de Ciencias Naturales) for the Scanning Electron

Microscopy services, and the excellent drawings that illustrate this work. We are

especially grateful to D. B. Wake who provided support during field trips, and B.

Sanchiz and A. Navas who provided facilities for the study of the material at the

Museo Nacional de Ciencias Naturales in Madrid. During part of this research

MG-P was supported with a postdoctoral fellowship of the Comunidad Aut6noma

de Madrid (Spain) and GP-O with a fellowship of the CONACyT (México).

LITERATURE CITED

Boving, A. G. & E C. Craighead. 1931. An illustrated synopsis of the principal larval forms of the

Order Coleoptera. Entomol. Amer. (n.s.), 11: 1-351.

Braun, E. L. 1955. The phytogeography of unglaciated eastern United States and its interpretation.

Bot. Rev., 21: 297-375.

Casey, T. L. 1907a. Notes on Chalcolepidius and the Zopherini. Can. Entomol., 39: 29-46.

Casey, T. L. 1907b. A revision of the American components of the tenebrionid subfamily Tentyriinae.

Proc. Wash. Acad. Sci., 9: 275-522.

2001 GARCIA-PARIS ET AL.: NEW SPECIES OF NOSERUS 155

Doyen, J. T. & J. E Lawrence. 1979. Relationships and higher classification of some Tenebrionidae

and Zopheridae (Coleoptera). System. Entom., 4: 333-377.

Garcia-Paris, M., G. Parra-Olea & M. Coca-Abia. 2000. First records of the genus Noserus LeConte

(Coleoptera: Zopheridae) in Mexico. Proc. Entomol. Soc. Wash., 102: 473-474.

Gebien, H. 1910. Tenebrionidae, X VIII, (I) pp. 1-166. Jn Junk, W. & S. Schenkling (eds.). Coleop-

terorum Catalogus, pars 15. W. Junk, Berlin.

Gebien, H. 1937. Katalog der Tenebrioniden (Col.: Heteromera). Teil I. Publicazioni del Museo En-

tomologico “Pietro Rossi”, 2: 505-883.

Horn, G. H. 1878. Contributions to the Coleopterology of the United States, No. 2. Trans. Amer.

Entomol. Soc., 7: 51-60.

Lawrence, J. F & A. EK Newton, Jr. 1995. Families and subfamilies of Coleoptera (with selected genera,

notes, references and data on family-group names). pp. 779-1006. Jn Pakaluk, J. & S. A.

Slipinski (eds.). Biology, phylogeny, and classification of Coleoptera: papers celebrating the

80th birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa.

Lawrence, J. EK 1994. The larva of Sirrhas variegatus, sp. nov. with notes on the Perimylopidae,

Ulodidae (stat. nov.), Zopheridae and Chalcodryidae (Coleoptera: Tenebrinoidea). Invert. Tax-

on., 8: 329-349.

LeConte, J. L. 1859. Catalogue of the Coleoptera of Fort Tejon, California. Proc. Acad. Nat. Hist.

Phil., 11: 69-90.

LeConte, J. L. 1862. Classification of the Coleoptera of North America. Smithsonian Misc. Coll., 136:

209-286.

Triplehorn, C. A. 1972. A review of the genus Zopherus of the World (Coleoptera: Tenebrionidae).

Smithsonian Cont. Zool., 108: 1-24.

Received 9 Dec 1999; Accepted 19 Feb 2001.

PAN-PACIFIC ENTOMOLOGIST

71(3): 156-160, (2001)

POLYTRICHAPHIS (HOMOPTERA: APHIDIDAE), A

NEWLY RECORDED GENUS FROM CHINA, AND

DESCRIPTION OF A NEW SPECIES

GEXIA QIAO AND GUANGXUE ZHANG

Institute of Zoology, the Chinese Academy of Sciences,

Beiing 100080 P. R. China

Abstract.—The aphid genus, Polytrichaphis Miyazaki, is newly recorded from China. A new

species, Polytrichaphis illicius Qiao and Zhang, is described. The type is deposited in the Zoo-

logical Museum, Institute of Zoology, the Chinese Academy of Sciences, Beijing, China.

Key Words.—Insecta, Aphididae, Macrosiphinae, Polytrichaphis, New Species, New Record,

China.

Some aphids infesting ///icium verum on young leaves of the tree were collected

in Beiliu County, Guangxi Autonomous Region, PR. China. Blackman and Eastop

(1984, 1994) listed no aphids on Jllicium verum. Miyazaki (1971) described one

new genus, Polytrichaphis collected from Illicium anisatum in Japan, which be-

longs to the member of Macrosiphinae. Careful examination revealed that the

present specimens belong to a new species, Polytrichaphis illicius. Because the

tree, [llicium verum is economically important in Guangxi Autonomous Region,

biological studies on the aphid will have important effects on agricultural pro-

duction of the tree.

To morphological characters, the genus Polytrichaphis Miyazaki is originally

described in the Macrosiphini because of the absence of lateral abdominal tuber-

cles and the presence of moderately developed antennal tubercles (Miyazaki 1971,

Remaudiére & Remaudiére 1997). However, Eastop (1979) thought that the genus

may even belong to the subtribe Aphidina but it seems more useful to include

than to exclude it. Therefore, regarding the systematic status of Polytrichaphis

much work needs to be done in the future. In this paper, the genus will be regarded

as a member of Macrosiphinae, because the distance between stigmal pori on 2nd

and 3rd abdominal segments is at least 3.10 times as long as the distance between

stigmal pori on Ist and 2nd abdominal segments (Eastop 1979).

The specimens were collected from Guangxi Autonomous Region (Beiliu City

130 m) by Jinyi Huang. All specimens were deposited in the Zoological Museum,

Institute of Zoology, the Chinese Academy of Sciences, Beijing, China.

The terminologies and descriptions follow that of Miyazaki (1971). The unit

of measurements is in millimeters (mm).

POLYTRICHAPHIS MIYAZAKI, 1971 NEW RECORD IN CHINA

Polytrichaphis Miyazaki, 1971, Ins. Mats., 34(2): 166.

Polytrichaphis Miyazaki: Eastop VE 1979, Syst. Ent. 4: 386; Remaudiére G and

M Remaudiere, 1997, Catalogue of the world’s Aphididae. Homoptera: Aphi-

doidea, 139.

Type Species.—Polytrichaphis fragilis Miyazaki, 1971, by original description.

