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New records & species, & taxonomic alterations in 
milliped family Paeromopodidae (Julida) R.M. Shelley, S.B. Bauer 


A new species of Oxygonia (Coleoptera: Cicindelidae) 
from Ecuador F. Cassola, M.G. Kippenhan 


An unusual new Pachychernes (Pseudoscorpionida: 
Chernetidae) from Panama & Mexico William B. Muchmore 


A new species of Janus (Hymenoptera: Cephidae) 
from Indonesia David R. Smith 


Notes on mating behavior of Rhipiphorus luteipennis 
(Coleoptera: Rhipiphoridae) A.G. Wheeler, Jr. 


Aquatic Coleoptera & Hemiptera of Organ Pipe 
National Monument, AZ . E. Larsen, C. Olson 


Report of a morphologically hermaphroditic flea 
(Siphonaptera) and other flea anomalies from 
Morocco Michael W. Hastriter 


Sister relationship of the Neoephemeridae and Caenidae 
(Ephemeroptera: Pannota) T.-Q. Wang, W.P. McCafferty, Y.J. Bae 


A South Dakota record for Chauliodes rastricornis 
(Megaloptera: Corydalidae) P.J. Johnson, K.D. Roush, X. Lin 


Two noteworthy collections of ticks (Acari: I[xodida: 
Ixodidae) from endangered carnivores in the Lao 
People’s Democratic Republic R.G. Robbins, W.B. Karesh, 
S. Rosenberg, N. Schonwalter, C. Inthavong 


New host record for Ooencyrtus kuvanae (Hymenoptera: 
Encyrtidae) R.W. Hofstetter, K.F. Raffa 


SCIENTIFIC NOTE: First host record for genus 
Larissimus (Hymenoptera: Braconidae) A.M. Penteado-Dias 


Bark beetle (Coleoptera: Scolytidae) outbreak in 
pine forests of Sierra de las Minas Reserve, 
Guatemala R.A. Haack, G. Paiz-Schwartz 


ANNOUNCEMENT 
SOCIETY MEETING OF OCTOBER 23, 1996 


1 


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24 


29 


34 


43 


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76 


ENTOMOLOGICAL NEWS is published bi-monthly except July-August by The American 
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Vol. 108, No. 1, January & February, 1997 1 


NEW RECORDS AND SPECIES, AND TAXONOMIC 
ALTERATIONS IN THE MILLIPED FAMILY 
PAEROMOPODIDAE (JULIDA)! 


Rowland M. Shelley2, Selena B. Bauer2 


ABSTRACT: Diagnoses are presented for two new paeromopodid milliped species from California 
— Paeromopus paniculus, from the Merced River Valley, Mariposa County, and Californiulus blechro- 
striatus, from the eastern slope of the Sierra Nevada and the desert of Inyo County. Paeromopus 
buttensis is reduced to subspecific status under P. angusticeps because of a newly discovered inter- 
grade male from central Sonoma County and reinterpretation of P. ocellatus in western Solano 
County, which is comprised of anatomically intermediate forms and placed in synonymy under P. 
a. buttensis, new status. New localities of Californiulus dorsovittatus and C.yosemitensis in Lassen, 
Modoc, and Siskiyou counties connect the formerly disjunct Wamer Mountain population of the 
latter to the main ranges of the genus and family. The southern area of C. yosemitensis, extending 
from Mariposa to Kern counties, is segregated from that from Placer County northward because the 
species has never been collected in El Dorado, Amador, and Calaveras counties. The distribution of 
C. euphanus in western Washington extends eastward into the western periphery of the Columbia 
Plateau. 


The endemic west-Nearctic diplopod family Paeromopodidae (order Julida) 
includes the longest millipeds on the North American continent and is com- 
prised of two genera, Paeromopus Karsch and Californiulus Verhoeff. Accord- 
ing to Shelley (1994), it occupies a continuous area along the Pacific Coast and 
the Sierra Nevada and Cascade Mountains from central California to northern 
Washington, and occurs in the Warner Mountains, on the contiguous corners of 
California, Nevada, and Oregon, and from northeastern Oregon and southeast- 
ern Washington to western Montana. Paeromopus is restricted to California and 
extends along the Pacific Coast from Humboldt to Monterey counties and the 
western slope of the Cascades and Sierras from Shasta to Mariposa counties; 
Californiulus, however, inhabits all six states but is absent from coastal Califor- 
nia. The recent discovery of substantial new material among unsorted millipeds 
in Six institutions, including a new species from the California desert, prompts 
publication of supplemental distributional data. With new records from western 
Modoc, eastern Siskiyou, and northern Lassen counties, the disjunct area of the 
family and Californiulus in the Warner Mountains now connects with the rest 
of the range in northern California; the ranges of these taxa also expand longitudi- 
nally in the south into the deserts of central Inyo County, California, and in the 
north into the western periphery of the Columbia Plateau, in Yakima County, 


1 Received June 14, 1996. Accepted July 2, 1996. 
2 North Carolina State Museum of Natural Sciences, P. O. Box 29555, Raleigh, NC 27626-0555. 


ENT. NEWS 108(1) 1-14, January & February, 1997 \\\1 HSON/Z 


NV 


MAR 2 6 1997 


SIBRARIESD 


2 ENTOMOLOGICAL NEWS 


Washington. As a male is now available, we diagnose a new species for the 
previously unassignable females of Paeromopus from Mariposa County, Cali- 
fornia, and we reduce P. buttensis Chamberlin to subspecific status under P. 
angusticeps (Wood) based on an anatomically intermediate male from Sonoma 
County and reinterpretation of P. ocellatus Loomis, which we place in syn- 
onymy under P. a. buttensis. No additional samples are available for P. eldoradus 
and P. cavicolens, both by Chamberlin, C. chamberlini (Brolemann), and C. 
parvior (Chamberlin), so their accounts are as published by Shelley (1994). 


Acronyms of sources of the new material are as follows: 


AMNH - American Museum of Natural History, New York, New York. 

BYU - Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah. 
CAS - California Academy of Sciences, San Francisco. 

FSCA - Florida State Collection of Arthropods, Gainesville. 

LACMNH - Los Angeles County Museum of Natural History, Los Angeles, California. 
NCSM - North Carolina State Museum of Natural Sciences, Raleigh. 

UCB - Essig Museum of Entomology, University of California at Berkeley. 

UCD - Bohart Entomological Museum, University of California at Davis. 


Paeromopus angusticeps (Wood) 


On the basis of new material, we interpret P. ocellatus and a male from a 
geographically intermediate locality in Sonoma County as intermediate between 
P. angusticeps and P. buttensis, so the two most widespread forms of Paeromopus 
are races of a single species. Consequently, the genus is comprised of one widely 
distributed and three localized species. 

Distribution. With this taxonomic change, the range of P. angusticeps con- 
sists of two branches, one extending along the Pacific Coast from Humboldt to 
northern Monterey counties and the other extending along the western slope of 
the Cascades and Sierra Nevada from Shasta to Tuolumne counties (Fig. 11). 


Paeromopus angusticeps angusticeps (Wood), NEW STATUS 
Figs. 1-2 


The southern subspecies, P. a. angusticeps occurs around San Francisco/ 
San Pablo and Monterey Bays. The distributional limits are Mill Valley in south- 
ern Marin County, Castro Valley, Alameda County, and the literature record 
from Pacific Grove, Monterey County (Chamberlin 1941). Material was exam- 
ined from the following new localities; the initials JSB and MRG in this and 
succeeding accounts denote samples collected by J. S. Buckett and M. R. 
Gardner. 


Vol. 108, No. 1, January & February, 1997 3 


CALIFORNIA: Santa Clara Co., 6 mi (9.6 km) SE Holy City, M, 29 December 1966, 
MRG, S. E. Harrison (UCD). Santa Cruz Co., 2 mi (3.2 km) NW Boulder Cr., M, 29 Decmber 
1966, MRG, S. E. Harrison (UCD), 3 mi (4.8 km) NNE Soquel, F, 21 April 1972, C. E. 
Griswold (UCB); Santa Cruz, Empire Cv. on Univ. Calif. Santa Cruz campus, F, 7 August 
1962, R. Graham (FSCA); and Mt. Madonna E of Watsonville, M, 4 July 1958, W. J. Gertsch, 
V. Roth (AMNH). 


Remarks. The female from Empire Cave, Santa Cruz County, was the sub- 
ject of the photo in Shelley (1994, fig. 2). At the time that work was published, 
the specimen had not been discovered in the FSCA holdings. 


Paeromopus angusticeps buttensis Chamberlin, NEW STATUS 
Figs. 7-8 


Paeromopus buttensis Chamberlin, 1954:232. Buckett, 1964: 18. a 
Paeromopus ocellatus Loomis, 1972:260, figs. 1-5. Shelley, 1994:187-188, figs. 27-28. NEW 
SYNONYMY. 


The northern subspecies, P. a. buttensis is the most wide spread representa- 
tive of the genus. The distribution is essentially as described by Shelley (1994), 
but the southern coastal limit is in northern Sonoma County, and records are 
now available from Lake, Nevada, Placer, and Yuba counties (Fig. 11). We place 
P. ocellatus in synonymy here because the subterminal branch of the midlength 
projection (Fig. 5, mlp) of its anterior gonopod is long and resembles the condi- 
tion in this subspecies. Material was examined from the following new locali- 
ties: 


CALIFORNIA: Amador Co., along Consumnes R., SE Latrobe, 2M, 5F, 26 November 1965, MRG 
(UCD). Butte Co., 6 mi (9.6 km) SE Stirling City, M, 14 May 1967, S.R. Kutcher (UCD); and 2 mi 
(3.2 km) S Camp Storrie, exact location unknown, M, 20 June 1964, JSB, MRG (UCD). Calaveras 
Co., Avery & 1 mi (1.6 km) E Avery, 30M, 27F, 26 March 1966, JSB, MRG (UCD). Ei Dorado 
Co., 1 mi (1.6 km) E & 1 mi (1.6 km) NE Pacific House, 4M, F, 21 March-2 May 1992, W. D. 
Shepard (NCSM); 6 mi (9.6 km) S El Dorado, 3M, 2F, 28 November 1964, JSB, MRG (UCD); 
Camino, F, 19 April 1969, K. Lorenzen (UCD); and Pollock Pines, M, 20 February 1966, R. Denno 
(UCD). Humboldt Co., Redcrest, M, F, juv., 20 March 1976, and M, 25 November 1977, A. K. 
Johnson (FSCA); 2.2 mi (3.5 km) N Willow Creek, along CA hwy. 96, F, 10 October 1076, A. K. 
Johnson (FSCA); and Richardson Grove St. Pk., M, 2F, 20 March 1976, A. K. Johnson (FSCA). 
Lake Co., 4 mi (6.4km) NW Middletown, M, F, 21 February 1965, JSB, MRG (UCD). Mendocino 
Co., 3 mi (4.8 km) N Branscomb, M, 17 May 1975, C. Kellner (UCB); ca. 6 mi (9.6 km) N 
Branscomb, F, 24 November 1974, T. L. Smith (FSCA); 7 mi (11.2 km) NW Yorkville, M, 2F, 21 
December 1964, JSB, MRG (UCD); 9 mi (14.4 km) N Laytonville, 3F, 9 March 1968, JSB, MRG 
(UCD); 4-6 mi (6.4-9.6 km) N Potter Valley, 4M, 2F, 28 January 1967, JSB, MRG (UCD); 2 mi (3.2 
km) W Willitts, 24M, 26F, 25 March 1977, A. K. Johnson (FSCA); and 10 mi (16 km) N Cloverdale, 
F, 27 January 1967, JSB, MRG (UCD). Nevada Co., Grass Valley, M, 27 May 1967, D. S. Horning 
(UCD); 1 mi (1.6 km) SW Grass Valley, M, 10 February 1968, JSB, MRG (UCD); and 5 mi (8 km) 
NW Alta, 11M, 2F, 11 May 1969, D. A. Mead (UCD). Placer Co., 9 mi (14.4 km) N Aubum, 2M, 


4 ENTOMOLOGICAL NEWS 


4 February 1968, MRG (UCD); 15 mi (24 km) N Auburn, 2F, 4 February 1968, MRG (UCD). 
Plumas Co., | mi (1.6 km) N Elephant Butte, exact location unknown, M, 12 July 1964, JSB, MRG 
(UCD). Shasta Co., Inwood, 2F, 26 May 1967, JSB, MRG (UCD); and 21 mi (33.6 km) W Redding, 
M, F, 21 December 1966, JSB, MRG (UCD). Sonoma Co., 6 mi (9.6 km) NE Cloverdale, M, 21 
December 1964, JSB, MRG (UCD). Trinity Co., 3 mi (4.8 km) SW Douglas City, F, 21 December 
1966, JSB, MRG (UCD). Yuba Co., | mi (1.6 km) W Strawberry Valley Ranger Sta., 2F, 6 May 
1980, J. T. Doyen (UCB). 


Paeromopus angusticeps intergrades 
Figs. 3-6 


In the Sonoma County male, the subterminal branch of the midlength pro- 
jection (Fig. 3, mlp) of the anterior gonopod is short and apically broad, an 
intermediate condition between the long, broad projection in P. a. buttensis 
(Fig. 7) and the short, uncinate structure in the nominate subspecies (Fig. 1). 
On the posterior gonopod, the subterminal process of the terminal projection 
(Fig. 4, tp) is faint and weakly sinusoid; this condition is somewhat intermedi- 
ate between that in the nominate subspecies (Fig. 2) and the dactyliform, strongly 
sinusoid process in P. a. buttensis (Fig. 8). Paeromopus ocellatus also occupies 
an intermediate geographical location in western Solano County. We believe it 
is comprised of intergrade forms because the subterminal branch of the anterior 
gonopod midlength projection typically combines features of the two races, 
being long with an uncinate lateral margin (Fig. 5, mlp); the terminal projection 
of the posterior gonopod is variable but typically weakly convex apically witha 
slightly sinusoid subterminal process (Fig. 6, tp). In the holotype of P. ocellatus, 
the distal projection of the anterior gonopod bends strongly anteriad at midlength 
and overhangs the midlength projection (see Shelley 1994. fig. 27, dp). This 
configuration has not been seen again and is not characteristic of the Solano 
County population; we consider it an artifact, possibly from Loomis’ original 
dissection. as the distal projection is fragile and easily bent. Material was ex- 
amined from the following localities: 


CALIFORNIA: Sonoma Co., 7 mi (11.3 km) NE Santa Rosa, Los Alamos Rd., M, 26 
November 1964, JSB (UCD). Solano Co., 2 mi (3.2 km) W Cordelia, 4M, 4F, 21 December 
1964, JSB, MRG (UCD); and 1.7 mi (2.7 km) W jet. Hwy. I-80 and CA hwy. 12, nr. Corde- 
lia. F, 23 December 1967, JSB (UCD). 


Vol. 108, No. 1, January & February, 1997 


‘ MTT pp 


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Figs. 1-8. Paeromopus angusticeps. 1-2, P. a. angusticeps. 1, right anterior gonopod of male from 
Stanford, Santa Clara County, anterior view. 2, left posterior gonopod of the same, caudal view. 
3-6, P. angusticeps intergrades. 3, right anterior gonopod of male from 7 mi (11.2 km) NE Santa 
Rosa, Sonoma County, anterior view. 4, left posterior gonopod of the same, caudal view. 5, right 
anterior gonopod of male from 2 mi (3.2 km) W Cordelia, anterior view. 6, left posterior gonopod 
of the same, caudal view. 7-8, Pa. buttensis. 7, right anterior gonopod of holotype, anterior view. 
8, left posterior gonopod of the same, caudal view. dp, distal projection; mlp, midlength projection; 
tp, terminal projection. Setation omitted on figs. 1-2 and 7-8. Scale line = 1.00 mm for all figs. 


6 ENTOMOLOGICAL NEWS 


Paeromopus paniculus, NEW SPECIES 
Figs. 9-10 


Type specimen. Male holotype (UCD) collected by J. S. Buckett & M. R. 
Gardner, 8 February 1969, 14.1 mi (22.6 km) E Briceburg (1.5 mi (2.4 km, W 
El Portal), Mariposa County, California. 


Diagnosis. Color pattern consisting of transverse blue gray bands, generally indistinct from base 
color but becoming slightly lighter and more distinct anteriad; anterior gonopod curved moderately 
laterad, apical margin sublinear, tooth very short and inconspicuous, barely noticeable in anterior 
view, only slightly elevated above gonopodal surface, subterminal projection of midlength projec- 
tion short, barely projecting beyond surface, broadly sublinear apically, angling dorsad, distal pro- 
jection upright, bowed laterad proximal to midlength, extending well beyond distal extremity of 
gonopod; posterior gonopod without basal spine on anterior surface medial to terminal projection, 
with tuft of spinules on caudomedial surface at base of latter, terminal projection upright, with 
torsion, without subterminal caudomedial lamina, apical lamellae slightly flared, forming calyx 
with longer medial margin, without subterminal process (Figs. 9-10). 


Variation. The tuft on the left posterior gonopod contains about nine spinules, 
while that on the right contains only three. 

Paeromopus paniculus is the species with the longest millipeds on the North 
American continent, and a 165 mm (6 1/2 in.) long female collected by the first 
author at Happy Isles in 1990 is the longest individual ever collected. Though 
possessing mature gonopods and the same total segment number (75), the male 
holotype may not be fully grown, as it has three legless segments and measures 
79.8 mm in length, in contrast to the females from Happy Isles, which average 
75 segments, none legless, and 155.6 mm in length. 

Ecology. The habitat of the holotype is not indicated on the vial labels, but 
individuals at Happy Isles occur under moist decaying logs (Shelley 1994). 

Distribution. The Merced River Valley from above Nevada Falls, Yosemite 
National Park, to the community of Briceburg, west of the Park along Califor- 
nia highway 140, a linear distance of some 18 mi (28.8 km). In addition to the 
holotype, the following specimens are known; for convenience we repeat the 
records cited as Paeromopus sp. by Shelley (1994) (Fig. 11). 


CALIFORNIA: Mariposa Co., Yosemite Nat. Pk., Hidden Meadow (Lost Lake), ca. | 
mi (1.6 km) N Nevada Falls, F, 17 April 1954, J. Gorman (FSCA), Vernal Falls trail, F, 29 
May 1953, J. Gorman (FSCA), Happy Isles, F, 2 April 1967, J. R. Helfer (UCD) and 2F, 23 
April 1990, R. M. Shelley (NCSM), and Yosemite Valley, F, July 1952, M. Hood (LACMNH). 

The following two literature records refer to P. paniculus: 

Mariposa Co., Vernal Falls, Yosemite Nat. Pk., and Briceburg, along CA hwy. 140, ca. 
15 mi (24 km) W Yosemite (Causey 1955, Buckett 1964). 


Remarks. We are pleased to confirm Causey’s conclusion, based on labels 
with the Yosemite specimens at the FSCA, which were formerly in her private 


Vol. 108, No. 1, January & February, 1997 7 


Figs. 9-10. Paeromopus paniculus holotype. 9, right anterior gonopod, anterior view. 10, left poste- 
rior gonopod, caudal view. Abbreviations as in figs. 1-8. Scale line = 1.00 mm for both figs. 


8 ENTOMOLOGICAL NEWS 


Fig. 11. Distribution of Paeromopus. Stars, P. a. angusticeps; dots, P. a. buttensis; X’s, P. angusticeps 
intergrades; question marks, unknown forms of P. angusticeps; inverted triangle, P. eldoradus; 
square, P. cavicolens; diamonds, P. paniculus. Open symbols denote literature records deemed 
valid. 


collection, that this, the southern most population of Paeromopus in the Sierras, 
does represent a new species. Shelley (1994) tentatively concurred because of 
the darkly banded color pattern, in contrast to the light bands of other forms of 
Paeromopus, but could not propose a name in the absence of males, which still 
have not been taken within Yosemite National Park. Paeromopus paniculus is 
closely related to P. eldoradus and comes off with this species at the second half 
of couplet 4 in the key to species by Shelley (1994). It is distinguished from P. 
eldoradus by the inconspicuous anterior gonopod tooth, the absence of the 
caudomedial lamina on the posterior gonopod terminal projection, and the pres- 
ence of the tuft of spinules at the base of the latter (compare figs. 9-10 with figs. 
29-30). The specific name means tuft or bush, and refers to the spinules. 


Vol. 108, No. 1, January & February, 1997 9 


Paeromopus sp. 


Additional female samples from Sonoma County cannot be assigned with 
confidence as either intergrades or a subspecies of P. angusticeps. We record 
them here and show them as question marks in fig. 9. 


CALIFORNIA: Sonoma Co., 3 mi (4.8 km) N Kenwood, along Adobe Cyn. Rd., 3F, 20 
December 1964, JSB, MRG (UCD); Rio Nido, F, 17 June 1916, W. Gaeggel (CAS); and 
Occidental, F, 29 October 1964, S. Rattoro (CAS). 


Californiulus dorsovittatus Verhoeff 


The distribution of C. dorsovittatus expands in the north and east with records 
from eastern Siskiyou County and central Lassen County. Material was exam- 
ined from the following new localities: 


CALIFORNIA: Lassen Co., Silva Lake, F, 16 June 1956, P. S. Bartholomew (CAS). 
Shasta Co., Moose Camp, exact location unknown, M, 6 July 1993, J. R. Pierson (BYU). 
Siskiyou Co., Lava Beds Natl. Mon., M, 28 August 1966, R. C. Gardner, S. E. Harrison 
(UCD); 21 mi (33.6 km) E McCloud, Dead Horse Summit, along CA hwy. 89 nr. Ponderosa, 
M, F, 7 August 1968, H. B. Leech (CAS) and M, 18 September 1961, W. J. Gertsch, W. Ivie 
(AMNH); and along Tate Cr., exact location unknown, F, 24 June 1954, H. P. Chandler (CAS). 
Tehama Co., along Deer Cr., generally E of Tehama but exact location unknown, F, 19 Sep- 
tember 1961, W. J. Gertsch, W. Ivie (AMNH). 


Californiulus yosemitensis Chamberlin 


New records from central Modoc and northeastern Lassen counties expand 
the area of the Warner Mountains population and connect it with the main area 
of the genus and family in the southern Cascades, although it is still disjunct 
from the main area of C. yosemitensis. A new record is available from Nevada 
County, but the continued absence of samples from E1 Dorado, Amador, and 
Calaveras counties (Fig. 16) indicates that this area constitutes a lacuna within 
both the genus and species, but not within the family because P. a. buttensis and 
P. eldoradus occur in this gap (Fig. 11). Therefore, the southern area of C. 
yosemitensis, extending from Mariposa and Mono to Kern counties, is disjunct 
from that from Placer County northward (Fig. 16). Material was examined from 
the following new localities. 


CALIFORNIA: Fresno Co., Kings River Canyon, along Roaring River, M, 4F, July- 
August, 1910, collector unknown (CAS). Kern Co., 2 mi (3.2 km) N L. Isabella, 2F, May 
1971, E. A. Kane (UCD). Lassen Co., Lost L., S. Warner Mts., F, 19 September 1964, A. B. 


10 ENTOMOLOGICAL NEWS 


Gurney (UCD). Modoc Co., 15 mi (24 km) N Alturas, along Joseph Cr., F, 17 June 1954, R. 
O. Schuster (FSCA); and Crowder Flat & Twin Spgs., ca. 7 mi (11.2 km) S Oregon border & 
11-12 mi (17.6-19.2 km) W Goose L., 2F, 25 June 1974, A. K. Johnson (FSCA). Nevada 
Co., Sagehen Cr., exact location unknown, 2M, 3F, 25 June 16-July 1960, J. J. Jarvis (UCD). 
Placer Co., Tacoma, exact location unknown, M, 18 August 1964, MRG (UCD). Plumas 
Co., 8 mi (12.8 km) NW Quincy, M, 21-23 June 1984, P. S. Ward (UCD); Johnsville, F, 20 
September 1964, JSB, MRG (UCD) and F, July 1971, H. Pini (UCD); Mt. Ingalls, M, 5F, 11 
July 1964, and M, F, 4 September 1967, JSB, MRG (UCD); and Bucks L., M, F, 4 July 1964, 
and M, 7 September 1964, L. L. Dunning (UCD). Tulare Co., Kings Cyn. Nat. Pk., Red- 
wood Cyn., Redwood Saddle, M, 17 August 1984, V. F. Lee, T. S. Briggs, D. Ubick (CAS); 
Sequoia Nat. Pk., nr. Crystal Cv., M, 13 July 1961, S. B. Peck (FSCA); Mineral King, M, 2F, 
date unknown, P. S. Bartholomew (CAS); and Quaking Aspen Cpgd., Sequoia Nat. For., M, 
9 September 1959, W. J. Gertsch, V. Roth (AMNH). Tuolumne Co., Pinecrest, M, 14 June 
1965, JSB (UCD). 


Californiulus euphanus (Chamberlin) 


The only new record of C. euphanus extends its range in western Washing- 
ton eastward into the western periphery of the Columbia Plateau (Fig. 16). 
Sample data are as follows: 


WASHINGTON: Yakima Co., 10 mi (16 km) N Yakima, juv. M, 8 May 1948, E. S. Ross 
(CAS). 


Californiulus blechrostriatus, NEW SPECIES 
Figs. 12-15 


Type specimens. Male holotype (CAS) collected by B. Banta, 5 April 1959, 
at an unknown site in Saline Valley, Death Valley National Park, Inyo County, 
California; one male and two female paratypes (CAS) taken by the same col- 
lector, 2 April 1960, along Grapevine Canyon Rd., Saline Valley. 


Diagnosis. Color pattern subuniformly dark brown, caudal margins of collum and 
pleuroterga 2-4 slightly darker but otherwise without trace of transverse banding; pleurotergal 
striation very faint and shallow, barely noticeable, exoskeleton appearing generally smooth; 
anterior gonopod ridge curvilinear, tooth moderately long, angling strongly mediad, over- 
hanging groove and nearly overhanging medial margin, distal projection either divided with 
short, irregular inner branch, or simple and expanded basally, with numerous spiculate pro- 
jections, margins serrate to jagged, extending to level of distal extremity of gonopod; subter- 
minal projection of posterior gonopod divided into 4 branches, with short, slightly expanded 
and laminate, accessory process arising ventrad below process ‘C’; process ‘A’ entire, not 
distally divided, with one termination; process ‘B’ broad and laminate, moderately expanded 
mediad, obscuring process ‘C’ in anterior view, marginally irregular with fine teeth and 
moderate indentations; process ‘C’ slightly expanded basally, falcate distad, tip well segre- 
gated from that of termination of ‘B’ (Figs. 12-15). 


Vol. 108, No. 1, January & February, 1997 11 


Figs 12-15. Californiulus blechrostriatus, holotype. 12, right anterior gonopod, anterior view. 13, 
distal extremity of right posterior gonopod, medial view. 14, the same, anterior view. 15, right 
anterior gonopod of male from Mt. Whitney, anterior view. A, process “A”; ap, accessory process; 


B, process “B”; C, process “C”; other abbreviations as in figs. 1-8. Scale line = 1.40 mm for figs. 12 
and 15, 1.00 mm for figs. 13-14. 


12 ENTOMOLOGICAL NEWS 


Fig. 16. Distribution of Californiulus. Inverted triangles, C. chamberlini; upright triangles, C. 
dorsovittatus; dots, C. yosemitensis; squares, C. euphanus; diamonds, C. parvior; stars, C. 
blechrostriatus. Open symbols denote literature records deemed valid, and the dot in Nevada County, 
California, represents the Sagehen Creek record of C. yosemitensis, whose exact location is un- 
known. The asterisk shows the one locality of Aprosphylosoma darcenae Hoffman 
(Aprosphylosomatidae), the only other species and genus in the superfamily Paeromopodoidea. 


Vol. 108, No. 1, January & February, 1997 13 


Variation. The male paratype agrees closely with the holotype, with only 
minor differences in the degrees of expansion and marginal spination of 
process ‘B’ of the posterior gonopod and of the intricacy of the spiculation of 
the anterior gonopod distal projection. The specimen from Mt. Whitney agrees 
with both Saline Valley males except for the anterior gonopod (Fig. 15). The 
apical process of its midlength projection (mlp) is shorter, narrower, and more 
obviously uncinate, and the distal projection (dp) is divided into two branches, 
instead of being expanded basally (Fig. 12), and the inner branch is short, with 
long, irregular, marginal teeth. Additionally, this male has an accessory spur on 
the medial surface of the ridge on the right anterior gonopod but not on the left 
one. Including the epiproct, the holotype has 67 segments, none legless, and 
measures 83.2 mm in length; a female paratype, the only intact and measurable 
specimen in this sample has 69 segments, one legless, and measures 74.9 mm 
in length; and the Mt. Whitney male has 73 segments, one legless, and mea- 
sures 93.2 mm in length. 

Ecology. Californiulus blechrostriatus has a marked affinity for arid envi- 
ronments. The only information on the vial labels with the types is the elevation 
of 4,200 ft. (1,260 m), but this is high desert, in the Basin and Range Physi- 
ographic Province, and Saline Valley is the next valley west of Death Valley, the 
lowest, and the hottest, driest place in North America. The Mt. Whitney male 
was discovered under a fallen, decaying pine long at 8,000 ft. (2,400 m) on the 
dry, eastern side of the Sierra Nevada. Californiulus yosemitensis can occur in 
relatively dry biotopes (Shelley 1994), but not to the extreme of those occupied 
by C. blechrostriatus, the easternmost representative of the family in California 
and the only one occurring in true desert. 

Distribution. Known only from Inyo County, California, where it extends 
eastward from Mt. Whitney to the Saline Valley, a linear distance of approxi- 
mately 38 mi (60.8 km). In addition to the type specimens, the following sample 
is known. . 


CALIFORNIA: Inyo Co., east side of Mt. Whitney, Inyo Nat. For., end of rd. to Whitney 
Portal, ca. 10.5 mi (16.8 km) WSW Lone Pine, M, 20 May 1970, E. A. Kane (UCD). 


Remarks. As they share an undivided process “A” of the posterior gonopod, 
C. blechrostriatus is closely related to C. yosemitensis. They differ in the color 
pattern, as the former shows no trace of the yellow middorsal stripe and medial 
black line diagnostic of the latter, nor does C. blechrostriatus display the trans- 
verse banding characteristic of C. euphanus, C. parvior, and the species of 
Paeromopus. The accessory, ventral process on the posterior gonopod (Fig. 14, 
ap) is also unique, but the most striking feature of C. blechrostriatus is the 
extremely faint striation, which contrasts with the condition in every other spe- 
cies and is best seen under oblique light under a stereomicroscope. This trait is 
the source of the specific name, which derives from the Greek word, “blechros,” 


14 ENTOMOLOGICAL NEWS 


meaning faint or weak, and “striae,” meaning grooves or channels. In the key to 
species of Californiulus (Shelley 1994), C. blechrostriatus goes to couplet 4 
because it lacks the yellow middorsal stripe, and it is readily distinguished from 
both C. euphanus and C. parvior by the faint striae, the accessory process of the 
posterior gonopod, and the simple, undivided process “A”. 


ACKNOWLEDGMENTS 


We thank the following curators and collection managers for loaning material from the 
indicated repositories: N. I. Platnick (AMNH); R. W. Baumann (BYU); C. E. Griswold (CAS), 
G. B. Edwards (FSCA); the late C. L. Hogue (LACMNH); C. B. Barr (UCB); and L. S. 
Kimsey (UCD). R. G. Kuhler, North Carolina State Museum Scientific Illustrator, assisted 
in preparation of figures 1-10 and 12-15. 


LITERATURE CITED 


Buckett, J. S. 1964. Annotated list of California Diplopoda. Simmons Publ. Co., Davis, CA, 34 pp. 

Causey, N. B. 1955. New records and descriptions of Californian Diplopoda. Proc. Biol. Soc. 
Wash., 68:87-94. 

Chamberlin, R. V. 1941. New western millipeds. Bull. Univ. Utah, 31(12) [Biol. Ser., 6(5)]:1-23. 

Chamberlin, R. V. 1954. Six new Californian millipeds. Proc. Biol. Soc. Wash., 67:231-234. 

Loomis, H. F. 1972. Some notes on the milliped family Paeromopodidae, with a description of a 
new species. Fla. Entomol., 55:259-262. 

Shelley, R. M. 1994. Revision of the milliped family Paeromopodidae, and elevation of the 
Aprosphylosomatinae to family status (Julida: Paeromopodoidea). Ent. Scand., 25:169-214. 


Vol. 108, No. 1, January & February, 1997 15 


A NEW SPECIES OF OXYGONIA FROM ECUADOR 
(COLEOPTERA: CICINDELIDAE)! 


Fabio Cassola2, Michael G. Kippenhan>>4 


ABSTRACT: A new species of Oxygonia, O. onorei, is described based on two male specimens 
from Las Pampas and La Otonga, Cotopaxi, Ecuador. Distinguishing characters are depicted. 


The genus Oxygonia Mannerheim, 1837 (type-species: O. schoenherri 
Mannerheim, 1837), is an interesting Neotropical tiger beetle group which so 
far includes approximately fifteen species, ranging from Costa Rica and Panama 
(1 species) to Colombia (3 species), Ecuador (11 species), Peru (5 species) and 
Bolivia (2 species) (Bates 1881-84, Horn 1926, Wiesner 1992). Most of these 
species occur in the Andean slopes of Ecuador and this country appears to be 
the center of the evolution and distribution of the genus Oxygonia. 

Specimens are uncommon in collections and little is known concerning their 
ecology and behavior. Buckley collected three species in the River Upano area 
near Macas in Ecuador over one century ago and found his specimens chiefly 
on mossy stones in the bed of rapid streams (Bates 1872). Recent observations 
by Pearson et al. (1995) confirm that Oxygonia specimens forage on mossy 
rocks and boulders in the middle of shallow white-water streams. The diurnal 
species roost at night on leaves of overhanging bushes and trees above the stream 
surface, while the nocturnal species spend the daytime under rocks and gravel 
along small to moderate-sized mountain streams and forage on rocks in stream 
at night. 

Most Oxygonia species show a remarkable sexual dimorphism in shape and 
body color, and there is often considerable difficulty identifying both sexes of 
the various species, especially when associated males and females are not avail- 
able. However, a systematic review of this genus is presently being made by the 
second author and there are no described species of which the male is unknown 
(Kippenhan, in prep.). The purpose of this paper is to describe an interesting 
new species collected in northwestern Ecuador. 


1 Received May 14, 1996. Accepted June 8, 1996. 
2 Via F. Tomassucci 12/20, 1-00144 Roma, Italy (Studies on Tiger Beetles, LX XXIII). 
3 15185 Deby Drive, Colorado Springs, Colorado 80921, U.S.A. 


4 Associate, C.P.Gillette Arthropod Biodiversity Museum, Colorado State University, Fort Collins, 
CO 80523, U.S.A. 


ENT. NEWS 108(1) 15-18, January & February, 1997 


16 ENTOMOLOGICAL NEWS 


Oxygonia onorei NEW SPECIES 
(Fig. 1 a, b, c) 


TYPE MATERIAL. Holotype (male) from Las Pampas, Cotopaxi, Ecuador, May 1988, 
G.Onore leg., deposited in the collection of the Carnegie Museum of Natural History, Pitts- 
burgh, Pennsylvania. One paratype (male) from La Otonga, 1800m, Cotopaxi, Ecuador, 25 
May 1996, A. Paucar leg., in first author’s collection. 

DIAGNOSIS. The species can be distinguished from all others by its small size (10.5 
mm), the testaceous color of the underside and the cleft aedeagal apex. 

COLOR. Dorsum metallic brownish bronze, with greenish or blue-violet reflections on 
head and pronotum. Sides and underside of head and pronotum metallic golden green, with 
some cupreous reflections; a yellow patch in the middle of pro-and mesosterna. Metasternum, 
abdominal sterna and elytral epipleura completely testaceous. White elytral markings com- 
prised of a narrow humeral lunule, a short transverse middle band behind front half of elytra 
and a subapical subtriangular spot. Female unknown. 

DESCRIPTION. Head: relatively small, metallic dark bronze above with some slight 
blue-green reflections, golden green below; strong cupreous reflections on genae, clypeus, 
frons and behind eyes. Surface entirely glabrous, a single sensory seta near middle edge of 
each eye; vertex very finely striated, some shallow transverse wrinkles on neck. Eyes bulg- 
ing outward and upward, with a rounded moderate excavation between. Labrum short, trans- 
verse, black with an unpigmented subsquare patch on either side of middle; anterior margin 
excavated in the middle with a fairly substantial central tooth protruding outwards and two 
small lateral bumps; two pairs of submarginal setae, first one at sides of central tooth, sec- 
ond one inside lateral apical angles of labrum. Mandibles black, rufous on the apical and 
inner teeth. Labial and maxillary palpi basally testaceous. tinged with metallic brown on last 
segments. Antennae rather long, reaching nearly the middle of the elytral length; extreme 
base of scape testaceous, scape and articles 2-4 black with some violaceous reflections, gla- 
brous, antennomeres 5-11 brownish-black, evenly and finely pubescent. 

Prothorax: pronotum about as long as wide, rounded at sides, maximum width near the 
middle, with the front lobe very short in proportion; color metallic dark bronze with violaceous 
reflections, some golden green or cupreous reflections in front and hind transverse grooves. 
Surface completely glabrous, very finely transversely striate, the striae shallow, almost ef- 
faced on disk; midline fine but distinct, slightly excavated behind. Pro- and mesosternal 
pieces golden green, glabrous, with some cupreous reflections; a yellow patch in middle of 
pro- and mesosternum. Metasternum and metepisterna completely testaceous, glabrous, with 
scattered patches of pearly lustre. 

Elytra: purple-brownish shining bronze throughout, completely covered with shallow, 
evenly-spaced, small round punctures with metallic green or cupreous reflections; some larger 
foveae near base and front part of suture. Surface uneven, with a strong depression near 
suture on front third of disk, a second depression on inside part of middle spot, and a third 
one near apex. Elytral maculation white, consisting of a narrow humeral lunule (only poste- 
rior end visible from above), a subrectangular, transverse, middle band behind front half of 
elytra (extended from near side margins to middle of disk) and a subtriangular hind spot near 
the subapical margin of elytra (not reaching sutural angle) (Fig. la). Elytral apex ended ina 
strong protruding sutural spine; microserrations present. Epipleura yellow-testaceous. 

Abdomen: sterna completely testaceous, glabrous with scattered translucid patches; a 
single pair of sensory setae near hind edge of 3rd, 4th and Sth visible sterna. 

Legs (only pro legs and femur of right meso leg remain in the male holotype): coxae 
and trochanters yellow, the former with some white pilosity on sides; femora testaceous 
below. more or less tinged with metallic green above, with a dark black ring at “knees”. 
Tibiae and tarsi of forelegs shining black. 


Vol. 108, No. 1, January & February, 1997 17 


Fig. 1. Oxygonia onorei n.sp., male holotype from Las Pampas. Cotopaxi, Ecuador: a. habitus; b. 
aedeagus (right side); c. apex of aedeagus (dorsal view) . 


18 ENTOMOLOGICAL NEWS 


Genitalia: Male aedeagus rather small, tapering, evenly enlarged in the middle, with a 
short spatulate apical knob, slightly curved upwards (bi-lobate in dorsal and ventral views) 
(Fig. | b,c). 

Length: 10.5 mm (without labrum). 


ETYMOLOGY. The species is named after Professor Giovanni Onore, Di- 
rector of QCAZ (Instituto de Zoologia, Pontificia Universidad Catdlica, Quito, 
Ecuador), who collected the single male holotype and kindly submitted it for 
identification. 


DISTRIBUTION. Western Andean slopes of northern Ecuador. 


REMARKS. Only two male specimens of O. onorei n.sp. are available thus 
far. However, these specimens are distinctive enough to be described as a new 
species. The shape of the aedeagus and flagellum is similar to the group of O. 
vuillefroyi Chaudoir (Kippenhan, in prep.), but the cleft apex is unique within 
the genus. The color of the underside and other features (such as the apically 
knobbed aedeagus with small “J” shaped flagellum) suggest that this species is 
most closely related to O. nigricans W. Horn and O. oberthueri W. Horn 
(Kippenhan, in prep.) and can be placed between. 


LITERATURE CITED 


Bates, H. W. 1872. Notes on Cicindelidae and Carabidae, and descriptions of new species (No. 13). 
Ent. Monthly Mag., 8: 237-242. - 

Bates, H. W. 1881-84. Biologia Centrali-Americana. Insecta. Coleoptera. Vol. 1, Part 1. London. 

Horn, W. 1926. Carabidae: Cicindelinae. Jn: W. Junk (ed.), Coleopterorum Catalogus, Pars 86. 
Berlin, 345 pp. 

Mannerheim, M. 1837. Mémoire sur quelques genres et espéces de Carabiques. Bull. Soc. Natur. 
Moscou, 10: 3-49. 

Pearson, D. L., D. W. Brzoska and J. Buestan. 1995. Natural history observations on species of 
the tiger beetle genus Oxygonia in Ecuador (Coleoptera: Cicindelidae). Cicindela, 27: 45-50. 

Wiesner, J. 1992. Verzeichnis der Sandlaufkafer der Welt. Checklist of the Tiger Beetles of the 
World. Verlag Erna Bauer, Keltern, 364 pp. 


Vol. 108, No. 1, January & February, 1997 19 


AN UNUSUAL NEW PACHYCHERNES 
FROM PANAMA AND MEXICO 
(PSEUDOSCORPIONIDA: CHERNETIDAE)! 


William B. Muchmore2 


ABSTRACT: A new species, Pachychernes zehorum, is described on the basis of material collected 
in Panama and Mexico (Chiapas). It is unusual in the structure of the first leg of the male, which has 
modified setae on the tibia and also a unique depression on the dorsum of the tarsus. 


The genus Pachychernes was established by Beier (1932), based upon 
Chelifer (? Atemnus) subrobustus Balzan, from Venezuela. Since then, 7 more 
species have been assigned to the genus, distributed from Argentina to Florida 
(Harvey 1991: 611-612; unpublished observations). However, detailed knowl- 
edge of the type species is scanty, and the definition of the genus is unsatisfac- 
tory; there is good reason to believe that some of the species are improperly 
assigned. Nevertheless, 3 species are fairly well known and seem to form a 
coherent group in which males have highly modified setae on the first legs and 
females have distinctive spermathecae; these are P. baileyi Feio (1945), from 
Brazil (Bahia and Amazonas), P. shelfordi Hoff (1946), from Mexico (no fur- 
ther data), and P. attenuatus Muchmore (1990), from Mexico (Yucatan and 
Quintana Roo). A new species, related to these three, has been found in both 
Panama and Mexico (Chiapas). 


Genus Pachychernes Beier 


Pachychernes Beier, 1932: 114; Beier 1933: 516; Harvey 1991: 611. Type species: Chelifer 
(? Atemnus) subrobustus Balzan, 1892. Type locality: Caracas, Venezuela. 


Pachychernes zehorum, NEW SPECIES 
(Figs. 1-4) 


Description: Male and female generally similar, except for the genitalia and the con- 
spicuously modified tibia and tarsus of leg I of the male. Palps dark reddish brown, carapace 
and other parts lighter brown or tan. Carapace a little longer than broad; surface heavily 
granulate, with 2 shallow transverse furrows; 2 large eyespots; 60-65 short, finely denticu- 
late setae, usually 4 at anterior and 6 at posterior margin. Coxal area unremarkable. Abdomi- 
nal tergites 1-10 and sternites 4-10 divided; surfaces of anterior tergites granulate, posterior 
ones scaly; sternites smooth; pleural membranes granulostriate anteriorly, smoothly striate 


! Received June 24, 1996. Accepted August 1, 1996. 
2 Department of Biology, University of Rochester, Rochester, New York 14627. 


ENT. NEWS 108(1) 19-23, January & February, 1997 


20 ENTOMOLOGICAL NEWS 


posteriorly; setae finely denticulate or acuminate. Tergal chaetotaxy of holotype 8:9:10:10: 
12:13:13:13:14:12:T3T2T3T:2, others generally similar. Sternal chaetotaxy of holotype 
~50:[4-4]:(3)8(3):(1)10(1):20:19:20:22:21:20:T3T7T4T:2, other males similar; sternites 2- 
4 of allotype with 24:(3)10(3):(1)9(1) setae. Internal genitalia of male typically chernetid in 
structure, but very large and heavily sclerotized, much like those of P. baileyi (see Feio 
1945: fig.19). Spermathecae of female (Fig. 1) apparently like those of P. bailevi, P. shelfordi, 
and P. attenuatus (see Mahnert 1979: fig. 118; Muchmore 1975: fig. 9, 1990: fig. 6), but in 
the specimens examined here the entire course of the thin tubules could not be seen. 

Chelicera 0.33 as long as carapace; hand with 5 setae, bs and sbs long, denticulate, 
others long, acuminate; flagellum of 3 setae; galea of male slender, with 5-6 small rami, that 
of female heavier and with larger rami, 2 at middle and 4 more terminal. 

Palp (Fig. 2) rather slender, patella slightly longer than femur. L/B of trochanter 1.65- 
1.8, femur 2.6-2.9, patella 2.35-2.7, and chela(without pedicel) 2.75-3.1; L/D of hand(without 
pedicel) 1.7-1.9; movable finger L / hand L 0.8-0.9. Surfaces moderately to lightly granu- 
late, except chelal fingers smooth; most setae slender, finely denticulate. Trichobothria as 
shown in Fig. 3. Fixed finger with 55-60 and movable finger with 50-55 cusped teeth. Venom 
apparatus well developed in movable finger, nodus ramosus at, or just proximad of, level of 
trichobothrium ¢. 

Legs rather slender: leg IV with L/D of femur+patella 2.7-3.1, tibia 3.8-4.4, and tarsus 
3.8-4.4. Leg I sexually dimorphic: that of female typically chernetid in structure; that of 
male with tibia somewhat enlarged and unusually setaceous, and tarsus having an extensive, 
bare depression in the proximal half of the dorsum (Fig. 4). Tarsus of leg IV with a very long 
tactile seta 0.3 length of segment from proximal end. 

Ttritonymph (from Chiapas). Generally similar to adults, but without sexual modifica- 
tions. Carapace with 2 eyes, 2 faint transverse furrows, and about 50 finely denticulate or 
acuminate setae. Tergal chaetotaxy 8:8:8:10:11:12:12:12:-. Chelicera with 5 setae on hand 
and 3 setae in flagellum. Palp with L/B of femur 2.5, patella 2.25, and chela (without pedicel) 
2.9; L/D of hand (without pedicel) 1.65; movable finger L / hand L 0.85. Legs as in adults; 
tarsus with very long tactile seta 0.3 length of segment from proximal end. 


Measurements (mm): Male. Figures given first for holotype, followed in parentheses 
by ranges for 4 paratypes. Body L 3.84 (3.19-3.66). Carapace L 1.25 (1.12-1.23). Chelicera 
L 0.43 (0.39-0.42). Palp: trochanter 0.67 (0.63-0.69) / 0.385 (0.36-0.385); femur 1.16 (1.04- 
1.10) / 0.405 (0.36-0.39); patella 1.24 (1.11-1.23) / 0.47 (0.415-0.46); chela (without pedicel) 
1.85 (1.76-1.83) / 0.635 (0.56-0.605); hand (without pedicel) 1.07 (0.975-1.07) / 0.62 (0.54- 
0.60); pedicel L 0.13 (0.13); movable finger L 0.89 (0.865-0.91). Leg I: femur+patella L 
0.815 (0.76-0.85); femur 0.39 (0.38-0.435) / 0.26 (0.235-0.265); patella 0.635 (0.585-0.66) 
/ 0.22 (0.21-0.23); tibia 0.59 (0.55-0.62) / 0.18 (0.17-0.18); tarsus 0.615 (0.58-0.64) / 0.13 
(0.12-0.125). Leg IV: femur+tpatella 1.04 (0.96-1.10) / 0.36 (0.36-0.39); tibia 0.88 (0.815- 
0.88) / 0.20 (0.20-0.21); tarsus 0.59 (0.57-0.605) / 0.14 (0.13-0.14). 

Female. Figures given first for allotype, followed in parentheses by ranges for 3 paratypes. 
Body L 4.11 (3.85-4.87). Carapace L 1.15 (1.21-1.36). Chelicera L 0.39 (0.415-0.43). Palp: 
trochanter 0.585 (0.64-0.69) / 0.325 (0.355-0.41); femur 0.985 (1.04-1.12) / 0.35 (0.385- 
0.43); patella. 1.03 (1.10-1.19) / 0.39 (0.445-0.51); chela (without pedicel) 1.70 (1.84-1.91) 
/ 0.55 (0.63-0.69); hand (without pedicel) 0.985 (1.06-1.13) / 0.52 (0.615-0.65); pedicel L 
0.11 (0.12-0.14); movable finger L 0.815 (0.85-0.89). Leg I: femur+patella L 0.695 (0.75- 
0.83); femur 0.35 (0.38-0.415) /0.215 (0.235-0.265); patella 0.54 (0.585-0.62) / 0.19 (0.21- 
0.235); tibia 0.52 (0.55-0.585) / 0.13 (0.14-0.16); tarsus 0.465 (0.49-0.525) / 0.105 (0.105- 
0.12). Leg IV: femur+patella 0.96 (1.05-1.16) / 0.31 (0.36-0.415); tibia 0.79 (0.85-0.925) / 
0.185 (0.20-0.23); tarsus 0.54 (0.555-0.615) / 0.13 (0.14-0.16). 

Tritonymph. Body L 3.23. Carapace L 0.93. Chelicera L 0.32. Palp: femur 0.73/0.29; 
patella 0.74 / 0.325; chela(without pedicel) 1.31 / 0.45; hand(without pedicel) 0.74 / 0.445; 


Vol. 108, No. 1, January & February, 1997 21 


movable finger L 0.63. Leg IV: femur+patella 0.75/0.275; tibia 0.615/0.155; tarsus 0.45/ 
0.11. 

Type data: Holotype male (WM7097.01001), allotype female (WM7098.01001), and 8 
paratypes (3 males, 4 females, | tritonymph) from under bark of live trees along Pipeline 
Road, Parque Nacional Soberania, PANAMA, 16 January 1988, J. A. Zeh; all but 1 male, | 
female and tritonymph mounted on slides. Three paratypes (2 males, | tritonymph) from 
under bark of trees along trail to ruins, Palenque, Chiapas, MEXICO, 24 January 1976 & 22 
January 1985, C. R. Hignutt; mounted on slides. All types deposited in the Florida State 
Collection of Arthropods, Gainesville, Florida. 


Etymology: The species is named in honor of Jeanne A. Zeh and David 
W. Zeh, who collected and sent me the type specimens from Panama. 


Diagnosis: Easily distinguished from all other members of the genus 
by the first leg of the male, which has both long and short modified setae 
on the tibia and a unique, distinct depression on the dorsal side of the 
tarsus. 


Figs. 1-4. Pachychernes zehorum, new species. 1. Spermathecae of female. 2. Right palp, dorsal 
view. 3. Left chela, lateral view. 4. Leg I of male. 


DISCUSSION 


The striking modifications of leg I of the male of Pachychernes zehorum 
are undoubtedly related to courtship and mating. Though no direct evidence 
exists to support this contention, it seems reasonable, inasmuch as many pseu- 
doscorpions of the family Chernetidae have complex sexual behaviors, during 


22 ENTOMOLOGICAL NEWS 


which the male grasps and manipulates the female (see Weygoldt 1969). 

Although it is common in pseudoscorpions of the family Cheliferidae, sexual 
modification of the first legs of males is rare in the Chernetidae. Modified first 
legs have been found in only 9 species other than P. zehorum among the 600+ 
described species in the Chernetidae — Pachychernes baileyi Feio (1945) from 
Brazil, P. shelfordi Hoff (1946) from Mexico, P. attenuatus Muchmore (1990) 
from Mexico, Bituberochernes mumae Muchmore (1974) from Florida and Cuba, 
B. jonensis Muchmore (1979) from the Virgin Islands, Orochernes nepalensis 
Beier (1968) from Nepal, O. sibiricus Schawaller (1986) from Siberia, and 2 
new species of a new genus from California and Arizona (Muchmore, in prep.). 
The relationships among these species will be interesting to investigate (in 
progress). 

As mentioned above, the spermathecae in the female of Pachychernes 
zehorum are similar to those of some other species in the genus, namely, P. 
baileyi, P. shelfordi, and P. attenuatus; no information is available concerning 
the spermathecae of other species assigned to Pachychernes (see Harvey 1991: 
611-612). Also, these spermathecae are similar to those of the two known spe- 
cies of Bituberochernes Muchmore, namely, B. mumae (see Dumitresco & 
Orghidan 1977: fig. 12B) and B. jonensis (Muchmore 1979: fig. 4). It should be 
noted that the spermathecae of Chelanops (Neochelanops) peruanus Mahnert 
(1984: figs. 28-29) appear like the foregoing, even though the species is not 
otherwise similar to Pachychernes or Bituberochernes. 

It seems probable, from the similarity of the first pedal tarsi of males and 
the spermathecae of females, among other characters, that the genera 
Pachychernes and Bituberochernes are closely related; a more detailed exami- 
nation of this situation is in progress. 

The occurrence of this species in Panama and Mexico strongly suggests 
that it is widely distributed through Central America. 

All of the specimens of P. zehorum were collected from under loose bark of 
standing trees. Concerning those from Panama, D. W. Zeh comments (in litt.), 
“We did see clear evidence that this species feeds on ants. Several of the indi- 
viduals we sent you were collected from under bark in close association with 
ant corpses.” The identity of the ants upon which these pseudoscorpions pre- 
sumably feed has not yet been ascertained, but they are not the same ants preyed 
upon by Paratemnus elongatus (Banks) (Zeh & Zeh 1990) [Note: the genus of 
this species is now Paratemnoides Harvey 1991: 469]. 


ACKNOWLEDGMENTS 


I am grateful to Charlotte R. Hignutt and David W. Zeh for sending me the specimens 
upon which this study is based. Also, I am much indebted to Ms. Hignutt for the excellent 
drawings presented herein, and to Uzi Nur, David W. Zeh, Jeanne A. Zeh, and two anony- 
mous reviewers for constructive comments on the manuscript. 


Vol. 108, No. 1, January & February, 1997 23 


LITERATURE CITED 


Beier, M. 1932. Pseudoscorpionidea II. Subord. C. Cheliferinea. Tierreich 58: 1-294. 

Beier, M. 1933. Revision der Chernetidae (Pseudoscorp.). Zool. Jahrb., Syst. 64: 509-548. 

Beier, M. 1968. Ein neues Chernetiden-Genus (Pseudoscorp.) aus Nepal. Khumbu Himal 3: 17-18. 

Dumitresco, M. & T. N. Orghidan. 1977. Pseudoscorpions de Cuba. Rés. Expéd. Biospéol.Cubano- 
Roum. Cuba 2: 99-122. 

Feio, J. L. de Araujo. 1945. Novos pseudoscorpides da regiao neotropical (com a descri¢ao de 
uma subfamilia, dois géneros e sete espécies). Bol. Mus. Nac., Rio de Janeiro, n.s. Zool. 44: 
1-47. 

Harvey, M. S. 1991. Catalogue of the Pseudoscorpionida. Manchester University Press, Manches- 
ter, England. 726 pp. 

Hoff, C. C. 1946. Descripcion de una especie nueva del género Pachychernes Beier, 1932 
(Pseudoscorpionida). Ciencia, México 7: 13-14. 

Mahnert, V. 1979. Pseudoskorpione (Arachnida) aus dem Amazonas-Gebiet (Brasilien), Rev. Suisse 
Zool. 86: 719-810. 

Mahnert, V. 1984. Pseudoscorpions (Arachnida) récoltés durant la mission spéologique espagnole 
au Pérou en 1977. Rev. Arachnol. 6: 17-28. 

Muchmore, W. B. 1974. Pseudoscorpions from Florida. 2. Anew genus and species Bituberochernes 
mumae (Chernetidae). Florida Entomol. 57: 77-80. 

Muchmore, W. B. 1975. Use of the spermathecae in the taxonomy of chernetid pseudoscorpions. 
Proc. 6th Int. Arachnol. Congr. 1974: 17-20. 

Muchmore, W. B. 1979. Pseudoscorpions from Florida and the Caribbean area. 8. A new species 
of Bituberochernes from the Virgin Islands (Chernetidae). Florida Entomol. 62: 313-316. 

Muchmore, W. B. 1990. Pseudoscorpionida. pp. 155-173 Jn: D. Navarro L. & J. G. Robinson 
(eds.). Diversidad Biologica en la Reserva de la Biosfera de Sian Ka’an, Quintana Roo, México. 
Centro de Investigaciones de Quintana Roo, México. 

Schawaller, W. 1986. Pseudoskorpione aus der Sowjetunion, Teil 2. (Arachnida: Pseudoscorpiones). 
Stuttgart. Beitr. Naturk. (A) 396: 1-15. 

Weygoldt, P. 1969. Biology of the pseudoscorpions. Harvard University Press, Cambridge, Massa- 
chusetts. 145 pp. 

Zeh, J. A. & D. W. Zeh. 1990. Cooperative foraging for large prey by Paratemnus elongatus 
(Pseudoscorpionida, Atemnidae). J. Arachnol. 18: 307-311. 


24 ENTOMOLOGICAL NEWS 


A NEW SPECIES OF JANUS (HYMENOPTERA: 
CEPHIDAE) FROM INDONESIA! 


David R. Smith2 


ABSTRACT: Janus ecarinatus, n. sp., is described from western Kalimantan, Indonesia. This is 
the first record of the family Cephidae for Indoncsia, the most southern record in Asia, .and one 
of the few known from the Southern Hemisphere. The absence of a genal carina and the long cer- 
cus are two of the significant characters that separate it from other known species of Janus. 


The Cephidae, stem or twig borers as larvae, are primarily a northern 
temperate, holarctic group of about 100 species in 15 genera. Representatives 
are very rare in tropical regions. The southernmost record in the Western 
Hemisphere is for Hartigia mexicana (Guérin) from the State of Chiapas, Mexico 
[ca.17°N] (Smith, 1988). One genus, Achetocephus Benson, with two species, is 
known from Madagascar [ca. 20°S] (Benson, 1946). In Asia, a species of Janus 
Stephens and one of Urosyrista Maa have been recorded from as far south as 
Burma [ca. 22°N] (Benson, 1946). 

The species described below as Janus ecarinatus from western Kaliman- 
tan, Indonesia, now represents the most southern record of Cephidae in Asia,the 
second genus and third species from the Southern Hemisphere, and the first 
record of the family from Indonesia. 

About ten species of Janus are known from Eurasia and North America. 
Larvae are twig borers, and recorded hosts are species of Malus, Populus, Pyrus, 
Ribes, Quercus, Salix, and Viburnum. Adults are distinguished from other cephid 
genera by the tarsal claw with the inner tooth longer and stouter than outer 
tooth and with an acute basal lobe; antenna not thickened apically but gradually 
thickened after the second flagellar segment and the third antennal segment 
longer than the fourth; left mandible lacking a central tooth, outer tooth simple 
and inner tooth simple with a basal shoulder on lower inner surface; apical 
maxillary palpal segment originating near the base of the penultimate segment; 
forewing with vein 2A adjacent to the posterior margin of the wing and vannal 
fold, not separated from them by more than twice the width of the vein; and one 
or two preapical spines on the hindtibia. 

Several characters typical for Janus, however, are different in J. ecarinatus. 
All previously known species of Janus have a genal carina, cercus one third or 
less the length of the sawsheath, the hindbasitarsus shorter than the following 
tarsal segments combined, and serrulae of the lancet truncate at their apices. In 
J. ecarinatus, the genal carina is absent, the cercus is nearly as long as the 


1 Received March 8, 1996, Accepted Apmnil 3, 1996. 


2 Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of 
Agriculture, c/o National Museum of Natural History, MRC-168, Washington, D.C. 20560. 


ENT. NEWS 108(1) 24-28, January & February, 1997 


Vol. 108, No. 1, January & February, 1997 25 


sheath (Figs. 2, 4), the hindbasitarsus is longer than the following tarsal seg- 
ments combined, and the serrulae of the lancet are pointed at their apices (Fig. 
5). regard these differences as more significant for species separation than for 
generic distinction, and I believe J. ecarinatus belongs in the same lineage as 
other Janus species because it shares many derived character states with them. 
The absence of a genal carina also occurs elsewhere in Cephidae: in Achetocephus 
and in one species of Pachycephus Stein (Benson, 1946), a genus of about five 
species found in the Mediterranean area and Eurasian steppes. 


Janus ecarinatus Smith, NEW SPECIES 
(Figs. 1-5) 

Female.— Length, 13 mm. Antenna with scape and pedicel yellow, flagellum black. Head yel- 
low with dorsum between eyes black and with black extension through ocellar area anteriorly to 
antennae; broad black band on occiput surrounding occipital foremen; narrow anterior margin of 
clypeus, apex of mandible, and apical maxillary palpal segment black. Thorax yellow with anterior 
half of pronotum, anterior margin of mesopleuron, line separating mesopleuron and mesepimeron, 
spot on lower angle of mesepimeron, and anterior half of mesonotum (prescutum, anterior halves 
of lateral lobes, and anterior corner of scutellum) black. Abdomen yellow with brown bands on 
apical half to two thirds of terga 2-7, narrower on 8; cercus and sheath black. Legs yellow with 
hindfemur (more yellow at base), extreme apex of hindtibia lightly, and apical two tarsal segments 
black. Wings slightly uniformly yellowish; veins and stigma light brown. 

Antenna 23- to 24-segmented, with scape slightly longer than pedicel and third segment about 
1.6X longer than fourth segment. Anterior clypeal margin rounded. Distance between antennae 
subequal to distance from antenna to tentorial pit, 0.5X distance from antenna to anterior margin of 
clypeus, and 1.6X distance between antenna and inner margin of eye. Malar space less than diam- 
eter of anterior ocellus; genal carina absent; apical maxillary palpal segment originating from base 
of penultimate segment; distance between posterior ocelli and posterior margin of head 2.3X dis- 
tance between anterior ocellus and antennae and 3.3X distance between posterior ocellus and eye; 
fine shallow punctures on head with shiny interspaces. Thorax shining with very fine scattered 
punctures, mesonotum more punctate with fine surface sculpture; abdomen shining without surface 
sculpture or punctures. Hindbasitarus 1.4X longer than length of remaining tarsal segments com- 
bined; midtibia with one preapical spine; hindtibia with 2 preapical spines; foretibia with one long, 
simple apical spine, 0.4 X,length of forebasitarsus. Vein Ir of forewing reaches stigma. Cercus 
nearly reaching to apex of sawsheath; sawsheath about 0.6X length of basal plate. Serrulae of 
lancet pointed at apices, each with several small anterior subbasal teeth. 

Male.— Unknown. 

Holotype.— Female, labeled “Indonesia: W. Kalimantan, Gunung Palung Nat. Pk., June 15 - 
August 15, 1991, Darling, Rosichon, Sutrisno, 11S 910116,” “Cabang Panti Res. Sta. 1°15'S, 110° 
S'E, 1° rainforest, Malaise trap head, Sandstone - light gap.” Deposited in the Museum Zoologu 
Bogor, Bogor, Indonesia. 


Etymology. The species name is from Latin, referring to the lack of a genal 
carina. 

Discussion.— This species differs from all other species of Janus by the 
absence of a genal carina, long hindbasitarsus, long cercus, and pointed serrulae 
of the lancet, as discussed above. The predominantly yellow coloration with 
black primarily on top of the head, anterior half of the mesonotum, and bands 
on the abdomen are also distinctive. 

Benson (1946) described a species of Janus from Burma, Maa (1949, 1950) 
described two species and gave keys to the species of China, and Muche (1981) 


26 ENTOMOLOGICAL NEWS 


gave a key to the world species. The differences cited above, however, preclude 
J. ecarinatus as a previously described species. 


Figs. 1-2. Janus ecarinatus. 1, Head, dorsal view. 2, Apical 6 or 7 abdominal segments, sheath, and 
ovipositor, lateral view. 


Vol. 108, No. 1, January & February, 1997 27 


Fig. 3-5. Janus ecarinatus. 3, Lateroventral view. 4, Cercus and sheath, lateral view. 5, Central 
serrulae of lancet. 


28 ENTOMOLOGICAL NEWS 


ACKNOWLEDGMENTS 


I thank D. Christopher Darling, Royal Ontario Museum, Toronto, Ontario, for bringing this 
specimen to my attention. Thanks also to the following for review of this manuscript: H. Goulet, 
Agriculture Canada, Ottawa; N.M. Schiff, Bee Research Laboratory, USDA, Beltsville, Maryland; 
and F.C. Thompson and R.W. Hodges, Systematic Entomology Laboratory, USDA, Washington, 
DIC. 


LITERATURE CITED 


Benson, R.B. 1946. Classification of the Cephidae (Hymenoptera Symphyta). Trans. Roy. Ento- 
mol. Soc. London 96: 89-108. 

Maa, T.-C. 1949. A synopsis of Chinese sawflies of the superfamily Cephoidea (Hymenoptera). 
Chinese Jour. Zool. 3: 17-42. 

Maa, T.-C. 1950. New or little known Chinese sawflies and woodwasps (Hymenoptera). Quarterly 
Jour. Taiwan Museum 3: 14-21. 

Muche, H. 1981. Die Cephidae der Erde (Hym., Cephidae). Dt. Entomol. Zeitschr., N.F. 28: 239- 
295. 

Smith, D.R. 1988. A synopsis of the sawflies (Hymenoptera: Symphyta) of America south of the 
United States: introduction, Xyelidae, Pamphiliidae, Cimbicidae, Diprionidae, Xiphydnidae, 
Siricidae, Orussidae, Cephidae. Syst. Entomol. 13: 205-261. 


Vol. 108, No. 1, January & February, 1997 29 


NOTES ON MATING BEHAVIOR OF 
RHIPIPHORUS LUTEIPENNIS (COLEOPTERA: 
RHIPIPHORIDAE)! 


A. G. Wheeler, Jr. 2 


ABSTRACT: The generally ephemeral adults of rhipiphorid beetles (subfamily Rhipiphorinae) 
offer few chances to observe their courtship and mating behavior. A large population of Rhipiphorus 
luteipennis in a sand plain in Oneida County, New York, during mid-August 1992 provided an 
opportunity to observe several aspects of its sexual behavior, including precopulatory swarming by 
males. 


The Rhipiphoridae, often called wedge-shaped beetles, are interesting bio- 
logically: larvae are hypermetamorphic and exhibit both an endoparasitic 
(endophagous) and an ectoparasitic (ectophagous) phase. Hosts have been re- 
ported only for about 15% of an estimated 300 species worldwide (Clausen 
1940, Linsley et al. 1952, Selander 1991). Known rhipiphorid hosts are bees 
and wasps (Rhipiphorinae), cockroaches (Rhipidiinae), and wood-boring beetles 
(Hemirhipidiinae and Pelecotominae) (Svacha 1994). 

Adults are infrequently seen, especially males, although most of the ap- 
proximately 44-54 North American species (Arnett 1985, Z. H. Falin pers. 
comm.) are not actually rare. Many rhipiphorid beetles are poorly represented 
in collections chiefly because of their short lifespan. Males of rhipiphorines 
typically live no longer than a day; females may be similarly short-lived but 
tend to emerge over a longer period than do males (Clausen 1940, Linsley and 
MacSwain 1951, Linsley et al. 1952). 

Rhipiphorus, the largest rhipiphorid genus in North America, contains about 
36, mostly western, species. Of the nine or ten species whose hosts are known 
(Linsley et al. 1952, Hurd 1979), all parasitize the immature stages of anthophorid 
or halictid bees. Habits of Rhipiphorus adults are little known except for their 
attraction to plants visited by host bees. Females often oviposit on flower buds, 
where the phoretic triungulin first instar eventually attaches to a flower-visiting 
adult bee and is carried to its provisioned nesting cell. After feeding internally, 
the rhipiphorid larva exits the host larva and begins to feed externally. The host 
larva remains alive until feeding by the parasitoid is nearly completed. After a 
pupation period of about two weeks, the free-living, ephemeral adults emerge 
from nesting sites of their hosts (Clausen 1940, Linsley et al. 1952). 

Opportunities to observe sexual activity in rhipiphorines are undoubtedly 
rare. Courting and mating pairs of R. dammersi Barber have been observed in 


1 Received April 12, 1996, Accepted May 31, 1996. 


2 Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110. 
Present address: Department of Entomology, Clemson University, Clemson, SC 29634. 


ENT. NEWS 108(1) 29-33, January & February, 1997 


30 ENTOMOLOGICAL NEWS 


California (Barber 1939), but premating and mating behavior of North Ameri- 
can species have been described only for R. smithi Linsley & MacSwain in 
California (Linsley et al. 1952). Here, I give notes on mating behavior of R. 
luteipennis (LeConte), including precopulatory male swarms observed in New 
York. 

Described from New York by LeConte (1865), R. luteipennis is also known 
from Maine, New Jersey, and Alabama in the eastern United States (Rivnay 
1929) and ranges west to Idaho, Montana, California, and New Mexico (Hatch 
1965, Linsley and MacSwain 1950, Arnett 1983). Despite its wide distribution, 
R. luteipennis is uncommon in collections, and its hosts and other biological 
information remain unknown (Linsley et al. 1952, Staines 1983). 


STUDY SITES AND METHODS 


Observations were made in the sand plains east of Holland Patent (Oneida 
Co.), N.Y. The study site is a hanging delta of glacial origin; the loose, sandy 
soil is characterized as Merrimac fine sand (Maxon et al. 1915). Scattered colo- 
nies of a panic grass, Panicum lanuginosum var. lindheimeri (Nash) Fern., and 
poverty oatgrass, Danthonia spicata (L.) F. Beauv., occur in large open patches 
of sand. Red pine, Pinus resinosa Ait., and Scotch pine, P. sylvestris L., have 
been planted within the sand plain. Small trees growing at the edge of the mainly 
open sandy area (c. 0.5 hectare) include American beech, Fagus grandifolia 
Ehrh.; gray birch, Betula populifolia Marsh.; and red maple, Acer rubrum L. 

Observations were made on 15 August 1992 between 1515 and 1645 hours. 
During this period it was about 22°C and partly cloudy, with a light breeze. The 
rhipiphorid population was encountered by chance during fieldwork on other 
insects; only a notebook was available for recording behavioral observations. 
Voucher specimens (28) of R. luteipennis are deposited in the Cornell Univer- 
sity Insect Collection, Ithaca, N.Y. 


OBSERVATIONS 


Accustomed only to encountering single or a few Rhipiphorus or Macro- 
siagon adults on flower buds and flowers of various composites and other plants, 
I was struck by the number of R. /uteipennis adults — probably several hundred 
— in the Oneida Co. sand plain. Adults were common on low vegetation (0.5- 
1.0 m), including a panic grass plant with 21 individuals on stems and inflores- 
cences: 9 unpaired adults (sex not determined) and 6 pairs with male atop fe- 
male. Whether the paired individuals were actually in copula or were exhibiting 
some postinsemination association (Alcock 1994) is unknown. Other adults, 
mostly females, were observed on stems of poverty grass or on foliage of Ameri- 
can beech saplings or a low Rubus sp. (c. 1m high). Occasionally only a single 


Vol. 108, No. 1, January & February, 1997 31 


male or female was found on a plant. Adults were never observed on larger 
plants at the periphery of the open sand. Two dead adults were on panic grass 
stems, and several others were in spider webs near ground level. 

Other adults were in flight. Two males hovered over a sapling red maple 
(c.0.5 m high), whereas groups of 5-10 males swarmed a few centimeters above 
bare sand. Swarms consisted of males and a female that apparently entered or 
became part of the swarm. A swarming male hit the sand, struggled to right 
itself, and took 10-15 seconds before resuming flight. Males could be caught by 
hand, but, when disturbed, they were capable of darting away like a bee or 
syrphid fly. 

I observed four or five unsuccessful attempts at copulation. One pair flew 
from a swarm to a grass stem where the male mounted the female; they dropped 
to the sand and grappled for several seconds before separating. Other unsuc- 
cessful pairings were observed, the males and females falling to the sand, grap- 
pling 5-10 seconds and separating without flying to vegetation. I watched another 
pair crawl over the sand with male atop female, ascend a grass stem, and sepa- 
rate without mating. Several of the brief struggles on the sand appeared to in- 
volve males. 

One apparently successful mating included nearly four minutes of grap- 
pling on the sand, followed by copulation that lasted about 1.5 minutes before 
the pair abruptly separated. Another grappling pair was observed in copula on 
the sand, but in both cases, behavior that preceded copulation was not observed. 
Twice I observed a male on top of a mating pair. 

I returned to the sand plain on 22 August, but there was no sign of the 
species. Not even a dead individual could be found in litter at the base of panic 
grass colonies or under other plants. 


DISCUSSION 


The foregoing observations on R. luteipennis, although fragmentary, invite 
comparison with those on R. smithi, the only North American rhipiphorid whose 
sexual behavior has been detailed. Linsley et a/. (1952) reported intense activ- 
ity in males of R. smithi: swarming 0.3-0.6 meters above the ground in search 
of females emerging from nests of their bee hosts. Mating typically occurred 
upon females’ emergence. Two or more males usually attended a female, the 
other males flying away when copulation was effected. Males moved upwind 
toward females, and a caged female attracted numerous males within 15 min- 
utes. These observations, coupled with that of the female’s raising her abdomen 
intermittently, suggest the presence of a female sex pheromone. The male’s 
feathery, fan-shaped (biflabellate) antennae tend to support this hypothesis 
(Linsley et al. 1952, White 1983). 

My observations on R. luteipennis, though less extensive than those of Linsley 
et al. (1952) on R. smithi, are somewhat similar. Males of both species swarm 


32 ENTOMOLOGICAL NEWS 


near the ground and occasionally attempt to copulate with one another. I did not 
observe males moving upwind toward an emerged female, but this may merely 
have reflected my inability to identify emergence of female R. luteipennis from 
nests of a presumed hymenopteran host. 

Not only is the host of R. luteipennis undiscovered and its seasonal history 
unknown, but numerous other biological data are also lacking, especially de- 
tails of the reproductive behavior. Assuming a female sex pheromone is present, 
does it operate at relatively long range, or does it serve as a contact pheromone 
within male swarms? Is copulation necessarily preceded by swarming, or cana 
female, calling from a plant, attract a male and mate? Do males and females 
mate only once or multiple times? Does mate guarding occur in this species? 

Given the ephemeral adult life of rhipiphorines, information on their repro- 
ductive behavior most likely will continue to accumulate slowly. It is hoped 
that the adaptive significance of various components of their mating behavior 
can eventually be elucidated and that enough comparative data will become 
available to allow reproductive behavior in Rhipiphorus, and in this poorly known 
family, to be placed in a phylogenetic context. 


ACKNOWLEDGMENTS 


1 am grateful to E. R. Hoebeke (Cornell University, Ithaca, N.Y.) for identifying R. 
luteipennis, Z. H. Falin (University of Kansas, Lawrence) for confirming the identification 
and providing helpful comments on the manuscript, P. H. Adler (Clemson University, Clemson, 
S.C.) for his valuable review of the manuscript, C. L. Staines (Maryland Department of 
Agriculture, Annapolis) for sending copies of several relevant papers, and R. Dirig (Cornell) 
for information on the study site. 


LITERATURE CITED 


Alcock, J. 1994. Postinsemination associations between males and females in insects: the mate- 
guarding hypothesis. Annu. Rev. Entomol. 39: 1-21. 

Arnett, R. H., ed. 1983. Checklist of the beetles of North and Central America and the West Indies. 
Vol. 6 Family 122 Rhipiphoridae. Flora & Fauna, Gainesville, Fla. 3 pp. 

Arnett, R. H. 1985. American insects: a handbook of the insects of America north of Mexico. Van 
Nostrand Reinhold, New York. 850 pp. 

Barber, H. S. 1939. A new parasitic beetle from California (Ripiphondae). Bull. Brooklyn Entomol. 
Soc. 34: 17-20. 

Clausen, C. P. 1940. Entomophagous insects, McGraw-Hill, New York. 688 pp. 

Hatch, M. H. 1965. The beetles of the Pacific Northwest. Part IV: Macrodactyles, Palpicornes, and 
Heteromera. University of Washington Press, Seattle. 168 pp. 

Hurd, P. D., Jr. 1979. Superfamily Apoidea. pp. 1741-2209. In: K. V. Krombein, P. D. Hurd, Jr., D. 
R. Smith, and B. D. Burks, eds. Catalog of Hymenoptera in America north of Mexico. Vol. 2 
Apocrita (Aculeata). Smithsonian Institution Press, Washington, D. C. 

LeConte, J. L. 1865. Note on the species of Myodites Latr. inhabiting the United States. Proc. 
Acad. Nat. Sci. Philadelphia 1865: 96-98. 

Linsley, E. G. and J. W. MacSwain. 1950. New westem species of Rhipiphoridae (Coleoptera). 
Wasmann J. Biol. 8: 229-239. 


Vol. 108, No. 1, January & February, 1997 33 


Linsley, E. G. and J. W. MacSwain. 1951. The Rhipiphoridae of California (Coleoptera). Bull. 
Calif. Insect Surv. 1(3): 79-88. 
Linsley, E. G., J. W. MacSwain, and R. F. Smith. 1952. The life history and development of 


Rhipiphorus smithi with notes on their phylogenetic significance (Coleoptera, Rhipiphoridae). 
Univ. Calif. Publ. Entomol. 9: 291-314. ; 

Maxon, E. T., M. E. Carr, and E. H. Stevens. 1915. Soil survey of Oneida County New York. 
Cornell Univ. Agric. Exp. Stn. Bull. 362. 59 pp. 


Rivnay, E. J. 1929. Revision of the Rhipiphoridae of North and Central America (Coleoptera). 
Mem. Am. Entomol. Soc. 6: 1-68. 

Selander, R. B. 1991. Rhipiphoridae (Tenebrionoidea). pp. 509-512. Jn F. W. Stehr, ed. Immature 
insects. Vol. 2. Kendall/Hunt, Dubuque, Iowa. 

Staines, C. L., Jr. 1983. The Rhipiphoridae (Coleoptera) of Maryland. Maryland Entomol. 2: 53- 
Sif: 

Svacha, P. 1994. Bionomics, behaviour and immature stages of Pelecotoma fennica (Paykull) (Co- 
leoptera: Rhipiphoridae). J. Nat. Hist. 28: 585-618. 

White, R. E. 1983. A field guide to the beetles of North America. Houghton Mifflin, Boston. 368 
Pp. 


44 ENTOMOLOGICAL NEWS 


AQUATIC COLEOPTERA AND HEMIPTERA OF 
ORGAN PIPE CACTUS NATIONAL 
MONUMENT, ARIZONA! 


Eric Larsen2, Carl Olson 


ABSTRACT: Aquatic Coleoptera and Hemiptera were collected in Organ Pipe Cactus National 
Monument (ORPI), Arizona, from March, 1940 through 1995. The Monument, a Man in the Bio- 
sphere Preserve, is located in the Sonoran Desert on the border with Sonora, Mexico, Most aquatic 
habitats in ORPI are tinajas, scoured out basins in the bedrock of ephemeral streams in the Ajo and 
Bates Mountains. Quitobaquito Springs, its associated pond, several concrete stock troughs, and 
Armenta Tank are the aquatic habitats of the valley floor. Prior to 1940 there had been no system- 
atic collection of aquatic insects. The collection consisted of five species, three beetles and two 
bugs. By 1983 the recorded fauna consisted of 61) species of aquatic Coleoptera and Hemiptera, To 
date, 42 species of aquatic Coleoptera from 5 families, and 21 species of aquatic Hemiptera from 8 
families have been collected 


Aquatic insect communities are not as well known as their terrestrial coun- 
terparts and aquatic environments in desert regions are even less well known 
than those in more temperate regions. Even so, the invertebrate faunas of streams 
have a rich history of study, due in part to an early interest by anglers in the 
invertebrate prey of fish. In recent years the fauna of ephemeral pools have 
received some attention in community studies (e.g., Larson 1985, Nilsson 1984, 
1986, Ranta 1982, 1985), and as model systems in the study of competition 
(e.g., Juliano and Lawton 1991, Larson 1985, Pajyunen 1977,1979a, 1979b, 1982, 
Vepsalainen 1978), 


‘This report on the aquatic Coleoptera and Hemiptera of Organ Pipe Cactus 
National Monument (ORPI) in southwestern Arizona contains information on 
species collected within the Monument over the past 15 years. The Monument, 
a Man in the Biosphere Preserve, is well within the boundaries of the Sonoran 
Desert (Fig. 1) and receives about 6" of rainfall per year during a winter and 
summer “rainy” season, The winter rains are generated by storms from the 
Pacific Ocean that have enough energy to continue past the Sierras of Califor- 
nia to deposit small amounts of moisture in the Sonoran Desert. This precipita- 
tion tends to come from broad based weather patterns and results in gentle rain 
fall. Conversely, moisture for the summer rains (monsoons) comes from the 
Gulf of Mexico, This rain is produced by local, intense, short-lived convection 
storms, usually within individual valley systems. These storms may deposit 


! Received November 2 1, 1995, Accepted January 7, 1996 
2 Biology Department, Villanova University, Villanova, PA 19085-1699 
4 Department of Entomology, University of Arizona, Tucson, AZ 65721 


ENT, NEWS 108(1) 34-42, January & February, 1997 


Vol. 108, No. 1, January & February, 1997 35 


BATES 
& 2) 
MOUNTAINS 


DRIPPING 


SPRINGS ~ Sx 


SS) 
500 km 


Figure |. Organ Pipe Cactus National Monument and its location in Arizona with respect to the 
boundaries of the Sonoran Desert (inset). The concrete troughs (open circles), associated with wells 
in some cases, are no longer reliable sources of water within the Monument. Dripping Springs is 
the only semi-permanent spring within the Monument. Perennial aquatic habitats are found in the 
major drainages of the Ajo Mountains and Quitobaquito Springs. Topographic lines (feet) coarsely 
outline the major features of the Monument. 


36 ENTOMOLOGICAL NEWS 


over two centimeters of precipitation in less than 30 minutes. It is perhaps this 
type of storm that scoured out the basins in the bedrock of ephemeral streams 
and produced the tinajas in the arid mountains of the Southwest (Bryan, 1920). 
These aquatic habitats are ephemeral by definition. The larger, more protected 
tinajas may persist for many years, drying up in only the most severe droughts. 


METHODS 


This report is the result of extensive surveys for aquatic beetles (Coleoptera) 
and bugs (Hemiptera) that were conducted from 1980 through 1988, with occa- 
sional surveys through 1995. There were 105 days of collecting from 1980 to 
1988, but 85 were within the first two years of the survey. Additional records 
were obtained from the ORPI insect collection and the insect collection at the 
University of Arizona. 

The water column, sides of tinajas and stock troughs, and the bottom of 
water sources were sampled. Most species were collected in the water column 
or on the gravel/muck substrate. Several of the smaller beetle species were col- 
lected by sifting through the substratum of the tinajas. Sweep samples were 
made with hand nets. Collections were preserved in 70% ethanol and trans- 
ported to the laboratory for identification. 

Of the aquatic habitats in the Monument, tinajas in Alamo Canyon and Bull 
Pasture of the Ajo Mountains and concrete stock troughs on the valley floor 
were censused most frequently. Aquatic insects were collected at Dripping 
Springs and the pond at Quitobaquito Springs on only a few occasions. At the 
time surveys for aquatic insects were initiated, Monument personnel maintained 
water in seven concrete stock troughs (Bates Well, Bonito Well, Gachado Well, 
Armenta Well, Pozo Solado, Alamo Canyon, and Blankenship Well). That prac- 
tice was abandoned in the mid 1980’s and these habitats hold water only fol- 
lowing rain. 