Diagnosis.—Body oval or elliptical. Median front slightly developed; frontal

2001 QIAO & ZHANG: POLYTRICHAPHIS FROM CHINA 157

tubercles distinct, Antenna 6-segmented, longer than body; without secondary

rhinaria in apterous viviparous female; in alate viviparous females with round,

variable in size secondary rhinaria on 3rd—S5th segments. Mesosternal furca with

arms separated from each other. All first tarsal segments bearing 2 setae which

are much removed from apex of the segment. Media vein of fore wing 2-branched,

hind wing with two oblique veins; wing veins bordered with black bands. Si-

phunculus sub-cylindrical, weakly imbricated, with slightly spinulose shorter

stripes and distinct flange at apex. Cauda elongated tongue-shaped, non-constrict-

ed.

Host Plants.—Aphids in this genus feed on plants in the Illiciaceae.

Distribution.—China, Japan.

The genus is represented by two species as follow: Polytrichaphis fragilis.

Miyazaki, and new species P. illicius Qiao and Zhang. The genus is newly re-

corded in China, and there is one new species described from China in this genus.

POLYTRICHAPHIS ILLICIUS, NEW SPECIES

(Figs. 1-10)

Types.—Holotype, apterous viviparous female (Figs. 1-7), deposited in Zoo-

logical Museum, Institute of Zoology, the Chinese Academy of Sciences, P.R.

China, data: CHINA, GUANGXI AUTONOMOUS REGION, BEILIU Co.:

110.3° E, 22.7° N, 130 m, 27 Mar 1997, on JIlicium verum, by Jinyi Huang (No.

Y8243-1-5-2); paratypes: 14 apterous viviparous females and 18 alate viviparous

females, other data same as holotype.

Description of Apterous Viviparous Females——Body oval, black in life, 1.63—1.81 mm long, 0.93—

1.13 mm wide. In mounted specimens: distal of 3rd antennal segment, tip and base of 4th—Sth seg-

ments, basal of 6th segment darkly brown; basal 4%—% of siphunculi darkly brown, others pale brown.

Frontal tubercle slightly developed, antennal tubercles distinct, inner margins slightly diverging (Fig.

1). Dorsal hairs of body fewer, slightly thick and sharp. Head with 1 pair of cephalic hairs, 2 pairs of

antennal tubercular hairs and 6 or 7 dorsal hairs (Fig. 1). Length of cephalic hairs 0.05—0.07 mm,

1.85X-—2.4X as long as widest diameter of 3rd antennal segment. Antennae 6-segmented (Fig. 2),

slender, 1.51—-1.97 mm long, 0.88X-—1.15X as long as body; 3rd—6th segments with weak imbrications.

Proportion of 1st—6th segments: 26:17:100:91:97:40+ 157; processus terminalis 3.33 xX—4X as long as

basal part. Antennal hairs short and sharp, Ist-6th segments each with 5 or 6, 4, 4-7, 2-5, 3-5, 2+0

or | hairs, respectively; apex of processus terminal with 2 or 3 hairs; length of hairs on 3rd antennal

segment 0.02 mm, 0.63X as long as widest diameter of this segment. Primary rhinaria small rounded,

ciliated. Rostrum reaching mid-coxae; ultimate rostral segment wedge-shaped (Fig. 3), 1.71 X—2.18X

as long as its basal diameter, 1.7 as long as 2nd segment of hind tarsi, with 4-6 accessory hairs

among them. Thorax tergum membranous. Pronotum with 2 or 3 spinal, 1 pair of pleural and 1 pair

of marginal hairs; mesonotum with 2-3 pairs of spinal, 1 pair of pleural and 2 pairs of marginal hairs;

metanotum with 2 pairs of spinal, 1 pair of pleural and 2 pairs of marginal hairs. Hind femur 0.59—

0.71 mm long, 1.68X-—2X as long as 3rd antennal segment. Hind tibia 1.03-1.24 mm long, 0.6x-—

0.72X as long as body. Hairs on legs sparse, short and sharp; length of hairs on hind tibia 0.03—0.05

mm, 0.92X-—1.38X as long as middle diameter of the segment. First tarsal chaetotaxy: 2, 2, 2. Me-

sosternal furca with two arms separated, length of single arm, 0.08—0.11 mm, 1.33X-—1.83X as long

as 2nd antennal segment. Abdomen tergum membrane (Fig. 4); dorsal hairs on abdominal tergites

sparse and sharp; 1st—Sth abdominal tergites each with 3-5, 3 or 4, 2—4, 2 or 3 and 1-3 spinal hairs,

respectively; 1st—4th abdominal tergites each with 2-5, 4 or 5, 3-5, and 4-6 pleural hairs, respectively;

5th tergite without pleural hairs; 1st—Sth tergites each with 1 pair of marginal hairs, 6th tergite with

2 pairs of spino-pleural and 1 pair of marginal hairs; 7th tergite with | pair of pleural and 1 pair of

marginal hairs; 8th tergite with 4 or 5 hairs. Length of marginal hairs on Ist tergite and dorsal hairs

on 8th tergite 0.05—0.06 mm, 0.05-0.07 mm, 1.85X—2.22X and 1.85X-—2.41X as long as widest

diameter of 3rd antennal segment, respectively. Spiracles round and open, spiracular plates long oval,

158 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

Figures 1-7. Apterous viviparous females of Polytrichaphis illicius.

Figure 1. Dorsal view of head.

Figure 2. Antenna.

Figure 3. Ultimate rostral segment.

Figure 4. Dorsal view of abdomen.

Figure 5. Siphunculus.

Figure 6. Cauda.

Figure 7. Anal plate.

Figures 8-10. Alate viviparous females of Polytrichaphis illicius.

Figure 8. Third and fourth antennal segments.

Figure 9. Dorsal hair on abdominal tergites and hair-bearing sclerite.

Figure 10. Fore wing.

Scale bar: Figs. 1-8, 10 = 0.1 mm, Fig. 9 = 0.05 mm.

brown. Siphunculi sub-cylindrical (Fig. 5), with weakly spinulose short imbrications; 0.34—0.37 mm

in length, distal diameter 0.06 mm, length 3x—4.13%X as long as its basal width, 2X-2.12X as long

as cauda, 0.2X—0.22X as long as body. Cauda elongated tongue-shaped (Fig. 6), non-constricted, with

spinulose short imbrications, 0.16—0.18 mm in length, 1.42x-1.6X as long as its basal diameter, with

7-11 hairs. Anal plate circular at apex (Fig. 7), with spinulose short imbrications and 15-20 hairs.

Genital plate with weakly spinulose transverse imbrications and 12 or 13 hairs, 4 anterior hairs among

them. Gonapophyses three, each with 4 or 5 short sharp hairs.

Description of Alate Viviparous Females——Body long elliptical, black in life, 1.43-1.88 mm long,

0.53-0.95 mm wide. In mounted specimens: Head, thorax, Ist antennal segment, apex of rostrum,

2001 QIAO & ZHANG: POLYTRICHAPHIS FROM CHINA 159

Table 1. Measurements of apterous viviparous females (mm).

Body Body Hind Hind Ant. Ant. Ant. Ant.

No length width femur tibia III IV Vv VI Siphunculi Cauda

1 1.70 1.05 064 109 036 031 O33 014+ 052 035 0.16

pi 181 1.13 O65 1.13 0.36 033 O35 0.15 +058 0.36 0.18

3 1.75 1.06 0.67 1.24 036 032 035 O014+059 0.34 0.16

4 1.78 1.03 066 1.14 O35 O37 £420.38 0.15+059 037 0.16

5 1.63 0.95 0.59 103 0.30 0.26 027 £0.12 + 0.41 0.35 0.16

6 1.70 096 066 1.13 0.38 0.33 O36 O15 +056 0.36 0.18

7 1.73 0.93 0.66 1.14 0.36 0.31 0.33 0.14 + 0.55 036 =60..18

8 1.70 100 066 1.08 0.38 0.30 036 0.15 + 0.57 0.36 0.16

9 1.75 1.03 O71 1.19 0.38 0.32 O37 40.15 +060 0.36 0.18

10 163 0.98 061 103 031 O30 O31 O15 +054 0.34 0.16

Average 1.71 1.01 O65 1.12 O35 O32 034 O014+055 0.36 1.68

coxae, trochanters, basal % of siphunculi and veins dark brown, 7th-8th abdominal tergites with cross

bands across the segment; others pale brown.

Frontal tubercle slightly developed, antennal tubercles distinct, inner margins slightly diverging.

Head with 1 pair of cephalic hairs, 2 pairs of antennal tubercular hairs and 6 or 7 dorsal hairs; Antennae

6-segmented, 1.74—2.15 mm long, 1 X—1.25X as long as body, Ist antennal segment dark brown; 3rd—

6th segments with weak imbrications (Fig. 8); proportion of 1st—6th segments: 21: 18: 100: 80: 85:

34+ 139; processus terminalis 3.53xX—4.46X as long as its basal part. Antennal hairs much short and

sharp, lst—Sth segments each with 5 or 6, 4, 3-6, 2—4, 3-6 hairs, respectively; base of 6th segment

with 1 or 2 hairs, processus terminalis each with 0 or 1 hair; apex of processus terminalis with 3 hairs.

Length of hairs on 3rd segment 0.01—0.02 mm, 4%X—%X as long as widest diameter of this segment.

Secondary rhinaria rounded, variable in size, 3rd segment with 23-31 rhinaria and 4th segment with

10-19 rhinaria along their whole length, 5th segment with O or 1 rhinarium. Rostrum reaching me-

sothorax, ultimate rostral segment wedge-shaped, 0.1—-0.12 mm long, 1.57X-—2.2X as long as its basal

width, 1.43 X—1.71X as long as 2nd segment of hind tarsi, with 2-3 pairs of accessory hairs. Pronotum

each with 1 pair of spinal, pleural, and marginal hairs. Hind femur 0.52—0.67 mm long, 1.27xX—1.63 Xx

times as long as 3rd antennal segment. Hind tibia 0.98—1.19 mm long, 0.57 X-—0.69%X as long as body.

Hairs on legs sparse, short sharp. Length of hairs on hind tibia 0.02—0.03mm, as long as middle width

of the segment. First tarsal chaetotaxy: 2, 2, 2. Fore wing 2.25—2.85mm long, 1.31 X-—1.66X as long

as body (Fig. 10). Media vein 2-branched; hind wing with two oblique veins. Wing veins bordered

with black bands. 7th-8th abdominal tergites with spinulose short imbrications. Dorsal hairs of body

sparse, slightly short sharp, which on abdominal tergites with distinct hair-bearing sclerites (Fig. 9);

lst—7th abdominal tergites each with 4, 2, 2 or 3, 2 or 3, 2, 1 or 2 and 2 or 3 spinal hairs, respectively;

1st—4th tergites each with 2—4, 3-6, 4, and 5 or 6 pleural hairs, respectively, 6th tergite with 2 pleural

hairs, Sth and 7th tergites without pleural hairs; 1st—7th tergites each with 1 pair of marginal hairs;

8th tergite with 4 or 5 hairs. Length of marginal hairs on abdominal Ist tergite and dorsal hairs on

8th tergite 0.03—0.05 mm, 0.05—0.06 mm, 1X-—1.67X and 1.67X-—2xX as long as widest diameter of

3rd antennal segment, respectively. Siphunculi with weakly spinulose imbrications; 0.26—0.31 mm

long, 3.16—3.76X as long as its basal diameter, 1.86X—2.21X as long as its basal width. Cauda 0.13-

0.15 mm long, with spinulose short imbrications and 7-9 hairs. Anal plate with spinulose short im-

brications and 16-18 hairs. Genital plate with weakly spinulose imbrications and 13 or 14 hairs, 4

anterior hairs among them. Other data same as apterous viviparous females.

Diagnosis.—The new species is near to Polytrichaphis fragilis Miyazaki, but

differs from it as follow: basal 4%—% of siphunculi dark (the latter: dark brown in

wholly length); black in life (the latter: yellow to pale yellowish green in life);

8th tergite with 4 or 5 hairs (the latter: 9-13 hairs); genital plate with 4 anterior

hairs (the latter: 2); dorsal hairs on body shorter and fewer (the latter: tergum

abounding with long flagellate hairs) (Miyazaki 1971).

Etymology.—The new species is named for its infesting on Jllicium verum.

160 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

Table 2. Measurements of alate viviparous females (mm).

Body Body Fore Hind Hind Ant. Ant. Ant. Ant. Siphuncu-

No length width wing femur tibia Il IV Vv VI hi Cauda

1 1.70 0.70 2.25 0.61 1.10 0.38 0.34 035 0.15 +056 0.31 0.13

2 1.88 0.73 2.53 064 1.19 044 O41 O37 O.15 + 0.59 0.30 0.14

3 143 0.53 2.50 052 0.98 0.36 0.27 0.31 0.12 +053 0.28 0.13

4 185 095 2.85 0.67 1.13 043 0.39 O37 O15 +055 0.29 0.14

5 1.83 0.75 2.78 0.65 1.15 047 0.39 0.37 0.15 + 0.60 0.30 0.15

6 1.80 0.78 2.65 0.62 1.15 045 040 0.38 0.144 0.62 0.31 0.15

ei 165 065 2.40 057 41.01 O39 0.31 O31 O14 +054 0.26 0.13

8 1.71 0.78 2.50 0.62 1.12 037 0.36 0.37 0.13 + 0.60 0.28 0.15

9 1.70 0.75 2.55 0.64 1.11 040 0.33 O35 0.14 +053 0.26 0.14

10 163 0.75 245 062 1.13 036 038 0.36 014+ 0.59 0.30 0.14

Average 1.72 0.74 255 0.62 1.11 O41 O33 O35 014+ 057 0.29 1.14

ACKNOWLEDGMENT

We thank Jinyi Huang, for collecting specimens. The work is supported by a

grant from National Natural Science Foundation from P. R. China (Grant No.