RESULTS 


The current aquatic beetle and bug fauna of Organ Pipe Cactus National 
Monument comprises 63 species: 42 beetle species (1 Dryopidae, 24 Dytiscidae, 
1 Gyrinidae, 1 Hydraenidae, 15 Hydrophilidae), and 21 bug species (2 
Belostomatidae, 5 Corixidae, | Gerridae, 1 Mesoveliidae, 3 Naucoridae, | 
Nepidae, 7 Notonectidae, | Veliidae,) (Table 1). 

Only five aquatic insects were in the ORP1 collection prior to a survey of 
aquatic insects conducted by a graduate class from the University of Arizona in 
1980. Twenty-seven species were added from that survey. Since that time, an 
additional 31 species have been found within ORP1 over a 3 year period. As 
there have been no new species collected in ORP1 in the last decade of inter- 


Vol. 108, No. 1, January & February, 1997 37 


mittent sampling, it is reasonable to assume that at least the common or regular 
residents of the Monument have been documented. Indeed, of the 63 species 
collected so far, only 11 are represented by a single specimen (E. Larsen, un- 
published data). 

Of the current aquatic habitats represented in the Monument, more species 
of beetles have been collected in the tinajas of the Ajo Mountains than in the 
other habitats (Table 2). Species collected in the other habitats are, for the most 
part, a subset of those found in the Ajo Mountains, with 7 exceptions: 4 species 
of dytiscids and 3 species of hydrophilids. In all cases, these species were rare 
in the Monument (E. Larsen, unpublished data). Obviously, the absence of some 
species in collections from the pond at Quitobaquito Springs may be due to a 
lesser sampling effort than expended in the Ajo Mountains, but the absence of 
some dytiscid species (e.g., Hydreana circulate, some species of Laccophilus, 
and Copelatus spp.) is probably due to a preference for gravel substrate not 
present in the pond. 

Virtually all of the specimens of beetles from Dripping Springs were col- 
lected in the outflow, a muddy trickle several meters long that appears when 
outflow exceeds evaporation. As this occurrence is relatively rare and the 
majority of the basin that collects water is nearly completely hidden from above 
by overhanging rock, few aquatic insects were ever found here on any particu- 
lar collecting trip. 

Heteropteran species do not show the same nested pattern of occurrence 
exhibited by the coleopteran species. Species found in the pond at Quitobaquito 
Springs are not a subset of species found in the tinajas (Table 3). Twelve of 18 
species were found in either the pond or the tinajas, but not both. Of the species 
found in both habitats, 3 skate or walk across the surface of the water in pursuit 
of prey (the gerrid, the mesoveliid, and the veliid) and perhaps are less restricted 
by the nature of the habitat substratum than the fully aquatic insects. 


DISCUSSION 


The aquatic insect fauna of ORPI is a mixture of temperate and tropical 
insects with a few Southwest endemics. This is illustrated by two genera of 
Dytiscidae. Of the six species of Laccophilus, L. pictus, L. vaecensis, and L. 
mexicanus are largely tropical, L. fasciatus is widespread, and L. sonorensis is 
endemic to the Southwest (Zimmerman 1970). Five species of Deronectes have 
been collected in ORPI; D. striatellus is widespread, D. roffiand D. coelamboides 
are Southwest endemics, and D. aequinoctialis is tropical, reaching the north- 
ern extent of its range in Arizona (Zimmerman and Smith 1975). Among the 
bugs, the Notonectidae exhibit a similar pattern. Seven species of backswimmers 
have been collected within the Monument. Notonecta indica, N. kirbyi, B. 
margaritacea, and B. scimitra have temperate distributions, N. lobata and B. 


38 ENTOMOLOGICAL NEWS 


arida are endemic to the Southwest, and B. albida and B. arizonis are tropical 
(Truxal 1953, Hungerford 1933). Both of the tropical species reach the north- 
ern-most portions of their ranges in Arizona. Among the naucorids, Pelocoris 
biimpressus biimpressus, has been recorded from Guatemala, Mexico, and sev- 
eral States in the Southwest (LaRivers, 1956, Polhemus and Sites 1995). 
Ambrysus occidentalis is endemic to the Southwest and A. pulchellus reaches 
the most northern extent of its range in southern Arizona, southern New Mexico, 
and southwest Texas (La Rivers, 1951). 

Little evidence for habitat selection between the pond and the tinajas was 
observed in the beetle assemblages (Table 2). The species collected only at the 
pond were rare, represented by only one or two specimens over the past 15 
years (E. Larsen, unpublished data). In contrast, of the species unique to the 
Ajo Mountains (i.e., Agabus lugens, Copelatus spp., Deronectes aequinoctialis, 
Desmopachria portmani, Dineutus sublineatus, and Enochrus carinatus), only 
Dineutus sublineatus is represented by a single observation (E. Larsen unpub- 
lished data). 

All families of bugs represented by more than one species in ORPI exhibit 
nearly mutually exclusive distribution patterns between the tinaja and pond habi- 
tats (Table 3). The nature of the habitat preferences is unknown, but may reflect 
habitat choice at the moment of colonization (e.g., some species will not colo- 
nize a small habitat such as that represented by a tinaja relative to a pond) or 
physiological tolerances (e.g., the tinajas would have greater temperature fluc- 
tuations than the pond due to their smaller size). 


New Records 

All species recorded in this report are new site records for Arizona. The 
following three species are new Arizona records. The dytiscid, Copelatus 
distinctus, has few records in the U.S., but is known to occur throughout Mexico 
(Young, 1963). It was abundant in the gravel bottoms of tinajas in the Ajo Moun- 
tains during the time course of these surveys. The closest published records for 
the mesoveliid, Mesovelia mulsanti White, are from California and Texas (Smith 
1988). The notonectid, Buenoa albida, has been reported from Texas and Mexico 
(Truxal 1953). In addition to this record, B. albida has been collected just out- 
side of ORPI in Armenta Tank, as well as other locales in Pima and Yuma Coun- 
ties (Larsen, 1986 and C. Olson, unpublished data). 

The environment of Organ Pipe Cactus National Monument is harsh. For 
aquatic insects to gain a foot-hold in this rugged environment they must be able 
to find the isolated tinajas, survive the often elevated temperatures in tinajas 
unshaded by canyon walls, and successfully reproduce. The number of aquatic 
species reported here demonstrates that finding the tinajas is not a problem. The 
presence of beetle larvae and bug nymphs in the peak of the Sonoran Desert 
summer (E. Larsen, personal observation) is strong evidence that the physical 
environment is not beyond the physiological capabilities of these insects. 


Vol. 108, No. 1, January & February, 1997 39 


Table 1. Aquatic Coleoptera and Hemiptera collected within the boundaries of Organ Pipe 


Cactus National Monument. Arizona. 


COLEOPTERA 
Dryopidae 
Helichus immsi Hinton 


Dytiscidae 
Agabus lugens (LeConte) 
Copelatus chevrolati renovatus Guignot 
C. distinctus Aube 
Cybister ellipticus LeConte 
Deronectes aequinoctialis (Clark) 
D. coelamboides Fall 
D. decemsignatus (Clark) 
D. roffi (Clark) 
D. striatellus (LeConte) 
Desmopachria portmani Clark 
Eretes sticticus (Linnaeus) 
Hydroporus vilis LeConte 
Hygrotus fraternus LeConte 
H. medialis (LeConte) 
Laccophilus fasciatus terminalis Sharp 
L. maculosus shermani Leech 
L. mexicanus mexicanus Aube 
L. pictus coccinelloides Regimbart 
L. sonorensis Zimmerman 
L. vaecensis Young 
Rhantus atricolor (Aube) 
R. gutticollis Say 
Thermonectus nigrofasciatus (Aube) 
Uvarus sp. 


Gyrinidae 
Dineutus sublineatus Cheverolat 


Hydraenidae 
Hydreana circulate Perkins 


Hydrophilidae 
Berosus infuscatus LeConte 
B. miles LeConte 
B. punctatissimus LeConte 
B. rugulosus Horn 
B. salvini Sharp 
B. stylifer Horn 
Chaetarthria pallida (LeConte) 
Enochrus carinatus fuscatus (Horn) 
E. cristatus (LeConte) 
E. pygmaeus pectoralis (LeConte) 
Helochares normatus (LeConte) 
Helophorus brevipalpus Bedel 
Hydrophilus triangularis (Say) 
Tropisternus ellipticus (LeConte) 
T. lateralis LeConte 


HEMIPTERA 
Belostomatidae 
Belostoma flumineum Say 
Lethocerus medius (Guerin-Meneville) 


Corixidae 
Corisella edulis (Champion) 
Graptocorixa abdominalis (Say) 
G. serrulata (Uhler) 
Hesperocorixa laevigata (Uhler) 
Trichocorixa sp. 


Gerridae 
Aquarius remigis (Say) 


Mesoveliidae 
Mesovelia mulsanti White 


Naucoridae 
Ambrysus occidentalis LaRivers 
A. pulchellus Montandon 
Pelocoris biimpressus biimpressus Montandon 


Nepidae 
Ranatra quadridentata Stal 


Notonectidae 
Buenoa albida (Champion) 
B. arizonis Bare 
B. margaritacea Torre-Bueno 
B. scimitra (Bare) 
Notonecta indica Linnaeus 
N. kirbyi Hungerford 
N. lobata Hungerford 


Veliidae 
Microvelia beameri McKinstry 


40 ENTOMOLOGICAL NEWS 


Table 2. Habitats for aquatic Coleoptera collected in Organ Pipe Cactus National Monument, 
Arizona. 


Ajo Quitobaquito Dripping Concrete 
Mountains Springs Springs Troughs 


Dryopidae 
Helichus immsi 


Pas 


X 


Dytiscidae 
Agabus lugens 
Copelatus c. renovatus 
C. distinctus 
Cybister ellipticus 
Deronectes aequinoctialis 
D. coelamboides 
D. decemsignatus 
D. roffi 
D. striatellus 
Desmopachria portmani 
Eretes sticticus 
Hydoporus vilis 
Hygrotus fraternus 
H. medialis 
Laccophilus f. terminalis 
L. m. shermani 
L. mexicanus 
L. p. coccinelloides 
L. sonorensis 
L. vaecensis 
Rhantus atricolor 
R. gutticollis 
Thermonectus nigrofasciatus 
Uvarus sp 


x KR «K 
*< xxx OK 


HK KKK KK OK OK OK OK OK OK OK OK OK 
KKK Mm KM KKK OK 


Gyrinidae 
Dineutes sublineatus X 


Hydreanidae 
Hydreana circulata > 4 


~ 


Hydrophilidae 
Berosus infuscatus xX 
B. miles 
B. punctatissimus 
B. rugulosus 
B. salvini 
B. stylifer 
Chaetarthria pallida X 
Enochrus c. fuscatus 
E. cristatus 
E. p. pectoralis 
Helochares normatus 
Helophorus brevipalpus 
Tropisternus ellipticus 
T. lateralis 


xx mK 
KKM mK 


exe KKK 


xx KK 


Vol. 108, No. 1, January & February, 1997 4] 


Table 3. Habitats for aquatic Heteroptera collected in Organ Pipe Cactus National Monu- 
ment, Arizona. 


Ajo Quitobaquito Dripping Concrete 
Mountains Springs Springs Troughs 


Belostomatidae 
Belostoma flumineum xX 
Lethocerus medius Xx 


~< 


Corixidae 
Corisella edulis 
Graptocorixa abdominalis 
G. serrulata 
Hesperocorixa laevigata X 
Trichocorixa sp. 4 


mx mK 
PKK OM OS 


Gerridae 
Aquarius remigis x x 


~< 


Mesoveliidae 
Mesovelia mulsanti Xx Xx 


Naucoridae 
Ambrysus occidentalis 
A. pulchellus 
Pelocoris b. biimpressus 


x KX 


Nepidae 
Ranatra quadridentata x 


~< 
Pas 


Notonectidae 
Buenoa albida 
B. arizonis Xx 
B. margaritacea ? 
B. scimitra 
Notonecta indica 
N. kirbyi x 
N. lobata Xx 


xx Kx 


xxx KM 


Veliidae 
Microvelia beameri Xx X X 


Ps 


ACKNOWLEDGMENTS 


This work was carried out with the cooperation and assistance of the staff of Organ Pipe 
Cactus National Monument. Terry Peters, the former Natural Resources Specialist, and 
Caroline Wilson, Chief Naturalist, deserve special thanks for logistic support. Voucher speci- 


42 ENTOMOLOGICAL NEWS 


mens have been deposited in the University of Arizona Entomology collection and the col- 
lection of Organ Pipe Cactus National Monument. We thank Gene Hall, David Madison, J.T. 
Polhemus, Dan Polhemus, and an anonymous reviewer for many useful comments on the 
manuscript. 


LITERATURE CITED 


Bryan, K. 1920. Origin of rock tanks and charcos. Am. J. Sci. 50: 188-206. 

Hungerford, H.B. 1933. The genus Notonecta of the world (Notonectidae-Hemiptera). Univ. Kans. 
Sci. Bull. 21: 5- 195. 

Juliano, S.A., and J.H. Lawton. 1990. The relationship between competition and morphology: I. 
Morphological patterns among co-occurring dytiscid beetles. J. Anim. Ecol . 59:403 -419. 
LaRivers, I. 1951. A revision of the genus Ambrysus in the United States. Univ. Calif. Publ. Ent. 8: 

277-338. 

LaRivers, I. 1956. A new subspecies of Pelocoris shoshone from the Death Valley drainage. Wash. 
J. Biol. 14: 155- 158. 

Larsen, E. 1986. Community structure of backswimmers (Hemiptera, Notonectidae) in the South- 
west: A group of predaceous aquatic insects. Ph.D. dissertation. Univ. Arizona, Tucson. 

Larson, D.J. 1985. Structure in temperate predaceous diving beetle communities (Coleoptera: 
Dytiscidae). Holarctic Ecol. 8:18-32. 

Nilsson, A. N. 1984. Species richness and succession of aquatic beetles in some kettle-hole ponds 
in northern Sweden. Holarctic Ecol. 7: 149-156. 

Nilsson, A. N. 1986. Community structure in the Dytiscidae (Coleoptera) of a northern Swedish 
seasonal pond. Ann. Zool. Fenn. 23:39-47. 

Pajunen, V.I. 1977. Population structure in rock pool corixids (Hemiptera: Corixidae). Ann. Zool. 
Fennici | 4:26-47. 

Pajunen, V.I. 1979a. Competition between rock pool corixids. Ann. Zool. Fennici 16: 138-143. 
Pajunen, V.I. 1979b. Quantitative analysis of competition between Arctocorixa carinata (Sahlb.) 
and Callicorixa producta (Reut.) (Hemiptera: Corixidae). Ann. Zool. Fennici 16: 195-200. 
Pajunen, V.I. 1982. Replacement analysis of non-equilibrial competition between rock pool corixids 

(Hemiptera: Corixidae). Oecologia 52:153-155. 

Polhemus, J.T., and R.W. Sites. 1995. The identity of Pelocoris biimpressus Montandon and syn- 
onymy of Pelocoris species in the southwestern United States (Heteroptera: Naucoridae). Proc. 
Entomol. Soc. Wash. 97: 654-658. 

Ranta, E. 1982. Animal communities in rock pools. Ann. Zool. Fenn. 19:337-347. 

Ranta, E. 1985. Communities of water beetles in different kinds of waters in Finland. Proc. Acad. 
Nat. Sci. Phil. 137:33-45. 

Smith C.L. 1988. Family Mesoveliidae Douglas and Scott, 1867. pp. 247-248. Jn T.J. Henry and 
R.C. Froeschner, eds. Catalog of the Heteroptera, or True Bugs, of Canada and the Continental 
United States. E.J. Brill, NY. 

Truxal, F.S. 1953. A revision of the genus Buenoa (Hemiptera: Notonectidae). Univ. Kans. Sci. 
Bull. 35: 1351-1523. 

Vepsalainen, K. 1978. Coexistence of two competing corixid species (Heteroptera) in an archi- 
pelago of temporary rock pools. Oecologia 37: 177-182. 

Young, F. N. 1963. The Nearctic species of Copelatus Erickson. Quart. J. Florida Acad. Sci. 26: 56- 
Tie 

Zimmerman, J.R. 1970. A taxonomic revision of the aquatic beetle genus Laccophilus (Dytiscidae) 
of North America. Mem. Amer. Ent. Soc. 26: 1-275. 

Zimmerman, J.R., and A.H. Smith. 1975. A survey of the Deronectes (Coleoptera: Dytiscidae) of 
Canada, the United States, and northern Mexico. Trans. Amer. Ent. Soc. 101:651-722. 


Vol. 108, No. 1, January & February, 1997 43 


REPORT OF A MORPHOLOGICALLY 
HERMAPHRODITIC FLEA (SIPHONAPTERA) AND 
OTHER FLEA ANOMALIES FROM MOROCCO! 


Michael W. Hastriter2 


ABSTRACT: A case of morphological hermaphrodism in Nosopsyllus henleyi henleyi and anoma- 
lous conditions in specimens of Xenopsylla ramesis, Leptopsylla segnis, and Stenoponia tripectinata 
megaera from Morocco are described and illustrated. 


Numerous authors have reported a variety of anomalies among Siphonaptera. 
Bartkowska (1968), Beaucournu et al. (1988), Goncharov (1972a), Haas (1965; 
1983b), Holland (1943), Jordan (1921), Lopes et al. (1992), Mead-Briggs (1964), 
Sharma and Joshi (1961), Smit (1949a; 1949b), and Worms (1973) discussed 
cases of double spermathecae. Triple spermathecal conditions are reported by 
Beaucournu et al. (1988), Benton (1967), and Stark (1953). Various anomalous 
conditions to include castration of male and/or female fleas were reported by 
Bartkowska (1966; 1968), Beaucournu (1969), Benton (1967), Brinck-Lindroth 
and Smit (1973), Claassens (1967), Goncharov (1972b), Haas (1972; 1983a), 
loff and Tiflov (1940), Li and Wang (1973), Smit (1952; 1953), and Wagner 
(1931). Holland (1959) and Jordan (1921) suggested possible evolutionary sig- 
nificance of reoccurring anomalous conditions. Deformities of flea appendages 
were reported by Sanjeans and Travis (1955), Schwan and Dobkin (1981), and 
Smit (1981). Akofyan (1961), Brink-Lindroth and Smit (1973), Goncharov 
(1972b), and Haas (1983a) each reported descriptive cases of intersexuality 
defined by Brink-Lindroth and Smit as “dominantly male..., but also the struc- 
tures of secondary sexual characters, such as head, antennae, antesensilial 
setae, and postsensilial region, which are then more or less intermediate be- 
tween those of male and female”. Beaucournu and Launay (1987) described 
and illustrated a case of morphological hermaphrodism in Nosopsyllus henleyi 
and distinguished it from cases of intersexuality as, “co-existence of male geni- 
talia (segment [X and phallosome) and female genitalia (vagina, duct and sperma- 
theca)”. The author does not consider Brink-Lindroth and Smit’s definition of 
intersexuality to be the same as hermaphrodism. Based on the definition of 
Beaucournu and Launay (1987), a second hermaphrodite, noted in Hastriter 
and Tipton (1975), is herein described with significant biological similarities to 
their report. 


! Received March 12, 1996. Accepted June 19, 1996. 


2 Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, P.O. Box 20200, 
Provo, Utah 84602-0200. 


ENT. NEWS 108(1) 43-51, January & February, 1997 


44 ENTOMOLOGICAL NEWS 


MATERIALS AND METHODS 


During 1969-70, 9,936 fleas were collected from small mammals in 
Morocco (Hastriter and Tipton, 1975). Among these fleas were five anomalies. 
All specimens were processed using KOH, dehydrated in serial alcohols, cleared 
in oil-of wintergreen, and mounted in Canada balsam. Terminologies follow 
those of Rothschild and Traub (1971). 


OBSERVATIONS AND DISCUSSION 


A single specimen identified as Nosopsyllus henleyi henleyi (Rothschild, 
1904) collected 15 kilometers southwest of Missour, Fes Province, Morocco, 
from female Meriones libycus, 28 Jun 1970 by L.W. Robbins had primary and 
secondary male and female sexual characteristics commensurate with a true 
morphological hermaphrodite. 


Head: Setation of the head is similar in males and females, although males 
possess a single stout seta that females do not have, located ventral to the anten- 
nal fossa and anterior to the ocular row. This seta is not present in the hermaph- 
rodite. The specimen has a normal male occipital groove, but it does not extend 
forward as deeply toward the interantennal suture (falx). The dorsal antennal 
fossae do not completely merge forming a definite interantennal suture or falx 
as in normal males; however, the separation is not as great as in females. The 
antennae bear important secondary sexual characteristics of both sexes. Bilater- 
ally, the scape is typically male, the second segment bears long setae character- 
istic only of females, and the clubs are male. The left antenna has two missing 
terminal segments leaving seven, and does not extend onto the thoracic depres- 
sion (antennal fossa) of the proepisternum. 

Thorax: The thorax of Nosopsyllus henleyi henleyi does not significantly 
differ in males and females except for the proepisternal region. In males, the 
heavy ridge separating the dorsal and ventral pleural region (hereafter termed 
the propleural strut) is strongly arched ventrad, extending the cephalic portion 
of the antennal fossa to accommodate the longer male club. In females, this 
propleural strut bows slightly upwards as the antennal fossa does not extend 
onto the proepisternum below the cervical link plate (Figs. 1-3). The antennal 
fossa of the proepisternum is less developed than normal males and the last two 
segments of the male right antenna are abnormally bent to conform to this poorly 
developed groove. This anomalous condition may have attributed to the loss of 
the two terminal segments on the left side, as no protection was provided. The 
pronotal combs of male and female N. h. h. do not exhibit sexual dimorphism as 
in some species. 

Legs and Abdomen: Apparent secondary sexual features are not associ- 
ated with the legs, or first six segments of the abdominal tergites and sternites. 


Vol. 108, No. 1, January & February, 1997 45 


Figs. 1-3. Nosopsyllus h. henleyi prosternosomes. Fig. 1. Normal female. Fig. 2. Normal male. Fig. 
3. Hermaphrodite. 


Figs. 4-6. N.h.h. antepygidial bristles, eighth tergite (T-8), and dorsal/ventral anal lobes. Fig. 4. 
Normal female. Fig 5. Normal male. Fig. 6. Hermaphrodite — rudimentary anal stylet (an.sty.) 
shown enlarged. 


46 ENTOMOLOGICAL NEWS 


Tergum VII - Normal males and females possess two and three antepy gidial 
bristles, respectively, arranged as in Figures 4-5. This specimen has the same 
number and arrangement as typical females with exception of the ventral bristle, 
which is slightly shorter (Fig. 6). 

Tergum VIII - The shape of T-8 of the anomaly is characteristically male, 
but differs slightly in contour from either sex and has the setal pattern of neither 
(Figs. 4-6). 

Tergum IX - The 7th abdominal spiracle is situated within the sensilial plate 
as in normal males. Female spiracles are not elongate in lateral view as are 
males. The spiracle is slightly deformed and situated anteriorly more than nor- 
mal males (Figs. 4-6). The P1 and movable process of the clasper, apodeme of 
T-9, and manubrium are slightly deformed, but well developed (Fig. 7). 

Dorsal and Ventral Anal Segments - These terminal segments are malformed 
and sexually intermediate. The dorsal anal segment of the hermaphrodite 
resembles the normal female as it bears an almost normal anal stylet (Fig. 6). It 
also bears a reduced, but clear rudiment of a female anal stylet bilaterally on the 
ventral anal lobe, which usually has no such appendages. It is of interest to note 
that Goncharov (1972b) illustrates cases of rudimentary anal stylets in numer- 
ous specimens of Nosopsyllus laeviceps (Wagner, 1909) arising from what he 
terms the eleventh segment. The author does not wish to discuss the numerical 
interpretation of terminal segments, but points out that the rudimentary anal 
stylets illustrated by Goncharov occupy the same position on the ventral anal 
segment as the hermaphrodite herein described. These two closely related flea 
species evidently display similar and possibly atavistic expression of “anoma- 
lous” traits when exposed to teratogenic agents. 

Sternum VII - The S-7 is typical of normal males. The S-8, although very 
reduced in normal specimens of WN. h. h., is completely nondiscernable in this 
specimen. 

Sternum IX - The S-9 appears in duplicate, although the more posterior 
ventral of the two pairs are grossly deformed. The anterior dorsal pair are 
essentially normal except for malformations of the proximal arms and apodemal 
rods (Fig. 7). 

Aedeagus - The aedeagus does not significantly differ from normal males, 
although the crochets are bluntly deformed and the penis rods are vastly shorter 
than normal males whose penis rods coil three times like a watch spring. The 
penis rods do not exceed a half coil (Fig. 7). The paired Wagner’s gland was 
clearly visible. Testicular tissues were not visible as preparation procedures had 
undoubtedly destroyed them. 

Female Genital Features - The primary sexual characteristics that define 
this specimen as a true hermaphrodite morphologically include a small and 
deformed spermatheca, complete with strigillae, a cribriform area surrounding 
the bulge orifice, and a duct leading from the spermatheca that broadens into a 


Vol. 108, No. 1, January & February, 1997 47 


100y 


Figs 7-8. Nosopsyllus h. henleyi hermaphrodite. Fig. 7. Male clasper, anterior aedeagus (terminal 
aedeagus not illustrated, dual ninth sternite (S-9), bursa copulatrix, spermatheca and duct. Fig. 8. 
Enlargement of spermatheca (sper.), bursa copulatrix (b.c.), and pars dilatata (p.d.). 


48 ENTOMOLOGICAL NEWS 


nearly non-discernible pars dilatata at the juncture of the bursa copulatrix (Fig 
8). The pars dilatata extends from a grossly deformed and anatomically 
inverted sclerotized duct of the bursa copulatrix (Fig. 7). The position of the 
bursa copulatrix may have been anatomically altered during processing. A vaginal 
canal is not visible with the imposition of heavily chitinized male structures. 

The hermaphrodite, N. h. henleyi, described by Beaucournu and Launey 
(1987) is remarkably like this specimen not only morphologically, but biologi- 
cally. Their specimen was collected in Laghouat, Algeria approximately 375 
miles east of Missour, Fes Province, Morocco from the same host species, 
Meriones libycus. The nominate subspecies of Nosopsyllus henleyi occurs on 
M. libycus and M. caudatus (some consider the latter to be a subspecies of M. 
libycus) in arid lowland areas of Morocco east of the Middle Atlas Mountains. 
The common ecological attributes shared by the only two reported hermaphro- 
ditic Siphonaptera removed from the same host species with similar geographi- 
cal habitats can likely be attributed to an entomophagous Mermethid nematode 
identified coincidentally in N. h. h. by Deunff (1984). It should be noted that 
there is wide, uniform geographical distribution for these mammalian host spe- 
cies and their parasites (nematodes and Siphonaptera) from Morocco to 
Laghouat, Algeria and across North Africa. 


Miscellaneous Anomalies: Three specimens had single anomalous sper- 
mathecae with dual hillae. They include: Leptopsylla segnis (Schénherr, 1811) 
(3 kilometers north of Settat, Settat Province, Ex: female Rattus norvegicus, 23 
Apr 1970 by L.W. Robbins) (Figs. 9-10); Xenopsylla ramesis (Roths., 1904) (8 
kilometers south of Tiznit, Agadir Province, Ex: male G. campestris, 18 Dec 
1969 by L.W. Robbins) (Figs. 11-12); and Stenoponia tripectinata megaera 
Jordan, 1958 (16 kilometers west of Aoulouz, Agadir Province, Ex: female 
Gerbillus campestris, 14 Mar 1970 by R.E. Vaden) (Figs. 13-14). 

A single male specimen of X. ramesis (9 kilometers southeast of Al Hoceima, 
Al Hoceima Province, Ex: male G. campestris, 31 May 1970 by M.G. Hearst) 
bears an extra P1 of the clasper only on the left side. It is distinct, but smaller 
than the anatomically normal adjacent P1 (Figs. 15-16). 

It is interesting to note that of the 9,936 fleas removed from 38 mammal 
species (Hastriter and Tipton, 1975), three of the four miscellaneous anomalies 
were all taken from G. campestris. The inferred teratogenic etiology of a com- 
mon entomophagous nematode parasite among the G. campestris population is 
probable. Studies of nematode parasites infesting murine rodents and their fleas 
in the region warrant further investigation similar to those of Brink-Lindroth 
and Smit (1973) in northern Scandinavia. In addition, numerous authors have 
observed and reported entomophagous nematode parasites in the coelomic cav- 
ity of adult fleas, but the infestation of pupal and prepupal stages have not been 
reported. Surely the occurrences of dual anatomical structures and coexisting 


Vol. 108, No. 1, January & February, 1997 49 


Nena hihi v 


Figs 9-16. Respectively, normal and dual tailed spermathecae of: Leptopsylla segnis (Figs. 9-10), 
Xenopsylla ramesis (Figs. 11 12), and Stenoponia tripectinata maqaera (Figs. 13-14). Figs. 15-16. 
Processes of clasper of X. ramesis. Fig. 15. Normal P1, P2 and P3 of clasper. Fig. 16. Duel P1, 
normal P2 and P3 of clasper. 


50 ENTOMOLOGICAL NEWS 


male/female organs are likely to be produced by parasitic trauma (physical and/ 
or chemical) during earlier developmental stages. The methods of parasite 
invasion, timeliness of infection, and effects of parasitosis all deserve addi- 
tional inquiry. 


ACKNOWLEDGMENTS 


The author expresses his appreciation to Richard W. Baumann, Curator of Insects, Monte L. 
Bean Museum, Brigham Young University for kindly providing space and equipment. Also, special 
thanks to Glenn E. Hass and Harold E. Stark for reviewing the manuscript and providing helpful 
suggestions. 


LITERATURE CITED 


Akofyan, M.M. 1961. On the problem of parasitic castration of fleas. Vest. Mikrobiol., Epidem. 
Parazit. 3: 562-67. 

Bartkowska, K. 1966. A case of monstrosity in Ctenophthalmus assimilis (Tasch.) (Aphaniptera). 
Annls. Zoologici. 23(24): 535-39. 

Bartkowska, K. 1968. Cases of monstrosities in Ctenophthalmus assimilis (Tasch.) (Siphonaptera). 
II. Annls. Zoologici. 26(16): 355-61. 

Beaucournu, J.C. 1969. Quelques cas de teratologie chez les Siphonaptera. Annls. Parasitol. Hum. 
Comp. 44(2): 173-96. 

Beaucournu, J.C. and H. Launay. 1987. Description du premier cas d’*hermaphrodisme chez un 
Siphonaptere. Annls. Soc. Entomol. Fr. 23(3): 273-77. 

Beaucournu, J.C., H. Launay and A. Sklair. 1988. Les anomalies des spermatheques et des con- 
duits genitaux chez les Siphonapteres (Insecta) - Revue bibliographique et cas personnels. Annls. 
Parasitol. Hum. Comp. 63(1): 64-75. 

Benton, A.H. 1967. A case of teratology in Monopsyllus vison (Baker). Jour. N.Y. Entomol. Soc. 
75(1): 31-33. 

Brinck-Lindroth, G. and F.G.A.M. Smit. 1973. Parasitic nematodes in fleas in northern Scandinavia 
and notes on intersexuality and castration in Amphipsylla sibirica Wagn. Entomol. Scand. 4: 
302-22. 

Claassens, A.J.M. 1967. Morphological anomalies in fleas (Siphonaptera). Jour. Entomol. Soc. 
South Afr. 29: 124-34. 

Deunff, J. 1984. Parasites de Siphonaptéres. Etude de la systématique, de La biologie et du pouvoir 
pathogéne des Tylenchides (Nematodea) dans une perspective de Lutte biologique. Thése Doct. 
Etat Sc. Pharm., Univ. Rennes, 406 pp. 

Goncharov, A.I. 1972a. On the female of Ceratophyllus styx riparius Jordan & Rothschild, 1920 
with two spermathecae (Siphonaptera: Ceratophyllidae). Parazitologiya. 6: 133-36. 

Goncharov, A.I. 1972b. On the basic structure of the abdomen in Siphonaptera, based on studies of 
castrated males (Siphonaptera: Ceratophyllidae, Leptopsyllidae). Parazitologiya 6: 465-68. 

Haas, G.E. 1965. Another specimen of Opisocrostis bruneri (Baker) with two spermathecae 
(Siphonaptera). Jour. Med. Entomol. 2(2): 140. 

Haas, G.E. 1972. Partial castration in Monopsyllus vison (Siphonaptera). Entomol. News. 83(10): 
275-78. 

Haas, G.E. 1983a. Intersexes and abnormal genitalia in Alaskan fleas (Siphonaptera). Jour. Ari- 
zona-Nevada Acad. Sci. 18: 4-12. 

Haas, G.E. 1983b. Anomalies of the head, thorax and abdomen in the Order Siphonaptera. Adv. 
Biosci. 2: 133-44. 


Vol. 108, No. 1, January & February, 1997 51 


Holland G.P. 1943. A remarkable instance of retention of a double spermatheca in a Dolichopsyillid 
flea, Opistocrostis bruneri (Baker). Can. Entomol. 75: 175-76. 

Holland, G.P. 1959. An unusual case of teratology in Siphonaptera. Can. Entomol. 91: 703-09. 

loff, I.G. and V.E. Tiflov. 1940. Material for the study of fleas. IV. Additonal notes concerning the 
genus Coptopsylla. Flea castration by parasitic Nematodes. Vest. Mikrobiol. Epidem. Parazit. 
19(1): 98-103. 

Jordan, K. 1921. A link between the double and single receptacula seminis of Siphonaptera. Ecto- 
parasites. 1(3): 127 28. 

Li Kuei-chen and Wang Tun-ch’ing. 1973. A study on the teratology of fleas. Acta Entomol., 
Sinica. 16:154-62. 

Lopes, C.M.L., P.M. Linardi and J.M. Botelho. 1992. Duplication of the spermatheca in Polygenis 
tripus (Jordan)(Siphonaptera: Rhopalopsyllidae). Jour. Med. Entomol. 29(1): 111-12. 

Mead-Briggs, A.R. 1964. Structural abnormalities in the spermathecal system of two specimens of 
Spillopsyllus cuniculi (Dale) (Siphonaptera). Entomol. Gaz. 15(1): 35-38. 

Rothschild, M. and R. Traub. 1971. A revised glossary of terms used in the taxonomy and mor- 
phology of fleas. The British Museum (Natural History), London, pp. 7-85, 24 figs., 13 plates. 

Sanjeans, J. and B.V. Travis. 1955. An eight-legged flea, Orchopeas howardi howardi (Baker). 
Jour. Parasit. 41: 636-37. 

Schwan, T.G. and D.S. Dobkin. 1981. An unusual example of teratogenesis in the flea Thrassis 
fotus from Colorado (Siphonaptera: Ceratophyllidae). Proc. Entomol. Soc. Wash. 83(1): 93-98. 

Sharma, M.I.D. and G.C. Joshi. 1961. An abnormal form of female rat flea, Xenopsylla cheopis 
Roths. Nature. 191(4789): 727. 

Smit, F.G.A.M. 1949a. Monstrosities in Siphonaptera. Tiidschr. Entomol. 90: 35-42. 

Smit, F.G.A.M. 1949b. Monstrosities in Siphonaptera II. Entomol. Bericht. 12(293): 436-37. 

Smit, F.G.A.M. 1952. Monstrosities in Siphonaptera III. Entomol. Bericht. 14: 182-187. 

Smit, F.G.A.M. 1953. Monstrosities in Siphonaptera IV, more cases of castration. Entomol. Bericht. 
14: 393-400. 

Smit, F.G.A.M. 1981. A Case of tarsal trichotomy in Siphonaptera. Entomol. Bericht. 41:14. 

Stark, H.E. 1953. An unusual occurrence of three spermthecae in a specimen of Hystrichopsylla 
dippiei (Siphonaptera). Pan-Pacif. Entomol. 29: 135-37. 

Wagner, J. 1931. Ein fall der “geschlechtslosigkeit” bei Insekten. Acta Soc. Entomol. Jugoslav. 5- 
6: 30-38. 

Worms, M.J. 1973. Partial duplication of the reproductive system of female Ceratophyllus species 
(Siphonaptera). Entomol. Gaz. 24: 327-29. 


a2 ENTOMOLOGICAL NEWS 


SISTER RELATIONSHIP OF THE 
NEOEPHEMERIDAE AND CAENIDAE 
(EPHEMEROPTERA: PANNOTA)!2 


T.-Q. Wang?, W. P. McCafferty3, Y. J. Bae4 


ABSTRACT: A consistent structural characteristic has been found to be unique to the mayfly fami- 
lies Neoephemeridae and Caenidae. It is termed a sutural ommation and is present on the adult 
mesonotum. The structure is described and illustrated. Its unique presence supports the hypothesis 
of a sister relationship of Neoephemeridae and Caenidae among the pannote mayflies, a cladistic 
arrangement that has been somewhat debatable in the past. The Baetiscidae and Prosopistomatidae, 
which previously have been hypothesized to be closely related to either Neoephemeridae or Caenidae 
by various authors, are considered to constitute an aberrant clade of mayflies with still dubious 
relationships within the Ephemeroptera. 


Neoephemerid mayflies, the “large squaregills” (McCafferty 1981), were 
considered among the burrowing mayflies in the first half of this century. For 
example, Traver (1935) considered them as one of the subfamilies of 
Ephemeridae (= families of Ephemeroidea), and Ulmer (1939) considered them 
in the family Potamanthidae. This association was based on the common pos- 
session of basally arched MP? and CuA veins in the forewings. Edmunds and 
Traver (1954) placed the neoephemerids with the Caenidae, the “small 
squaregills” (McCafferty 1981), in a separate superfamily Caenoidea. This 
association was based on larval morphology, in particular the similar gill struc- 
ture. Since that time, all workers, with the exception of Demoulin (1958), have 
grouped Neoephemeridae with Caenidae rather than Ephemeroidea. McCafferty 
and Edmunds (1979) placed the Caenoidea among other mayflies that possess 
the apomorphic characteristic of more or less fused developing wingpads and 
are known as the pannote mayflies. 

Given the general relationships of large groupings of mayfly families (e.g., 
see McCafferty 1991), it appears that adult similarities (wing venation) of 
neoophemerids and burrowing mayflies (Ephemeroidea) were present in their 
immediate common ancestor, and of the pannote mayflies, only retained in the 
neoephemerids. On the other hand, the larval similarities of neoephemerids and 
other pannotes, including caenids, (e.g., fused wingpads) appear to be derived 
in these groups. Edmunds (1965), McCafferty (1972), and McCafferty and 
Edmunds (1976) have discussed how differential rates of evolution in the larval 
and adult stages can lead to such disparate stage characterization in certain 


1 Received May 6, 1996. Accepted August 9, 1996. 
2 Purdue Agriculture Research Program Journal Number 15040. 
3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 


4 Department of Biology, Seoul Woman’s University, 126 Kongung 2-dong, Nowon-gu, Seoul 139- 
744, Korea. 


ENT. NEWS 108(1) 52-56, January & February, 1997 


Vol. 108, No. 1, January & February, 1997 53 


Ephemeroptera taxa (i.e., with relatively ancestral characterization in one stage 
and relatively derived characterization in another stage). 

Among the pannote mayflies, three major historical schemes of familial 
relationships are noteworthy. McCafferty and Edmunds (1979) recognized Neo- 
ephemeridae as a phyletic sister group of Caenidae. Landa and Soldan (1985) 
recognized Neoephemeridae as a sister group of Baetiscidae, with the two 
derived with another lineage consisting of Caenidae and Prosopistomatidae. 
McCafferty (1991) followed Landa and Soldan (1985) in recognizing 
Neoephemeridae as a sister group of Baetiscidae, but derived this lineage from 
near the base of the Pannota. He also included Prosopistomatidae and Caenidae 
as sister groups (see also Tshernova 1970). None of the previous arguments 
supporting these various schemes have been compelling. 

The Landa and Soldan (1985) scheme was based on data from internal 
anatomy, especially the ureterlike (their term) characteristics of Malpighian 
tubules in the case of Neoephemeridae and Baetiscidae, and the arrangement of 
the alimentary canal in the case of Prosopistomatidae and Caenidae. Although 
these internal anatomical data provided valuable descriptions for certain pannote 
mayflies, any interpretation of them as synapomorphic is suspect because of 
not only the very small number of representatives sampled from families (Landa 
1969) but also the current inability to substantiate cladistic polarity with respect 
to them. 

McCafferty (1991) suggested that the operculate gills of abdominal seg- 
ment 2 in Neoephemeridae and Caenidae were fundamentally different. How- 
ever, based on our recent comprehensive morphological studies, these gills are 
indeed essentially the same with respect to shape, ridge development and arma- 
ture. McCafferty (1991) also stated that the carapacelike development of the 
larval thorax was not similar in Baetiscidae and Prosopistomatidae. Nonethe- 
less, among Ephemeroptera, the carapace is found only in the Baetiscidae and 
Prosopistomatidae. We have additional data (Wang and McCafferty unpublished) 
showing that underlying gill morphology and certain mouthpart structures are 
similar and unique in these two families. These data strongly suggest that the 
carapace was commonly derived in the Baetiscidae and Prosopistomatidae. Kluge 
et al. (1995), without stating any reason, removed these latter two families 
from the Pannota and considered them sister families in a separate suborder of 
Ephemeroptera. 

Although we have not been able to find any larval characteristics common 
to both Neoephemeridae and Caenidae that are, without a doubt, apomorphic 
and unique (not subject to homoplasy), we have recently discovered a stable 
adult characteristic that strongly supports the hypothesis of a sister relationship 
of the two families. We present this characterization herein. 


54 ENTOMOLOGICAL NEWS 


NEW CHARACTERIZATION AND DISCUSSION 


A small, elongate eye-shaped membrane located medially on the mesonotum 
of adult caenids was noted and illustrated by Provonsha (1990). This appears as 
a short, primordial split along the medial suture, comprised of a clear mem- 
brane (Figs. 1, 3). We have found that this structure also occurs in adults through- 
out the Neoephemeridae (Figs. 2, 4-10), but in no other mayflies (obviously all 
possible outgroups) that we know of. It differs somewhat in the Neoephemeridae 
in terms of how membranous the structure is; sometimes it is not membranous. 
We call this structure the sutural ommation because of its eye shape. We 
hypothesize that it evolved in the immediate common ancestor of the 
Neoephemeridae and Caenidae: its uniqueness among all mayflies is strongly 
suggestive of a sister relationship of the two families. Since we cannot surmise 
a likely function of this ommation, it is highly possible that it is non-adaptive. If 
that is the case, it may be an excellent stable character for not only defining the 
Neoephemeridae + Caenidae clade but diagnosing the Edmunds and Traver 
(1954) and McCafferty and Edmunds (1979) concept of the superfamily 


Nt 


3 4 5 6 7 


Figs. 1-10. Sutural ommation of the adult mesonotum. 1. Caennis latipennis. 2. Neoephemera youngi. 
3. Brachycercus nasutus. 4-5. Potamanthellus amabilis. 6-7. P. chinensis. 8-9. N. purpurea. 10. P. 
compressa. 


Vol. 108, No. 1, January & February, 1997 55 


Caenoidea in the adult stage. 

The relative phylogenetic position of the hypothesized Neoephemeridae + 
Caenidae clade among the Pannota is still not clear. None of the previously 
published proposals are convincing because of the use of characteristics with 
unsubstantiated polarity. In any case, we are now convinced that the two fami- 
lies form a distinct clade within the Pannota. 

Baetiscidae and Prosopistomatidae, which have been variously associated 
with Neoephemeridae or Caenidae previously, appear to be sister lineages that 
form a separate clade based on the carapace and certain mouthpart and gill 
characteristics as mentioned above (Wang and McCafferty ms). The clade is 
aberrant, and its relationship to the Pannota is unclear at this time. For example, 
if the shared trait of the A; vein of the forewings ending in the outer margin (see 
Fontaine 1958) is plesiomorphic in Prosopistomatidae + Baetiscidae, then all 
other mayflies constitute a separate grouping because they would share the apo- 
morphic modified forewings with a reduced anal area and shortened A. This 
would also indicate that the carapace of Prosopistomatidae + Baetiscidae is not 
derived in common with the fused wingpads of the Pannota, including the Ephe- 
merellidae, Leptohyphidae, Tricorythidae, Neoephemeridae, and Caenidae. 
Obviously, more cladistic research is needed before these further questions can 
be completely resolved. 


LITERATURE CITED 


Demoulin, G. 1958. Nouveau schema de classification des Archodonates et des Ephéméroptéres. 
Bull. Inst. Roy. Sci. Nat. Belg. 34: 1-19. 

Edmunds, G. F., Jr. 1965. The classification of Ephemeroptera in relation to the evolutionary 
grade of nyrmphal and adult stages. Proc. Internat. Congr. Entomol., London 1964: 112. 

Edmunds, G. F., Jr. and J. R. Traver. 1954. An outline of reclassification of the Ephemeroptera. 
Proc. Entomol. Soc. Wash. 56: 236-240. 

Fontaine, J. 1958. Les affinités systématiques du genre Prosopistoma Latreille (Ephémeéroptéres). 
Deuxiéme note: le genré Baetisca Walsh: les formes ailees. Bull. Men. Soc. Linn. Lyon 27: 
133-140. 

Kluge, N., D. Studemann, P. Landolt, and T. Gosner. 1995. A reclassification of Siphlonuroidea 
(Ephemeroptera). Bull. Soc. Entomol. Suisse 68: 103-132. 

Landa, V. 1969. Comparative anatomy of mayfly larvae (Ephemeroptera). Acta Entomol. Bohem. 
60: 289-316. 

Landa, V. and T. Soldan. 1985. Phylogeny and higher classification of the order Ephemeroptera: a 
discussion from the comparative anatomical point of view. Academia, Prague. 

McCafferty, W. P. 1972. Pentageniidae: a new family of Ephemeroidea (Ephemeroptera). J. Geor- 
gia Entomol. Soc. 7: 51-56. 

McCafferty, W. P. 1981. Aquatic entomology. Jones and Bartlett, Boston. 

McCafferty, W. P. 1991. Toward a phylogenetic classification of the Ephemeroptera (Insecta): a 
commentary on systematics. Ann. Entomol. Soc. Am. 84: 343-360. 

McCafferty, W. P. and G. F. Edmunds, Jr. 1976. Redefinition of the family Palingeniidae and its 
implications for the higher classification of Ephemeroptera. Ann. Entomol. Soc. Am. 69: 486- 
490. 


56 ENTOMOLOGICAL NEWS 


McCafferty, W. P. and G. F. Edmunds, Jr. 1979. The higher classification of the Ephemeroptera 
and its evolutionary basis. Ann. Entomol. Soc. Am. 75: 5-12. 

McCafferty, W. P. and T.-Q. Wang. 1995. A new genus and species of Tricorythidae 
(Ephemeroptera: Pannota) from Madagascar. Ann. Limnol. 31: 179-183. 

Provonsha, A. V. 1990. A revision of the genus Caenis in North America (Ephemeroptera: Caenidae). 
Trans. Am. Entomol. Soc. 116: 801-884. 

Traver, J. R. 1935. Part II. Systematic, Jn: The biology of mayflies with a systematic account of 
North American species (J. G. Needham, J. R. Traver, and Y-C. Hsu, eds.), pp. 239-739. Comstock 
Publ. Co., Ithaca, New York. 

Tshernova, O. A. 1970. On the classification of fossil and recent Ephemeroptera. Entomol. Obozr. 
49: 124-145. (in Russian). 

Ulmer, G. 1939. Eintagsfliegen (Ephemeropteren) von den Sunda-Inseln, Arch. Hydrobiol., Suppl. 


16: 442-692. 


ANNOUNCEMENT 


The Humboldt (formerly Eagle Hill) Field Research Institute is again offering a wide variety 
of natural history seminars and workshops, June-September, 1997. Three topics of particular inter- 
est to entomologists are: Aquatic Entomology, Aug. 10-16; Intro. to Forest Insect Biodiversity, 
Aug. 10-16; and Intro. to Biology of Spiders, Aug. 17-23. Further information and a detailed bro- 
chure is available from the Humboldt Field Research Inst., PO Box 9, Steuben, ME 04680-0009; 
207-546-2821; FAX 3042; Email humboldt@nemaine.com 
http://maine.maine.edu/~eaglhill 


Vol. 108, No. 1, January & February, 1997 57 


A SOUTH DAKOTA RECORD FOR CHAULIODES 
RASTRICORNIS (MEGALOPTERA: CORYDALIDAE)! 


P.J. Johnson, K.D. Roush, X. Lin2 


ABSTRACT: Chauliodes rastricornis is reported from Bennett County, South Dakota. This is the 
first confirmation of this species in the state. The collection locality is the most westward locality in 
the northern Great Plains region and for the species. This species may be introduced to the region. 


Corydalidae, or fishflies and dobsonflies, have not been well surveyed in 
the northern Great Plains region. Parfin (1956) provided a checklist and some 
ecological notes for the species of Minnesota. Tarter et al. (1976) presented a 
distributional synopsis of selected megalopterans, including Chauliodes spe- 
cies in the region. Apparently, the only published South Dakota record was 
provided by Tarter et al. (1976) who reported C. rastricornis Rambur from the 
state, without further information. These authors also presented records for 
Nebraska, from Cherry and Antelope counties, in the northcentral and north- 
eastern portions of the state. Smith (1925) reported this species from Douglas 
County, Kansas, but this record is in the east-central portion of the state and 
appears continuous in distribution through neighboring states to the east, and 
eastern Oklahoma and Texas (Hazard 1960; Tarter et al. 1976). 

The larvae of Chauliodes spp. can readily be distinguished from those of 
other genera of megalopterans by the presence of elongate respiratory tubules 
on abdominal segment 8, these extending beyond the anal prolegs (Cuyler 1958). 
Larvae of the two species in this genus, C. rastricornis and C. pectinicornis 
(Linnaeus), can be differentiated by the color of the ecdysial cleavage line on 
the abdomen. Chauliodes rastricornis has a black cleavage line and C. pectini- 
cornis has a yellow cleavage line (Cuyler 1958; Hazard 1960). 

We collected four C. rastricornis larvae from the outflow area of an anthro- 
pogenic sportfishing pond in southwestern South Dakota, approximately 8 air 
miles south-southeast of Martin. Specifically, these larvae are labelled as fol- 
lows: South Dakota, Bennett County, Lacreek National Wildlife Refuge, trout 
pond #2 outflow, T36N R37W sec. 25 NW1/4, 24.ix.1995, ca. Long. 10138W, 
Lat. 4304N, P.J. Johnson, X. Lin & K.D. Roush collrs. The recovery of these 
specimens confirms the presence of C. rastricornis in South Dakota and pro- 
vides the first specific record of occurrence in the state. 

Available distributional information for C. rastricornis from Parfin (1952), 
Hazard (1960), Tarter et al. (1976), and A. Contras-Ramos, University of Min- 


! Received April 10, 1996. Accepted May 4, 1996. 


2 Address for all authors: Insect Research Collection, Box 2207A, South Dakota State University, 
Brookings, SD 57007, U.S.A. 


ENT. NEWS 108(1) 57-59, January & February, 1997 


58 ENTOMOLOGICAL NEWS 


nesota (pers. comm., 1995) indicates that the previous confirmed westernmost 
records of this species in the northern Great Plains region were from Cherry 
and Antelope counties in the northcentral and northeastern portions of Nebraska. 

Otherwise, the nearest recorded sites are in Winona County, Minnesota, in the 
~ southeastern corner of the state, and Story County in central Iowa. Thus, the 
discovery of this species from southwestern South Dakota provides a further 
westward range extension. 

All specimens reported herein were collected by sweeping the waters, sedi- 
ments and vegetation at shoreline and near-shore areas < 0.3 m in water depth 
with a triangular dipnet. The pond is incompletely shaded by only scattered 
sapling peachleaf willow (Salix amygdaloides). Cattail (Typha angustifolia) and 
waterweed (Elodea canadensis) dominated the aquatic flora and vegetation, 
with only a few arrowhead (Sagittaria latifolia), sedge (Carex comosa) and 
burreed (Sparganium eurycarpum); plant names follow Larson (1993). The 
sample site was 2-4 m from the entrance to a 0.5 m diameter outflow culvert but 
possessed a slow flow rate due to a dense cattail stand before the culvert. Surfi- 
cial bottom sediments included a deep layer of unconsolidated fine organic par- 
ticulates overlain with macrophyte leaves and stems in various stages of decay. 
No larvae were recovered in flowing water immediately below the culvert. This 
site agrees with generalized habitats for C. rastricornis (Neunzig and Baker, 
19915). 

Records of C. rastricornis from Nebraska, as cited by Tarter et al. (1976) 
and noted above, are from counties included in the Nebraska Sandhills physi- 
ographic region. The new South Dakota site is literally at the northern margin 
of the Sandhills. Together, these records may suggest a natural distribution 
through the Sandhills and that this distribution is part of the greater range of the 
species through adjacent states to the east and south. The disjunct pattern of 
occurrence could reflect land and water use changes, or local extinction of popu- 
lations during the last century and prior to adequate faunal sampling. Chauli- 
odes rastricornis is typically a species of lentic waters (Neunzig and Baker, 
1991), though slow stream waters are also acceptable (Parfin 1952). Appropri- 
ate lentic habitats in most of South Dakota and Nebraska were few prior to 
extensive impoundment and pond building since the late 1800’s. This suggests 
that a natural distribution of this species to the Lacreek Refuge through south- 
ern South Dakota would require this species to have occupied portions of rivers 
that drain the refuge and vicinity, including the Missouri River, White River, 
and Little White River. Similarly, distribution through the Nebraska Sandhills 
would have included the Niobrara, Elkhorn, and Loup river systems. Available 
distribution and collection information does not include any of these rivers as 
habitats for C. rastricornis. 

Anthropogenic environmental changes may be implicated in the apparent 
range expansion of C. rastricornis. The pond from which the South Dakota 


Vol. 108, No. 1, January & February, 1997 59 


specimens were collected is regularly stocked with rainbow trout (Oncorhynchus 
mykiss (Walbaum) derived from a state fish hatchery located near Rapid City, 
South Dakota; but no Chauliodes species is known from Rapid City or the Black 
Hills and vicinity. 

Pond building and stocking since the late 1800’s for support of livestock, 
wildlife, food fish, and sportfishing brought the introduction of non-native fishes 
and attendant food resource organisms. From 1899 through 1901 many species 
of food and game fish were rail transported in bulk and introduced into lentic 
and lotic waters throughout the northern Great Plains (R. Smith, U.S. Fish & 
Wildlife Service, Spearfish, pers. comm. 1996). Many of these game fishes de- 
rived from source areas in more eastern portions of the United States and within 
the natural range of C. rastricornis. Additionally, intentional and unintentional 
introductions of both fish and aquatic insects continue to occur in the region. 
For example, in October 1995, two larvae of Corydalus cornutus (Linnaeus) 
were received for identification from a pet store in Sioux Falls, South Dakota. 
These specimens were included with tropical fish reared and shipped from 
Florida, an apparently frequent event in the hobbyist fish trade. 

Competing hypotheses suggesting either nativeness or introduction of C. 
rastricornis cannot be fully reconciled at this time. However, a cursory exami- 
nation of the history of fisheries development in the northern Great Plains sug- 
gests that introduction of C. rastricornis is a strong possibility. 

Specimens reported herein are deposited in the Insect Research Collection, 
South Dakota State University. This is report #2915 of the South Dakota Agri- 
cultural Experiment Station. 


LITERATURE CITED 


Cuyler., R.D. 1958. The larvae of Chauliodes Latreille (Megaloptera: Corydalidae). Ann. Entomol. 
Soc. Amer., 51: 582-586. 

Hazard, E.I. 1960. A revision of the genera Chauliodes and Nigronia. Unpubl. M.S. thesis, Ohio 
State University, Columbus. 

Larson, G.E. 1993. Aquatic and wetland vascular plants of the northern Great Plains. U.S. Dept. 
Agric., Forest Service, General Technical Report RM-238. 

Neunzig, H.H. and J.R. Baker. 1991. Order Megaloptera. /n F.W. Stehr, Immature Insects, Vol. 2. 
Kendall/Hunt Publishing Co., Dubuque. 

Parfin, S.I. 1952. The Megaloptera and Neuroptera of Minnesota. Amer. Midl. Nat., 47(2): 421- 
434. 

Smith, R.C. 1925. The Neuroptera and Mecoptera of Kansas. Bull. Brook. Entomol. Soc., 20: 165- 
7 

Tarter, D.C., W.D. Watkins, M.L. Little, and J.T. Goodwin. 1976. New state records of fishflies 
(Megaloptera: Corydalidae). Entomol. News, 87(7-8): 223-228. 


60 ENTOMOLOGICAL NEWS 


TWO NOTEWORTHY COLLECTIONS OF TICKS 
(ACARI: IXODIDA: IXODIDAE) FROM 
ENDANGERED CARNIVORES IN THE LAO 
PEOPLE’S DEMOCRATIC REPUBLIC! 


Richard G. Robbins2, William B. Karesh3, Susan Rosenberg?, 
Nancy Schonwalter4, Chanthaviphone Inthavong> 


ABSTRACT: Adults of the ixodid ticks Haemaphysalis asiatica and Rhipicephalus 
haemaphysaloides and nymphs of an undetermined species of Amblyomma are reported from a 
Laotian specimen of the Asian golden cat, Catopuma temminckii, and an adult of Ixodes ovatus is 
reported from a Laotian specimen of Owston’s palm civet, Chrotogale owstoni. These are the first 
published records of any ticks from either carnivore and the first record of /. ovatus from the Lao 
People’s Democratic Republic. 


The escalating human assault on the world’s biomes will, in the near future, 
obscure or obliterate host-parasite relationships that have taken eons to evolve 
(Windsor 1990, Rozsa 1992, Durden and Keirans 1996). In acarology, major 
patterns of tick-host coevolution are now relatively clear (Hoogstraal and 
Aeschlimann 1982, Hoogstraal and Kim 1985), but a remarkable number of 
terrestrial vertebrates have seldom or never been examined for these interesting 
and biomedically important arthropods. In January 1996, one of us (WBK) was 
summoned to the village of Lak Sao (18.10N, 104.55E), Khammouane Prov- 
ince, on the eastern border of the Lao People’s Democratic Republic, to provide 
care for recently captured specimens of two endangered carnivores: a young 
adult female Asian golden cat, Catopuma temminckii (Vigors and Horsfield, 
1827) (Felidae), and an adult male Owston’s palm civet, Chrotogale owstoni 
Thomas, 1912 (Viverridae). Both animals were also examined for ticks, and 
complete collections from each were immediately shipped to the senior author 
for identification and evaluation. 

The tick collection from Ca. temminckii comprises the following: Haema- 
physalis asiatica (Supino, 1897), 10 males; Rhipicephalus haemaphysaloides 
(Supino, 1897), 1 female; and Amblyomma sp., 3 nymphs; all collected 22 Janu- 


1 Received July 8, 1996. Accepted September 12, 1996. 


2 Armed Forces Pest Management Board, Walter Reed Army Medical Center, Washington, DC 
20307-5001. 


3 Field Veterinary Program, Wildlife Health Center, Wildlife Conservation Society, 185th Street 
and Southern Boulevard, Bronx, NY 10460-1099. 


4 Carnivore Preservation Trust. 1940 Hanks Chapel Road, Pittsboro, NC 27312. 


5 Director, Center for Protected Areas and Watershed Management, Ministry of Agro-forestry, 
Vientiane, Lao P.D.R. 


ENT. NEWS 108(1) 60-62, January & February, 1997 


Vol. 108, No. 1, January & February, 1997 61 


ary 1996. Haemaphysalis asiatica is a morphologically striking species that 
cannot be confused with any other member of its genus in Southeast Asia. 
Hoogstraal and Trapido (1966) redescribed all stages of this tick and summa- 
rized the literature on hosts (chiefly viverrids and felids) and distribution. 
Rhipicephalus haemaphysaloides occurs throughout the Oriental zoogeographic 
region, where it parasitizes a wide range of indigenous and introduced mam- 
mals, wild and domestic, as well as various birds (Anastos 1950, Tanskul et al. 
1983). Of the world’s approximately 103 species of Amblyomma, 16 occur in 
the Oriental zoogeographic region and at least 11 are found in continental South- 
east Asia (Toumanoff 1944, Tanskul et al. 1983, Keirans 1992, Petney and 
Keirans 1995). However, immatures of many species remain poorly known and 
are generally “identified” by association with adults. We have been unable to 
locate published records of these or any other ticks from Ca. temminckii, a cat 
that is declining over much of its range, is classified as endangered by the U.S. 
Department of the Interior, and is on appendix 1 (species threatened with ex- 
tinction that are or may be affected by trade) of the Convention on International 
Trade in Endangered Species of Wild Fauna and Flora (Nowak 1991). This tick 
collection constitutes accession AGC1, MEDARKS (Medical Archives) 96-0406, 
Field Veterinary Program, Wildlife Conservation Society, on long-term loan to 
RGR. 

The tick collection from Ch. owstoni consists of a single partly engorged 
female of Ixodes ovatus Neumann, 1899, collected 22 January 1996. Ixodes 
ovatus 1s the type of the monotypic subgenus Partipalpiger of Ixodes (Hoogstraal 
et al. 1973) and is both common and widespread in Asia, ranging from Japan to 
northeastern India and Nepal. In continental Southeast Asia, /. ovatus is known 
from Vietnam, Thailand, and Myanmar (formerly Burma) (Petney and Keirans 
1994). Adults of this distinctive species usually parasitize larger mammals, in- 
cluding humans, but there appear to be no published records from the Lao P.D.R. 
and, again, no records of this or other tick species from Ch. owstoni, an exceed- 
ingly rare civet that is under considerable pressure from hunters (Schreiber et 
al. 1989, Nowak 1991). Indeed, the Lak Sao civet was being sold as food in the 
village market and is the only known living representative of its species. This 
tick collection constitutes accession OPC], MEDARKS 96-0407, Field Veteri- 
nary Program, Wildlife Conservation Society, also on long-term loan to RGR. 

The Wildlife Conservation Society, founded in 1895 as the New York Zoo- 
logical Society, is arguably the world’s premier conservation research institu- 
tion, with more than 270 on-the-ground projects in 51 nations (Conway 1994). 
Since 1992, the senior author has been privileged to identify and provide eco- 
logical information for over 3,000 tick specimens shipped from Society research 
stations in Asia, Africa, and Latin America (e.g., Calle et al. 1994). We antici- 
pate publishing additional host and distribution records for exotic ticks as speci- 
mens come to hand. 


62 ENTOMOLOGICAL NEWS 


ACKNOWLEDGMENTS 


For their constructive comments on an earlier version of this work, we thank our col- 
leagues at the Armed Forces Pest Management Board: Colonel Phillip G. Lawyer, U. S. 
Army, and Captain Craig H. Forcum, U. S. Air Force. The opinions and assertions advanced 
herein are those of the authors and are not to be construed as official or reflecting the views 
of the U. S. Departments of the Army or Defense. 


LITERATURE CITED 


Anastos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Entomol. Am. (new series) 30:1- 
144. 

Calle, P. P., J. Rivas, M. Munoz, J. Thorbjarnarson, E. S. Dierenfeld, W. Holmstrom, W. E. 
Braselton, and W. B. Karesh. 1994. Health assessment of free-ranging anacondas 
(Eunectes murinus) in Venezuela. J. Zoo Wildl. Med. 25: 53-62. 

Conway, W. 1994. President’s letter. For the children of Brooklyn, for the children of Papua New 
Guinea. pp. 3-4 Jn: Annual report, Wildlife Conservation Society. Bronx, New York. 

Durden, L. A. and J. E. Keirans. 1996. Host-parasite coextinction and the plight of tick conserva- 
tion. Am. Entomol. 42: 87-91. 

Hoogstraal, H. and A. Aeschlimann. 1982. Tick-host specificity. Mitt. Schweiz. Entomol. Ges./ 
Bull. Soc. Entomol. Suisse 55: 5-32. 

Hoogstraal, H. and K. C. Kim. 1985. Tick and mammal coevolution, with emphasis on 
Haemaphysalis. pp. 505-568 In: K. C. Kim (ed.). Coevolution of parasitic arthropods and mam- 
mals. John Wiley & Sons, New York. 

Hoogstraal, H. and H. Trapido. 1966. Studies on Southeast Asian Haemaphysalis ticks (Ixodoidea, 
Ixodidae). Species described by Supino in 1897 from Burma, with special reference to H. 
(Rhipistoma) asiaticus (= H. dentipalpis Warburton and Nuttall). J. Parasitol. 52: 1172-1187. 

Hoogstraal, H., C. M. Clifford, Y. Saito, and J. E. Keirans. 1973. [Ixodes (Partipalpiger) ovatus 
Neumann, subgen. nov.: identity, hosts, ecology, and distribution (Ixodoidea: Ixodidae). J. Med. 
Entomol. 10: 157-164. 

Keirans, J. E. 1992. Systematics of the Ixodida (Argasidae, Ixodidae, Nuttalliellidae): an over- 
view and some problems. pp. 1-21 Jn: B. Fivaz, T. Petney, and I. Horak (eds.). Tick vector 
biology. Medical and veterinary aspects. Springer-Verlag, Berlin. 

Nowak, R. M. 1991. Walker’s mammals of the world. 5th ed. Johns Hopkins University Press, 
Baltimore. 

Petney, T. N. and J. E. Keirans. 1994. Ticks of the genus /xodes in South-east Asia. Trop. Biomed. 
11: 123-134. 

Petney, T. N. and J. E. Keirans. 1995. Ticks of the genera Amblyomma and Hyalomma from 
South-east Asia. Trop. Biomed. 12: 45-56. 

Rozsa, L. 1992. Points in question: endangered parasite species. Int. J. Parasitol. 22: 265-266. 

Schreiber, A., R. Wirth, M. Riffel, and H. Van Rompaey. 1989. Weasels, civets, mongooses, and 
their relatives. An action plan for the conservation of mustelids and viverrids. Union Conserv. 
Nat., Gland, Switzerland. 

Tanskul, P., H. E. Stark, and I. Inlao. 1983. A checklist of ticks of Thailand (Acari: Metastig- 
mata: Ixodoidea). J. Med. Entomol. 20: 330-341. 

Toumanoff, C. 1944. Les tiques (Ixodoidea) de |’ Indochine. Instituts Pasteur de I’ Indochine, S.I-L.I., 
Saigon. 

Windsor, D. A. 1990. Heavenly hosts. Nature (London) 348: 104. 


Vol. 108, No. 1, January & February, 1997 63 


NEW HOST RECORD FOR OOENCYRTUS 
KUVANAE (HYMENOPTERA: ENCYRTIDAE)! 


Richard W. Hofstetter, Kenneth F. Raffa2 


ABSTRACT:. Eggs of the pine tussock moth, Dasychira pinicola (Lepidoptera: Lymantriidae), 
were successfully parasitized in the laboratory by the gypsy moth egg parasitoid Ooencyrtus kuvanae 
(Hymenoptera: Encyrtidae). This is the first report of O. kuvanae parasitizing this species. The pine 
tussock moth is a pest species in Wisconsin and may be considered as an alternate host for O. 
kuvanae. 


Ooencyrtus kuvanae Howard (Hymenoptera: Encyrtidae) was introduced 
into New England from Japan for biological control of the gypsy moth, Lymantria 
dispar (L.) (Lepidoptera.: Lymantriidae) (Crossman 1917). It successfully es- 
tablished, and has since spread west with the expanding gypsy moth popula- 
tion, including Wisconsin. O. kuvanae has relatively few known hosts around 
the world outside of the gypsy moth. It has been reared from field collected 
Dendrolimus spectabilis Butler (Lepidoptera: Lasiocampidae), Eriogyna 
pyretorum Westwood (Lepidoptera: Saturniidae), Euproctis chrysorrhoea L. 
(Lepidoptera: Lymantriidae) eggs and two gypsy moth parasitoids, Cotesia 
melanoscela Ratzeburg (Hymenoptera: Braconidae) and Anastatus disparis 
Ruschka (Hymenoptera: Eupelmidae), and has been reared in the laboratory on 
8 other species of Lepidoptera: 3 Lymantriidae, 3 Saturniidae, 2 Lasiocampidae 
(Brown 1984, Crossman 1925, Ni etal. 1994, Tadi¢ & Binéev 1959). O. kuvanae 
is multivoltine and may utilize alternative hosts throughout the year. 

The pine tussock moth, Dasychira pinicola Dyar (Lepidoptera: Lyman- 
triidae), is one of the most damaging defoliators of jack pine, Pinus banksiana 
Lamb, in northern Wisconsin (Sreenivasam et al. 1972). D. pinicola can also 
feed on red pine, eastern white pine, spruce, and fir. D. pinicola overwinter as 
early instar larvae, resume feeding in the spring, pupate in mid summer and 
emerge as adults in late July and August. Eggs are present throughout August. 

The objective of this study was to determine whether O. kuvanae could 
successfully parasitize pine tussock moth eggs. 


MATERIALS AND METHODS 


The laboratory culture of Ooencyrtus kuvanae originated as a field collec- 


l Received May 28,1996. Accepted June 29, 1996. 
2 University of Wisconsin-Madison, Department of Entomology, Madison WI 53706. 


ENT. NEWS 108(1) 63-65, January & February, 1997 


64 ENTOMOLOGICAL NEWS 


tion on gypsy moth (Lymantria dispar L.) egg masses from Virginia. Parasi- 
toids were collected by S. Barth and R. Cameron, Beneficial Insects Introduc- 
tion Research Lab, USDA ARS, Newark, DE, in the spring of 1993. It has been 
cultured since that time in the laboratory on L. dispar egg masses provided by 
USDA APHIS, Otis Air National Guard Base, Cape Cod, Massachusetts. Rear- 
ing conditions were 22"C, 60% - 70% RH, and 14L:1OD. Experiments began 
in August of 1994 after 12 to 14 generations on gypsy moth egg masses in the 
laboratory. 

Voucher specimens of O. kuvanae have been deposited in the University of 
Wisconsin, Madison Insect Research Collection. 

Egg masses of the pine tussock moth, Dasychira pinicola were field col- 
lected in Douglas Co., WI in late July (1994) by Shane Weber, Wisconsin De- 
partment of Natural Resources. Individual females were placed on single 2 week 
old intact egg clusters, and allowed to oviposit for four days. The number and 
sex of emerging offspring were recorded. Also the number of emergence holes 
were recorded for each egg mass. Newly emerged offspring were mated and 
placed on L. dispar egg masses and allowed to oviposit. 


RESULTS AND DISCUSSION 


Female O. kuvanne successfully parasitized pine tussock moth egg masses 
(Table 1). The mean number of O. kuvanae offspring was 7.0 (4.8 std) per egg 
mass and 10.7 (0.58 std) per female. In comparison, parasitoids placed on gypsy 
moth egg masses under these conditions produce an average of 30 (14 std) 
offspring per egg mass and per female (Hofstetter & Raffa 1996). O. kuvanae 
progeny emerged from three of the four D. pinicola egg masses tested. In two 
egg masses, there were more offspring than number of holes, suggesting that 
several parasitoids emerged from one or more individual eggs. Of the egg masses 
that were parasitized, 35.2% (12.0 std) of the eggs per mass were successfully 
parasitized. 

Female offspring successfully mated and parasitized gypsy moth egg masses. 
Progeny were observed to emerge from the gypsy moth egg masses. 

The pine tussock moth is in the same family as the gypsy moth, so it is not 
surprising that O. kuvanae can successfully parasitize these eggs. The eggs are 
spherical, whitish, and approximately Imm in diameter. They are deposited in 
loose clusters, one to two layers thick, mostly on pine needles near midcrown 
but also on the trunk, dead twigs, and similar vegetation. 

The pine tussock moth could potentially serve as an alternate host for O. 
kuvanae when primary hosts are scarce. In addition, O. kuvanae could poten- 
tially add to the biological control of D. pinicola. 


Vol. 108, No. 1, January & February, 1997 65 


Table 1. Number of Ooencyrtus kuvanae progeny and percent male offspring when placed on 
Dasychira pinicola egg masses. 


No. of eggs No. emergence Parasitoid % Male 
Mass # per egg mass holes offspring Offspring 
1 47 11 11 36.4 
2 19 9 11 36.4 
3 21 0 0 a 
4 23 8 10 30.0 
ACKNOWLEDGMENTS 


We thank personnel of the USDA ARS, Beneficial Insects Introduction Laboratory, par- 
ticularly R. Fuester, S. Barth, and R. Cameron, for collecting and shipping O. kuvanae, and 
Shane Weber, Wisconsin DNR, for collecting and shipping pine tussock moth egg masses. 
Funding was provided by the Wisconsin Department of Natural Resources, The UW-Madi- 
son Graduate School, McIntire Stennis, and UW-Madison College of Agricultural and Life 
Sciences. 

LITERATURE CITED 


Brown, M.W. 1984. Literature review of Ooencyrtus kuvanae (Hymenoptera: Encyrtidae), an egg 
parasite of Lymantria dispar (L.) (Lepidoptera: Lymantriidae). Entomophaga 29:249-265. 
Crossman, S.S. 1917. Some methods of colonizing imported parasites and determining their 
increase and spread. J. Econ. Entomol. 10:177-193. 

Crossman, S.S. 1925. Two imported egg parasites of the gypsy moth, Anastatus bifasciatus Fonsc. 
and Schedius kuvanae Howard. J. Agric. Res. 30:643-675. 

Hofstetter, R.W. & K.F. Raffa. 1996. Endogenous and exogenous factors affecting parasitism by 
Ooencyrtus kuvanae (Hymenoptera: Encyrtidae). Entomol. Exp. Appl. (submitted). 

Ni, L.X., X.W. Tong, X.M. Lao. 1994. Influence of multiparasitism of egg parasitoids of pine 
lasiocampids on efficacy of biological control. Acta Entomologica Sinica 37:145-152. 

Sreenivasam, D.D., D.M. Benjamin & D.D. Walgenback. 1972. The bionomics of the pine tus- 
sock moth. University of Wisconsin School of Natural Resources. Res. Bull. 282. 

Tadié, M. and B. Binéev. 1959. Ovencyrtus kuvanae How. in Yugoslavia. Zastita bilja 10:52-53. 


66 ENTOMOLOGICAL NEWS 


SCIENTIFIC NOTE 


FIRST HOST RECORD FOR THE GENUS LARISSIMUS 
(HYMENOPTERA: BRACONIDAE)! 


Angélica Maria Penteado-Dias2 


Larissimus cassander was originally described based on material collected by Fritz 
Plaumann in Seara (Nova Teutonia), Santa Catarina, Brazil. 

The only described species in the genus, it is one of the larger Microgastrinae, with 
strong coloration similar to some Braconinae (Nixon, 1965) (e.g. yellow or orange and black 
with banded wings). At the time of its description, the species was classified in the tribe 
Microgastrini. Mason (1981) proposed a reclassification to Microgastrinae and considered 
Larissimus to belong to the Cotesiini.Besides the detailed description of the female, Mason 
(1981) provided a figure of its hind wing. At that time hosts and larvae were unknown. 

In August, 1989, one female was reared from larvae of Bertholdia sp. (Lepidoptera: 
Arctiidae) collected in a forested area of the Canchim farm (Embrapa, Sao Carlos, SP, Bra- 
zil) on Croton floribundus Spreng (Euphorbiaceae). This specimen has been deposited in the 
taxonomic collection of the Departamento de Ecologia e Biologia Evolutiva da Universidade 
Federal de Sao Carlos (DCBU) . Croton floribundus is a very common plant in the secondary 
forest areas in Rio de Janeiro, Minas Gerais, Sao Paulo and Parana states and several other 
species of Lepidoptera larvae are found on it. 


ACKNOWLEDGMENTS 


I gratefully acknowledge the help of Manoel Martins Dias Filho (UFSCar) by providing 
the name of the lepidopteran host. This work was supported by Conselho Nacional de 
Desenvolvimento Cientifico e Tecnol6gico (CNPq) from Brazil. 


LITERATURE CITED 


Mason, W.R.M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae): 
A phylogeny and reclassification of Microgastrinae. Mem. Ent. Soc. Canada no. 115. 147 pp. 

Nixon, G.E.J. 1965. A reclassification of the Tribe Microgastrini (Hymenoptera: Braconidae). Bull. 
Brit. Mus. (Natural History) Entomol. Suppl. 284pp. 


1 Received June 15, 1996. Accepted July 5, 1996. 


2 Departamento de Ecologia e Biologia Evolutiva, Universidade Federal de Sao Carlos. CP 676. 
CEP 13 565-905. Sao Carlos, SP, Brasil. 


Vol. 108, No. 1, January & February, 1997 67 


BARK BEETLE (COLEOPTERA: SCOLYTIDAE) 
OUTBREAK IN PINE FORESTS OF THE SIERRA DE 
LAS MINAS BIOSPHERE RESERVE, GUATEMALA! 


Robert A. Haack2, Gerardo Paiz-Schwartz> 


ABSTRACT: From 1992 to 1995 an outbreak of pine-infesting bark beetles occurred in eastern 
Guatemala, primarily along the southern slopes of the Sierra de las Minas Mountains, between 500 
and 2100 m elevation, in the Departments of El Progreso and Zacapa. Based on examination of 
bark beetle-killed pine (Pinus) trees at several sites, Dendroctonus frontalis was the principal mor- 
tality agent and Pinus oocarpa was the most affected tree species. Egg galleries similar to those of 
D. frontalis were also found on dead Pinus maximinoi and Pinus tecunumanii trees. Infested pine 
stands were typically unmanaged, overstocked, and located on steep slopes. In addition, most out- 
break areas had experienced frequent ground fires and overgrazing. The ultimate stressing agent 
that apparently triggered this outbreak was a severe regional drought in 1992. Besides D. frontalis, 
other scolytids collected along the lower trunk of P. oocarpa trees were Dendroctonus parallelocollis, 
Ips calligraphus, Ips grandicollis, Pityophthorus confusus, and Xyleborus intrusus. At lower eleva- 
tions, Ips calligraphus was the only scolytid collected from the lower trunk of infested Pinus caribaea 
trees. 


Bark beetle (Coleoptera: Scolytidae) outbreaks in the pine (Pinus spp.) for- 
ests of Guatemala have been recorded since the late 1800s, with outbreaks this 
century occurring during the 1930s, 1950s, 1970s, and 1980s (Alvarado 1939, 
Schwerdtfeger 1955, Haack 1995). Most of these outbreaks occurred in the 
western highlands of Guatemala, primarily in the Departments of Huehuetenango 
and Totonicapan, but during the outbreaks of the 1970s and 1980s, pines in 
several other western departments were affected. Although several species of 
bark beetles in the genera Dendroctonus and Ips were encountered during these 
past outbreaks, Dendroctonus adjunctus Blandford was recognized as the prin- 
cipal mortality agent (Becker 1955, Schedl 1955, Schwerdtfeger 1955, 1956, 
1959, 1961). During these outbreaks, Pinus rudis Endlicher was the most se- 
verely affected pine species, but also attacked were Pinus maximinoi H.E. Moore 
(= Pinus tenuifolia Bentham), Pinus montezumae Lambert, Pinus oocarpa 
Schiede, and Pinus pseudostrobus Lindley. In this paper, we use the pine no- 
menclature given in Perry (1991). Few outbreaks have been reported in pine 
forests of eastern Guatemala, which consists largely of P. oocarpa and Pinus 
caribaea Morelet. However, several major outbreaks have occurred this cen- 


1 Received March 28, 1996. Accepted April 28, 1996. 


2 USDA Forest Service, North Central Forest Experiment Station, 1407 South Harrison Road, East 
Lansing, Michigan 48823, USA. 


3 Fundacién Defensores de la Naturaleza, Avenida Las Americas 20-21, Zone 14, Guatemala City, 
Guatemala. 


ENT. NEWS 108(1) 67-76, January & February, 1997 


68 ENTOMOLOGICAL NEWS 


tury in the P. oocarpa forests of neighboring Honduras; the most severe out- 
break took place during the 1960s, when about 20% of the pines in Honduras 
were killed, primarily by Dendroctonus frontalis Zimrnermann (Beal et al. 1964, 
Ketcham and Bennett 1964, FAO 1968, Wilkinson and Haack 1987). In both 


GUATEMALA 


| wih 


16° 
Lake Izabal 
Sierra de las Minas 
Biosphere Reserve 
ie 


CA) Lake Atitlan 


14° 
0 100 
nt et — jt ht ht} 
Kilometers 
g9° 


13° 
92° 91° 90° 


Fig. 1. Outline map of Guatemala, showing location of the Sierra de las Minas Biosphere Reserve 
(shaded area). Insert map shows outline of Mexico and Central American countries, with Guate- 
mala shaded. 


Vol. 108, No. 1, January & February, 1997 69 


SIERRA DE LAS MINAS 
BIOSPHERE RESERVE 


MOTAGUA RIVER 0 20 30 
KILOMETERS 
CORE ZONE RECOVERY ZONE 


MULTIPLE USE Mi BUFFER ZONE 
ZONE 


90° $9 45° 89 30° 8915 


Fig. 2. Outline map of the Sierra de las Minas Biosphere Reserve, indicating the location of the four 
management zones and the major bark beetle outbreak sites (squares, either black or white). 


Guatemala and Honduras, it appears that poor forest management, overmature 
stands, frequent fires, overgrazing, and drought were the major contributing 
factors in these outbreaks (Beal et al. 1964, Ketcham and Bennett 1964, 
Schwerdtfeger 1955, Wilkinson and Haack 1987). 

This paper deals with a bark beetle outbreak that occurred during 1992- 
1995 in the Sierra de las Minas Biosphere Reserve in eastern Guatemala (Figs. 
1-2). This site, recognized by UNESCO as a Biosphere Reserve in 1993, is 
administered by Fundacion Defensores de la Naturaleza, a non-governmental 
organization located in Guatemala. The senior author visited the outbreak area 
in August 1995 to help evaluate the situation and recommend management op- 
tions (Haack 1995). 

The Sierra de las Minas is a chain of mountains in eastern Guatemala that 
runs mostly east to west (Figs. 1 -2). This mountain range is about 150 km long, 
10-30 km wide, and varies in elevation between 150 m and 3015 m (Dix 1996). 
There are 63 rivers that drain the Sierra de las Minas, feeding into the Polochic 
River to the north and the Motagua River to the south (Fig. 2). Mean annual 
rainfall varies from 0.5 to 4.0 m. The area’s vegetation is diverse, including 
seven Holdridge life zones that range from dry forests to cloud forests (Holdridge 


70 ENTOMOLOGICAL NEWS 


et al. 1971, Dix 1996). There are 13 species of conifers in the Sierra de las 
Minas, representing 6 genera: Abies, Cupressus, Juniperus, Pinus, Podocarpus, 
and Taxus (Dix 1996). Of these 13 species, there are 6 Pinus species; P. oocarpa 
is found at elevations between 500 and 2750 m, P. maximinoi between 1100 and 
2400 m, P. tecunumanii Schw. above 1500 m, P. caribaea below 800 m, P. 
pseudostrobus above 1600 m, and P. ayacahuite Ehrenb. above 2000 m. 