39700015), a grant for special fund of systematic and evolutionary biology, the

Chinese Academy of Sciences to Yang Xingke, and a biological and technical

innovation grant from the Chinese Academy of Sciences (Grant No. C2999084).

LITERATURE CITED

Blackman, R. L. & V. E Eastop. 1984. Aphids on the world’s crops. An identification guide. The

Pitman Press, Ltd., Bath, Avon. John Wiley & Sons, VII: 466 pp.

Blackman, R. L. & V. E Eastop. 1994. Aphids on the world’s trees. An identification and information

guide. University Press, Cambridge, Center for Agriculture and Biosciences International VIII:

1024 pp.

Eastop, V. E 1979. Key to the genera of the subtribe Aphidina (Homoptera). Syst. Ent., 4: 379-388.

Heie, O. E. 1992. The Aphidoidea (Hemiptera) of Fennoscandia and Denmark. IV. Scandinavian

Science Press Ltd., Denmark, Fauna Ent. Scand., 25: 1-189.

Miyazaki, M. 1971. A revision of the tribe Macrosiphini of Japan (Homoptera: Aphididae: Aphidinae).

Ins. Mats., 34: 1-247.

Remaudiere, G. & M. Remaudieére. 1997. Catalogue of the world’s Aphididae. Homoptera: Aphidoidea.

Paris, Institut Natural de la Recherche Agronomique, 473 pp. (in French.)

Received 2 Sep 2000; Accepted 13 Feb 2001.

PAN-PACIFIC ENTOMOLOGIST

77(3): 161-167, (2001)

ABUNDANCE AND MATING BEHAVIOR OF ORIENTAL

FRUIT FLIES (DIPTERA: TEPHRITIDAE) NEAR

METHYLEUGENOL-BAITED TRAPS

ANDREI V. ALYOKHIN!?, RUSSELL H. MESSING?, & JIAN J. DUAN?24

‘Department of Entomology, University of Hawaii, Honolulu, Hawaii 96822

*Kauai Agricultural Research Center, University of Hawaii,

Kapaa, Hawaii 96746

Abstract—Methyleugenol-baited traps are used for Oriental fruit fly control through male an-

nihilation, as well as for detection and monitoring of fly populations. However, if the males

which come to these traps emit sex pheromones, attract females from the surrounding vegetation,

and mate with them before being killed, then using such traps might in fact increase levels of

fruit infestation. In the present study, we monitored fly abundance in an experimental orchard

before, during, and after methyleugenol-baited trap deployment. We also recorded the numbers

of flies recruited to the trees with and without traps, and quantified their sexual activity. The

males attracted by methyleugenol in our experiments fed on the poisoned baits almost imme-

diately upon their arrival, and did not attempt to emit pheromones or attempt copulations before

entering the traps. No changes in female abundance in the vicinity of deployed traps were

recorded. Because of their high specificity, low cost, and environmental safety, methyleugenol-

baited traps might be a valuable tool for integrated management of Oriental fruit fly populations.

Key Words.—Insecta, Bactrocera dorsalis, behavioral control, methyleugenol, mating, para-

pheromone.

Oriental fruit fly, Bactrocera dorsalis (Hendel), is one of the most important

insect pests of tropical horticulture, which causes direct damage to more than 150

species of fruits and vegetables (Christenson & Foote 1960, Haramoto & Bess

1970). Male (but not female) Oriental fruit flies are strongly attracted to methyl-

eugenol (4-allyl-1,2-dimethoxybenzene) (Steiner 1952). Feeding on this com-

pound significantly increases male mating success (Shelly & Dewire 1994). Meth-

yleugenol-baited traps have been widely used for Oriental fruit fly control through

male annihilation, as well as for detection and monitoring of fly populations (Cun-

ningham 1989). Male annihilation using methyleugenol has been successful in

eradicating Oriental fruit fly from several isolated geographic areas, including the

Marianas Islands (Steiner et al. 1965), the Amami Islands (Ushio et al. 1982),

and the Okinawa Islands (Koyama et al. 1984). It has also been used to uproot

the small “‘bridgehead”’ populations of the papaya fruit fly, Bactrocera papayae

Drew and Hancock, which invaded Queensland, Australia in 1995 (Lloyd et al.

1998, Meats 1998, Hadwen et al. 1998). However, eradication of the Oriental fruit

fly in Hawaii is hindered by the existence of large well-established fly populations,

often in hard-to-reach natural areas (Vargas et al. 1989, 1990). Because of the

high economic and environmental costs of fruit fly eradication, the goals of fruit

fly control programs in Hawaii are beginning to shift from complete annihilation

towards suppression of fly populations below economic thresholds in selected

fruit-growing areas (Mitchell & Saul 1990, Vargas et al. 2000).

With a more targeted approach to the Oriental fruit fly management, we should

3 Present Address: Department of Biological Sciences, University of Maine, Orono, Maine 04469.

4 Present Address: Monsanto Company, St. Louis, Missouri 63167.

162 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

anticipate an increased use of methyleugenol-baited traps for smaller-scale fly

control within individual orchards. Cunningham & Suda (1986) achieved a 48%

reduction in fruit infestation within a 63 ha papaya orchard by using only 9 small

fiberboard blocks saturated with a mixture of methyleugenol and malathion per 1

ha of orchard area. However, using methyleugenol-baited traps within even small-

er orchards (e.g., several ha) presents a potential danger. Male Oriental fruit flies

attract females by secreting airborne sex pheromones (Kobayashi et al. 1978).

They also display specialized courtship behaviors, expressed as rapid wing fan-

ning and rear leg movements over abdomen, probably in an attempt to facilitate

pheromone dispersal in the air (Arakaki et al. 1984, Shelly & Kaneshiro 1991).

Therefore, if males which come to traps emit sex pheromones, attract females

from surrounding vegetation, and mate with them before being killed, the use of

methyleugenol traps might in fact increase levels of fruit infestation in localized

areas. In the present study, we performed a series of experiments to investigate

the possibility of this occurring in small orchards in Hawaii.

METHODS AND MATERIALS

Experimental Sites—The study was conducted between 28 Apr. and 13 Jun.,

1999 at two sites on the Kauai Agricultural Research Center (Kapaa, island of

Kauai, Hawaii). The two sites were isolated from each other by approximately

700 m of predominantly non-host vegetation. Site 1 was a small orchard of 75

grapefruit (Citrus paradisi Macf.) and eight mandarin (C. nobilis Lour.) trees, and

Site 2 was a cluster of two shaddock (C. grandis (L.)), two grapefruit, and one

mandarin trees adjacent to a coffee grove. The grapefruit trees at Site 1 were

fruiting throughout the study, while no fruit was observed on any of the trees at

Site 2 after 15 May.

Trap Design.—The methyleugenol-baited traps used in our experiments con-

sisted of white plastic buckets (20 cm high, 20 cm in diameter) with 4 entry holes

(2.5 cm in diameter) each. A single cotton wick (15.5 cm in length, 1 cm in

diameter) was soaked in 10 ml of methyleugenol and 5 ml of a 10% water sus-

pension of malathion and hung by wire from the inner side of each bucket’s lid.

Buckets were hung on the trees approximately 1.6 m above the ground.

Experiment 1.—The objective of our first experiment was to determine if de-

ploying methyleugenol-baited traps leads to an increase in fly numbers in the

immediate surrounding area. Twelve 22.5 by 13.75 cm yellow plastic rectangles

(Trécé Inc., Salinas, California) covered with Tanglefoot were placed on trees at

random locations within Site 1 (1 rectangle per tree). On the 7th day after rect-

angle deployment, 1 methyleugenol-baited trap was placed on a tree in the middle

of the orchard for another 6 days, and then removed and taken to the laboratory

for examination. The rectangles were replaced and checked for flies every 2 days

for 18 consecutive days (6 days before placing the methyleugenol-baited trap in

the orchard, 6 days during which the trap was deployed, and 6 days after the trap

was removed). The numbers of captured male and female flies were recorded.

The location of each replacement rectangle was selected at random. The experi-

ment was repeated three times. Since the data did not follow a normal distribution

(Wilk-Shapiro test, n = 216, W = 0.5538, P < 0.05 for the number of males,

and W = 0.8680, P < 0.05 for the number of females), they were transformed

using rank transformations (Conover & Iman 1981). Transformed data were an-

2001 ALYOKHIN ET AL.: ORIENTAL FRUIT FLY BEHAVIOR NEAR TRAPS — 163

alyzed using a two-way ANOVA (Analytical Software 1996), with the period

during which the flies were captured (e.g., before, during, or after trap deploy-

ment) and replicates treated as main effects. When interaction terms were statis-

tically significant, we also analyzed treatment effects separately within each rep-

licate using one-way ANOVA.

Experiment 2.—In our second experiment, we investigated if there was any

increase in courtship or mating activity of Oriental fruit flies in the areas imme-

diately adjacent to methyleugenol-baited traps. One such trap was placed on a

tree within one of the two sites. The other site was used as a control. The number

of male and female flies, number of calling males, and number of mating pairs

were recorded on the tree with a trap, as well as on the 4 trees immediately

surrounding it. Observations began approximately 1 hour before sunset and con-

tinued for 40 minutes, thus covering the period of peak mating activity reported

for the Oriental fruit fly (Arakaki et al. 1984, Shelly & Kaneshiro 1991). Obser-

vations were repeated 10 times. Each site was used five times as a treatment, and

five times as a control. The numbers of flies recruited to the sites with and without

a trap were compared using two-sample Student f-tests. One-tailed Fisher exact

test (Zar 1999) was used to test the null hypothesis that the sex ratio of flies

encountered around the trap was not skewed towards males. Two-tailed Fisher

exact tests were used to test the null hypotheses that the sex ratio of flies observed

in the control site was not different from 1:1 (Analytical Software 1996).

Experiment 3.—The goal of the third experiment was to determine if Oriental

fruit fly males responding to methyleugenol arrive directly to the source of its

odor. Two neighboring fruit trees were selected at random for observations. The

distance between the trees was approximately 6.18 m (SE = 0.74). The number

of flies on these trees was visually observed immediately before the beginning of

the experiment, and their number was recorded. A methyleugenol-baited trap was

then placed on one tree. The number of flies on the trees with and without the

trap was then counted every 15 min for 1 h. Flies located on the outer or inner

trap walls were not included in the count. The experiment was repeated five times

at each of the two sites. Since the data did not follow a normal distribution (Wilk-

Shapiro test, nm = 100, W = 0.8301, P < 0.05), they were transformed using rank

transformations (Conover & Iman 1981). Transformed data were analyzed using

a three-way ANOVA (Analytical Software 1996), with site, time since trap place-

ment, and presence of the trap treated as main effects. Means and standard errors

were calculated from the non-transformed data only.

We also estimated the period of time between male arrival in the vicinity of a

methyleugenol-baited trap and subsequent entry into the trap. Since all the trees

within sites 1 and 2 had large canopies, preventing us from following the move-

ments of individual flies, a methyleugenol-baited trap was placed on a potted

guava tree (1.75 m tall) located approximately 10 m from Site 2. Time between

fly landing on the guava tree and its entering the trap was recorded for 10 flies.

The observations were repeated 4 times, with the behavior of a total of 40 flies

observed.

RESULTS

Experiment ].—Experimental results and ANOVA statistics are presented in

Table 1. Placing a methyleugenol-baited trap in the orchard significantly reduced

164 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

Table 1. Abundance of male and female Oriental fruit flies within Site 1 before, during, and after

deploying of a methyleugenol-baited trap.

Mean no. of males Mean no. of females

Replicate 1 Replicate 2 Replicate 3. Overall Replicate 1 Replicate 2 Replicate 3 Overall

Before trap placement 17.13 11.96 3.79 10.96 5.96 6.83 TAQ 6.86

(2.48) (1.71) (0.48) (1.20) (0.83) (0.98) (1.11) (0.56)

During trap placement 14.29 4.46 pA lg 6.97 5.50 10.79 5.88 F239

(1.51) (0.53) (0.33) (0.82) (0.81) (1.52) (0.85) (0.69)

After trap placement 15.33 4.88 3.46 7.89 3.88 8.63 7.13 6.54

(5.44) (0.73) (0.45) (1.91) (0.75) (1.19) (1.13) (0.63)

ANOVA

Df. 2, 69 2,69 2, 69 2207 2.69 25:60 2.69" 2,207

F 2.93 12.86 4.91 10.64 2.62 2.81 0.98 0.89

sm 0.0584 0.0001 0.0102 0.0001 0.0782 0.0653 0.3809 0.4148

the number of Oriental fruit fly males captured by yellow rectangles. There was

substantial variation in the extent of this influence as indicated by a highly sig-

nificant interaction term (ANOVA, df = 4, 207, F = 4.49, P = 0.0018). However,

the general trend towards a reduction in male numbers due to trap deployment

remained consistent within each replication. However, when the male population

in the orchard was the highest (Replicate 1), the difference in the number of males

caught before, during, and after trap deployment was not statistically different.