The Sierra de las Minas Biosphere Reserve, 130 km long and 15-30 km 
wide, covers much of the Sierra de las Minas Mountains. The Reserve is 236,300 
ha in size and is divided into four management zones, 1.e., the core zone (105,700 
ha), recovery zone (4200 ha), multiple use zone (34,600 ha), and buffer zone 
(91,800 ha; Fig. 2). In the core zone of the Sierra de las Minas Biosphere Re- 
serve, no economical activities are allowed and the natural resources are pro- 
tected. In the recovery zone, restoration activities are conducted. In the multiple 
use zone, sustainable resource use is allowed, including the extraction of wood 
products. In the buffer zone, communities are established and agriculture is 
allowed. Typically, the buffer zone of a Biosphere Reserve supports forestry, 
low-intensity agriculture, recreation, and scattered small settlements (Gregg 
1991). 


METHODS 


Several outbreak areas were visited in late 1994 and early 1995 by the jun- 
ior author and various cooperators. During these visits, data were taken on el- 
evation, presence or absence of bark beetles, size of the affected area, and pine 
species affected. During August 1995, both authors visited nine outbreak sites 
throughout the affected area and recorded information on elevation, pine spe- 
cies affected, and the current status of the outbreak. When we encountered bark- 
beetle killed pines that had been cut or had fallen, we noted the within-tree 
distribution of the various groups of bark beetles along the trunk and branches. 
This determination was based on the distinctive egg galleries that are created by 
Dendroctonus, Ips, and other bark beetles (Wood | 982, Kirkendall 1 983, Cibrian 
etal. 1995). Occasionally, we found dead adults in the egg galleries of attacked 
trees; such adults were used to make positive associations between a particular 
bark beetle species and its egg gallery pattern. For several pines that were cur- 
rently under attack, the bark was removed from the lower trunk and several 
scolytid specimens were collected. A few currently infested pines were felled 
and inspected for bark beetles. Scolytid specimens were identified by Stephen 
L. Wood. In the present paper, we use the scolytid names as given in Wood 
(1982) and Wood and Bright (1992). All required permits for this study were 
obtained from CONAP (Cosejo Nacional de Areas Protegidas = Guatemalan 
National Council for Protected Areas). Voucher specimens were deposited in 
the museum of the Universidad del Valle, Biology Department, Guatemala City, 
Guatemala. 


Vol. 108, No. 1, January & February, 1997 71 


RESULTS AND DISCUSSION 


Host Trees and Associated Scolytids. Bark beetle outbreak sites were lo- 
cated along the southern slopes of the Sierra de las Minas Mountains in the 
Departments of El Progreso and Zacapa (Fig. 2). Most outbreak sites occurred 
within the buffer zone of the Biosphere Reserve or just outside of it (Fig. 2). 
The outbreak sites occurred at elevations between 500 and 2100 m, primarily in 
lower montane moist forests dominated by P. oocarpa. For 43 individual out- 
break sites from separate watersheds that were visited in late 1994 and early 
1995, the average outbreak area was 16 ha (range 1-95 ha) and the average 
elevation was 1200 m (range 500-2100 m). By August 1995, more than 50 
outbreak sites had been recorded in the Sierra de las Minas Mountains, cover- 
ing more than 900 ha. 

The vast majority of the bark beetle-killed pines in the Sierra de las Minas 
were P. oocarpa. The diameter at breast height (1.4 m) of most bark beetle- 
killed P oocarpa trees ranged from 10 to 40 cm. The second most affected pine 
species was P. maximinoi, and the third most often attacked was P. tecunumanit. 

Specimens of six scolytid species were collected from the lower trunk of 
currently infested P. oocarpa trees in August 1995, including Dendroctonus 
frontalis, Dendroctonus parallelocollis Chapuis, [ps calligraphus (Germar), Ips 
grandicollis (Eichhoff), Pityophthorus confusus Blandford, and Xyleborus 
intrusus Blandford. Species of Dendroctonus, Ips, and Pityophthorus are phloem 
feeding bark beetles, while X. intrusus is a xylem-infesting ambrosia beetle. 

In dead P. maximinoi and P. tecunumanii trees, from which the bark beetles 
had already exited, egg galleries similar to those of D. frontalis dominated along 
the entire trunk. It is possible that other species of Dendroctonus, such as D. 
mexicanus Hopkins or D. vitei Wood, could have made some of the galleries, 
especially at higher elevations within the outbreak area. Adult size and egg 
gallery pattern are similar among D. frontalis, D. mexicanus, and D. vitei (Vite 
et al. 1974, Renwick et al. 1975, Wood 1982, Lanier et al. 1988), and thus 
identification by egg galleries alone is difficult. In areas of Mexico where both 
D. frontalis and D. mexicanus occur, D. frontalis is found at elevations below 
2000 m, while D. mexicanus is found between 1800 and 2500 m (Lanier et al. 
1988). In Honduras, D. frontalis was collected from P. oocarpa trees between 
1400 and 1700m (Wilkinson and Haack 1987). Dendroctonus vitei was first 
recognized in 1974 (Wood 1974) and is so far known from Guatemala and 
southern Mexico (Wood and Bright 1992); it has been collected between 2000 
and 2500 m in Guatemala (Vite et al. 1974, 1975). Dendroctonus mexicanus 
has been reported from P. oocarpa, but D. vitei has not (Wood and Bright 1992). 

At the eastern edge of the Sierra de las Minas near Lake Izabal (Fig. 2), a 
few bark beetle-killed P. caribaea trees were found at elevations below 800 m. 
These trees were not part of the main outbreak area. Ips calligraphus was the 


72 ENTOMOLOGICAL NEWS 


only scolytid that we found on the lower trunk of currently infested P. caribaea 
trees. 

There are few reports of D. frontalis in Guatemala (Moser et al. 1974, Lanier 
et al. 1988, Wood and Bright 1992). According to Wood and Bright (1992), D. 
frontalis occurs from the southeastern United States to Honduras. Adults, 2.0 to 
3.2 mm long, attack several species of pine, including P. maximinoi, P. oocarpa, 
and P. tecunumanii in Mexico and Central America (Wood 1982, Cibrian et al. 
1995). In the genus Dendroctonus, females initiate attack, produce an aggrega- 
tion pheromone, and are monogynous (i.e., joined by a single male). In Hondu- 
ras, it is estimated that D. frontalis may complete as many as nine generations 
per year (Beal et al. 1964). When reading the earlier Dendroctonus literature it 
is important to note that Wood (1963) synonymized D. mexicanus under D. 
frontalis, but later he (Wood 1974) recognized them again as two distinct 
species. 

Dendroctonus parallelocollis ranges from Mexico to Honduras (Wood and 
Bright 1992, Cibrian et al. 1995). Adults, 4.9 to 6.3 mm long, have been re- 
corded from four species of pine, including P. oocarpa (Becker 1955, Schedl 
1955, Wood 1982, Cibrian et al 1995). Attack by this beetle occurs along the 
lower trunk and major roots of pine trees and is usually associated with D. 
frontalis and D. mexicanus (Wood 1982, Cibrian et al. 1995). Care must be 
used when reading the literature since Wood and Bright (1992) state that refer- 
ences to D. parallelocollis from 1909 to 1969 actually refer to Dendroctonus 
approximatus Dietz. 

Ips calligraphus occurs in eastern North America, the southern and south- 
western United States, Mexico south to Nicaragua, and the Caribbean islands 
of the Dominican Republic, Bahamas, and Jamaica (Lanier 1972, Yates 1972, 
Garraway 1986, Haack etal. 1989, Bunce and McLean 1990, Wood and Bright 
1992). Adults are 3.8 to 5.9 mm long and have six spines along their elytral 
declivity. [ps calligraphus attacks several species of pine, including P. caribaea, 
P. maximinoi, and P. oocarpa (Sched! 1955, Wood 1982, Cibrian et al. 1995). In 
the genus /ps, males initiate attack, produce an aggregation pheromone, and are 
polygynous (i.e., joined by two or more females). In Florida and the Dominican 
Republic, /. calligraphus can complete 8 to 12 generations per year (Haack 
1985, Haack et al. 1989). It is important to note that /ps interstitialis (Eichhoff) 
was synonymized under the name /. calligraphus by Wood (1982), and that 
most early papers on Guatemalan six-spined /ps mention only J. interstitialis 
(Becker 1955, Sched] 1955, Schwerdtfeger 1956). 

Ips grandicollis occurs in eastern North America; Mexico south to Nicara- 
gua; and various Caribbean islands (Wood and Bright 1992). Adults are usually 
2.9 to 4.6 mm long and have five spines along their elytral declivity. They at- 
tack several species of pine, including P. caribaea and P. oocarpa (Wood 1982, 
Cibrian et al. 1995), and their biology is very similar to that of I. calligraphus. 
Readers should note that early papers on Guatemalan five-spined Jps usually 


Vol. 108, No. 1, January & February, 1997 73 


mention only /. cribricollis (Eichhoff) (Becker 1955, Schedl 1955, Schwerdtfeger 
1956); however, I. cribricollis was placed in synonymy with /. grandicollis by 
Wood (1977). Although Lanier (1987) stated that these two Ips species were 
distinct and that both occur in Central America, Wood and Bright (1992) con- 
tinue to regard /. cribricollis as a synonym of I. grandicollis. - 

Pityophthorus confusus occurs from the southern United States to Nicara- 
gua (Wood and Bright 1992). Adults, 1.9 to 2.9 mm long, have been recorded 
on several species of pine, including P. oocarpa (Becker 1955, Sched] 1955, 
Bright 1981, Wood 1982, Wilkinson and Haack 1987). Like Dendroctonus and 
Ips bark beetles, P. confusus breeds in the phloem (inner bark). Males initiate 
attack and are polygynous, usually with 2 to 5 females joining each male. Gal- 
leries may extend in any direction. Pityophthorus confusus usually infests the 
trunk and branches of pine trees that are already under attack by species of 
Dendroctonus or Ips (Bright 1981, Wilkinson and Haack 1987). 

Xyleborus intrusus has a geographic range that extends from Canada to 
Honduras, including certain Caribbean islands (Bunce and McLean 1990, Wood 
and Bright 1992). Adults, 2.2 to 2.7 mm long, have been recorded from several 
species of pine, but no specific host records were given for Guatemala (Wood 
1982, Wood and Bright 1992). The occurrence of X. intrusus on P. oocarpa is 
apparently a new host record. Little is known about the biology of X. intrusus. 
In general, beetles in the genus Xyleborus construct their many branched galler- 
ies in the sapwood of the host tree. An ambrosial fungus is cultivated on the 
gallery walls, and the larvae feed on this fungus. Males are rare in the genus 
Xyleborus, and when they do occur, they are flightless. Xyleborus females typi- 
cally mate with their brothers before leaving the host tree. Wood (1982) reports 
that X. intrusus usually attacks the base of recently killed, standing pine trees. 

Bark Beetle Attack Sequence and Within-Tree Distribution. Based on 
our inspection of the egg and larval galleries in dozens of already dead P. oocarpa 
trees, D. frontalis was, the first bark beetle to attack, and was found from 
groundline into the lower crown branches. Jps and Pityophthorus galleries were 
found along the lower trunks of these same trees, but they were most common 
along the upper trunk and main branches. Galleries of D. parallelocollis were 
found only along the lower 2-3 meters of the trunk. These observations on within- 
tree distribution are similar to those made for bark beetles on P. oocarpa in 
Honduras (Wilkinson and Haack 1987). 

Possible Contributing Factors and Current Conditions. Pine forests in 
the Sierra de las Minas are subjected to many environmental stresses. Most 
outbreak areas consisted of mature, overstocked stands, on steep slopes with 
shallow soils. In addition, most affected stands had a history of frequent ground 
fires and overgrazing. Such conditions have been listed as contributing factors 
to pine bark beetle outbreaks in Guatemala, Honduras, and the Dominican 
Republic (Schwerdtfeger 1955, Beal et al. 1964, Ketcham and Bennett 1964, 


74 ENTOMOLOGICAL NEWS 


FAO 1968, Wilkinson and Haack 1987, Haack et al. 1989). Another factor that 
likely favored the current bark beetle outbreak in Guatemala was a severe drought 
throughout the Motagua River Valley in 1992. Outbreaks of pine bark beetles 
have often been reported during or soon after periods of severe drought (Mattson 
and Haack 1987), although the link between drought and outbreaks of D. frontalis 
in the United States is not clearcut (Turchin et al. 1991). 

In August 1995, very few pines were currently under attack and the out- 
break appeared to be collapsing. The vast majority of the bark beetle-killed 
pine trees had died during 1993, 1994, and the first half of 1995. The 1995 rainy 
season was among the wettest on record in the Motagua River Valley, and this 
likely improved tree resistance (Mattson and Haack 1987). In addition, in 
August 1995, natural enemies of bark beetle were very common under the bark 
of currently infested P. oocarpa trees, including larvae of predatory beetles 
(Cleridae, Histeridae, Tenebrionidae, Trogositidae), larvae of predatory flies 
(Dolichopodidae), and signs of parasitism by various Hymenoptera (exit holes 
and cocoons in larval bark beetle galleries). Pressure from these natural 
enemies likely aided in the collapse of the current D. frontalis outbreak, as has 
been suggested for D. frontalis outbreaks in the United States (Turchin et al. 
1991, Reeves et al. 1995). Lists of predators, parasitoids, and other associated 
arthropods of pine bark beetles have been previously reported for Guatemala 
(Becker 1955, Sched] 1955, Moser et al. 1974) and Honduras (Clark 1974). 


ACKNOWLEDGMENTS 


We thank the USDA Forest Service and Fundacion Defensores de la Naturaleza for finan- 
cial and logistical support; the US AID office in Guatemala for logistical support; Stephen 
Wood, Brigham Young University, for identifying the bark beetles; and Peter Lorio, USDA 
Forest Service, and Kenneth Raffa, University of Wisconsin, for reviewing an earlier version 
of this paper. 


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Scolytidae), a taxonomic monograph. Great Basin Nat. Mem. 6: 1-1359. 

Wood, S.L. and D.E. Bright. 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: 
Taxonomic index. Great Basin Nat. Mem. 13: 1-1553. 

Yates, H.O., III. 1972. Bark beetles attacking Caribbean pine in northeastern Nicaragua. FAO 
Plant Prot. Bull. 20: 25-27. 


SOCIETY MEETING OF OCTOBER 23, 1996 
ECOLOGICAL WANDERINGS IN THE ORIENT 


Paul W. Schaefer 
Beneficial Insects Introduction Research Laboratory, USDA, Newark, DE 


Through years (beginning in 1975) of having foreign exploration experiences in Japan, Korea, 
People’s Republic of China, Hong Kong and Taiwan, Dr. Schaefer presented a brief summary of the 
various projects that he has worked on in the Orient. These include principally the gypsy moth, 
Lymantria dispar and its close relatives, Lymantria spp. (Lepidoptera: Lymantriidae), but other 
explorations involved the Mexican bean beetle, Epilachna varivestis (Coleoptera: Coccinellidae), 
Japanese beetle, Popillia japonica (Coleoptera: Scarabaeidae), Larch casebearer, Coleophora 
laricella (Lepidoptera: Coleophoridae), beneficial coccinellids, including Harmonia axyridis (Co- 
leoptera: Coccinellidae) and the chestnut gall wasp, Dryocosmus kuriphilus (Hymenoptera: 
Cynipidae). Inasmuch as many of these projects have involved the recovery of very small, non- 
descript, generally uninteresting insect species to most people, Dr. Schaefer focused his camera 
lens on some other insects encountered incidental to his main research projects. This was often 
done through opportunities to observe his host Japanese, Chinese and Taiwanese scientists con- 
ducting their individual research projects, including such activities as black lighting and phero- 
mone trapping for moths, sampling various insect populations, and frass and/or biomass collec- 
tions. A series of color illustrations of insects was shown as a means of illustrating a small part of 
the diversity of Asian insect life. The magnificent diversity of Oriental species as well as the great 
variation in form, color and function was the underlying theme for the evening. 


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OL 


7 Gl 
§3/ 
ENT 
Sta.us Of some species of Enallagma (Odonata: Coen- 
agrionidae) Michael L. May 77 
Perlesta xube, a new stonefly species from Nebraska 
(Plecoptera: Perlidae) B. P. Stark, H. A. Rhodes 92 


Description of adults of Labiobaetis apache (Ephem- 
eroptera: Baetidae) with additions and corrections 
to inventory of Colorado mayflies R.S. Durfee, B.C. Kondratieff 97 


Position of Hemicaecilius (Psocoptera: Lachesillidae) 
and description of a new species from Venezuela 
A. N. G. Aldrete, E.L. Mockford 102 


Descriptions of female, nymph, egg, and redescription 
of male of Amphinemura mockfordi (Plecoptera: 
Nemouridae) Charles H. Nelson 107 


First adult description of the unusual baetid mayfly 

genus Echinobaetis (Ephemeroptera: Baetidae) 

C. R. Lugo-Ortiz, W. P. McCafferty 113 
Mating behavior of Pelecinus polyturator (Hymenoptera: 

Pelecinidae) A. P. Aguiar 117 
Rate of species descriptions in Odonata K. J. Tennessen 122 
Activity and reproduction of Calosoma frigidum (Coleop- 

tera: Carabidae) in northern Michigan forests 

R. M. Snider, R. J. Snider 197 
New generic synonymies in Baetidae (Ephemeroptera) 
R. D. Waltz, W. P. McCafferty 134 
Critical commentary on the genus Siphlonisca (Ephem- 

eroptera: Siphlonuridae) W. P. McCafferty, G. F. Edmunds, Jr. 141 
Establishment of Aleiodes (= Rogas) indiscretus (Hymen- 

optera: Braconidae), an introduced parasite of 

gypsy moth in North America’ R. F. W. Schroder, A. M. Sidor 148 
A new North American species of Cluzobra (Diptera: 

Mycetophilidae ) EdwardI.Coher 151 
First record of Anthopotamus verticis and new records 

of A. distinctus (Ephemeroptera: Potamanthidae) in 

New England Steven K. Burian 155 
First Alabama record of the pale-bordered cockroach, 

Pseudomops septentrionalis (Dictyoptera: Blat- 

tellidae) T. H. Roulston, A. G. Appel 159 


BOOK REVIEW 91 
ANNOUNCEMENT: Increase in page charges 101 


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Vol. 108, No. 2, March & April, 1997 77 


THE STATUS OF SOME SPECIES OF ENALLAGMA 
(ODONATA: ZYGOPTERA: COENAGRIONIDAE)! 


Michael L. May 


ABSTRACT: I have investigated the identity and generic placement of five little known species of 
coenagrionid damselflies usually assigned to Enallagma. Of these, E. camerunense is shown not to 
belong to Enallagma but probably to be an aberrant Pseudagrion. Enallagma kauderni, commonly 
regarded as a subspecies of E. nigridorsum, appears to be as well-differentiated from the latter as 
either is from E. vansomereni, so I consider E. kauderni to be a full species. Examination of the 
type of Enallagma melanotum demonstrated it to be identical with Agrion (now Cercion) sexline- 
atum. Enallagma psuedalongatum has been incorrectly placed as a synonym of E. elongatum in 
recent catalogs, probably owing to misinterpretation of Fraser’s (1947) comparison of these dis- 
tinct species. Finally, E. strouhali is apparently identical with the earlier described E. risi; I also 
discuss the possible relations of these taxa to E. cyathigerum and E. boreale. 


The genus Enallagma, as currently understood, has been characterized largely 
by default, 1.e., on the basis of plesiomorphic characters shared with numerous 
other coenagrionid genera (e.g., anal vein separating from the wing margin 
before Ac, R3 separating from R> near the Sth and 4th postnodal crossveins in 
the fore- and hindwings respectively, pterostigmata similar in all wings of both 
sexes, a rounded frons, postocular spots present, and females with a vulvar spine 
on the sternum of abdominal segment 8 [Selys, 1876; Munz., 1919; Kennedy, 
1920b]) and by the absence of the derived characters that distinguish presum- 
ably related genera (i.e., the dorsoapical prominence of abdominal segment 10 
of male /schnura). Appropriate synapomorphies have not been recognized that 
serve to diagnose Enallagma unambiguously. 

As a consequence, Enallagma is a large and unusually diverse genus with 
strong, disjunct centers of diversity in eastern North America and subsaharan 
Africa, and a scattering of species in the Palearctic and Oriental regions 
(Haritonov & Belyshev, 1980). This strong concentration of species in two ar- 
eas that have been geologically separated for at least 90 MY (and perhaps up to 
170 MY; Windley, 1984) suggests the independent radiation of two clades, iso- 
lated either by vicariance or, more probably, by dispersal in these regions. No 
generally acceptable characters have been proposed to distinguish them, how- 
ever. Kennedy (1920a) separated the African species into three genera, 
Africallagma (type species, Agrion glaucum Burmeister), schnallagma (type, 
Ischnura elongata Martin), and Proischnura (type, Enallagma subfurcatum 


! Received July 5, 1996. Accepted August 20, 1996. 


: Department of Entomology, New Jersey Agricultural Experiment Station, Cook College, Rutgers 
University, New Brunswick, NJ 08903, U.S. A. 


ENT. NEWS 108(2): 77-91, March & April, 1997.5 \ S 
) Pp PWITH ON 


MAY 19 1997 > 


LIBRARIES 


78 ENTOMOLOGICAL NEWS 


Selys), but these also were ill-defined and have not been widely accepted (e.g., 
Longfield, 1936; Bridges, 1994). 

It is hardly surprising, then, that of the 70 extant species of Enallagma listed 
by Bridges (1994) at least a dozen are of uncertain identity, mostly because 
their descriptions are not decisive as to generic placement or possible synonymy 
with other species and because the species have not been clearly identified or 
redescribed subsequently. In this paper I attempt to clarify the status of five of 
these. 


MATERIALS AND METHODS 


Visual observations and drawings were made with a Wild™ stereomicro- 
scope equipped with a camera lucida. When possible, structural details, espe- 
cially of the penes, were examined using an Hitachi™ S510 electron microscope 
after coating with gold-palladium. Descriptive terminology for venation is that 
of Tillyard and Fraser (1938) and for the male caudal appendages that of Snod- 
grass (1954). Collections from which material was obtained are designated by 
the following acronyms: CM — Carnegie Museum of Natural History (Pitts- 
burgh); CU — Cornell University Collection; IRSN — Institut Royal des Sci- 
ences Naturelles (Brussels); MLM — collection of M.L. May; MNB — Museum 
fiir Naturkunde der Humboldt Universitat, Berlin; NMNH — National Museum 
of Natural History (Smithsonian Institution, Washington); NHMV — 
Naturhistorisches Museum, Vienna; RU — Rutgers University Collection. 

Synonymies are not exhaustive but include selected works bearing directly 
on the taxonomic status of the species. Recent general catalogs (Bridges, 1994; 
Davies & Tobin, 1985; Tsuda, 1991) are omitted because all three of these 
include all the listed names. 


RESULTS AND DISCUSSION 


Enallagma camerunense Karsch 


Enallagma camerunense: Karsch 1899, p. 8 (male descr.); Pinhey 1962, p. 29 (taxon. and distrib. 
note). 


Besides the type series, comprising two males from Cameroon, this species 
has apparently been reported only by Pinhey (1962), who examined a specimen 
from Ghana. He considered it “not a true Enallagma’’, but gave no reason for 
this opinion. I obtained the types, both now lacking abdominal segments 5-10, 
from the MNB. Their colors are very well preserved, and virtually identical, but 
one has the mesepisternum broken as though by a pinhole, and from this speci- 
men (date label “15/5 96”) I dissected the penis; I here take the opportunity to 


Vol. 108, No. 2, March & April, 1997 79 


designate the other, more intact specimen as the lectotype and have added a 
label with this designation (white, orange bordered, printed with “E. 
camerunense, Lectotype, M. May 1996”; above this are the following labels: 
blue, printed “N. Kamerun, Johann Albrechtshohe, 18 VIII-30 IX 96, L. Conradt 
S.”; red, printed “Type” with “Syn-” handwritten in front; pale yellow, printed, 
”Zool. Museum, Berlin”). 

The color pattern was described in some detail by Karsch ( 1899), but only 
verbally, and minor discrepancies exist between description and specimens, so 
the head, thorax, and two basal abdominal segments are illustrated in Fig. 1. 
Most of the pale areas are orange or yellowish, but the postocular spots are blue 
green, the dorsal abdominal pale areas bright blue (these become yellowish 
laterally), and an indistinct metepisternal stripe pale bluish; these areas were 
described by Karsch as “gelb”. Otherwise, however, the color pattern, hindwing 
measurements, and dates agree exactly with the description, so I believe these 
are correctly identified as the types, although they bear no determination label. 


Figure 1: Color pattern of Enallagma camerunense Karsch, lectotype (A-C) and Pseudagrion sub- 
lacteum (Karsch), Kenya, L. Victoria, NMNH, ident as P. pseudomassaicum rusingae by Pinhey 
(D).- A: head and thorax, dorsal view; - B, D: thorax, lateral view: - C: first and second abdominal 
segments, dorsal view. Heavy stippling - black; intermediate - blue (note that mes- and metepimeral 
blue stripes are much paler than other blue areas and merge gradually with surrounding cream to 
yellow-green areas); light - bright orange; none - pale orange to cream. Scale bar = 2.0 mm for A- 
C, 3.0 mm for D. 


ENTOMOLOGICAL NEWS 


Vol. 108, No. 2, March & April, 1997 81 


Figure 2: Penes of: Enallagma camerunense Karsch, paratype (A, B); Enallagma cyathigerum 
Charpentier), Canada, B.C., nr Haney, RU (C, D); Enallagma antennatum (Say), U.S.A., Nebraska, 
Dodge Co., RU (E, F); Enallagma glaucum (Burmeister), South Africa, Bloemfontein, CM (G, H); 
Pseudagrion sudanicum LeRoi, Zimbabwe, Masuma R., NMNH, ident. as P. rubroviridis by Pin- 
hey. - A, C, E, G, I: ventral views (note that in A only, the apex points upward); - B, D, F, H, J: lateral 
views. Scale is equal for all figures (scale bar = 0.5 mm). 


In particular, the narrowing of the orange antehumeral stipes is distinctive and 
not found in other Enallagma. This is a result of marked inward expansion of 
the black humeral stripes in the vicinity of the shallow depression (the 
mesopleural fossa) at the upper end of each mesopleural suture, a feature of 
coloration present but much less strongly developed in many other coenagrionids. 

In the absence of the caudal appendages, the penis seems to offer the best 
opportunity to determine the affinities of the specimens. This is shown in Fig. 2, 


82 ENTOMOLOGICAL NEWS 


where it is compared to those of the North American Enallagma cyathigerum 
(Charpentier) and E. antennatum Say and the African E. glaucum (Burmeister). 
It is dissimilar to all of these (e.g., in lacking an internal fold and well-devel- 
oped laterodistal lobes) and to other African species (Fig. 3) but somewhat 
resembles that of, e.g., Pseudagrion sudanicum LeRoi (Fig. 21, J). In addition. 
the thoracic color pattern is reminiscent of that of P. sudanicum and even more 
so of P. sublacteum (Karsch) (Fig. 1 D), especially in the combination of dorsal 
orange and lateral bluish coloring and the encroachment of the humeral on the 
antehumeral stripe near the mesopleural fossa. The latter is not present in all 
Pseudagrion but occurs in several other species I examined at the NMNH (and 
see Pinhey, 1973, his Fig. 3). The venation of both camerunense specimens is 
like that of Enallagma in that the anal vein diverges from the wing margin well 
before Ac. However, although in Pseudagrion the separation of the anal vein 
generally occurs at or barely before Ac, Pinhey (1964) noted that this separa- 
tion can be as far before Ac as in Enallagma. Based on penile morphology, 
coupled with its unusual color pattern, camerunense clearly appears not to be 
an Enallagma and is most likely to belong in Pseudagrion. It is not identical to 
any Pseudagrion species that I have examined or that Pinhey (1964, 1973) keyed 
or illustrated, although the penis is also somewhat similar to P. sj6stedti, a spe- 
cies showing great variability in coloration; the color pattern is rather like P. 
serrulatum but the penis of that species is quite different. For now I suggest it 
stand as Pseudagrion camerunense (Karsch), especially since that name would 
have priority over any likely synonyms. 


Enallagma kauderni (Sj6stedt) 


Ischnura kauderni Sj6stedt 1917, p. 4 (female descr.; figs.). 

Enallagma nigridorsum forma b: Ris 1921, p. 323 (male, female descr.; figs.). 
Enallagma nigridorsum kauderni: Schmidt 1951. p. 207 (male, female descr.; figs.). 
Enallagma kauderni: Lieftinck 1965, p. 253 (listed without comment). 


Although described by Sjéstedt (1917) in Ischnura, E. kauderni was recog- 
nized by Ris (1921) as being closely related to Enallagma nigridorsum Selys 
and has been placed as a subspecies of the latter by most recent authorities 
(Davies & Tobin, 1985; Tsuda, 1991; Bridges 1994) except Lieftinck (1965). It 
is difficult to resolve the true relationship of these species, since kauderni ap- 
parently is confined to Madagascar and the Comoros Islands and hence is no- 
where sympatric with nigridorsum. Nevertheless, I believe E. kauderni should 
be recognized as a full species. Enallagma nigridorsum, E. vansomereni Pin- 
hey, and E. kauderni form a compact species group (May, unpublished data; E. 
somalicum Longfield is also close). Figs. 3 and 4 compare the caudal append- 
ages and penes of these three species. The cerci of all three have a smooth 


Vol. 108, No. 2, March & April, 1997 83 


Figure 3: Caudal appendages of Enallagma kauderni (Sjostedt), Madagascar, NMNH (A, B); E. 
nigridorsum Selys, Tanzania, Arusha, NMNH (C, D); and E. vansomereni Pinhey, Uganda, Paimol, 
Aicholi, NMNH (E, F). - A, C, E: tenth abdominal segment and caudal appendages, lateral view 
(note that the dorsum is to the right in A, E, to the left in C); - B, D, F: caudal appendages, oblique 
dorsolateral view (from left in B, F, from right in D). Scale bars = 0.5 mm. 


84 ENTOMOLOGICAL NEWS 


Figure 4: Penes of Enallagma kauderni (A), E. nigridorsum (B), and E. vansomereni (C), lateral 
view; specimens same as in Fig. 3. Scale bars = 0.2 mm. 


dorsal hook with its edge more or less dorsoventrally elongated, a somewhat 
flattened ventral lobe, and between these a central swelling. As the figure shows, 
in profile the cerci are all distinctive in shape, while the dorsal arm of each 
paraproct of E. kauderni is relatively longer and more slender than in the other 
two species. The dorsal cercal hook of E. kauderni is different in shape and the 
ventral lobe less abrupt, while the central swelling protrudes much more promi- 
nently backward than in E. nigridorsum or E. vansomereni. Finally, the penis, 
especially, the shape of the terminal lobes, is much closer in E. kauderni to E. 
vansomereni than to E. nigridorsum. In general it appears that the taxa are all 
about equally well differentiated and thus should all be regarded as distinct 
species. 


Enallagma melanotum Selys 


Enallagma melanotum: Selys 1876. p. 538 (male descr.); Kirby 1890. p. 147 (catalog); Needham 
1930, p. 266 (key); St. Quentin 1962, p. 16: Haritonov & Belyshev 1980, p. 81 (distrib.). Agrion 
sexlineatum: Selys 1883, p. 135 (female descr.). 

Coenagrion sexlineatum: Kirby 1890, p. 150 (catalog); Ris 1916, p. 35; Needham 1930, p. 275 
(male, female descr.; figs.; key). 

Cercion sexlineatum: Asahina 1950, p. 140 (figs.). 


Two males from China were doubtfully placed in Enallagma by Selys (1876) 
in his original description of melanotum. As far as I can determine, no speci- 
mens have subsequently been referred to this species, although the name was 
noted by Needham (1930), St. Quentin (1962), and Haritonov & Belyshev (1980) 
at least. In the syntypes, loaned by IRSN, the head and thoracic patterns are 


Vol. 108, No. 2, March & April, 1997 85 


essentially identical, but the entire abdomen is missing from one of the two 
specimens. That of the other has been broken and repaired, but twisted about 90 
degrees; nevertheless, the abdomen seems correctly associated with the thorax 
and head. This specimen is designated herewith as the lectotype (it bears the 
following labels: small, yellow, hand written “Chine, Hou.” [?]: white, hand- 
written except “det.” printed “det. de Selys, Enallagma ?, melanotum”; white, 
handwritten except “det.” printed “Agrion, sexlineatum ?, det. Asahina 1953”). 

As indicated by both color pattern and morphology of the caudal append- 
ages (Fig. 5), this specimen proved apparently to be identical to Cercion sex- 
lineatum (Selys). Cercion, although not very clearly defined, is distinguished 
from Enallagma by the absence of a vulvar spine in females and by the form of 
the male caudal appendages; sexlineatum is the only Cercion with the dorsolat- 
eral lobes of the cerci so short and the ventromedial arms closely appressed for 
most of their length to the caudal surface of segment 10 (Asahina, 1950, 1961). 
The only unusual feature in the type of E. melanotum is the presence of tiny 
dorsal denticles on the cerci. The penis of the E. melanotum type could not be 
extruded but was largely visible within the genital fossa. The slight difference 
in shape from C. sexlineatum (Fig. 5) is evidently due to postmortem changes 
resulting from compression within the fossa. Dr. Asahina suggested (personal 
communication, 1993) that E. melanotum might actually be synonymous with 
C. v-nigrum (Needham). I studied the type of the latter at CU, however, and 
found that the cerci protrude markedly backward and each have two easily vis- 
ible, nearly coplanar branches, while the paraprocts each have large, white, 
setose medial lobes, just as illustrated by Needham (1930) and quite unlike E. 
melanotum. | have not examined the type of C. sexlineatum, but it is a wide- 
spread and distinctive species in Japan and China that has been illustrated 
repeatedly. I therefore feel confident in concluding that E. melanotum is identi- 
cal to sexlineatum. Although the name melanotum has rarely been applied in 
taxonomic practice, it clearly has priority over sexlineatum, and Needham (1930) 
and St. Quentin (1962) gave comparisons, albeit very brief ones, to other spe- 
cies. Therefore, Enallagma melanotum should be transferred to Cercion and 
C. sexlineatum must fall as a junior subjective synonym of melanotum. 


Enallagma pseudelongatum Longfield 


Enallagma elongatum: Ris 1921, p. 324 (male descr.). 
Enallagma pseudelongatum: Longfield 1936. p. 474 (male, female descr.; figs), Fraser 1947, 
p. 143 (synonymy; figs.). 


This species has been erroneously treated as a synonym of E. elongatum 
(Martin) (Davies & Tobin, 1985; Tsuda, 1991; Bridges, 1994), probably 
because of the misleading title of Fraser’s 1947 paper (see Literature Cited). 
Fraser certainly regarded the two species as distinct, however, and illustrated 
the differences in the male caudal appendages quite clearly. Based on my 


86 ENTOMOLOGICAL NEWS 


Figure 5: Color patterns and reproductive structures of male Enallagma melanotum Selys, lecto- 
type (A, B, D-F) and Cercion sexlineatum (Selys), Japan, det. Asahina, FSCA (C, G-I). — An'G: 
thorax, lateral view, horizontal hatching is internal surface of cuticle exposed by break around pin; 
-B: distal abdominal segments, dorsal view. — D, G: caudal appendages and tenth abdominal seg- 
ment, posterolateral view; — E. H: caudal appendages and tenth abdominal segment, lateral view: — 
F, I: distal portion of penis, ventral view. Scale bar = 2.0 mm, except 1.0 mm in F, I. 


Vol. 108, No. 2, March & April, 1997 87 


examination of specimens at NMNH identified by E. Pinhey and compared by 
me to Fraser’s drawings, I concur fully with his opinion. The “synonymy” of 
his title refers not to the fact that the two names are synonyms but merely indi- 
cates that he included the synonymic history of each. 


Enallagma risi Schmidt 


Enallagma risi: Schmidt 1961, p. 409 (male, female descr.; figs.); Dumont 1975, p. 244 (male, 
female descr.; figs.); Harttonov 1975, p. 11 (distrib.); Asahina 1978, p. 116 (larva); Haritonov & 
Belyshev 1980 (distrib.). 

Enallagma strouhali: St. Quentin 1962, p. 16 (male, female descr.; figs.); Haritonov 1975, pp. 11, 
19 (distrib.; key); Haritonov & Belyshev 1980, p. 81 (distrib.). 


Among the most difficult and confusing species groups within Enallagma 
is that including the type, E. cyathigerum. Bridges (1994) listed seven species 
that I include here: belyshevi Haritonov, boreale (Selys) (with 2 subspecies and 
1 synonym listed), cyathigerum (2 subspecies, 9 synonyms), deserti (Selys) (1 
subspecies), nigrolineata Belyshev & Haritonov, risi, and strouhali St. Quen- 
tin. The actual status of several of the supposed synonyms and subspecies is 
open to question. For example, Donnelly (1989) placed E. vernale Gloyd as a 
subspecies of E. cyathigerum because apparently intermediate specimens exist, 
but the relationship cannot be that of subspecies in the usual sense because the 
two forms co-occur over a wide geographic area. Jurzitza (1975) regarded E. 
boreale as a synonym or subspecies of E. deserti on the basis of the essentially 
identical morphology of the caudal appendages, but this was provisionally 
rejected by Garrison (1984) and Westfall & May (1996) pending a thorough 
study of related forms. Variation in color pattern apparently is quite extensive 
among North American populations of E. boreale and E. cyathigerum (Walker, 
1953), yet many of the named infraspecific forms differ mainly in peculiarities 
of coloration as, indeed, do the supposed species, E. belyshevi and E. nigroline- 
ata, for example. 

Unravelling this tangle will require close study of abundant material over 
an enormous geographic span. Here I attempt only to straighten a few threads. 
The greatest proliferation of recognized taxa is in a great arc, from southwest- 
ern to central Asia, through southern Siberia, and on to the Kuriles and northern 
Japan, where these are the only species of Enallagma (Haritonov, 1975; 
Haritonov & Belyshev, 1980; Belyshev and Haritonov, 1982). Schmidt (1961) 
described E. risi based on types from Afghanistan. He compared these to E. 
deserti, but emphasized both verbally and in his illustrations the presence of a 
medially directed terminal point of the dorsal arm of the cerci; except for this, 
the illustrations are, unfortunately, not very detailed. Schmidt also noted and 
illustrated the caudal appendages of males from central Mongolia that he con- 
sidered to be the same species, although he did not include them in the type 


88 ENTOMOLOGICAL NEWS 


series. A year later St. Quentin (1962) described E. strouhali from Manchuria. 
This he considered closest to E. calverti Morse, normally regarded as a syn- 
onym of E. boreale. He compared it also to E. cyathigerum, E. deserti, and E. 
risi, but from these he thought the caudal appendages “completely different” 
(“durchaus verschieden”), although he noted that the male from Mongolia 
illustrated by Schmidt could be the same species. I directly compared the 
holotype male of E. strouhali (NMHV) to a series of supposed E. risi from 
Mongolia (MNB), including the three identified and listed by Schmidt, and 
found them to be essentially identical (Fig. 6). I have not located the type of 
E. risi, but Dumont’s (1975) illustrations of specimens from Afghanistan also 
appear identical, and Prof. Dumont has confirmed (personal communication, 
1996) that my illustrations are not distinguishable from his specimens. This 
includes that of the female mesostigmal plates, which are distinctive by virtue 
of the anterior margin of each being raised into a prominent, erect ridge (Fig. 


<) 


Tr... 


Figure 6: Caudal appendages and tenth abdominal segment of Enallagma strouhali St. Quentin, 
holotype (A-C) and E. risi Schmidt, Zentral-Mongolai, Ugai-nor, 27-IV-1954, det. E. Schmidt, 
MNB (D, E); mesostigmal plates of E. risi, Mongolia, Chara-us-nur, 28 June 1964, MNB (F); and 
cerci of Enallagma sp. nr. belyshevi, China, Chahar Prov. [now Inner Mongolian Autonomous 
Region], Yang-Kia-ping, NMNH (G), E. boreale circulatum. Japan, Hokkaido, Kushiro Marsh, 
MLM (H), and E. b. boreale, U.S.A., VT, Windsor Co., MLM (I). — A: dorsal view; — B, D: lateral 
views; — F: dorsal view; — C, E, G-I: right cercus, dorsomedial view. Scale bar = 1.0 mm except for 
C and E, for which bar = 0.7 mm. 


<> 


Vol. 108, No. 2, March & April, 1997 89 


6F). Thus I think there can be little doubt that the two species are the same and 
that E. risi thus extends at least from Afghanistan to Manchuria (Prof. Dumont 
also has specimens in his collection from northern Pakistan and Tajikistan). 
Enallagma cyathigerum mongolicum Benedek (Dumont, 1975), E. deserti 
continentale Belyshev, and E. beleyshevi might also be synonyms of E. risi, but 
this remains to be confirmed. 

Beyond this, the similarity of the caudal appendages of E. risito E. b. boreale 
and E. b. circulatum Selys. and thus also to E. deserti, is quite close (Fig. 6C, E, 
H, I). The apical hook is more obviously expanded in an anterodorsal to 
posteroventral direction, and it is more strongly recurved (thus less projecting 
in lateral view) in E. boreale than in E. risi. However, cerci from a specimen 
from northern China (Fig. 6G) appear more or less intermediate. This is from 
an area where belyshevi or circulatum as well as risi could occur; the specimen 
shows no sign of the lateral black abdominal markings that characterize typical 
circulatum but is similar in markings and apparently in the shape of the caudal 
appendages to belyshevi. A female from the same locality has mesostigmal plates 
slanting upward so that the anterior margins are distinctly elevated, but they do 
not form an abrupt ridge as in the Mongolian E. risi female; in this respect the 
Chinese specimen is more like North American E. cyathigerum and E. boreale. 
It appears possible, although again far from certain, that E. risi may not be 
specifically distinct from EF. boreale. If that were to prove so, then quite con- 
ceivably only two fully differentiated species make up this group, since E. 
nigrolineata seems only weakly distinct from E. cyathigerum and, as noted 
above, E. boreale may be synonymous with E. deserti. Alternatively, several 
infraspecific or heretofore unrecognized forms may actually be good species or 
semispecies. This group of taxa cries out for molecular and/or quantitative 
morphometric study. 