Unlike the numbers of captured males, female captures were not affected by

trap presence. There was a significant interaction between treatment and replica-

tion (ANOVA, df = 4, 207, F = 2.74, P = 0.0294), indicating different fluctuation

patterns in the numbers of females captured during each of the replications. How-

ever, separate analysis of treatment effects within each replication still did not

reveal any significant influence of deploying the methyleugenol-baited trap on the

number of females captured by yellow rectangles.

Experiment 2.—On average, 5.0 flies (SE = 0.98) encountered around meth-

yleugenol-baited traps were identified as males, and 2.6 flies (SE = 1.01) en-

countered around the traps were identified as females. At the control sites, ap-

proximately 3.0 flies (SE = 1.22) were identified as males, and approximately

2.7 flies (SE = 1.04) were identified as females. Sex ratio was significantly

skewed towards predominance of males in the presence of the trap (Fisher exact

test, x? = 3.89, df = 1, P = 0.0352), but did not differ from 1:1 in the absence

of the trap (Fisher exact test, x? = 0.08, df = 1, P = 0.8515).

Calling behavior was recorded for an average of 1.6 males (SE = 0.43) within

the treatment sites, and an average of 1.4 males (SE = 0.64) within the control

sites. The difference between the treatment and control sites was not statistically

significant (Student f-test, t = —0.26, df = 18, P = 0.7970). Only 4 mating pairs

were recorded during the entire study, with three of them being observed on a

single day. All the observed mating took place within the treatment sites. How-

ever, when we included both mating and calling males in our analysis, the dif-

ference between the treatment and the control sites still remained statistically

insignificant (Student f-test, t = —0.69, df = 18, P = 0.5001). No statistically

2001 ALYOKHIN ET AL.: ORIENTAL FRUIT FLY BEHAVIOR NEAR TRAPS — 165

significant difference in the number of recruited females was recorded between

the treatment and the control sites (Student t-test, t = 0.07, df = 18, P = 0.9460).

Experiment 3.—Experimental results are shown in Fig. 1. Overall, significantly

more flies were encountered at Site 2 than at Site 1 (ANOVA, df = 1, 80, F =

32.21, P = 0.0001), with the number of flies being significantly affected both by

the presence of a trap (ANOVA, df = 1, 80, F = 210.56, P = 0.0001), as well

as by the time since trap placement on a tree (ANOVA, df = 4, 80, F = 11.20,

P = 0.0001). Time trends were similar at both sites, as indicated by the absence

of a statistically significant interaction between the site and the time (ANOVA,

df = 4, 80, F = 0.94, P = 0.4451). At the same time, the interaction between

the time and the trap was highly significant (ANOVA, df = 4, 80, F = 12.07, P

= 0.0001), with a rapid increase in the fly numbers on the trees with a trap soon

after trap placement, and virtually no change on the trees without a trap (Fig. 1).

A three-way interaction among site, trap, and the time since the trap placement

was not statistically significant (ANOVA, df = 4, 80, F = 1.23, P = 0.3057).

On average, flies spent 66.37 seconds (SE = 10.05) on the tree before entering

the trap.

DISCUSSION

Results of the present study once again demonstrate that methyleugenol-baited

traps are highly attractive to Oriental fruit fly males. However, we did not find

any increase in fly sexual activity in the vicinity of the deployed traps. It appears

that the majority of males which are attracted to the odor of methyleugenol ap-

proach directly the source of the odor. Arrival rate curves observed in the present

study for the flies landing on the tree containing a trap are similar to the curves

reported by Nishida & Vargas (1990) for flies landing on their methyleugenol-

baited traps. The recruited males feed on methyleugenol almost immediately upon

their arrival and do not attempt to attract females and copulate before entering

the trap. Such behavior is not unexpected, since male Oriental fruit flies incor-

porate methyleugenol metabolites into their sex pheromone (Nishida et al. 1988),

and mating success of males fed on methyleugenol is significantly higher than

mating success of males deprived of this compound (Shelly & Dewire 1994).

Therefore, it is probably adaptive for males to ingest methyleugenol before trying

to attract a mate. Actually, 40-50% of males respond to methyleugenol baits even

before reaching full sexual maturity (Wong et al. 1989). Since there was no sig-

nificant increase in male calling around the traps, and by itself methyleugenol is

not very attractive to females (Steiner 1952), it is not surprising that we also did

not detect any changes in female numbers following the deployment of methyl-

eugenol-baited traps.

From a pest management perspective, it appears that using male lures on small

farms will not attract additional Oriental fruit fly females from the surrounding

vegetation, nor will it facilitate mating among the resident flies. Therefore, our

original concern that the use of methyleugenol traps might in fact increase the

level of fruit infestation appears to be unjustified. Still, it is unlikely that male

annihilation alone can provide a satisfactory level of Oriental fruit fly control on

small farms. This species is highly polygamous (Cunningham 1989). Therefore,

even very few surviving males will be able to fertilize a substantial number of

females, and each female can produce over 1000 eggs (Vargas et al. 1984). How-

166 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

Mean number of flies encountered

0 15 30 45 60

Time since trap deployment (min)

—¥— Tree without a trap

—@ Tree witha trap

Figure 1. Effect of methyleugenol-baited traps on recruitment of Oriental fruit flies to fruit trees.

The methyleugenol-baited trap consisted of a white plastic bucket with 4 entry holes and a single

cotton wick soaked in methyleugenol-malathion mixture and hung by wire from the inner side of the

bucket’s lid. Error bars represent standard errors of measurement among 5 replications.

ever, low cost, high specificity, and environmental safety of methyleugenol-baited

traps make them highly compatible with other management techniques, such as

sterile male releases, biological control, bait sprays of environmentally friendly

insecticides (e.g. spinosad), or visual lure-and-kill traps. Therefore, even though

male lures alone are unlikely to solve the Oriental fruit fly problem, they might

still be a valuable component of the future IPM systems.

ACKNOWLEDGMENT

We thank Bonnie Thorsby for providing technical assistance. This work was

supported in part by USDA-ARS Cooperative Agreement No. 59-5320-6-809.

This is Publication No. 4523 of the Hawaii College of Tropical Agriculture and

Human Resources Journal Series, University of Hawaii.

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eradication by male annihilation. J. Econ. Entomol., 58: 961-964.

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Received 5 Aug 2000; Accepted 13 Feb 2001].