ACKNOWLEDGMENTS 


It is a pleasure to acknowledge the advice and assistance of S. Asahina and H. Dumont 
in determining the identities of Enallagma melanotum and Enallagma strouhali, respectively. 
I also wish to thank Nancy Adams and O. S. Flint (NMNH), U. Aspock (NHMV), P. Grootaert 
(IRSN), K. K. Gunther and G. Peters (MNB), and M. J. Westfall (FSCA) for the loan of 
specimens in their care, K. Maramorasch for translating papers from Russian, and F. L. Carle, 
H. B. White and an anonymous reviewer for helpful comments on the manuscript. This is 
New Jersey Agricultural Experiment Station Publication #D-08002- 13-96, supported by State 
funds. 


LITERATURE CITED 
Asahina, S., 1950. Odonata. Jn S. Asahina, T. Ishihara, & K. Yasumatsu, [Eds.], Iconagraphia 


Insectorum Japonicorum, editio secunda reformata, Vol. 3, pp. 131-168. Hokuryukan, Tokyo. 
(In Japanese). 


90 ENTOMOLOGICAL NEWS 


Asahina, S., 1961. Contributions to the knowledge of the odonate fauna of central China. Tombo 4: 
1-17. 

Asahina, S., 1978. A new and some known species of Odonata from Kashmir. Senckenbergiana 
Biol. 59: 115-120. 

Belyshey, B. F., & A. Y. Haritonov, 1982. Areal Enallagma cyathigerum Charp. and the problem 
of Bering Land faunistic connections. Izv. Sib. Otdel. Akad. Nauk. SSSR (Biol.) 1982(3): 84- 
86. (In Russian). 

Bridges, C. A., 1994. Catalogue of the Family-group, Genus-group and Species-group Names of 
the Odonata of the World (Third Edition). Urbana, IL, privately published. 

Davies, D. A. L., & P. Tobin, 1985. The Dragonflies of the World: A Systematic List of the Extant 
Species of Odonata, Vol. 1, Zygoptera, Anisozygoptera. Utrecht, S.1.0. Rapid Comm. (Supp!.) 
S127 ipp: 

Donnelly, T. W. 1963. Possible phylogenetic relationships among North and Central American 
Enallagma. Proc. N. Central Branch, Entomol. Soc. Amer. 18: 116-119. 

Donnelly, T. W. 1989. The status of Enallagma cyathigerum (Charp.) and E. vernale Gloyd in 
south-central New York (Zygoptera: Coenagrionidae). Odonatologica 18: 373-378. 

Dumont, H.J. 1975. A note on some dragonflies from Afghanistan. Odonatologica 4: 243-248. 

Fraser, F. C., 1947. The synonymy of Enallagma elongatum (Martin) and E. pseudelongatum 
Longfield and their differentiation from a new African species (Order Odonata). Proc. R. Ento- 
mol. Soc. Lond. 16: 143-148. 

Haritonovy, A. Y., 1975. North-Eurasian species of the genus Enallagma Charp. (Insecta: Odonata). 
New Little-known Spec. Sib. Fauna 9: 99-20. (In Russian). 

Haritonoyv, A. Y., & B. F. Belyshev, 1980. Peculiarities in the geographic distribution of the genus 
Enallagma Charp., 1840 (Insecta. Odonata). Trudy Biol. Inst. Sib.Otdel. Akad. Sci. SSSR 40 
80-84. (In Russian). 

Jurzitza, G., 1975. Rasterelektronenmicroscopische Untersuchungen an den Appendices und den 
Laminae mesostigmales einiger Enallagma-Arten (Odonata. Zygoptera). Forma et Functio 8: 
33-48. 

Karsch, F., 1899. Odonaten von Johann-Albrechts-Hohe (Nord-Kamerun), gesammelt von Herrn 
Leopold Conradt. Ent. Nachrichten 25: 161-176. 

Kennedy, C. H., 1920a. Forty-two hitherto unrecognized genera and subgenera of Zygoptera. Ohio 
J. Sci. 21: 83-88. 

Kennedy, C. H., 1920b. The phylogeny of the zygopterous dragonflies as based on the evidence of 
the penes. Ohio J. Sci. 21: 19-29. 

Lieftinck, M.A., 1965. Notes on Odonata of Madagascar, with special reference to the Zygoptera 
and with comparative notes on other faunal regions. Verhand. Naturf. Ges. Basel 76: 229-256. 

Longfield, C., 1936. Studies on African Odonata, with synonymy and descriptions of new species 
and subspecies. Trans. R. Entomol. Soc. Lond. 85: 467-498. 

Munz, P. A. 1919. A venational study of the suborder Zygoptera (Odonata), with keys for the 
identification of genera. Mem. Amer. Entomol. Soc. 3: 1-78. 

Needham, J. G. 1930. A Manual of the Dragonflies of China. A Monographic Study of the Chinese 
Odonata. Vol. 11 in Zoologica Sinica. Ser. A, Invertebrates of China. Peking, Fan Memorial 
Institute of Biology. 399 pp. 

Pinhey, E., 1962. Some records of Odonata collected in tropical Africa. J. Entomol. Soc. S. Afr. 25: 
20-50. 

Pinhey, E., 1964. A revision of the African members of the genus Pseudagrion Selys (Odonata) 
Revta. Ent. Mocamb. 7: 5-196. 

Pinhey, E., 1973. Notes on some African representatives of the genus Pseudagrion Selys 1876, 
with descriptions of two new species (Zygoptera: Coenagrionidae). Odonatologica 2: 317-327. 

Ris, F., 1916. H.-Sauter’s Formosa Ausbeute. Odonata (Mit Notitzen tiber andere ostasiatische 
Odonaten). Supplta. Ent. 5:1-81. 

Ris, F., 1921. The Odonata or dragonflies of South Africa. Ann. S. Afr. Mus. 18: 245-452. 


Vol. 108, No. 2, March & April, 1997 91 


St. Quentin, D., 1962. Eine neue Enallagma aus der Mandschurei (Odonata, Zygoptera). Ann. 
Naturhist. Mus. Wien 65: 241-243. 

Schmidt, E., 1951. The Odonata of Madagascar, Zygoptera. Mem. Inst. Sci. Madagascar 6A: 115- 
283. 

Schmidt, E., 1961. Ergebnisse der Deutschen Afghanistan-Expedition 1956 der Landessammlungen 
Karlsruhe sowie der Expeditionen J. Klapperich, Bonn 1952-53 und Dr. K. Lindberg, Lund 
(Schweden) 1957-60. Beitr. Naturk. Forsch. SW-Deutschl. 19:399-435. 

Selys-Longchamps, E. De, 1876. Synopsis des agrionines, cinquiéme légion: Agrion (suite). Le 
genre Agrion. Bull. Acad. Sci. Belg. (2) 41:247-322, 496-539, 1233-1309. 

Selys-Longchamps, E. De, 1883. Les Odonates du Japon. Ann. Soc. Entomol. Belg. 27:82-143. 

Sjostedt, Y., 1917. Odonaten aus Madagascar eingesammelt von Dr. W. Kaudem 1911-1912. Ark. 
Zool 11(13): 1-12. 

Snodgrass, R. E. 1954. The dragonfly larva. Smith. Misc. Coll. 123: 1-38. 

Tsuda, S., 1991. A Distributional List of World Odonata. Osaka. privately published. 362 pp. 

Walker, E. M., 1953. The Odonata of Canada and Alaska. Vol 1. Toronto, Univ. Toronto Press. 292 


pp. 

Westfall, M. J., & M. L. May. 1996. Damselflies of North America. Gainesville, FL. Scientific 
Publishers. 649 pp. 

Windley, B. F., 1984. The Evolving Continents, 2nd edition. Wiley, New York. 399 pp. 


BOOK REVIEW 


DAMSELFLIES OF NORTH AMERICA. Minter J. Westfall, Jr. and Michael 
L. May. 1996. Scientific Publishers, Gainesville, FL. 650pp; ISBN 0-945417- 
93-4, $69.50 Cloth. 


The order Odonata (dragonflies) includes the suborders Anisoptera and Zygoptera. How- 
ever, in North America, the term “dragonfly” normally refers only to the Anisoptera. Thus, 
in 1955 when James G. Needham and Minter J. Westfalt Jr. published their A Manual to the 
Dragonflies of North America, it did not include the damselflies. For more than 40 years 
“The Manual” has awaited its companion often referred to as “the book” among those inter- 
ested in the Odonata. Wait no more. Damselflies of North America appeared in 1996 and is 
excellent. It surely will become the standard reference in the field supplanting Volume One 
of Edmund Walker’s The Odonata of Canada and Alaska (1953) and Sidney Dunkle’s Dam- 
selflies of Florida, Bermuda, and the Bahamas (1990), excellent regional works that have 
served in the absence of a continental resource. 

In a sign of the times, the imminent publication of Damselflies of North America was 
announced last June 6 by an e-mail message from the International Odonata Research Insti- 
tute in Gainesville, Florida to members of the Dragonfly Society of the Americas. By now, 
likely all have purchased a personal copy of this monograph that covers each of the 161 
species of damselflies known to occur in North America, northern Mexico, and the Greater 
Antilles. Unlike field guides that focus on adult insects and cater to novices with common 
names and color photographs for identification, this volume is a more scholarly work. While 
there are color photographs of almost 20% of the fauna; thorough keys, detailed descrip- 


(Continued on page 96) 


o2 ENTOMOLOGICAL NEWS 


PERLESTA XUBE, A NEW STONEFLY SPECIES 
FROM NEBRASKA (PLECOPTERA: PERLIDAE)! 


Bill P. Stark2, Howard A. Rhodes» 


ABSTRACT: Perlesta xube, anew stonefly species, is described from male, female, egg and nymphal 
stages, and a holotype male is designated. Adults are similar to Perlesta adena but differ most 
significantly in aedeagal shape and pattern of armature. The new species is known from a single 
location in northwestern Nebraska. 


For most of this century, Perlesta placida (Hager) was regarded as a com- 
mon, but variable, species found throughout eastern North America. Stark (1989) 
recognized twelve members in a complex of species differentiated primarily on 
the basis of internal male genitalia and egg morphology. In this study we call 
attention to an additional member of this complex found in Nebraska. Termi- 
nology and methods follow Stark (1989). 


Perlesta xube, NEW SPECIES 


Male. Forewing length 7 mm. General color dark brown. Head and pronotum brown, 
patterned with dark brown, but with a pair of oval pale spots anterolateral of posterior ocelli 
(Fig. 1). Basal antennal segments pale, becoming dark brown beyond segment 8. Forefemora 
longitudinally striped in yellow and brown. Wing membrane and veins dark brown except 
for pale areas proximal to the arculus, in the mid-costal region, along the median vein ante- 
rior to the cord, and in the intercubital area (Fig. 7). Basal cereal segments banded, becom- 
ing dark brown beyond segment 8. Paraprocts in lateral aspect slender, curved forward and 
bearing a subapical tooth (Fig. 4). Tergum 10 mesal sclerite almost divided by membranous 
band; sensilla basiconica patch sparse (Fig. 3). Penis tube slender and sinuate, dorsal hair 
patch (DP) narrow subapically but broadly expanded apically around base of caecum (C); 
apex of caecum bare (Fig. 6). 

Female. Forewing length 9-10 mm. Color pattern similar to male. Subgenital plate lobes 
truncate and separated by a U-shaped notch. Plate dark laterally and membranous mesally 
(Fig. 5). 

Egg. Length ca. 0.5 mm, width ca. 0.4 mm. Collar obscure, buttonlike, not shown in 
orientation of Fig. 8. Chorion finely pitted around poles and coarsely pitted in mesal third 
(Fig. 8). Micropylar row in posterior region with pitted field (Fig. 9). 

Nymph. General color dark brown. Light colored M-line of head pattern distinct (Fig. 
2). Abdominal terga brown with dark brown pigment spots around intercalary setae. 


1 Received August 3, 1986. Accepted September 9, 1996. 
2 Biology Department, Mississippi College, Clinton, MS 39058. 
3 Department of Entomology, Colorado State University, Fort Collins, CO 80523. 


ENT. NEWS 108(2): 92-96, March & April, 1997 


Vol. 108, No. 2, March & April, 1997 93 


Figs. 1-6. Perlesta xube morphological features. 1. Adult head and pronotum. 2. Nymphal head and 
pronotum. 3. Male terminal abdominal segments, dorsal. 4. Male paraproct, lateral. 5. Female ter- 
minal abdominal segments, ventral. 6. Penis tube+sac, lateral (C = caecum, DP = dorsal patch). 


94 ENTOMOLOGICAL NEWS 


Types. Holotype CG, 30 paratype CO and 29 paratype Q specimens collected in 
Nebraska, Cherry Co., Dry Creek, Merriman, 26 June 1996, B. C. Kondratieff, H. Rhodes. 
Additional paratypes reared from nymphs collected at type locality, 6 June 1995,2 0,9 Q, 
B. C. Kondratieff. Holotype and one female paratype deposited in the National Museum of 
Natural History, other paratypes deposited in the C. P. Gillette Museum of Arthropod Diver- 
sity, Colorado State University and in the Stark collection, Mississippi College. 


Type locality. Dry Creek flows through Merriman, Nebraska, as a 
channelized stream with rather steep banks and a swift current. Cottonwoods 
form a riparian corridor along the stream. Robust populations of the mayflies 
Acerpenna pygmaea (Hagen), Baetis tricaudatus Dodds, Ephemerella inermis 
Eaton, and Heptagenia diabasia Burks were found along with the damselflies 
Calopteryx aequabilis Say and Hetaerina americana (Fabricius). The only stone- 
fly species collected with P. xube were Isoperla quinquepunctata (Banks) and 
Perlesta decipiens (Walsh). 


Etymology. The Native American Ponca lived in the region of northern 
Nebraska where this species was discovered. Their word xube means “super- 
natural power’’, which they believed all things possessed in varying amounts 
(Leitch 1979). 


Fig. 7. Forewing of Perlesta xube showing pale areas in the costal area (C), proximal to the arculus 
(A), along the median vein (M), and in the intercubital area (1). 


DISCUSSION 


Adult specimens of P. xube key to couplet 6 in Stark (1989). At that point an 
impasse occurs because the penis tube is short, slender and has the dorsal patch 
expanded at the apex (Fig. 6). In these features P. xube agrees with neither P. 
cinctipes nor P. adena, but the expanded dorsal patch is somewhat similar to 
that of P. adena. The egg is also quite similar to that of P. adena and isolated 
females might prove difficult to distinguish from that species. However, the 
wing pigmentation pattern of P. xube (Fig. 7) should be distinctive enough to 


Vol. 108, No. 2, March & April, 1997 95 


-_e—e—_ eo SSSSSSSSSSSSSSSSSmmmmmmhFheFeFeFeFFFfFFsfsfsfsses 


a 


188ym 


5.86 kV 


Figs. 8-9. Scanning electron micrographs of Perlesta xube egg features. 8. Lateral aspect. 9. Detail 
of micropylar region of posterior pole. 


96 ENTOMOLOGICAL NEWS 


permit identification. Nymphal specimens appear distinct from P. decipiens 
(Walsh) and P. cinctipes (Banks) by virtue of the darker pigment pattern of the 
occiput and anterior region of the frons (Fig. 2). 

With the addition of P. xube, 13 species of stoneflies are now known for 
Nebraska (Rhodes & Kondratieff 1996). 


ACKNOWLEDGMENTS 


We thank B. C. Kondratieff for providing specimens and biological notes of this inter- 
esting species for our study. 


LITERATURE CITED 


Leitch, B. A. 1979. A concise dictionary of Indian tribes of North America. Reference Publica- 
tions, Inc. Algonac, Michigan. 646 pp. 

Rhodes, H. A. and B. C. Kondratieff. 1996. Annotated list of the stoneflies (Plecoptera) of west- 
em Nebraska, U.S.A. J. Kansas Entomol. Soc. 69:191-198. 

Stark, B. P. 1989. Perlesta placida (Hager), an eastern nearctic species complex (Plecoptera: Per- 
lidae). Entomol. Scand. 20:263-286. 


(Continued from page 91) 


tions, scanning electron micrographs, and line drawings of critical taxonomic features serve 
to identify both larval and adult forms. In fact, a number of previously undescribed larval 
forms are included in the keys. The authors have not used the “common” names recently 
given to all North American damselflies. 

The book’s introduction deals with morphology, classification, fossil record, life cycle, 
adult and larval behavior, physiology, biogeography, habitats, and conservation. That is fol- 
lowed by color photographs mostly of specimens in natural poses and mostly with good 
color reproduction. The main part of the book (~80%) contains keys, figures and detailed 
descriptions of each species. The extensive glossary, bibliography, and index all make this a 
very accessible book for both novices and old timers who want to learn more about damselflies. 

Although as insects go, damselflies are fairly well known, there are still new species 
and new larval forms to describe, smoldering taxonomic problems to resolve, and much to 
be learned about behavior and distribution. A particularly useful aspect of this book is that it 
identifies many of these lacunae and thus provides the sense that much remains to be discov- 
ered. Students looking for projects can find one for almost every species. Damselflies of 
North America will be widely used and should further increase the demand for the next book 
in this area, a much needed field guide. 


Harold B. White 
University of Delaware 


Vol. 108, No. 2, March & April, 1997 97 


DESCRIPTION OF ADULTS OF LABIOBAETIS 
APACHE (EPHEMEROPTERA: BAETIDAE) 
WITH ADDITIONS AND CORRECTIONS TO THE 
INVENTORY OF COLORADO MAYFLIES! 


Richard S. Durfee, Boris C. Kondratieff2 


ABSTRACT: Male and female adults of Labiobaetis apache are described from specimens col- 
lected in northwestern Colorado, USA. Biological notes for this species are given and a key sepa- 
rating the male adults of the six North American species of Labiobaetis is provided. Additional 
Colorado distributions for L. dardanus and Baetis notos are given. Cinygmula tarda 1s reported for 
the first time in Colorado. 


McCafferty and Waltz (1995) elevated the subgenus Labiobaetis Novikova 
and Kluge to generic rank. They included five North American species for- 
merly placed in the Baetis propinquus species group (Morihara and McCafferty 
1979a) and described a new species, L. apache, based on larvae collected in 
Arizona and Utah. They also provided an illustrated key to the larvae of all six 
North American species of Labiobaetis. Previously, the larvae in this group 
were treated diagnostically by Morihara and McCafferty (1979b) and Soluk 
(1981). 

Originally reported as Baetis propinquus (Walsh) by McCafferty et al. (1993), 
the larvae collected from Douglas Creek in Rio Blanco Co., Colorado were 
reexamined and determined to be L. apache. Previous to its description, the 
larvae of L. apache would key to B. propinquus as characterized by Morihara 
and McCafferty (1979b). 

A key separating adult males of the five North American species now 
included under Labiobaetis was provided by Morihara and McCafferty (1979a). 
At that time L. apache was unknown. Herein, we describe the adults of L. apache, 
thus completing the adult taxonomy of all known species of this genus in North 
America. During July, 1996, we returned to the Douglas Creek site and col- 
lected a large series of mature larvae for rearing. The following description is 
based on reared adults recently preserved in alcohol. 


Labiobaetis apache McCafferty and Waltz 


Adult male. Body length 5.0-6.0 mm, forewings 5.0-5.5 mm, hindwings 1.0 mm, caudal filaments 
9-10 mm. Head brown; antennal scape and pedicel light brown, flagellum light brown basally, pale 
apically: turbinate eyes orange. Thoracic nota and sterna dark brown, intersegmental areas white, 


1 Received September 26, 1996. Accepted October 8, 1996. 
2 Department of Entomology, Colorado State University, Fort Collins, Colorado 80523. 


ENT. NEWS 108(2): 97-101, March & April, 1997 


98 ENTOMOLOGICAL NEWS 


some stained with orange: forelegs smoky, apical 1/5 of foretibia pale, middle and hindlegs pale, 
inner apical margin of Ist and 2nd tarsal segments produced into a spine, all claws slightly dark- 
ened: forewings hyaline, basal area stained with brown, veins colorless. stigmatic area cloudy; 
hindwings hyaline, with two longitudinal veins and lacking a costal projection. Abdominal terga 
light brown with distinctive brown markings (Fig. 1): first abdominal sternite stained with brown, 
remaining abdominal sterna pale: sterna 2-9 with prominent red-brown mark laterally (Fig. 1) 
sometimes only visible on 2-6. Caudal filaments with basal three segments stained with red-brown, 
articulations of segments 4-6 also stained with red-brown, remainder of filament pale. Forceps 
white, basal 1/3 of first segment often stained with brown; basal enlargement of second segment as 
wide as long, terminal segment short and rounded; ventral posteromedian projection between for- 
ceps bases covered with setae, shaped as in Fig. 2. 


2 


Figures 1-2 Labiobaetis apache. Fig. 1 Adult male abdomen, lateral view. Fig. 2, male genitalia, 
ventral view. 


Vol. 108, No. 2, March & April, 1997 99 


Adult female. Body length 5.5-6.0 mm, forewings 5.0-5.5 mm, hindwings 1.0 mm, caudal fila- 
ments 9-10 mm. Head tan: lateral ocelli ringed with dark brown: antennae as in male. Thoracic nota 
and sterna generally light brown, metanotum and parts of the metasternum darker brown; all legs 
pale, tarsal spines as in male but present on forelegs as well. Wings similar to male but with lightly 
shaded longitudinal veins. Abdominal terga tan; abdominal sterna pale. Dorsal and ventral abdomi- 
nal markings as in male (Fig. |). Caudal filaments as in male. 


Material examined. Colorado: Rio Blanco Co.. Douglas Creek at Rt. 64. 19 July 1996. B. Kondra- 
tieff and R. Durfee, 26 males 23 females and 10 male and 9 female subimagos (reared), deposited 
in the C. P. Gillette Museum of Arthropod Diversity, Colorado State University, Fort Collins, Colo- 
rado. 


Diagnosis: The abdominal markings (Fig. 1) of both the male and female adults 
of L. apache readily distinguish it from the other North American Labiobaetis. 
These markings are so distinctive that field identification of adults and subima- 
gos is possible. In alcohol preserved specimens these color patterns may fade, 
particularly the dorsal markings on the male. Male genitalia of L. apache are 
very similar to L. dardanus (McDunnough). Specimens of L. dardanus from 
the Arkansas River, Pueblo Co., Colorado and the Niobrara River, Sioux Co., 
Nebraska were compared to L. apache and no major differences in the male 
genitalia were noted. 


We modify the following key to the adult males of North American Labio- 
baetis by Morihara and McCafferty (1979a) to include L. apache. 


1. Ventral posteromedian projection between forceps bases covered with setae (Morihara and 


McGaffertys 1979 askigssi25 1S. 15 i0S) rye crscto ctetoisolocie nein Serene Sane ele 2 
Ventral posteromedian projection bare, often well sclerotized and pigmented (Morihara and 
McGaffertys 1979 av Fig: VA) c/o i6 evccoe ote ee coho sees ehor asus. o) lotet ounsohece 6, ouskerers L. ephippiatus 


2. Ventral posteromedian projection between forceps bases broadly rounded to broadly truncate 
(Morihara and McCafferty, 1979a; Figs. 12. 15. 18); abdominal segments 2-6 pale translucent, 


Opaque brownlor withidistincl pattems cise eta tetas eril rier sie itopale cite oleae ies 3 
Ventral projection slightly constricted basally (Morihara and McCafferty, 1979a; Fig. 13); ab- 
dominallsegzments}2=6 palejtranslucentiis.).5..21 1c staves) ie leroy rere te L. longipalpus 
3. Basal enlargement of forceps short and robust (Fig. 2); no chitinous internal spine between for- 
Geps) bases emcee Re Aes Ao Eee OE AON ete ere Re ete Oates COUR One 4 
Basal enlargement of forceps slender (Morihara and McCafferty, 1979a; Figs. 15, 18); chitinous 
infermalispine Usually, VASIDIC: im, eyeniois ne cies srepicints See e er eterere nes Tomei re crerien serra 5 
4. Abdominal tergites 2-6 with distinct pattern (Fig. 1)... ............ 000s ee eee L. apache 
Abdominal tergites 2-6 light brown to yellow without distinct pattem .......... L. dardanus 
5. Basal enlargement of forceps simple, subquadrate from ventral perspective (Morihara and 
McCafferty, 1979a; Fig. 15); abdominal segments 2-6 usually opaque brown .... L. frondalis 


Basal enlargement of forceps with distal, medially directed, acute projection (Morihara and 
McCafferty, 1979a; Fig. 18); abdominal segments 2-6 usually pale translucent . . . L. propinquus 


DISCUSSION 


McCafferty and Waltz (1995) speculated that the adults of L. apache may 
key out near L. propinquus if a close relationship between these two species 
could be assumed based on similar larval morphology. However, L. apache 


100 ENTOMOLOGICAL NEWS 


appears to be most closely related to L. dardanus based on similarities of the 
male genitalia. In addition, both L. apache and L. dardanus have broad overlap- 
ping distributions in western North America and may share a common ancestor. 
Labiobaetis propinquus is widely distributed throughout eastern North America 
(Morihara and McCafferty 1979b) and has been reported from as far west as 
Texas (Lugo-Ortiz and McCafferty 1995) and eastern New Mexico (W.P. 
McCafferty, personal communication). Peters and Edmunds (1961) reported 
this species from the Navajo Reservoir basin, New Mexico, however, that was 
before the species description of L. apache. We have not examined the material 
from this study, but based on the geographical proximity, we suspect this record 
to be either L. apache or L. dardanus. 

Douglas Creek becomes highly turbid during precipitation events. Substrate 
is primarily sand with some gravel. Most mayfly larvae were collected from 
debris snags near the stream margin. This general habitat description is very 
similar to the type locality of L. apache, the Little Colorado River near St. 
Johns, Arizona. Other mayflies collected with L. apache from Douglas Creek 
included Choroterpes inornata Eaton, Heptagenia elegantula Eaton, and Tri- 
corythodes minutus Traver. McCafferty and Waltz (1995) indicated that C. inor- 
nata and L. apache may be characteristic species of the Colorado Plateau. 

Two species of Labiobaetis are currently known from Colorado. L. apache 
and L. dardanus. In Colorado, L. apache has only been collected from Douglas 
Creek near Rangely and may be restricted to streams of the Colorado Plateau. 
Labiobaetis dardanus had previously been reported from the Green River in 
Moffat Co. as Baetis dardanus (Durfee and Kondratieff 1994). Additionally, 
the authors have examined specimens of L. dardanus from the Conejos River in 
Conejos Co, the Arkansas River in Pueblo Co., and the South Platte River in 
Douglas Co. Although not commonly collected, this species appears to be wide- 
spread throughout Colorado. 

Baetis notos Allen and Murvosh was previously reported in Colorado from 
the Gunnison River (McCafferty et al. 1993). We have examined specimens 
from Four Mile Creek in Fremont Co. and the Arkansas River in Pueblo Co. A 
southwestern species with Mexican affinities, B. notos apparently reaches its 
northernmost limit in Colorado. 

A male adult of Cingymula tarda (McDunnough) was recently collected 
from a mating swarm on the South Fork of the South Platte River in Park Co. 
(elev. 1840 m). Previously recorded from Alberta and Montana (McDunnough 
1929), Idaho (Jensen 1966) and northern Utah (Edmunds 1954), the discovery 
of this species in central Colorado extends its range southward. 

With the addition of C. tarda and the elevation of Labiobaetis to generic 
rank, a total of 44 genera and 102 species of mayflies are now recorded for 
Colorado. 


Vol. 108, No. 2, March & April, 1997 101 


ACKNOWLEDGMENT 
We thank Lynn Bjork for the illustrations. 
LITERATURE CITED 


Durfee, R. S. and B. C. Kondratieff. 1994. New additions to the inventory of Colorado mayflies 
(Ephemeroptera) Entomol. News 105: 222-227. 

Edmunds, G. F., Jr. 1954. The Mayflies of Utah. Proc. Utah Acad. Sci. Arts Lettr., 31: 61-64. 

Jensen, S. L. 1966. The mayflies of Idaho (Ephemeroptera). Unpublished MS Thesis, Univ. Utah, 
Salt Lake City. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1995. The mayflies (Ephemeroptera) of Texas and their 
biogeographic affinities. Pp. 151-169. /n Current directions in research on Ephemeroptera. L. 
D. Corkum and J. J. H. Ciborowski (eds). Canadian Scholars’ Press Inc., Toronto. 

McCafferty, W. P and R. D. Waltz. 1995. Labiobaetis (Ephemeroptera: Baetidae): New status, 
new North American species and related new genus. Entomol. News 106: 19-28. 

McCafferty, W. P., R. S. Durfee. and B. C. Kondratieff. 1993. Colorado mayflies (Ephemerop- 
tera): An annotated inventory. Southwest. Nat. 38: 252-274. 

McDunnough, J. 1929. Notes on North American Ephemeroptera with descriptions of new spe- 
cies, II. Can. Entomol. 61: 169-180. 

Morihara, D. K. and W. P. McCafferty. 1979a. Systematics of the propinquus group of Baetis 
species (Ephemeroptera: Baetidae). Ann. Entomol. Soc. Am. 72: 130-135. 

Morihara, D. K., and W. P. McCafferty. 1979b. The Baetis larvae of North America (Ephemerop- 
tera: Baetidae) Trans. Am. Entomol. Soc. 105: 139-221. 

Peters, W. L., and G. F. Edmunds. Jr. 1961. The mayflies (Ephemeroptera) of the Navajo Reser- 
voir Basin, New Mexico and Colorado. Univ. Utah Anthropol. Pap., 55: 107-111. 

Soluk, D A. 1981. The larva of Baetis dardanus McDunnough (Ephemeroptera: Baetidae). Ento- 
mol. News 92: 147-151. 


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H.P.B. 


102 ENTOMOLOGICAL NEWS 


THE POSITION OF HEMICAECILIUS 
(PSOCOPTERA: LACHESILLIDAE) 
AND DESCRIPTION OF A NEW SPECIES 
FROM VENEZUELA! 


Alfonso Neri Garcia Aldrete2, Edward L. Mockford> 


ABSTRACT: Hemicaecilius venezolanus n. sp. is described from a single female collected near 
Trujillo, northern Venezuela. It differs from the generotype and hitherto single species of the genus, 
H. bogotanus from Colombia, in an important venational character. We argue that the generotype 
specimen, now known only from a fore- and a hindwing, is anomalous in the character, and that 
numerous venational similarities mark the two species as congeneric. Structural characters, includ- 
ing the nature of the lacinial apex, pretarsal claw, gonapophyses, subgenital plate, ciliation and 
venation of the forewing indicate that Hemicaecilius belongs in the Lachesillidae, subfamily 
Lachesillinae, rather than in the Elipsocidae where some authors have placed it. 


The genus Hemicaecilius has been an enigma for psocidologists since its 
description by Enderlein in 1903. The description was based only on wing 
venational characters of a single specimen collected in Bogota, Colombia. No 
genitalic information about Hemicaecilius was provided, and at present all that 
remains of the type specimen is one fore- and one hindwing, mounted on a slide 
deposited in the Museum fiir Naturkunde der Humboldt-Universitat, in Berlin. 

Enderlein (1903) regarded Hemicaecilius as near Graphocaecilius Enderlein 
and noted its close similarity in wing ciliation with that genus. Hemicaecilius 
was included in the Elipsocidae: Lesneiinae by Smithers (1972, 1990), who 
noted that its position was uncertain and that there was little known of this 
genus other than its venation and number of tarsal segments. 

The present study of a female specimen collected near Trujillo in northern 
Venezuela, allowed us to determine more precisely the systematic position of 
Hemicaecilius in the Psocoptera (see Discussion, below). Comparison of the 
wings of this specimen (Fig. 1) with the wings of the holotype of H. bogotanus 
(Fig. 9) convinced us that it belongs in Hemicaecilius. Although Hemicaecilius 
was described as having only two median veins in the forewing, we feel this is 
an anomalous condition of the holotype of H. bogotanus. Wing venational 
anomalies are frequent in the Psocoptera. When an anomaly involving two in- 


l Received. May 10, 1996. Accepted August 1, 1996. 


2 Instituto de Biologia, Universidad Nacional Auténoma de México, Departamento de Zoologia, 
Apartado Postal 70-153, 04510 México, D.F. 


3 Department of Biological Sciences, Illinois State University, Campus Box 4120, Normal, Illinois 
61790-4120. 


ENT. NEWS 108(2): 102-106, March & April, 1997 


Vol. 108, No. 2, March & April, 1997 103 


stead of three median veins in the forewing occurs, it usually results in broaden- 
ing of the distal end of cell R5, as seen in the type of H. bogotanus (Fig. 9). If 
we assume that the normal condition of the forewing of H. bogotanus is three 
median veins and a more slender distal end of cell RS, then the forewing simi- 
larities between H. bogotanus and the new species become striking. Notable 
similarities include the following: 1) basal Rs segment gently curved outward; 
2) Rs fork stem gently curved anteriad in its basal three-fifths, posteriad in its 
distal two-fifths; 3) Rs-M crossvein ca. one-fourth length of basal Rs segment; 
4) point of branching of R2+3 and R4+5 very slightly distal to basal M branch- 
ing; 5) median stem gently curved posteriorly in its basal three-fifths; 6) sparse, 
short setae present on margin and veins except none present on vein Cu 2. 
The holotype of the new species was dissected and its parts were mounted 
in euparal for microscopic study. Standard measurements (FW = forewing; HW 
= hindwing; F = femur; T = tibia; t1, t2 = tarsomeres 1| and 2; cttl = number of 
ctenidia on tl; P4 = fourth segment of maxillary palp; IO = minimum distance 
between compound eyes; d = transverse diameter of compound eye), given in 
microns, were made on the parts mounted, with a filar micrometer. The measur- 
ing unit is 1.36 microns for wings, and 0.53 microns for other parts. The holo- 
type is deposited in the Insect Collection, Zoology Department, Instituto de 
Biologia, Universidad Nacional Aut6noma de México, México City. 


Group Homilopsocidea 
Lachesillidae, Lachesillinae 


Hemicaecilius venezolanus NEW SPECIES (female) 


Diagnosis. Ocelli present, lacinial tips bifid, outer cusp larger than inner one; pretarsal claw with 
one preapical denticle and slender pulvillus; forewing with small setae along margin and veins, Rs 
and M connected by crossvein; hindwing with row of small setae along margin in cell R3, one pair 
of gonapophyses present; subgenital plate with a slender posterior projection on each side. Differ- 
ing from H. bogotanus by greater wing length, shape of the pterostigma, slightly more sigmoid 
stem of the Rs fork, shape of the areola postica, Rs-M in the hindwing meeting at a point instead of 
a fusion, and presence of marginal setae in cell R3 of the hindwing. 

Color (in 80% alcohol). Body dark reddish brown. Compound eyes black, ocelli clear, 
without centripetal crescents. Antennae and maxillary palps brown. Tergal lobes of meso- 
and metathorax more pigmented than rest of the body. Legs pale brown. Wings reddish brown. 


Morphology. Vertex slightly obtusely concave. Epicranial sulcus without lateral arms. 
Compound eyes below level of vertex. Lateral ocelli larger than median one. Epistomal sul- 
cus well defined (Fig. 8). Lacinial tips bifid, inner cusp short, slender; outer cusp large, 
conical, apically blunt (Fig. 2). With five distal inner lateral sensilla, three placoids alternat- 
ing with two trichoids. P4 with two small thin walled sensilla near the base. Pretarsal claws 
with slender pulvillus (Fig. 3). Hind femur (Fig. 6) with row of long setae ventrally. Fore- 
wing with small setae along margin and veins, as illustrated (Fig. 1). Hindwing with small 
setae on margin, between R2+3 and R4+5. No trichobothria distally on hind femur. Subgenital 


104 ENTOMOLOGICAL NEWS 


Figures 1-8. Hemicaecilius venezolanus n.sp. (Q ). 1. Fore and hind wings. 2. Lacinial apex. 3. 
Pretarsal claw. 4. Gonapophyses and ninth sternum. 5. Subgenital plate. 6. Hind femur. 7. Right 
paraproct and epiproct. 8. Front view of head. Scales in mm. Fig. 3 to scale of Fig. 2. Figs. 5-7 to scale 
of Fig. 4. 


Vol. 108, No. 2, March & April, 1997 105 


gies mien rmay ninguna tgs 


Fig. 9. H. bogotanus Enderlein (holotype). Fore and hind wings. Scale in mm. 


plate broad, setose, with four mesal macrosetae; with two slightly curved, stout, blunt, pos- 
terior projections, with border between them jagged, as illustrated (Fig. 5). Gonapophyses: 
only V3 present, elongate, slender, basally rounded, with distal one-third pointed; with setae 
along outer edge. Ninth sternum pigmented, as illustrated (Fig. 4), projected posteriorly in 
the middle. Spermapore large, near anterior border of ninth sternum. Paraprocts elongate, 
setose; sensory fields with 10-11 trichobothria on basal rosettes, and a marginal one without 
basal rosette. Epiproct almost trapezoidal, setose (Fig. 7). 
Measurements. FW: 1599; HW: 1339; F: 307; P4: 84; 10: 289; d: 72; IO/d: 4.01. 


Type Locality: VENEZUELA. Trujillo, 2,387 m. Old road, 46 km E Trujillo, 6. VIII. 
1988, C.W. & L.B. O’Brien & G. Wibmer, Holotype 9. 


DISCUSSION 


Two problems concerning Hemicaecilius must be considered: 1) its family 
placement, and 2) its relationships within the family. The size, color, wing 
venation, nature of the lacinial tip, reduction of the ovipositor valvulae to a 
single pair (the third), and presence of four macrosetae on the subgenital plate 
are all characters common in family Lachesillidae, and this particular constel- 
lation of characters is not found outside of that family. Assuming that the new 


106 ENTOMOLOGICAL NEWS 


species truly represents Hemicaecilius, we assign the genus to family 
Lachesillidae. 

Mockford and Sullivan (1986) proposed a classification of the family 
Lachesillidae, with two subfamilies, the Eolachesillinae and Lachesillinae. We 
believe that Hemicaecilius stands close to Nadleria Badonnel and Garcia Aldrete 
(1979) in the subfamily Lachesillinae. Our basis for placing it here, as opposed 
to near Graphocaecilius and allied genera in the Eolachesillinae is as follows: 
1) presence in the new species of a simple outer tine of the lacinial tip (bidentate 
in most Eolachesillinae); 2) Rs-M crossvein present in forewing in both 
Hemicaecilius species and in Nadleria (Rs and M fused for a distance through- 
out Eolachesillinae); 3) fusion of the ovipositor valvulae with the ninth sternum 
in the new species and in most Lachesillinae (junction is via the stem of v-1 in 
the Graphocaeciliini and there is no junction in Eolachesilla Badonnel). Wing 
ciliation, absent in most species of Lachesilla, is shared with Nadleria, but may 
be a plesiomorphous character for the family. 


ACKNOWLEDGMENTS 


We thank Charles W. O’Brien (Florida A & M University, Tallahassee), for the specimen 
of H. venezolanus. We also thank K. K. Giinther (Museum fiir Naturkunde der Humboldt- 
Universitat zu Berlin) for loan of the holotype of H. bogotanus. We also thank Lauren Brown, 
Scott Sakaluk, and two unknown readers for critical reading of the manuscript resulting in 
useful comments. 


LITERATURE CITED 


Badonnel, A. and A. N. Garcia Aldrete. 1979. Nadleria, un nouveau genre de Lachesillidae 
(Psocoptera) du Brésil. Nouv. Rev. Entomol. 9: 3-8. 

Enderlein, G. 1903. Zur Kenntniss amerikanischer Psociden. Zool. Jahrb. Abt. f. Syst. 18: 351- 
364. 

Mockford, E. L. and D. M. Sullivan. 1986. Systematics of the graphocaeciliine psocids with a 
proposed higher classification of the family Lachesillidae (Psocoptera). Trans. Amer. Entomol. 
Soc 112: 1-80. 

Smithers, C. N. 1972. The Classification and Phylogeny of the Psocoptera. Memoirs Australian 
Museum, 14: 1-349. 

Smithers, C. N. 1990. Keys to the Families and Genera of Psocoptera (Arthropoda: Insecta). Tech- 
nical Reports Australian Museum, 2: 1-82. 


Vol. 108, No. 2, March & April, 1997 107 


DESCRIPTIONS OF THE FEMALE, NYMPH, EGG 
AND REDESCRIPTION OF THE MALE OF 
AMPHINEMURA MOCKFORDI 
(PLECOPTERA: NEMOURIDAE)! 


Charles H. Nelson2 


ABSTRACT: The adult female, nymph, and egg of Amphinemura mockfordi are described and 
illustrated for the first time. The male, previously known only from the holotype and paratype male, 
is re-described in greater detail. Characters serving to distinguish this species from closely related 
congeners A. delosa and A. nigritta are also provided. 


Amphinemura mockfordi (Ricker) is one of several closely related species 
that comprise the Amphinemura nigritta complex of eastern North America 
(Ricker, 1952). Presently, this species is known from only two males collected 
at an unspecified location in Monteagle, TN. Recently, the female, pre-emer- 
gent nymph, and egg of A. mockfordi were collected by me from the area of the 
type locality and are described below. To further facilitate identification of this 
species, the male is more fully described as well. Terminology is after Baumann 
(1975) and Stewart and Stark (1988). 


Amphinemura mockfordi (Ricker) 
(Figs 1-15) 


Nemoura (Amphinemura) mockfordi Ricker, 1952, holotype male, paratype male, Monteagle, 
Tennessee. 


Male. Macropterous. Body length 5.3 - 7.9 mm. Forewing length 7.5 - 8.5 mm. General 
body color brown. Epiproct in dorsal view rounded triangular (Figs. 1, 5), in lateral view with a 
narrow base expanding to greatest width at midline and then tapering to a rounded tip (Figs. 2, 6). 
Dorsal sclerite of epiproct with dorsal surface lightly sclerotized, covered for most of its distance 
with hair-like spinulae, lateral surface, below the narrow sclerotized arms that extend nearly to 
tip, bearing short stout spinulae, and tip exhibiting large stout spinulae (Fig. 14). Keel-like ven- 
tral sclerite in ventral view spear-point shaped (Fig. 3), bearing numerous large spines on ante- 
rior two-thirds (Figs. 2, 3). Paraprocts divided into 3 lobes (Figs. 4, 7): inner lobe of each para- 
proct narrow, tip bilobed; middle lobe with broad base narrowing to a bluntly rounded membra- 
nous tip bearing 5 - 6 stout spines, posteroventral sclerotized portion with 2 - 3 stout spines 
apically; outer lobe with large broad base tapering abruptly to become long and narrow and then 
abruptly expanding apically to form a boot-shaped tip, bearing 7 - 9 stout spines. Basal sclerites 
triangular with apices meeting the basal lateral corners of the epiproct (Figs. 5, 6). 


1 Received June 19, 1996. Accepted September 9, 1996. 


2 Department of Biological and Environmental Sciences, 615 McCallie Avenue, University of Ten- 
nessee at Chattanooga, Chattanooga, TN 37403. 


ENT. NEWS 108(2): 107-112, March & April, 1997 


108 ENTOMOLOGICAL NEWS 


Female. Macropterous. Body length 7.4 - 9.4 mm. Forewing length 8.6 - 9.8 mm. Gen- 
eral body color brown. Seventh sternite produced posteriorly, with broadly rounded poste- 
rior margin. Subgenital plate narrow, divided into four posteriorly produced vaginal lobes: 
inner two lobes broad and with posterior margin rounded, outer two lobes thin and nipple- 
like (Fig. 8). 

Nymph. Body length 6 - 6.5 mm. Two anterior thoracic gills on each side of ventral 
cervical region, each gill with 6 - 7 branches. Pronotum with lateral margins bearing a fringe 
of bluntly pointed bristles. Mesonotum and metanotum with medium sized bristles near an- 
terior margins and anterolateral corners. Mesonotum also with a patch of medium to long 
bristles located along anterior margin of wingpad base (Fig. 9). Abdominal segments sparsely 
covered with short bristles, with short to medium bristles, interspersed occasionally with 
longer hairs, forming a posterior marginal fringe (Figs. 10 - 12). Cercal segments with whorls 
of long spines along apical margins, spines of mesal segments about one third to one half 
length of segment, intercalary hairs becoming longer and more numerous toward apical seg- 
ments (Fig. 13). 

Egg. Length and width 0.34 - 0.39 mm. General shape spherical. Chorionic surface 
smooth (Fig. 15). 

Specimens Examined. TENNESSEE: Grundy Co., nr. Monteagle, small seep across Rt. 
41 approx. 0.7 mi NE ject Rts. 64, 41A and 41, C. H. Nelson, 29-IV-1993, 2 males; 5-V-1993, 
1 male, 1 female; 7-V-1993, 6 males, 7 females; 6-V-1996, | nymph, 1 exuvium; 10-V 1996;1 
male; 16-VI-1996; | female. 


DISCUSSION 


Amphinemura mockfordi is very similar to A. nigritta (Provancher) and A. 
delosa (Ricker). A. mockfordi males can be distinguished most easily by the 
apically expanded outer lobe of the male paraproct that in profile appears boot- 
shaped, whereas, in A. nigritta and A. delosa the sclerotized apical region of the 
outer lobe is a narrow band. The A. mockfordi female subgenital plate with its 
four vaginal lobes most closely resembles that of A. delosa. In general the pos- 
terior margins of the inner lobes of A. mockfordi exhibit a somewhat more 
rounded appearance than those of A. delosa. However, there is some overlap in 
the range of variation of this feature as some specimens of A. mockfordi have a 
more truncated posterior margin. The nymph of A. mockfordi is distinguished 
from this stage in A. nigritta and A. delosa by the paucity of long hairs compris- 
ing the marginal fringe of the abdominal segments. 

A. mockfordi is an inhabitant of small headwater streams or seeps of the 
Cumberland Plateau. Other stonefly species collected with this species were 
Amphinemura nigritta, Ostrocerca truncata (Claassen), Leuctra biloba Claassen, 
Leuctra rickeri James, and Diploperla robusta Stark and Gaufin. 


Vol. 108, No. 2, March & April, 1997 109 


Figs. | - 8. Amphinemura mockfordi, adult terminalia. 1. Epiproct, dorsal. 2. Epiproct, lateral. 3. 
Epiproct, ventral. 4. Ventral aspect of male right paraproct showing inner (il), middle (ml), and 
outer lobes (ol). 5. Male terminalia, dorsal. 6. Male terminalia indicating basal sclerite (bs), lateral. 
7. Male terminalia, ventral. 8. Female terminalia, ventral. Scale lines = 0.5 mm. 


110 ENTOMOLOGICAL NEWS 


an mn 


t M4 fp jt tt-|> 


4 


me fis 
Uj 


vy! 5 
427/54 
7 {ated 


Figs. 9 - 13. Amphinemura mockfordi, nymph. 9. Habitus, dorsal. 10. Male terminalia, dorsal. 11. 
Male terminalia, lateral. 12. Male terminalia, ventral. 13. Right cercus, dorsal. Scale lines = 1 mm. 


Vol. 108, No. 2, March & April, 1997 111 


Soum 669690 


Figs. 14 - 15. Amphinemura mockfordi, scanning electron micrographs. 14. Epiproct apex, biased 
dorsal. 15. Egg. 


112 ENTOMOLOGICAL NEWS 


ACKNOWLEDGMENTS 


I thank Ross Durham and Gene Van Horn for their critical reading of earlier versions of 
this manuscript. 


LITERATURE CITED 


Baumann,. R. W. 1975. Revision of the stonefly family Nemouridae (Plecoptera): a study of the 
world fauna at the generic level. Smithson. Contrib. Zool. 211: 1 - 74. 

Ricker, W. E. 1952. Systematic Studies in Plecoptera. Indiana University Publications Science 
Series 18: | - 200. 

Stewart, K. W. & B. P. Stark. 1988. Nymphs of North American stonefly genera (Plecoptera). 
Thomas Say Found., Entomol. Soc. Amer. 12, 460 pp. 


Vol. 108, No. 2, March & April, 1997 113 


FIRST ADULT DESCRIPTION OF THE UNUSUAL 
BAETID MAYFLY GENUS ECHINOBAETIS 
(EPHEMEROPTERA: BAETIDAE)! 2 


C. R. Lugo-Ortiz, W. P. McCafferty3 


ABSTRACT: The female adult of the Indonesian species Echinobaetis phagas (Ephemeroptera: 
Baetidae) is described. The adult of Echinobaetis has not been known previously. It is distinct from 
other baetid adults in that the forewings have an R3 sector, which is formed by the basal union of 
R3q and R3p, multiple marginal intercalaries between MP and A, and a CuA that is attached to CuP. 
In addition, the pronotum has a deep, V-shaped anteromedian emargination, and the meso- and 
metasternal sclerites are ventrally extended. 


Mol (1989) erected the genus E’chinobaetis (Ephemeroptera: Baetidae) for 
E. phagas Mol, a species from Sulawesi (formerly Celebes) known from the 
larvae only. The genus has been notable because the larvae have depressed, 
peripherally expanded heads with prognathous mouthpart orientation [Mol 
(1989): Figs. 1, 2], triads of elongate, posteriorly oriented spines on terga 1-8 
[Mol (1989): Figs. 22, 23}, and mouthparts modified for predation [Mol (1989): 
Figs. 3-17]. Although highly atypical of baetid mayflies in general, this “flat- 
headed” small minnow mayfly does possess the unique baetid epicranial suture 
with lateral branches anterior to the lateral ocelli [Wang and McCafferty (1996): 
Fig. 6] and ventral orientation of the dorsal apical lobe of the femora. These 
latter characteristics distinguish all baetid larvae. 

In our recent examination of baetid material from Sulawesi, we discovered 
the adult of E. phagas. Although female, it, like the larvae, is highly unusual for 
the family and therefore deserving of this special report. The material examined 
is housed in the Purdue Entomological Research Collection, West Lafayette, 
IN. 


Echinobaetis phagas Mol, 1989 


Female adult. Body length: 10.6 mm; wing length: 9.7 mm; caudal filaments length: 
25.3 mm. Head (Figs. 1, 2): Face pale brown to medium brown. Vertex with medium brown 
fleur-de-lis marking. Flagella, scapes, and pedicels whitish dorsally, pale brown ventrally. 
Compound eyes black. Thorax: Pronotum (Fig. |) pale brown to medium brown, with deep, 
V-shaped anteromedian emargination. Meso- and metanota dark brown. Sterna cream. Meso- 
and metasternal sclerites (Fig. 3) enlarged, extending ventrally. Legs cream. Midfemora (Fig. 
4) with two triangular processes distally. Tarsi each with ovoid claw and acute claw. Fore- 


! Received July 6, 1996. Accepted August 6, 1996. 
2 Purdue Agricultural Research Program Journal No. 15117. 
3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 


ENT. NEWS 108(2): 113-116, March & April, 1997 


114 ENTOMOLOGICAL NEWS 


wings (Fig. 5) dark brown between C and Ry and in most crossveins; 2.71x longer than 
wide; single marginal intercalary between Rj and Rj; area between R2 and MP with elon- 
gate double marginal intercalaries, except between IMA and MAg with three; area between 
MP and A with two, three, four, and five marginal intercalaries of various lengths, mostly 
elongate; R3,g and R3p basally attached, forming a fork; CuA basally attached to CuP. Hind- 
wings absent. Abdomen: Terga dark brown. Larval spine remnants on terga 1-8, barely visi- 
ble. Gill socket remnants on abdominal segments 1-7. Sterna medium brown to yellow-brown. 
Caudal filaments dark brown, turning cream distally. 


Material examined. SULAWESI, Utura Prov., Toraut R, Dumoga-Bone Nat. Park, 211 
m, 0°44°N, 123°54 E, IX-3-1985, J. T. and D. A. Polhemus, female adult (forewing mounted 
on slide). 


DISCUSSION 


The specimen upon which the above description is based was not reared. 
However, we are confident that it belongs to E. phagas because it possesses 
vestiges of the three spines that are present on terga 1-8 of the larvae as well as 
possessing spiny gill socket remnants on abdominal segments 1-7. Both char- 
acteristics are diagnostic of the species. Also, the head shape (Figs. 1, 2) is 
indicative of the nature of the larval head. The adult was collected from the 
same general area and season as the original material described by Mol (1989). 

The adult of E. phagas possesses characteristics that are unique or highly 
unusual among the Baetidae. Wings are narrow-elongate, with atypical vena- 
tion. A striking venational characteristic is the presence of an R3 sector in the 
forewings (Fig. 5). This sector, formed by the basal union of R3, and R3p, is 
not present in other baetid adults, and may possibly represent an ancestral con- 
dition within Baetidae because it is present in Siphlaenigmatidae [see Penniket 
(1962): Fig. 1], some members of the Heptagenioidea [see McCafferty (1991): 
Figs. 26, 28], and some Siphlonuroidea [see McCafferty (1991): Fig. 24]. 

The union of CuA to CuP (Fig. 5) is another unusual characteristic for 
Baetidae. Only one other baetid genus, Centroptiloides Lestage, has CuA at- 
tached to CuP [see Crass (1947): Fig. 29a]. This condition is variously wide- 
spread throughout the Ephemeroptera and is highly subject to homoplasy. Thus, 
even within Baetidae, Centroptiloides and Echinobaetis clearly belong to dis- 
parate phyletic clades. 

The presence of multiple, elongate marginal intercalary veins in the fore- 
wing (Fig. 5) is of considerable additional interest. All other baetid genera have 
relatively short marginal intercalaries. The relatively large number of intercalaries 
in each space (up to five), particularly between MP and A, and their retention 
appears related to the lengthening of the cubito-anal area, which is the major 
contributor to elongation of the forewing. 

Despite the unusual and distinctive venation of the forewings of Echinobaetis, 


Vol. 108, No. 2, March & April, 1997 115 


—— = TREN SSE op ” 


aa, 
Sry, ee 


Figs. 1-5. Echinobaetis phagas, female adult. 1. Head and pronotum. 2. Head (lateral). 3. Meso- 
and metasterna (pointers towards meso- and metasternal sclerites). 4. Left midfemur (pointers 
towards distal processes). 5. Right forewing. 


116 ENTOMOLOGICAL NEWS 


adults are still readily identifiable as Baetidae due to the basally detached IMA, 
MA2, IMP, and MP2, and three-segmented mid- and hindtarsi. Also, based on 
larvae, the male adults should have turbinate eyes, although it has been recently 
shown that this is not a universal characteristic of Baetidae (Lugo-Ortiz and 
McCafferty 1996). 

The deep anteromedian emargination of the pronotum (Fig. | ) and enlarged, 
ventrally extended meso- and metasternal sclerites (Fig. 3) of E. phagas are not 
present in other baetid adults. We consider them additional apomorphies defin- 
ing Echinobaetis. 

The only other flatheaded baetid mayfly genus known at present is Jubabaetis 
Miiller-Liebenau (1980). It is known from larvae only of one species from the 
Philippines. It is apparently related to Echinobaetis, and therefore its adults 
may demonstrate some of the adult characteristics that we have established for 
Echinobaetis. In any case, adults of J. pescadori Miiller-Liebenau should be 
readily distinguished from E. phagas because J. pescadori will possess small 
hindwings and should have a single tergal spine vestige on abdominal segments 
1-9, while E. phagas lacks hindwings and has three tergal spine vestiges on 
abdominal segments 1-8. 


ACKNOWLEDGMENTS 


The specimen upon which this study was based was collected by J. T. and D. A. Polhemus 
with support from a research grant from the National Geographic Society. The specimen was 
donated to the Purdue Entomological Research Collection by G. F Edmunds, Jr. (Salt Lake 
City, Utah). 


LITERATURE CITED 


Crass, R. S. 1947. The may-flies (Ephemeroptera) of Natal and the eastern Cape. Ann. Natal Mus. 
11: 37-110. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. Aturbina georgei gen. et sp. n.: a small minnow 
mayfly (Ephemeroptera: Baetidae) without turbinate eyes. Aq. Insects. 18: 175-183. 

McCafferty, W. P. 1991. The cladistics, classification, and evolution of the Heptagenioidea 
(Ephemeroptera). Pp. 87-102. Jn: Overview and strategies of Ephemeroptera and Plecoptera (J. 
Alba-Tercedor and A. Sanchez-Ortega, eds.). Sandhill Crane Press, Florida. 

Mol, A. W. M. 1989. Echinobaetis phagas gen. nov., spec. nov., a new mayfly from Sulawesi 
(Ephemeroptera: Baetidae). Zool. Medede. 63: 61-72. 

Miiller-Liebenau, I. 1980. Jubabaetis gen. n. and Platybaetis gen. n., two genera of the family 
Baetidae from the Oriental region. Pp. 103-114. Jn: Advances in Ephemeroptera biology (J. F. 
Flannagan and K. E. Marshall, eds.). Plenum Publishing Corp., New York. 

Penniket, J. G. 1962. Notes on New Zealand Ephemeroptera. III. A new family, genus and species. 
Rec. Canterbury (NZ) Mus. 8: 389-398. 

Wang, T.-Q. and W. P. McCafferty.1996. New diagnostic characters of the mayfly family Baetidae 
(Ephemeroptera). Entomol. News. 107: 207-212. 


Vol. 108, No. 2, March & April, 1997 117 


MATING BEHAVIOR OF PELECINUS POLYTURATOR 
(HYMENOPTERA: PELECINIDAE)! 


A.P. Aguiar2 


ABSTRACT: The mating behavior of Pelecinus polyturator is described and briefly discussed. It is 
suggested that the mating behavior of the male P. polyturator may have evolved in response to the 
female’s highly specialized gaster. 


In spite of its usually large size and peculiar morphology, Pelecinus poly- 
turator (Drury) has not attracted much attention and remains poorly known, 
notably with respect to its biology and behavior. This species is better known as 
a parasite of Scarabaeidae larvae (Davis 1919; Hammond 1944), but Clausen 
(1940) believes that the true hosts are probably Coleoptera larvae living in 
decaying wood. Hammond (op.c.) observed larval and pupal stages on Phyllo- 
phaga, and discussed the economic importance of this parasitism, and Lim et 
al. (1980) described the pupa. The development is internal, and pupation takes 
place outside the host (Clausen, op.c.). Adult males are extremely rare in North 
America, where females are believed to be mainly parthenogenetic (Brues 1928; 
Young 1990). In the Neotropics, however, males seem to be as common as 
females (Masner 1995). In any case, P. polyturator is not commonly collected, 
and no rearing technique is known for this species, making biological studies 
about it a task usually difficult to achieve. This study was performed after the 
collection of a male and female in the Atlantic Forest of southern Brazil (Lapa, 
Parana State, 25°46°S 49°44°W) on March 14, 1991. Both the male and female 
were placed in a 400ml jar and observed during the entire mating process, and 
an additional 20 minutes. Each was then independently mounted as similarly as 
possible to its respective observed copulatory posture, as reference material for 
illustrations. 


MATING BEHAVIOR 


The male mounted the female immediately after they were put together, 
grasping her first gastral segment with the apex of his fore tibiae (fig. 1). The 
basitarsal antennal brush and apical spurs apparently helped the male grip the 
female’s gaster. The male vibrated his wings in brief periods of less than one 
second, at intervals of 1-4 seconds, throughout the mating process. He vibrated 


1 Received April 22, 1996. Accepted May 31, 1996. 


2 Museum of Biological Diversity, Dept. of Entomology, 1315 Kinnear Rd., Columbus, OH 43212- 
1192. 


ENT. NEWS 108(2): 117-121, March & April, 1997 


118 ENTOMOLOGICAL NEWS 


his antennae, in a simple and steady alternate pattern, touching the female with 
them. The female assumed a passive position at this time, with antennae show- 
ing no movement, wings partially retracted, and thorax and gaster stationary, 
almost touching the substratum. After 1.5 minutes, the female touched the male 
gaster with her own, by elevating it gently. The male reacted by sliding back- 
wards along the female gaster, while still holding her with his tibiae. The tip of 
his gaster also slid over the female’s until the two tips met. The hook-like male 
genitalia was then exerted. Soon after copulation started (fig. 2), the male bent 
his gaster downwards, while the female slightly arched hers upwards. This first 
copulation ended 15 seconds later but the male kept holding the female while 
vibrating his wings and antennae. After a few seconds, the male dismounted, 
walked briefly around, and remounted the female. A second copulation took 
place 30 seconds after the first one, with male and female showing the same 
behavior. During the interval between the two copulations, the female kept her 
gaster laying sideways on the substratum curved in a “C” shape, and was appar- 
ently in a very relaxed state. After three more minutes of courtship, the female 
lashed energetically at the male with her gaster, causing an immediate dismount. 
The female then entered into a strongly quiescent state, laying sideways, com- 
pletely motionless, on the substratum during a period of three minutes, after 
which she recovered totally and started to fly inside the jar. The entire mating 
process lasted five minutes. During the following 20 minutes no more trials for 
copulation were observed, and the experiment was terminated. 


DISCUSSION 


Mounting in P. polyturator is singular among Apocrita Hymenoptera in that 
the male does not mount the female on her thorax as usually observed (e.g., 
Assem etal. 1980a, 1980b, and Assem & Povel 1973 for Pteromalidae, Camargo 
1972 for Apidae, Mertins 1980 for Bethylidae, Michener 1948 for Ants, and 
Vinson 1972 for Ichneumonidae). Nonetheless, gastral mounting in P. polyturator 
may improve efficacy in copulating, since the two genitalia are approximately 
40% closer than they would be in a head-over-head mounting, making it easier 
and/or quicker for the male to access the female’s genitalia. 

Wing and antennal vibration during courtship seems to be an almost univer- 
sal behavior of courting male Hymenoptera (Matthews 1975), though the latter 
can usually show considerable variation among Hymenoptera (e.g., Assem & 
Povel 1973). For P. polyturator, however, both antennal and wing vibration fol- 
low a very simple pattern, indicating a generalized courtship behavior. 

In spite of the female’s inactivity during most of the time, she seems to play 
a decisive role in starting (and ending, see further below) copulation, since the 
male slid backwards to copulate only after the female touched his abdomen 
with hers, apparently signaling her willingness. The need of an acceptance sign 


Vol. 108, No. 2, March & April, 1997 


\ ZY 
\A Os 
“ 
, \. id 
SS 
ape 0}, x Ss = 
ye Hoe i, WAH <I 
SSS: zoe Sinst Se = aod 


a ees : ; - ae r ] 


\ EE \ 
Lae \ J! 

VA —N B a of IW Se 
OC —— oss ¥ 
: RW aN ; — = a 
= a \ lice — 
bay AE == aes = : SSN SN 
Sey 2a rere! ee eet a, 


Mating behavior of Pelecinus polyturator. Fig. 1. Mounting. Fig. 2. Copulation 


119 


120 ENTOMOLOGICAL NEWS 


appears also in a number of Hymenoptera species, albeit it can be expressed in 
a variety of forms. In several species of Pteromalidae for example, the female 
signals by a tight flagellar and “chin” retraction (Assem & Povel 1973; Assem 
et al. 1980a). 

The possibility of the female simply retracting her gaster to copulate with 
the male mounted on her thorax (for example by bending it upwards, reducing 
its length by curving it in a bell-shaped fashion, retracting it through telescop- 
ing, or even combining these possibilities) may be a simple evolutionary solu- 
tion to the problem, but some important flexibility restrictions apply to the female 
gaster of P. polyturator. Mason (1984) discussed the complex mechanics of the 
gastral articulation in this species; the articulation between gastral segments 1 
and 2 allows only a vertical movement, and the posterior segment cannot 
deflect upwards by more than 10°-20°. Rotary and vertical movements are also 
restricted in the subsequent segments. In fact, the muscular and sclerotic struc- 
ture shows a high degree of specialization to a downwards action, used for 
digging the soil. Thus, if the restricted flexibility of the gaster, crucial for dig- 
ging the soil for larval hosts, was more valuable to the survival of the species 
than a more flexible gaster, useful for an “easier” mating, then P. polyturator 
must have had to overcome the “longer female’s gaster” problem in an alterna- 
tive way. The more immediate solution may have been a more active role of the 
male, by simply sliding backwards to copulate. Selection favoring males using 
the front legs for grasping the female’s gaster can easily follow this sequence, 
since this behavior helps reduce the chances of losing the female, especially 
during the male’s backward slide. 

The short copula suggests a great availability of receptive females, since 
more prolonged copulations tend to occur only when the probability of acquir- 
ing multiple mates is low (Thornhill & Alcock 1983). This apparently implies 
that females of P. polyturator are more numerous than males in the area where 
the specimens were collected. However, the availability of receptive females 
may also be increased by other factors, as male/female efficiency in locating 
each other, and female willingness in copulating with more than one male. 
Inferences on sex ratio are, therefore, premature. 

The firm refusal of the male in trying to copulate again after being lashed by 
the female gaster strongly indicates that this behavior may serve as a sign for 
the end of the mating. The succeeding quiescent state of the female was also 
observed for Tetrastichus incertus (Eulophidae, Miller 1966) and Laelius pedatus 
(Bethylidae, Mertins 1980), but not so intense as here observed for P. polytura- 
tor. Its significance is unknown. 

In conclusion, it can be suggested that most of the current mating behavior 
of P. polyturator may have evolved in response to the female’s acquisition of a 
highly specialized gaster. 


Vol. 108, No. 2, March & April, 1997 121 


ACKNOWLEDGMENTS 


My sincere thanks to Linda J. Mason, Clifford Sadof, John F. MacDonald, Luciana 
Musetti, John Wenzel, and two anonymous reviewers, for valuable suggestions, and to Arwin 
V. Provonsha for preparing the illustrations. 


LITERATURE CITED 


Assem, J. van den, F. Jachmann and P. Simbolotti. 1980a. Courtship behaviour of Nasonia 
vitripennis (Hym., Pteromalidae): Some qualitative, experimental evidence for the role of phero- 
mones. Behaviour, 75:301-307. 

Assem, J. van den, M. J. Gijswijt and B. K. NYbel. 1980b. Observations on courtship - and 
mating strategies in a few species of parasitic wasps (Chalcidoidea). Neth. J. Zool., 30 (2):208- 
221. 

Assem, J. van den and G. D. E. Povel. 1973. Courtship behaviour of some Muscidifurax species 
(Hym., Pteromalidae): A possible example of a recently evolved ethological isolating mecha- 
nism. Neth. J. Zool., 23 (4):465-487. 

Brues, C. T. 1928. A note on the genus Pelecinus. Psyche, 35 (4):206-209. 

Camargo, C. A. de. 1972. Mating of the social bee Melipona quadrifasciata under controlled 
conditions (Hymenoptera, Apidae). J. Kans. Entomol. Soc., 45 (4): 520-523. 

Clausen, C. P. 1940. Entomophagous Insects. McGraw-Hill Book Company, Inc. New York and 
London. 688 pp. 

Davis, J. J. 1919. Contributions to a knowledge of the natural enemies of Phyllophaga. Ill. Nat. 
Hist. Surv. Bull. 13, art. 5, pp. 86-87. 

Hammond, G. H. 1944. Economic importance and host relationsilip of Pelecinus polyturator Drury. 
Can. Entomol., 76:130. 

Lim, K. P., W. N. Yule and R. K. Stewart. 1980. A note on Pelecinus polyturator (Hymenoptera: 
Pelecinidae), a parasite of Phyllophaga anxia (Coleoptera: Scarabaeidae). Can. Entomol., 112 
(2):219-220. 

Masner, L. 1995. The proctotrupoid families, pp. 209-246. Jn: P. E. Hanson and I. D. Gauld (eds). 
The Hymenoptera of Costa Rica. Oxford University Press, The Natural History Museum, Lon- 
don. XX+893pp. 

Mason, W. R. M. 1984. Structure and movement of the abdomen of female Pelecinus polyturator 
(Hymenoptera: Pelecinidae). Can. Entomol., 1 16:419-426. 

Mathews, R. W. 1975. Courtship in parasitic wasps. /n Symposium on evolutionary strategies of 
parasitic insects and mites, ed. P. Price. New York, Plenum Press, pp. 66-86. 

Mertins, J. W. 1980. Life history and behaviour of Laelius pedatus, a gregarious bethylid 
ectoparasitoid of Anthrenus verbasci. Ann. Entomol. Soc. Am., 73 (6): 687-693. 

Michener, C. D. 1948. Observations on the mating behaviour of harvester ants. J. N. Y. Entomol. 
Soc., 56 (4):239-242. 

Miller, M. C. 1966. Emergence and mating of Tetrastichus incertus, a parasite of the alfalfa weevil. 
J. Econ. Entomol., 59 (6): 1532-1534. 

Thornhill, R. and J. Alcock. 1983. The Evolution of the Insect Mating Systems. Harvard Univer- 
sity Press. 547 pp. 

Vinson, S. B. 1972. Courtship behaviour and evidence for a sex pheromone in the parasitoid 
Campoletis sonorensis Hymenoptera: Ichneumonidae). Environ. Entomol., 1 (4):409-413. 
Young, D.K. 1990. Distribution of Pelecinus polyturator in Wisconsin (Hymenoptera: Pelecinidae), 
with speculations regarding geographical parthenogenesis. Gt. Lakes Entomol., 23 (1): 1-4. 


122 ENTOMOLOGICAL NEWS 


THE RATE OF SPECIES DESCRIPTIONS 
IN ODONATA! 


K. J. Tennessen2 


ABSTRACT: The rate of new species descriptions of Odonata over the last 150 years yields an 
essentially straight line, indicating that many species are yet to be discovered within the Order. 
More than 5,300 species are now known, and the rates of description in the suborders Anisoptera 
and Zygoptera have been relatively equal. However, a decline in the number of new species appear- 
ing in the three largest families over the last six decades, despite an increasing number of authors, 
indicates that the Odonata are now at least half known and that fewer than 10,000 species exist 
worldwide. 


Examining the trend curve of the rate at which species have been described 
indicates whether nearly all species in a particular group are known or whether 
many remain to be discovered. For example, Steyskal (1965) found that the 
curve for the butterflies of North America has leveled off, indicating that alpha 
taxonomy for this group is essentially complete. By contrast, the curves for 
fleas, wasps, and mosquitoes show that many species still are undescribed. The 
curves for Aedes mosquitoes (Zavortink 1990) and the stoneflies of the world 
(Steyskal 1976) show no slowing in descriptive rate. In this paper, I present the 
status of the trend curve for the world Odonata, an Order generally purported to 
be well-known. 


METHODS 


Using Bridges’ catalogue (1993), I tallied the number of valid species 
described in each decade, beginning with Linnaeus in 1758. Subspecies, forms 
and known synonyms were omitted from the counts. A decade was defined as 
extending from January | of the zero year to December 31 of the ninth year 
(e.g., the 1980s began on Jan. 1, 1980 and ended on Dec. 31, 1989). Exceptions 
were made for the first and last decades: the 1750s, from Jan. 1, 1758, to Dec. 
31, 1759 (2 years); and the 1990s, from Jan. 1, 1990 to Dec. 31, 1994 (5 years). 
I derived the number of descriptions appearing after Bridges (1993) by search- 
ing literature and abstracts. 

RESULTS 


Approximately 5,300 valid species have been described from Linnaeus’ time 


through 1994. A significant increase in the rate of Odonata descriptions did not 
begin until 70 years after Linnaeus (Fig. 1). Since then, the rate has been fairly 


! Received June 11, 1996. Accepted August 1, 1996. 
2 1949 Hickory Ave., Florence, AL 35630. 


ENT. NEWS 108(2): 122-126, March & April, 1997 


Vol. 108, No. 2, March & April, 1997 123 


6000 
5000 
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® 4000 
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n ~~ ~ ice) (oe) oe) [0 @) (oe) [o>) aD [op) [o>) fop) 


Figure 1. Trend curve of the number of world Odonata species described per decade since Linnaeus. 


124 ENTOMOLOGICAL NEWS 


constant. The decade of greatest activity so far was the 1930s, when 548 species 
were described. A relatively low number of new species appeared in the 1970s 
(208), indicating that the line might begin to level off. However, in the 1980s, 
the rate increased again. As Steyskal (1965) pointed out, “. . . if the curve 
appears to be leveling off, we have no assurance that we are approaching the 
stage at which all species are known.’ The steeper inclination of the line in the 
first half of the 1990s, during which more than 235 species were described, 
indicates that we are far from nearing completion of odonate species discovery. 

The number of species described to date within the two suborders, Anisoptera 
and Zygoptera, are remarkably similar (about 2,770 and 2,540 species, respec- 
tively). In the first half of the current decade, more Anisoptera (145) have been 
described than Zygoptera (90). a trend opposite that of the 1980s. In the last 
three decades, families in which the most species were described were 
Gomphidae, Coenagrionidae, Libellulidae, Corduliidae, Aeshnidae, Protoneu- 
ridae, Platycnemididae, and Megapodagrionidae, listed in descending order. 

The majority of new species described recently came from tropical areas of 
the world (Neotropical. 92: Oriental. 74; Australasian, 40). The fauna of 
Europe is almost certainly known, as over 95 percent of the European fauna 
was described prior to 1900, and only five new species descriptions have 
appeared since then (see Askew 1988). Knowledge of the Nearctic fauna 
appears to be nearing completion (Fig. 2), although 8 species have appeared 
thus far in the 1990s and a few new species still await description. 

The rate of new species discovery in the three largest families (each with 
approximately 1,000 species known) might be a clue to the status of knowledge 
for the Order as a whole. For example, in the Coenagrionidae and Libellulidae, 
the descriptive rate over the last 4 decades has slowed (Table 1). The future rate 
in Coenagrionidae will probably be higher than in the Libellulidae, however, as 
several colleagues have informed me that many undescribed species of these 
small, taxonomically-difficult damselflies exist in tropical areas. In contrast to 
these two families, the rate in the Gomphidae increased. Reasons that new spe- 
cies continue to appear in this family is that gomphid dragonflies are rather 
local in distribution, have relatively brief adult flight ranges, and are relatively 
secretive in their habits, factors which make them difficult to collect. In sum- 
mary, the declining rate in two of the largest families, coupled with the fact that 
significant numbers of new species are being found in only three of the six 
biogeographic regions, indicates that over half the Odonata are now known. 


Vol. 108, No. 2, March & April, 1997 125 


450 


400 


350 


300 


250 


200 


150 


Cumulative No. of Described Species 


100 


WSS) 
1779 
1799 
1819 
1839 
1859 
1879 
1899 
OTS 
1939 
1959 
USA) 
1994 


Fig. 2. Trend curve of species descriptions for North American Odonata. 


Furthermore, the increasing number of authors has not increased the rate of 
description, although odonatologists are not evenly distributed around the world, 
and there are few or none in many of the tropical countries where diversity is 
greatest. The combined circumstances of these three factors lead me to specu- 
late that over half the Odonata are known and thus there are fewer than 10,000 
species in the world. 

Higher classification within the Order is highly controversial and in need of 
revision. Phylogenetic studies of Odonata have lagged behind studies of other 
aquatic insect Orders. Progress in these two areas will continue to be slow 
because funding for such efforts is difficult to obtain. Most currently active 
Odonata taxonomists expend personal resources to support research at the 
species level and higher, including costs for collecting, curating, analysis, and 
publishing. 


126 ENTOMOLOGICAL NEWS 


Table 1. Change in number of new species descriptions per decade in the three largest families of 
Odonata from 1900 to 1994, and the associated percent increase or decrease in the last five decades 
of 20th century compared with first half of century. 


Gomphidae Libeilulidae Coenagrionidae 

Mean No. New Species 
per Decade (1900 to 1949) 66 70 89 
No. New Species/Decade 
1950-59 i 77 87 
1960-69 55 50 84 
1970-79 67 20 46 
1980-89 83 31 3i7/ 
1990-94 85 19 24 
% Increase (+) or Decrease (—) 
1950-59 -14 +10 -—2 
1960-69 -17 —29 -— 6 
1970-79 +2 -71 48 
1980-89 +26 —56 —58 
1990-94 +29 -73 -73 

ACKNOWLEDGMENTS 


My thanks to Sid Dunkle and Dennis Paulson for reviewing the initial version of the 
manuscript, and two anonymous reviewers for additional comments. 


LITERATURE CITED 


Askew, R. R. 1988. The dragonflies of Europe. Harley, England. 291 pp. 

Bridges, C. 1993. Catalogue of the family-group, genus-group and species-group names of the 
Odonata of the world (second edition). Charles A. Bridges, Urbana. Illinois. 806 pp. 

Steyskal. G. C. 1965. Trend curves of the rate of species description in zoology. Science 149:880- 
882. 

Steyskal, G. C. 1976. Notes on the nomenclature and taxonomic growth of the Plecoptera. Proc. 
Biol. Soc. Wash. 88:408-410. 

Zavortink, T. J. 1990. Classical taxonomy of mosquitoes — a memonial to John N. Belkin. J. Amer. 
Mosq. Control. Assoc. 6(4):593-599. 


Vol. 108, No. 2, March & April, 1997 127 


ACTIVITY AND REPRODUCTION OF CALOSOMA 
FRIGIDUM (COLEOPTERA: CARABIDAE) IN 
NORTHERN MICHIGAN FORESTS! 


Renate M. Snider2, Richard J. Snider 


ABSTRACT: From 1985 to 1991, activity patterns of Calosoma frigidum were assessed by means 
of pit-trapping at weekly intervals from early May to late October in two deciduous forest sites in 
northern Michigan. Females were dissected in order to document ovarian development. The spe- 
cies was spring-breeding, with activity of gravid females peaking in May-June, and with spent 
females present from late May to July and occasionally into August. In 1988, only one of 85 
females was gravid, a phenomenon apparently related to delayed emergence from hibernation. 
Asynchronism between time of emergence and early spring shifts from short-day to long-day (criti- 
cal for vitellogenesis) was postulated as the underlying cause for reproductive failure in 1988. In 
both sites, the species essentially disappeared after 1988. 


Results presented here stem from a long-term ecological study in Michigan’s 
Upper Peninsula. The study was designed to assess potential effects of 
Extremely Low Frequency (ELF) electromagnetic fields, created by the U.S. 
Navy’s ELF antenna, on soil and litter invertebrates. One of the project goals 
was to document activity patterns of forest-dwelling arthropods by pit-trapping 
prior to (1985-1988) and after (1989-1991) antenna activation in two sites: a 
Test site adjacent to the ELF antenna, and a Control site removed from its influ- 
ence (Snider & Snider 1987). Among the Carabidae obtained from 1985 to 
1991, Calosoma frigidum Kirby exhibited a reproductive anomaly. This anomaly 
occurred in 1988, the year prior to ELF antenna activation, and it occurred in 
both sites; i.e., it was unrelated to potential effects of electromagnetic fields. 


METHODS 


Both sites were located in hardwood forest dominated by Acer saccharum 
Marsh, and were divided into 20 (10 x 10 m) quadrats. Twenty pit-traps were 
installed per site, one each in the same corner of each sampling quadrat (site 
descriptions, sampling design and quadrat layout are given in Snider & Snider 
1987). Traps were made of opaque plastic and consisted of a permanently 
installed outer cup (9 cm diameter), and a funnel which snapped into its rim and 
led to an inner collection cup. When traps were not in use, only the outer cup, 
covered with a tight-fitting lid, remained in the field. Four 1 m lengths of green 
plastic garden edging, protruding approx. 8 cm above ground and abutting to 


1 Received July 29, 1996. Accepted August 24, 1996. 
2 Department of Zoology, Michigan State University, East Lansing, Michigan 48824. 


ENT. NEWS 108(2): 127-133, March & April, 1997 


128 ENTOMOLOGICAL NEWS 


the trap at 90° angles were used as barriers. At intervals of | week from early 
May to late October, inner collection cups partly filled with ethylene glycol 
were inserted at dusk and collected at dawn (= night catch), replaced, and col- 
lected again the following dusk (= day catch). Specimens were rinsed and stored 
in 95% ethyl alcohol. Intact females of C. frigidum (i.e., those which had not 
been cannibalized while in the trap) were later dissected, mature ova were 
counted, and the developmental state of each specimen was classified as fol- 
lows: T = teneral females, ovaries undeveloped, integument soft; W = females 
with well-developed, slightly distended ovaries, mature ova absent; G = gravid 
females, mature ova present; S = spent females. Results span seven seasons, 
from 1985 through 1991. 


RESULTS AND DISCUSSION 


Basic information on annual catch size, diurnality, and sex ratios of C. 
frigidum is summarized in Table 1. In both sites, activity-density as measured 
by total catches increased toward a peak in 1987, declined in 1988, and abruptly 
decreased to near-zero thereafter. In 1987, peak populations of C. frigidum, 
which feeds on a variety of defoliating insect larvae (Sanders & Frankenhuyzen 
1979; Crins 1980; Burgess & Collins 1917; Gidaspow 1959), coincided with 
particularly severe defoliation of sugar maples and other tree species. Site-spe- 
cific data on defoliators were not obtained. In the area at large, however, densi- 
ties of Bruce’s spanworm (Operophtera bruceata (Hulst)), fall cankerworm 
(Alsophila pometaria (Harris)), linden looper (Erannis tiliaria [Harris]), and 
forest tent caterpillar (Malacosoma disstria Hiibner) were recorded as unusu- 
ally high; large numbers of active Calosoma sp. were noted as well (Depart- 
ment of Natural Resources, Dr. Robert Heyd, pers. comm.). Within Test and 
Control sites, beetles were conspicuously active on the litter surface in 1987, 
indicating that the large numbers trapped (Table 1) were partly due to increased 
activity. Crins (1980) also reported high numbers and increased mobility of 
C. frigidum in maple-dominated forests in Ontario during a year of severe 
defoliation. 

Observations on other Calosoma species may aid interpretation of 1988 
data on C. frigidum. Male C. sycophanta L. become less active when prey is 
scarce (Weseloh 1985); a reduced proportion of males in trap catches would be 
the result, as observed in C. frigidum (Tabie 1). Reduced prey availability also 
diminishes C. sycophanta mobility in general (Weseloh 1985), and may have 
been instrumental in the 1988 decline in C. frigidum catches (Table 1), despite 
normal reproduction in 1987 (discussed below). In addition, severe rainfall 
deficiency and rapidly decreasing soil moisture from May through July 1988 
(Snider & Snider 1995) probably had negative effects on activity-density of C. 
frigidum (Epstein & Kulman 1990; Martel et a/. 1991). Drought, rather than 
prey density (as observed in C. calidum Fabricius by Jeffords & Case [1987]), 


Vol. 108, No. 2, March & April, 1997 129 


may also have influenced diel activity patterns (Table 1). 

Pronounced between-year fluctuations in carabid numbers have been docu- 
mented for other species as well (Desender & Alderweireldt 1990). However, 
the extreme scarcity of C. frigidum after 1988 requires further explanation. Basic 
patterns of activity and reproduction are evident in 1985-1987 data. The species 
was Clearly spring-active (Fig. 1) and spring-breeding (Fig. 2 shows combined 
Test and Control site data on the developmental state of females). Two teneral 
females were captured in the first week of May 1987, indicating that a small 
proportion of larvae may overwinter and mature in early spring. A few indi- 
viduals with slightly distended ovaries were occasionally present at the same 
time. One to two weeks later, virtually all females were gravid (Fig. 2), and 
their activity peaked mid-May to early June. On average, females carried 
approximately 14 to 17 mature ova (Table 2). Although these data are in agree- 
ment with an estimated 17 eggs per female produced in laboratory culture (Bur- 
gess and Collins 1917), actual fecundity was probably higher: in many females 
carrying 20-30 fully mature ova, ovarioles in intermediate stages of develop- 
ment were also present. A few spent females were trapped from late May to 
July, and occasionally in mid- or late August (data beyond the tenth trapping 
week not shown in Figs. | and 2). 