PAN-PACIFIC ENTOMOLOGIST

71(3): 168-189, (2001)

NEW AND RARE SPECIES OF

PHYLLOPHAGA (S. STR.) FROM MEXICO

(COLEOPTERA: MELOLONTHIDAE, MELOLONTHINAE)

MIGUEL ANGEL MORON

Departmento de Entomologia, Instituto de Ecologia, A.C. (SEP-CONACYT)

Apdo. Postal 63, Xalapa, Veracruz 91000 México

Abstract.—Seven new species of Phyllophaga (s.str.) are described from Mexican localities as

follows: P. cahitana NEW SPECIES, from subtropical lowlands of northern Sinaloa; P. quetzala

NEW SPECIES and P. regiomontana NEW SPECIES from pine-oak forests of Nuevo Leén and

Tamaulipas; P. quetzaloides NEW SPECIES from temperate montane forests of northwestern

San Luis Potosi; P. stzotzilana NEW SPECIES from deciduous tropical forests of northwestern

Chiapas; P. tsajumiana NEW SPECIES and P. papaloana NEW SPECIES from pine-oak forests

of northern Oaxaca. Phyllophaga pilula (Moser) and P. nisuens Saylor, previously known only

by unique type specimens are redescribed with specimens collected in Chiapas and Oaxaca,

respectively. Drawings of male genital capsules, and tarsal claws are provided.

Key Words.—Insecta, May beetles, Phyllophaga, taxonomy, Mexico.

Resumen.—Se describen siete especies nuevas de Phyllophaga (s. str.) procedentes de seis lo-

calidades mexicanas: P. cahitana NUEVA ESPECIE de las tierras bajas del norte de Sinaloa,

P. quetzala NUEVA ESPECIE y P. regiomontana NUEVA ESPECIE de los bosques de pino-

encino del oeste de Nuevo Leon y Tamaulipas; P. quetzaloides NUEVA ESPECIE de los bosques

de montafia al noroeste de San Luis Potosi; P. stzotzilana NUEVA ESPECIE del bosque tropical

caducifolio del noroeste de Chiapas; P. tsajumiana NUEVA ESPECIE y P. papaloana NUEVA

ESPECIE de los bosques de pino y encino del norte de Oaxaca. Con base en ejemplares reco-

lectados en Chiapas y Oaxaca se redescriben Phyllophaga pilula (Moser) y P. nisuens Saylor,

previamente solo conocidas por ejemplares tipo tinicos. Se incluyen ilustraciones de las capsulas

genitales masculinas y de las ufias tarsales de todas las especies citadas.

Curatorial work of Phyllophaga specimens from Mexico, deposited in Canadian

or United States collections, as well as from recent collection trips to poorly

known localities in northwestern and southeastern regions of that country, pro-

vided specimens of many undescribed or little known species. Many of these

possess diagnostic sets of external characters and male genital capsules that are

very different from the known groups of species proposed by Moron (1986).

Some of these may be arranged in future new groups of species, but others remain

isolated, with uncertain position in the subgenus Phyllophaga (s. str.). Because

many of these species are rare or scarce, it seems more appropriate to describe

the new species, and redescribe the rare species while the new groups are config-

ured. This paper gives descriptions of males and some females of seven new

species of Phyllophaga (s. str.), and redescriptions and distributions of two species

previously known only by unique type specimens. The characters and terms used

in the descriptions are those of Sanderson (1958) and Moron (1986). Drawings

were made with the aid of a camera lucida and Leica stereomicroscope and mea-

surements were obtained with ocular micrometer or caliper.

Depository Abbreviations.—American Museum of Natural History, New York

(AMNH); California Academy of Sciences, San Francisco (CASC); Canadian

National Collection, Ottawa (CNC); Colegio de la Frontera Sur, San Crist6bal de

Las Casas, Chiapas, México (ECOSUR); Instituto de Ecologia, Xalapa, México

2001 MORON: NEW AND RARE PHYLLOPHAGA SPECIES 169

Figures 1-5. Phyllophaga (s.str.) cahitana Mor6n.

Figure 1. Male protarsal claw.

Figure 2. Female protarsal claw.

Figure 3. Male genital capsule, lateral view.

Figure 4. Paramera, distal view.

Figure 5. Female genital plates, ventral view. Scale lines: Figs. 1-2, 5 = 0.5 mm; Figs. 3-4 = 1

mm.

(EXA); Los Angeles County Museum of Natural History, California (LACM);

The Natural History Museum, London (BMNH); University of Nebraska State

Museum, Lincoln, Nebraska (UNSM); Zoologisches Museum, Humboldt Univ-

ersitat zu Berlin, Germany (ZMHU); H. & A. Howden/Canadian Museum of

Nature, Ottawa (HAHC); G. Nogueira, Guadalajara, México (GNGC) and M.A.

Moron, Xalapa, México (MXAL).

PHYLLOPHAGA (PHYLLOPHAGA) CAHITANA MORON, NEW SPECIES

(Figs. 1-5)

Types.—(Described from 6 males and 5 females). Holotype, male; MEXICO.

SINALOA: Los Mochis, 19 July 1960, W.W. Gibson; deposited: Canadian Na-

tional Collection, Ottawa. Allotype, female; same data as holotype deposited:

M.A. Mor6n collection, Xalapa. Paratypes: same data as holotype (4 females)

(CNC); Sinaloa, Los Mochis, 29 August 1997, G. Nogueira (2 males) (MXAL);

Sonora: Ciudad Obregon, 14 July 1960, W.W. Gibson (1 male) (HAHC); Obregon,

29 July 1952, C. & P. Vaurie (1 male) (AMNH); same data except 3 August 1952

(1 male) (AMNH).

Holotype.—Male. Head reddish brown, pronotum, elytra, sterna, pygidium and legs yellowish

brown, with silky luster. Clypeus wider than long (3.1:1), anterior border widely sinuated with margins

170 THE PAN-PACIFIC ENTOMOLOGIST Vol. 77(3)

moderately elevated, surface slightly convex, with many shallow, circular punctures, without setae.

Fronto-clypeal suture clearly sinuated and finely impressed. Frons wider than long (1.7:1) convex,

densely punctate, without setae. Antenna 10 segmented, with 3 segmented club; lamellae 1.3 longer

than the six preceding segments combined, segments 4° or 5° as long as segment 3, segments 6° and

7° wider than long, with acute process directed forward. Frons 4.4X wider than each eye dorsal

diameter. Canthus narrowed, curved and rounded, with 6 short setae. Labrum deeply bilobed, with

scattered setae on the borders. Mentum widely concave, polished, with scarce setae, anterior border

briefly notched. Pronotum wider than long (1.6:1) and 2.3X wider than frons. Pronotal disk with

round, shallow punctures regularly dispersed, separated by 1—3 diameters, without setae; lateral borders

widely angulated, marginal bead fine and irregularly crenulate, with scarce, short setae; anterior angles

obtuse, slightly pointed out; posterior angles obtuse, pointed out. Scutellum 1.4 wider than long,

with 10 shallow, small punctures. Elytron 2.2 longer than wide, densely punctate, without macro-

scopic setae; epipleural border narrowed, extended along the complete margin, with scattered, minute

setae; humeral calla rounded, prominent; apical calla rounded, diffuse. Metathoracic wings completely

developed. Propygidium with silky luster, fine and sparcely punctate, without setae. Pygidium slightly

convex, with silky luster, sparce and shallowly punctate, glabrous; apical margin with 10 short setae;

basal margin effaced from side to side. Pterosternum with scarce, short, yellowish vestiture. Visible

abdominal sternites 2° to 4° convex at the midline with scattered, short setae, sternite 5° convex, with

some short setae at the middline; anal plate shortened, slightly convex, with some, small punctures,

and anterior border slightly thickened, with 6 scattered setae near the posterior border. Protibiae shorter

than protarsi (0.8:1), with two big teeth and one small tooth on external border, preapical spur long,

straight, acute, shorter than 2° protarsomerus. Mesotibiae with one oblique, strong, setiferous, trans-

verse carina across external side, and one point with 2 short spines along dorsal border; upper apical

spur nearly straight, narrow, 1.2 longer than lower spur. Metatibiae nearly as long as metatarsi (0.9:

1), with one oblique, strong setiferous transverse carina across external side, and one point with 2

short spines along dorsal border; apical spurs articulated with border, upper spur curved, with rounded

apex, slightly shorter than 2° metatarsomerus, and 1.1 longer than lower spur. Tarsomeres semicy-

lindrical, elongated, with enlarged apex, some setae around the apex and two irregular lines of some

setae ventrally. Tarsal claws symmetrical, similar on all legs, with short teeth located near the middle

of ventral border (Fig. 1).

Genital capsule with short, wide paramera, fused at their basis, distal half widened and slightly

compressed, with apex briefly sinuated and nearly fused toward mid-line. Tectum convex. Aedeagus

long, with sclerotized narrow, tube-like support (Figs. 3-4). Length of genital capsule from the apex

of parameres to the border of basal piece: 2.6 mm. Total body length: 7.2 mm. Humeral width: 4.8

mm.

Alloty pe-—Female. Similar to the male except as follows: antennal club as long as the preceding

five segments; distal half of elytra more rounded; pygidial shape more triangular, disk less convex;

visible abdominal sternite 5° convex, with scattered setiferous punctures; anal plate convex, shallowly

punctate, with 8 short setae near the posterior border. All tarsi shorter than tibiae. Apical spurs of

metatibiae more wide and curvated than in male, with rounded apex. Tarsal claws with small teeth

located before the middle of ventral border (Fig. 2). Ventral genital plates strongly sclerotized, nearly

symmetrical, convex, with shallow punctures toward the sides, without setae; dorsal genital plates

fused, with wide, rounded distal borders provided with 6—7 setae on each side (Fig. 5). Total body

length: 11.2 mm. Humeral width: 5.1 mm.

Type Locality—Between Los Mochis, state of Sinaloa and Ciudad Obreg6n, state of Sonora, México

(25°47'—27°29' N; 109°0’—109°57' W).

Variation—Male paratypes are similar to the holotype except as follows: body color dark reddish

brown to light yellowish brown; pronotal lateral borders more or less rounded; pygidium with more

punctures; total body length: 7.2-13.5 mm, humeral width: 4.6-6.0 mm. Female paratypes are similar

to the allotype except by total body length: 7.0-14.6 mm; humeral width: 5.2—6.0 mm.

Biological Data.—Specimens of P. cahitana were collected at lights in open

deciduous tropical forest, thorn forest and xeric shrubs located from 10 to 50 m

of altitude, with 25—26° C mean annual temperature and 307-320 mm total annual

rainfall, where Prosopis juliflora (Swartz) DC, Acacia cymbispina Spregue et

Riley, Cercidium torreyanum (Wats.) Sarg. and Pithecollobium sonorae Wats

2001 MORON: NEW AND RARE PHYLLOPHAGA SPECIES 171

(Caesalpinaceae) are the most common trees. Phenology: July (8), August (3).

The other species of Phyllophaga flying at the same time and places were P.

(s.str.) fucata (Horn), and one undescribed species of P. (Listrochelus).

Remarks.—Phyllophaga cahitana resembles some species in the species group

fucata (sensu Mor6én 1986), by the external characters, as well as the shape of

clypeus, pronotum and elytra, style of punctuation, reduced vestiture, shape of

male tarsal claws and female dorsal genital plates, however, the compressed shape

of parameres is quite different from the short tube-like parameres of P. fucata or

P. opaca (Moser). Also, the tooth of the tarsal claws in female P. cahitana are

smaller than in P. fucata, and the female tarsi of the latter are longer than in P.

cahitana.

Etymology.—Derived from the indigenous people named ‘“‘Cahita’’ that actu-

ally live in some localities of Sonora and Sinaloa, speaking the language “‘cahita”’

(Tarahumara-Cahita subgroup of Sonoran languages), also known as “*Yaqui’’ or

‘“Mayo” (Manrique 1988).

PHYLLOPHAGA (PHYLLOPHAGA) QUETZALA MORON, NEW SPECIES

(Figs. 6-14)

Types.—(Described from 26 males and 49 females). Holotype, male; MEXICO.

NUEVO LEON, Monterrey, Mesa Chipinque, 26—29 August 1960, H. E Howden;

deposited Canadian National Collection, Ottawa. Allotype female; same data as

holotype; deposited A. & H. Howden/Canadian Museum of Nature, Ottawa. Par-

atypes: same data as holotype, except ‘‘on walnut’’ (7 males, 19 females) (CNC;

MXAL); Mesa Chipinque, nr. Monterrey, 8 July 1963, light, A. T. Howden (1

male) (HAHC); same data except 30 July 1963 (1 male, 1 female) (HAHC); near

Monterrey, Mesa Chipinque, 5400’, 30 July 1963, H. Howden (6 females) (CNC);

6 mi S Monterrey, 17 July 1963, H. & A. Howden (1 male) (CNC); Chipinque,

22 June 1971, H.R Howden (1 male, 1 female) (HAHC); 10 km S Chipinque,

4500’, 2 June 1983, UV, K. Kaulbars & R. Anderson (9 males, 20 females)

(HAHC; MXAL); TAMAULIPAS, Gomez Farias, 29 June 1969, deciduous forest,

S. & J. Peck, (1 female) (HAHC); Ciudad Victoria, Rancho La Reja, 30 July / 1

August 1981, 6000-8000’ B. C. Ratcliffe & C. L. Messenger (6 males, 1 female)

(UNSM; MXAL).

Holotype.—Male. Head and pronotum, shiny dark brown, elytra, pygidium, sterna and legs shiny

reddish brown, slightly pruinose, with iridescent luster. Clypeus bilobed, wider than long (4:1), anterior

border deeply sinuated with margins slightly elevated (Fig. 11), disk surface convex, with dense, round

punctures, and erect, short setae. Fronto-clypeal suture slightly curved, clearly impressed. Frons wider