250 4 
1987 
a 
nae MMM TEST 
= | MMM CONTROL 
ao 
= 
ao 
& 150 
a 
= 
= 
= ae 
= 1988 
= ‘ 
O N 
E 1985 1986 
50 i 
\ 
Nh 
anK vii 1989 
, ARK vil 
0 Me yA wae Nh 
j aan 
5/7 /6 5/5 5/10 5/8 


TRAPPING DATES 


Fig. 1. Weekly catches of C. frigidum in Test and Control sites, from the first trapping date in May 
(arrows) to the 10th week (early July) of 1985 through 1989; later dates and years omitted due to 
low numbers trapped. Day and night catches, and males and females, combined for each site. 


130 ENTOMOLOGICAL NEWS 


eee eee 


In 1988, the temporal pattern of activity was reminiscent of that observed in 
1986, peaking in early June (Fig. 2). However, the developmental state of 
females differed sharply from all previous years. Of 85 females dissected, only 
one was gravid, and no spent individuals were ever found (Table 3). Lack of 
food, which can cause arrested ovarian development (Jeffords & Case 1987), 
and has been linked to curtailed reproduction in C. sycophanta (Weseloh 1985) 
may have been a contributing factor, although all individuals were obviously 
well-fed, with plump abdomens distended by extensive fat body development. 
Another explanation for undeveloped ovaries may be found in delayed emer- 
gence of imagines from hibernation. Although soil temperature data prior to 
May | are not available to support this hypothesis, onset of activity (May 24) 
and appearance of tenerals (May 24 to June 7) occurred 2 to 3 weeks later than 
in previous years. One could postulate that ovarian dormancy in C. frigidum is 
overcome by exposure to the photoperiodic shift from short-day to long-day, 
which triggers vitellogenesis in other spring-breeding species (Thiele 1971, 
1977). Prolonged hibernation thus may have caused C. frigidum to “miss” the 
period of shifting day-length critical for vitellogenesis. 


1987 

a 
w 
a 
: mom [ Je 
x 7/6 
i 

26 60 

] 

cx 
ro 1986 
a 204 ig ‘4 1988 
= 
=) 
2 46+ - 30 

1077 = = 204 

6+ - i = 107 

ol ZA T T Ss! i ° + rl 

5/7 7/7 6/10 7/1" 
TRAPPING DATE TRAPPING DATE 


["]TENERAL Q@AwWELL-DEVELOPED Mill GRAVID (__] SPENT 


Fig. 2. Numbers of C. frigidum females in each developmental stage captured during the first 10 
weeks of 1985 through 1988. Control and Test site data, and day and night catches, combined. 


Vol. 108, No. 2, March & April, 1997 131 


Virtually complete reproductive failure in 1988 was undoubtedly a main 
factor in population reduction over the subsequent 3 years (Table 1). Burgess 
and Collins (1917) stated that adult C. frigidum survive into a second or third 
year in the field. In the present study, a small proportion of the 1988 adults may 
have survived to 1989 and beyond, but population recovery was not achieved 
during the 3 years following reproductive failure. 


Table 1. Annual summaries of trap catches of C. frigidum, 1985 to 1991: total number of females 
and males, sex ratios (% males), and percent captured during the day (% diurnal); T and C = Test 
and Control sites. 


YEAR SITE Females Males TOTAL % males % diurnal 
1985 T 18 49 67 1B) 98.5 
(e 6 23 29 79.5 86.2 
1986 4h 34 106 140 IDS 85.6 
(@ 28 83 111 74.8 86.0 
1987 a0 59 372 431 86.3 80.8 
€ 33 160 193 82.9 TEA 
1988 1p 74 57 131 43.5 56.1 
Cc 14 13 27 48.1 38.7 
1989 an 6 6 12 = — 
Cc 3 1 4 — is 
1990 1h ] 1 2 — = 
Cc 1 0 1 — — 
199] 1h 1 1 2 — — 
Cc 0 0 0 = = 


132 ENTOMOLOGICAL NEWS 


Table 2. Number of mature ova carried by C. frigidum: means + SD, range, and N gravid females 
dissected in parentheses. 


YEAR 1985 1986 1987 1988 1989 
Test site 
mean + SD 13.9 + 8.0 15.9+7.4 17.7 + 6.1 12.0 10.7 + 3.1 
range 3-35 4-34 4-30 = 8-14 
(N) (17) (29) (43) (1) (3) 


Control site 
mean + SD 15.8 + 10.2 16.3 + 6.0 17.3 + 6.1 — — 
range 3-31 6-31 4-28 — — 
(N) (6) (27) (28) = = 


Table 3. Total annual number of female C. frigidum in each developmental category (T = teneral; W 
= non-gravid, ovaries slightly distended; G = gravid; S = spent) 


TEST SITE CONTROL SITE 

TF W G S 4) W G S 
1985 0 l 7s 0 0 0 6 0 
1986 0 2 29 3 0 0 27 0 
1987 1 0 43 5 l 0 28 2 
1988 1 69 l 0 3 11 0 0 
1989 0 l 3 2 0 0 1 2 
1990 0 0 l ) 0 0 0 l 
199] 0 0 0 I 0 0 0 0 

ACKNOWLEDGMENTS 


Support for this research was provided by the Naval Electronic Systems Command 
through a subcontract to IIT Research Institute under contract NO0039-81-C-0357. 


Vol. 108, No. 2, March & April, 1997 133 


LITERATURE CITED 


Burgess, A.F. and C.W. Collins. 1917. The genus Calosoma: including studies of seasonal histo- 
ries, habits, and economic importance of American species north of Mexico and of several 
introduced species. USDA Bull. 417: 1-124. 

Crins, W.J. .1980. Two aggregations of Calosoma frigidum (Coleoptera: Carabidae) in Ontario 
during 1976. Entomol. News 91: 155-158. 

Desender, K. and M. Alderweireldt. 1990. Yearly and seasonal variation of carabid diel activity in 
pastures and cultivated fields. Rev. Ecol. Biol. Sol 27: 423-433. 

Epstein, M.E. and H.M. Kulman. 1990. Habitat distribution and seasonal occurrence of carabid 
beetles in East-central Minnesota. Am. Midl. Nat. 123: 209-225. 

Gidaspow, T. 1959. North American caterpillar hunters of the genera Calosoma and Callisthenes 
(Coleoptera, Carabidae). Bull. Am. Mus. Nat. Hist. 116: 225-344. 

Jeffords, M.R. and L.J. Case. 1987. Effect of prey density on diurnal activity and ovarian devel- 
opment in Calosoma calidum (Coleoptera: Carabidae): implications for biological control of 
the gypsy moth, Lymantria dispar (Lepidoptera: Lymantriidae) in the Midwest. Great Lakes 
Entomol. 20: 93-97. 

Martel, J., Y. Maufette and S. Tousignant. 1991. Secondary effects of canopy dieback: the epigeal 
carabid fauna in Quebec Appalachian maple forests. Can. Entomol. 123: 851-859. 

Sanders, C.J. and K. Frankenhuyzen. 1979. High populations of a carabid beetle associated 
with spruce budworm. Bi-mon. Res. Notes 35: 21-22. 

Snider, R.J. and R.M. Snider. 1987. ELF ecological monitoring in Michigan. I. Description of 
sites for soil biological studies. Pedobiologia 30: 241-250. 

Snider, R.J. and R.M. Snider; 1995. ELF Communications System Ecological Monitoring Pro- 
gram: Soil Arthropods and Earthworms. Final Report. IIT Research Institute Technical Report 
D06212-7, 286 pp. Available from National Technical Information Service, U.S. Dept. of Com- 
merce. 

Thiele, H.U. 1971. Die Steuerung der Jahresrhythmik von Carabiden durch exogene und endogene 
Faktoren. Zool. Jb. Syst. 98: 341-371. 

Thiele, H.U. 1977. Measurement of day-length as a basis for photoperiodism and annual periodic- 
ity in the carabid beetle Prerostichus nigrita F. Oecologia 30: 331-348. 

Weseloh, R.M. 1985. Changes in population size, dispersal behavior, and reproduction of Calosoma 
sycophanta (Coleoptera: Carabidae), associated with changes in gypsy moth, Lymantria dispar 
(Lepidoptera: Lymantriidae), abundance. Environ. Entomol. 14: 370-377. 


134 ENTOMOLOGICAL NEWS 


NEW GENERIC SYNONYMIES IN BAETIDAE 
(EPHEMEROPTERA)! 


R. D. Waltz2, W. P. McCafferty 


ABSTRACT: Several substantive errors in the classification of the family Baetidae have resulted 
from revisions of mayflies identified as Baetis. Revised synonymies of the valid genera Alainites, 
Baetiella, Baetis, Diphetor, Labiobaetis, Nigrobaetis , and Takobia are provided and include neces- 
sary new synonyms as follows: Alainites [= Baetis (Acerbaetis), n. syn.); Baetiella [= Baetis (Tenui- 
baetis), n. syn.); Baetis sensu stricto [= Baetis (Tatubaetis), n. syn.}; Labiobaetis [= Baetis (Muller- 
baetis), n. syn.]; and Nigrobaetis [= Baetis (Margobaetis), n. syn.]. Species name 
emendations include Alainites atagonis, n. comb., Alainites clivosus, n. comb., Alainites 
chocoratus, n. comb., Alainites florens, n. comb., Alainites talasi, n. comb., Alainites yehi, n. comb., 
Alainites yoshinoensis, n. comb.; Baetiella ardua, n. comb., Baetiella inornata, n. comb. (and 
corrected orthography), Baetiella pseudofrequenta, n. comb.; Labiobaetis morus, n. comb.; Nigro- 
baetis facetus, n. comb., Nigrobaetis gombaki, n. comb., Nigrobaetis gracilentus, n. comb., 
Nigrobaetis mirabilis, n. comb., Nigrobaetis mundus, n. comb., Nigrobaetis numidicus, n. 
comb., Nigrobaetis taiwanensis, n. comb., Nigrobaetis terminus, n. comb.; Takobia acuticostalis, 
n. comb., Takobia kogistani, n. comb., and Takobia solangensis, n. comb. 


Kang et al. (1994) published five new subgenera and 13 new species of 
Baetidae from Taiwan that they placed in the genus Baetis Leach. Novikova 
and Kluge (1994) provided several new recombinations and descriptions of two 
new species within their concept of Baetis (Nigrobaetis). Baetis has been the 
subject of other revisionary efforts that have included Miiller-Liebenau (1970) 
(Europe); Morihara and McCafferty (1979a), Waltz and McCafferty (1987a), 
and McCafferty and Waltz (1990) (North America); Novikova and Kluge (1987) 
(Palearctic); and Waltz et al. (1994) and McCafferty and Waltz (1995) (world). 

Possession of the villopore, located at the base of the larval femora was 
found by Waltz and McCafferty (1987a, 1987b) and McCafferty and Waltz (1990) 
to be a significant synapomorphy uniting a number of genera of Baetidae. This 
monophyletic grouping of genera is presently referred to as the Baetis complex 
(see e.g., Waltz et al. 1994, Lugo-Ortiz and McCafferty 1996) and includes 
Acentrella Bengtsson, Baetiella Uéno, Baetis, Barbaetis Waltz and McCafferty, 
Cymulabaetis McCafferty and Waltz, Gratia Thomas, Heterocloeon 
McDunnough, Labiobaetis Novikova and Kluge, Liebebiella Waltz and 
McCafferty, Platybaetis Miiller-Liebenau, many (but not all) Pseudocloeon sensu 
auctt., and Tanzaniella Gillies. In addition to having the unique villopore, all 


1 Received July 29, 1996. Accepted August 3, 1996. 


2 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Room W-290, 
Indianapolis, IN 46204. 


3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 


ENT. NEWS 108(2): 134-140, March & April, 1997 


Vol. 108, No. 2, March & April, 1997 135 


members of the Baetis complex have lost the plesiotypic cluster of bristles 
located between the incisors and mola of the mandibles. Species that were for- 
merly classified in Baetis, but that are not members of the Baetis complex have 
obviously required reclassification into other genera, including some new gen- 
era. Highly distinctive and monophyletic lineages within the Baetis complex, 
many with species that were also once known as Baetis, have been recognized 
as the various genera listed above. 

Kang et al. (1994) were evidently not aware of concepts and genus group 
names that were already established, beginning in 1987, for elements of the 
formerly broad and polyphyletic concept of Baetis. Thus, most species described 
from Taiwan, in various new subgenera of Baetis, by Kang et al. (1994) are 
actually members of other presently recognized genera. All subgenera described 
by Kang and Yang in Kang et al. (1994) are junior synonyms of nominal gen- 
era, including both Baetis complex and non-Baetis complex genera. Novikova 
and Kluge (1994) retained a highly conservative and polyphyletic concept of 
the genus Baetis, including a broad subgeneric grouping they identified as the 
subgenus Nigrobaetis, which incorporated elements of Nigrobaetis s.str, 
Alainites Waltz and McCafferty, Diphetor Waltz and McCafferty, and Takobia 
Novikova and Kluge. The bases of each of these genera were addressed by 
Waltz et al. (1994). Necessary corrections to the Kang et al. (1994) and Novi- 
kova and Kluge (1994) works by way of new generic synonymies and short 
discussions of the pertinent genera follow. 


Alainites Waltz and McCafferty 


Baetis gracilis group Miiller-Liebenau, 1970:174, in part. 

Baetis muticus group Miiller-Liebenau, 1974:34. 

Alainites Waltz and McCafferty, in Waltz et al., 1994:34 [type, Baetis muticus Linn., 1758]. 

Baetis (Acerbaetis) Kang and Yang, in Kang et al., 1994:35 [type, Baetis clivosus Chang and Yang, 
1994], n. syn. 

Baetis (Nigrobaetis) muticus group: Novikova and Kluge, 1994: 630. 


Alainites was characterized by Waltz et al. (1994) and separated from other 
genera of the Baetis complex by tergal setae and armature characteristics previ- 
ously discussed by Miiller-Liebenau (1970) and by the apomorphic develop- 
ment of special paraproct prolongation (see Waltz et al., 1994:34). Species 
included in Acerbaetis Kang and Yang possess all of these characteristics and 
clearly belong to Alainites. Alainites atagonis (Gose), n. comb., A. chocoratus 
(Gose) n. comb., A. clivosus (Chang and Yang), n. comb., A. florens (Imanishi), 
n. comb., A. talasi (Novikova and Kluge), n. comb., A. yehi (Kang and Yang), 
n. comb., and A. yoshinoensis (Gose), n. comb., should be added to the list of 
Alainites species given by Waltz et al. (1994). Placement of A. atagonis is based 


136 ENTOMOLOGICAL NEWS 


on adult morphology. Alainites is widely distributed throughout the Palearctic, 
the Mediterranean/northern Africa area, and parts of southeast and east Asia, 
including southern China and Taiwan. 


Both the Kang et al. (1994) and the Waltz et al. (1994) papers bear Decem- 
ber, 1994 publication dates. Based on International Commission of Zoological 
Nomenclature (ICZN) convention, the priority of names published with con- 
temporaneous issue dates must be resolved on the basis of date of availability. 
The genus Alainites description in the Bulletin de la Société d’ Histoire Natu- 
relle de Toulouse, with an issue date of December, 1994 (which by ICZN con- 
vention is regarded as December 31, 1994), was distributed and met the criteria 
of availability on January 15, 1995, as confirmed by editors of the journal. This 
was prior to the verifiable availability date of April 15, 1995, for the Acerbaetis 
description in the Journal of the Taiwan Museum, with an issue date of Decem- 
ber 31, 1994. This was confirmed by the Assistant Curator of the Taiwan 
Museum. The name Alainites therefore met the criteria of availability approxi- 
mately three months prior to the date of availability of the name Acerbaetis. 
(see also Acknowledgments). 


Baetiella Uéno 


Baetiella Uéno 1931:220 [type, Acentrella japonica Imanishi, 1930). 

Pseudocloeon Klapalek, in part: Bogoescu and Tabacaru, 1957:483. 

Pseudocloeon (Baetiella): Kazlauskas, 1963:318 (English version pagination). 

Neobaetiella Miller-Liebenau, 1985:103 [type, Neobaetiella uenoi Miiller-Liebenau, 1985], syn- 
onymized by Waltz and McCafferty 1987b: 561. 

Baetis (Baetiella): Novikova and Kluge, 1987:16. 

Baetiella: Waltz and McCafferty, 1987b:561. 

Baetis (Tenuibaetis) Kang and Yang, in Kang et al., 1994:26 [type, Baetis pseudofrequentus Miiller- 
Liebenau, 1985], n. syn. 


The genus Baetiella was most recently characterized by Waltz and McCaf- 
ferty (1987b) to include those species of the Baetis complex that have an elon- 
gate segment 2 and conical segment 3 of the labial palps, among other distinctive 
characteristics. Species placed in Tenuibaetis by Kang et al. (1994) possess the 
characteristics of Baetiella. Species included in the genus were listed by Waltz 
and McCafferty (1987b). Baetiella ardua (Kang and Yang), n. comb., B. inor- 
nata (Kang and Yang), n. comb., and B. pseudofrequenta (Miiller-Liebenau), n. 
comb., should be added to that list. The species name Baetis (Tenuibaetis) inor- 
naturs Kang and Yang, was a lapsis calami, obvious from the etymology given 
for the species and figure citations accompanying the description (Kang et al. 
1994). Baetiella is Palearctic and Oriental in distribution. 


Vol. 108, No. 2, March & April, 1997 137 


Baetis Leach 


Baetis Leach, 1815:137 [type, Ephemera fuscata Linn., 1761). 

Brachyphlebia Westwood, 1840:25 [type, Ephemera fuscata Linn., 1761). 

Baetis (Tatubaetis) Kang and Yang, in Kang et al., 1994:23 [type, Baetis tatuensis Miiller-Liebe- 
nau, 1985], n. syn. 


The genus Baetis is a member of the Baetis complex, and because B. fus- 
catus is the type species, the genus is most typified by the B. fuscatus species 
group (Miiller-Liebenau 1970). Baetis also includes species of the rhodani and 
vernus groups in the Holarctic region (see Miiller-Liebenau 1970 and Morihara 
and McCafferty 1979a) as well as possibly certain species from non-Holarctic 
parts of the world that have yet to be associated with any species groups. Based 
on body coloration, mouthpart characteristics (esp. of the labium and maxil- 
lae), tergal armature, and setation of the legs, B. tatuensis is clearly a member 
of the B. fuscatus group. Baetis tatuensis is therefore a member of Baetis sensu 
stricto and should not be placed in a separate subgenus Tatubaetis as was done 
by Kang et al. (1994). Species currently classified in Baetis from sub-Saharan 
Africa, South America, and Australia require additional study before they can 
be confirmed to be members of the genus. As a result, the only recently up to 
date and confirmed listing of Baetis species for a large geographic area is for 
North America (see McCafferty 1996). 


Diphetor Waltz and McCafferty 


Diphetor Waltz and McCafferty, 1987:669 [type, Baetis hageni Eaton, 1885]. 
Baetis (Nigrobaetis): Novikova and Kluge, 1994:627. 


The distinctiveness of Diphetor from other described taxa was presented by 
Waltz et al. (1994). The genus is known from three species in North America 
and Algeria as listed by Waltz et al. (1994). The placement of species of Diphetor 
(a non- Baetis complex genus) in a subgenus of Baetis by Novikova and Kluge 
(1994) is untenable. 


Labiobaetis Novikova and Kluge 


Baetis atrebatinus group Miiller-Liebenau, 1970:150. 

Baetis propinquus group Morihara and McCafferty, 1979b: 130. 

Baetis molawinensis group Miiller-Liebenau, 1984:260. 

Baetis (Labiobaetis) Novikova and Kluge, 1987:13 [type, Baetis atrebatinus Eaton, 1870). 

Labiobaetis: McCafferty and Waltz, 1995:20. 

Baetis (Miillerbaetis) Kang and Yang, in Kang et al., 1994:32 [type, Baetis molawinensis Miiller- 
Liebenau, 1982], n. syn. 


138 ENTOMOLOGICAL NEWS 


The genus Labiobaetis is a member of the Baetis complex of genera and 
was most recently characterized by McCafferty and Waltz (1995). Synapo- 
morphies defining the genus include, among others, the excavate tip of the max- 
illary palps and usually the notched antennal segment |. McCafferty and Waltz 
(1995) indicated that the Oriental Baetis molawinensis group, originally recog- 
nized by Miiller- Liebenau (1984), clearly belonged to Labiobaetis. Kang et al. 
(1994) considered 11 previously described Oriental species in the subgenus 
Miillerbaetis (type, B. molawinensis). These are all species of Labiobaetis and 
were listed as such by McCafferty and Waltz (1995), along with all other spe- 
cies of the genus. The Taiwan species L. morus (Chang and Yang), n. comb., 
should be added to the list of known species of this Holarctic and Oriental 
genus. 


Nigrobaetis Novikova and Kluge 


Baetis niger group Miiller-Liebenau, 1970:163. 

Baetis gracilis group Miiller-Liebenau, 1970:174, in part. 

Baetis (Nigrobaetis) Kazlauskas: Novikova and Kluge, 1987:8 [type, Ephemera nigra Linn., 1761). 

Nigrobaetis Novikova and Kluge: Waltz, McCafferty and Thomas, 1994:34. 

Baetis (Margobaetis) Kang and Yang, in Kang et al., 1994:11 [type, Baetis mundus Chang and 
Yang, 1994], n. syn. 

Baetis (Nigrobaetis): Novikova and Kluge, 1994: 627, in part. 


Nigrobaetis is a non-Baetis complex genus that was most recently charac- 
terized by Waltz et al. (1994). Species from Taiwan assigned to Margobaetis by 
Kang et al. (1994) demonstrate Nigrobaetis generic characteristics, and thus, 
Margobaetis must be placed as a junior synonym of Nigrobaetis. The genus is 
known from the Holarctic and Oriental regions, and species included were listed 
by Waltz et al. (1994). To that list the following species should now be added: 
N. facetus (Chang and Yang), n. comb., N. gombaki (Miiller-Liebenau), n. comb., 
N. gracilentus (Chang and Yang), n. comb., N. mirabilis (Miiller-Liebenau), n. 
comb., N. mundus (Chang and Yang), n. comb., N. numidicus (Soldan and Tho- 
mas), n. comb., N. tatwanensis (Miiller-Liebenau), n. comb., and N. terminus 
(Chang and Yang), n. comb. Novikova and Kluge (1987,1994) incorrectly 
ascribed the name Nigrobaetis to Kazlauskas when in fact, by rules of nomen- 
clature, they are recognized as the inadvertent authors of the genus group name. 
Novikova and Kluge (1994) also incorrectly placed species of Diphetor and 
certain species of Alainites and Takobia in Nigrobaetis. That concept is poly- 
phyletic because it includes species of both the Baetis complex and non-Baetis 
complex. In any case, Nigrobaetis is not a subgenus of Baetis. 


Vol. 108, No. 2, March & April, 1997 139 


Takobia Novikova and Kluge 


Baetis (Takobia) Novikova and Kluge 1987:10 [type, Centroptilum maxillare Braasch and Soldan, 
1983]. 

Takobia: Waltz, McCafferty, and Thomas, 1994:35. 

Baetis (Nigrobaetis) maxillaris group Novikova and Kluge, 1994:630. 


The genus Jakobia and its systematic status was discussed by Waltz et al. 
(1994). Novikova and Kluge (1994) regarded it as part of a subgenus Nigro- 
baetis of the genus Baetis. We regard Takobia as a distinct taxon at the genus 
level. Additional species to those listed by Waltz et al. (1994) include Takobia 
acuticostalis (Dubey), n. comb., 7: kogistani (Novikova and Kluge), n. comb., 
and T. solangensis (Dubey), n. comb. The genus Jakobia is known from Cen- 
tral Asia including the western Himalayas. The terminal segment of the male 
forceps is elongated and not spherical. This is a correction to the description of 
the male genitalia reported in Waltz et al. (1994) and should be noted. 


ACKNOWLEDGMENTS 


We thank A. Thomas (Toulouse) for confirming dates of availability at the Paris 
Museum and with the publisher of Bulletin de la Société Histoire Naturelle de Toulouse; 
Robert Skarr (Senior Reference Librarian, Smithsonian Institution Libraries, Washington, 
D.C.), for providing dates of receipt of the journals at the Smithsonian Institution; Chia- 
Hsiang Wang (Associate Curator of the Taiwan Museum) for confirming availability dates of 
the Bulletin of the Taiwan Museum based on date of first distribution; T.-Q. and Y. Wang 
(Purdue University) for their assistance in obtaining information from Taiwan. This paper 
has been assigned Purdue ARP Journal No. 15082. 


LITERATURE CITED 


Bogoescu, C. and I. Tabacaru. 1957. Etude comparée des nymphes d’Acentrella et de Pseudoclo- 
eon, considérations phylogénétiques concernant la famille Baetidae (Ephemeroptera). Beitr. 
Entomol. 7: 483-492. 

Kang, S.-C., H.-C. Chang, and C.-T. Yang. 1994. A revision of the genus Baetis in Taiwan (Ephe- 
meroptera, Baetidae). J. Tatwan Mus. 47: 9-44. 

Kazlauskas, R. 1963. New and little-known mayflies (Ephemeroptera) from the USSR. Entomol. 
Obozr. 42: 582-593. (1964. Entomol. Rev. 42: 317-321 English translation) 

Leach, W. E. 1815. Entomology. Brewster’s Edinburgh Encyclopedia 9: 57-172. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. Aturbina georgei gen. et sp. n.: a small minnow 
mayfly (Ephemeroptera: Baetidae) without turbinate eyes. Aquat. Insects 18: 175-183. 

McCafferty, W. P. 1996. The Ephemeroptera species of North America and index to their complete 
nomenclature. Trans. Am. Entomol. Soc. 122: 1-54. 

McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) 
of North and Middle America. Trans. Am. Entomol. Soc. 116: 769-799. 

McCafferty, W. P. and R. D. Waltz. 1995. Labiobaetis (Ephemeroptera: Baetidae) new status, 
new North American species, and related new genus. Entomol. News 106: 19-28. 


140 ENTOMOLOGICAL NEWS 


Morihara, D. K. and W. P. McCafferty. 1979a. The Baetis larvae of North America (Ephemerop- 
tera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221. 

Morihara, D. K. and W. P. McCafferty. 1979b. Systematics of the propinquus group of Baetis 
species (Ephemeroptera: Baetidae). Ann. Entomol. Soc. Am. 72: 130-135. 

Miiller-Liebenau, I. 1970. Revision der Europaischen arten der gattung Baetis Leach, 1815 (Insecta, 
Ephemeroptera). Gewass. Abwass. 48/49 (1969): 1-214. 

Miiller-Liebenau, I. 1974. Baetidae aus Siidfrankreich, Spanien und Portugal (Insecta, Ephemerop- 
tera). Gewass. Abwass. 53/54: 7-42. 

Miiller-Liebenau, I. 1984. New genera and species of the family Baetidae from West-Malaysia 
(River Gombak) (Insecta: Ephemeroptera). Spixiana 7: 253-284. 

Miiller-Liebenau, I. 1985. Baetidae from Taiwan with remarks on Baetiella Uéno, 1931 (Insecta, 
Ephemeroptera). Arch. Hydrobiol. 104: 93-110. 

Novikova, E. A. and N. Kluge. 1987. Systematics of the genus Baetis (Ephemeroptera, Baetidae) 
with description of a new species from Middle Asia. Vestn. Zool., 1987 (4): 8-19. 

Novikova, E.A. and N. Kluge. 1994. Mayflies of the subgenus Nigrobaetis. (Ephemeroptera, Baeti- 
dae, Baetis Leach, 1815). Entomol. Oborz. 73: 623-644. (1995. Entomol. Rev. 74: 16-39- En- 
glish translation). 

Uéno, M. 1931. Contributions to the knowledge of Japanese Ephemeroptera. Annot. Zool. Japon. 
13: 189-226. 

Waltz, R. D. and W. P. McCafferty. 1987a. New genera of Baetidae for some Nearctic species 
previously included in Baetis Leach (Ephemeroptera). Ann. Entomol. Soc. Am. 80: 667-670. 

Waltz, R. D. and W. P. McCafferty. 1987b. Systematics of Pseudocloeon, Acentrella, Baetiella, 
and Liebebiella, new genus (Ephemeroptera: Baetidae). J. New York Entomol. Soc. 95: 553- 
568. 

Waltz, R. D., W. P. McCafferty, and A. Thomas. 1994. Systematics of Alainites n. gen., Diphetor, 
Indobaetis, Nigrobaetis n. stat., and Takobia n. stat. (Ephemeroptera, Baetidae). Bull. Nat. Hist. 
Soc., Toulouse 130: 33-36. 

Westwood, J. O. 1840. An introduction to the modern classification of insects, founded on the 
natural habits and corresponding organization of the different families. Longman, Orme, Brown, 
Green, and Longman, London. 


Vol. 108, No. 2, March & April, 1997 141 


CRITICAL COMMENTARY ON 
THE GENUS SIPHLONISCA 
(EPHEMEROPTERA: SIPHLONURIDAE)!2 


W. P. McCafferty, George F. Edmunds, Jr4 


ABSTRACT: Data are presented that contravene certain unfounded statements that have been pub- 
lished regarding Siphlonisca aerodromia, a siphlonurid mayfly from extreme northeastern U.S.A. 
and eastern Canada. The species is shown to be locally common in restricted habitats within its 
geographic range. A large number of mayfly species in North America are known from fewer locale 
records and specimens. Paleontological data indicate that Siphlonisca is not represented in the 
fossil record, the Holarctic family Siphlonuridae sensu stricto is not represented prior to the Ter- 
tiary, and S. aerodromia should not be considered a living fossil. Cladistic data place Siphlonisca as 
a sister group of the genus Parameletus. The presence of lateral abdominal flanges in the adults are 
independent adaptations in a number of distantly related mayfly lineages and are related to gill 
position and type in the larval stage. The presence of laterally expanded abdomens in some Paleo- 
zoic insects does not necessarily denote close relationships with particular lineages of extant may- 
flies. The fact that Siphlonisca is monotypic is not unique among North American mayfly genera. 


In an appendix to the 1908 Report of the State Entomologist of New York, 
James G. Needham described the northeastern Nearctic siphlonurid mayfly 
Siphlonisca aerodromia. The description of the new genus and species was ini- 
tially drafted for a single female adult from Sacandaga Park, Johnstown, New 
York, collected by C. P. Alexander. However, based on additional material of 
both male and female adults also collected by Alexander on June 6, 1909, and 
also from Sacandaga Park, the male description was also included. The date 
ascribed to this species has often been 1908 (e.g., Traver 1935; Burian and 
Gibbs 1988, 1991; McCafferty 1996). The date of the species name should be 
1909, as listed by Edmunds and Allen (1957) and Edmunds (1962) because the 
name was not published or available until 1909 and because the description was 
in part based on 1909. data. Incidentally, for the same reason, the name Pota- 
manthus inequalis Needham, which appeared in the same appendix, should 
also be dated 1909, as has been historically recognized in the literature. With 
respect to this somewhat awkward chronology, Bae and McCafferty (1991), in 
their comprehensive revision of the family Potamanthidae, referenced the name 
P. inequalis under the synonymy of Anthopotamus myops as “Potamanthus 
inequalis 1909 Needham Jn: Needham 1908. . .” 

The larval stage of S. aerodromia was originally described by Clemens (1915) 


1 Received August 20, 1996; Accepted September 9, 1996. 

2 Purdue Agricultural Research Program Journal No. 15146. 

3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 
= Department of Biology, University of Utah, Salt Lake City, UT 84112. 


ENT. NEWS 108(2): 141-147, March & April, 1997 


142 ENTOMOLOGICAL NEWS 


from larvae collected by Alexander at the species’ type locality. Alexander’s 
collecting notes were included in the paper by Clemens. Various other works 
essentially reviewed data from the original materials of this species from 
Sacandaga Park up until the time that Magnin and Harper (1970), Fiance (1978), 
and Hutchinson (1989) reported it from parts of eastern Canada; Gibbs (1980) 
and Burian and Gibbs (1991) reported specimens from several localities in Maine; 
Gibbs and Mingo (1986) provided some life history and feeding data; Burian 
and Gibbs (1988) thoroughly redescribed the species; and Gibbs and Siebenmann 
(1996) most recently critically studied the biology of the species. In the latter 
paper, another record from New York was mentioned, but with no data associ- 
ated with it. No other species of Siphlonisca have been discovered, and the 
genus remains monotypic and restricted to northeastern North America. 

Two primary conceptions associated with this relatively large sized and strik- 
ing species seem to have preoccupied both researchers who have recently worked 
with the species and others who have written about it. The first is the presence 
of unusually large lateral flanges on the abdomen of the adults that are con- 
spicuous in flight. The other is the continually repeated idea that S. aerodromia 
is one of the rarest mayflies. Both of these conceptions have now apparently 
been linked with an idea that S. aerodromia is unique among recent mayflies 
and is somehow a “living fossil” (Anonymous 1995). We argue herein that these 
conceptions are not valid. Abdominal flanges are found in a number of pisciform 
mayflies, including both siphlonurids and non-siphlonurids, but are more com- 
mon outside North America (abdominal flanges are also independently evolved 
in pannote mayflies and others). Siphlonisca appears to be very closely related 
to another more widespread genus of the family Siphlonuridae. And, based on 
its abundance and studied populations recently found of the species, S. 
aerodromia should be considered locally common and is not rare compared to 
many other mayflies in North America. 


DISCUSSION 


Alexander (pers. comm. with GFE) stated of his 1909 collections of S. 
aerodromia, that “they were abundant and they kept high in the air where they 
were conspicuous by reason of the wide abdomen (lateral expansions of seg- 
ments 5-9).” The noted insect morphologist G. C. Crampton was intrigued by 
the resemblance of the abdominal flanges to similar expansions on the thorax 
or abdomen of certain Paleozoic insects (Alexander, pers. comm.). This led to 
additional collecting trips to Sacandaga Park, where Alexander and, later, 
Crampton obtained more specimens (see Edmunds et al. 1976). Needham (1908) 
compared the abdomen of S. aerodromia to that of the New Zealand Onisci- 
gaster wakefieldi McLachlan (Oniscigastridae), one of the other species known 
at the time to have abdominal flanges. We know of no suggestion by Crampton 


Vol. 108, No. 2, March & April, 1997 143 


that the abdominal flanges of Siphlonisca represented a retention of Paleozoic 
characteristics or a suggestion that Siphlonisca was more primitive than other 
extant pisciform mayflies. Furthermore, Siphlonisca was not compared with 
Paleozoic insects in Crampton’s insect morphology notes (read by GFE in a 
course given at the University of Massachusetts by J. F. Hanson). 

A small, rather curious article about S. aerodromia recently appeared in Fly 
Fisherman Magazine (Anonymous 1995). The article was accompanied by a 
photograph of a female adult mayfly taken in western Pennsylvania that was 
labeled as “An adult mayfly that may be a rare carnivorous Tomah mayfly.” 
(“Tomah mayfly” is a vernacular name given to S. aerodromia, alluding to the 
stream in Maine where most published scientific studies of the mayfly have 
been conducted; but the photographed mayfly from Pennsylvania is most cer- 
tainly not Siphlonisca, based on its obvious non-siphlonuroid wing venation.] 
We take exception to certain impressions conveyed by statements about S. aero- 
dromia given in that article because the article will most certainly be seen by 
some entomologists, and erroneous or unfounded information regarding the 
species may be cited in the literature. Obviously, popular articles must be taken 
with “ a grain of salt,” because errors in fact can be common; however, this 
article is additionally vexing because an ephemeropterist is evidently the source 
of the information. We are particularly concerned with impressions of S. aero- 
dromia that are conveyed with the statements: “. . . it’s so rare—in effect a 
flying fossil dating from the Paleozoic Era — that it disappeared for 50 years and 
was considered extinct” and that it “has been called the rarest mayfly in the 
world.” Moreover, the article indicated that in the Spring, 1995 issue of the 
Maine Legacy, the magazine of the Maine Nature Conservancy, the mayfly was 
cited as standing apart from other North America mayflies by not sharing its 
genus with any other. 

From existing published records of S. aerodromia (see references above), it 
has become apparent that the species — while restricted somewhat geographi- 
cally to the extreme northeastern United States and eastern Canada, and by 
habitat availability (stream backwaters and temporary floodplains) — is none- 
theless locally common and often abundant where it occurs. Despite this, a 
recent scientific article continued to promulgate the misconception of rareness 
by using the phrase “the rare mayfly Siphlonisca aerodromia” in its title (Gibbs 
and Siebenmann 1996). We estimate that at least 100 North American mayfly 
species, representative of most North American families, are presently known 
from fewer specimens and fewer locale records. In many of these cases, the 
“rarity” of such species is a collecting artifact. In other words, efficient sam- 
pling historically has not been possible because usual collecting techniques 
have not been amenable to capturing the species in their peculiar habitats or 
with respect to their peculiar life histories. A few examples of North American 
species that are not generally collected as larvae for these reasons include deep- 


144 ENTOMOLOGICAL NEWS 


water riverine species such as the heptageniids Acanthomola pubescens Whit- 
ing and Lehmkuhl, Anepeorus simplex (Walsh), and Raptoheptagenia cruentata 
(Walsh); fast-swimming psammophilous species such as Analetris eximia 
Edmunds (Acanthametropodidae) and Pseudiron centralis McDunnough (Pseu- 
dironidae); very small species such as the baetids Apobaetis indeprensus Day 
and Paracloeodes minutus (Daggy); certain clay-burrowing species such as 
Pentagenin vittigera (Walsh) (Ephemeridae) and Tortopus primus (McDun- 
nough) (Polymitarcyidae); and early season emergers whose developed larvae 
are only available in late fall, winter and very early spring, such as the more 
northern temperate species of Metretopodidae. 

Siphlonisca aerodromia should not be considered a fossil any more than 
any other extant mayfly species. The oldest fossils that have been associated 
with the family Siphlonuridae sensu lato date from the Jurassic (see discussion 
of McCafferty 1990). However, such data are based on the broadest historical 
concept ever given to that family (see McCafferty and Edmunds 1979, 
McCafferty 1990). The actual clade to which Siphlonisca belongs is a much 
more restricted Holarctic grouping [Siphlonuridae sensu McCafferty (1991) 
and Kluge et al. (1995)], and includes along with Siphlonisca, the genera 
Edmundsius Day, Parameletus Bengtsson, Siphlonurus Eaton, and provision- 
ally Siphluriscus Ulmer (see McCafferty and Wang 1994a). Fossils that are 
possibly assignable to this particular clade include members of the genera 
Aphelophlebodes Pierce from the Miocene of California and Baltameletus 
Demoulin and Siphlonurus from Eocene Baltic amber. Siphlonuridae sensu 
stricto is not an archaic group and is comparable in age to most other extant 
mayfly families. 

Neither Siphlonisca and its close relatives (other Siphlonuridae), nor any 
other extant group of mayflies are fossils “dating from the Paleozoic.” Most 
extant families of mayflies (McCafferty 1991) are also represented by fossils 
from the Tertiary as is Siphlonuridae, but only seven of them, not including 
Siphlonuridae, apparently have representative fossils from the Mesozoic, and 
no extant families have representative fossils from the Paleozoic, although there 
are strictly fossil families of mayflies from that era (see Hubbard 1987, 
McCafferty 1990). 

Although cladistic analyses of all lineages of pisciform mayflies have not 
yet been performed satisfactorily to reach comprehensive conclusions regard- 
ing the entire group, all siphlonuroid genera have been studied independently 
by both of us, and we have reached the same conclusion regarding the phyloge- 
netic position of Siphlonisca. Siphlonisca apparently is the sister genus of Para- 
meletus. Illustrative of this hypothesized relationship are the very similar 
apomorphic male genitalia and short hindtarsi shared by adults of the two gen- 
era. Of further possible consequence in this regard, Parameletus larvae in Utah 
are found in flooded Carex (Edmunds 1957), very similarly to that which has 
been reported for Siphlonisca in Maine by Gibbs (1980). Like Siphlonisca, 


Vol. 108, No. 2, March & April, 1997 145 


Parameletus has been a genus with species that are often difficult to find and 
collect. 

It is evident that, Siphlonisca’s erroneous relationship with the Paleozoic 
has been associated with the observation that certain Paleozoic insects pos- 
sessed somewhat similar laterally expanded abdomens. A number of extant 
mayflies have a larval abdomen that is variously expanded laterally. In every 
instance where this occurs, the abdominal gill lamellae originate dorsally rather 
than laterally on the abdomen and tend to overlay the lateral abdominal flanges. 
This feature is typical of the entire subgroup of mayflies known as pannote 
mayflies (McCafferty and Edmunds 1979). In genera with highly developed 
flanges, e.g., those of the Timpanoga complex of the family Ephemerellidae 
(McCafferty and Wang 1994b), the larval flanges remain evident in the adults. 
In the non-pannote and non-pisciform burrowing mayfly subfamily Palingeniinae 
of the family Ephemeridae, flanged larval abdomens are also carried over into 
the adults. Besides Siphlonisca, other examples of this phenomenon among the 
pisciform mayflies include the genera Oniscigaster McLachlan and Siphlonella 
Needham and Murphy (family Oniscigastridae) from the Southern Hemisphere. 
Given the wide variety of mayflies from disparate Ephemeroptera lineages dem- 
onstrating broad lateral flanges and the fact that it is apparently related to larval 
gill adaptation, we can not assume from such data, alone, any common deriva- 
tions. This includes deducing a primordial relationship of Siphlonisca with Pa- 
leozoic forms having similar modifications. 

We do not know if the adult abdominal flanges are entirely non-adaptive 
vestiges of the larval structures or whether there may be some advantage (e. g., 
aerodynamic) accrued to the adult by their retained presence that explains their 
nearly full retention in such mayflies as Siphlonisca. Only very slight lateral 
expansions are found in larvae of other Siphlonuridae (with no appreciable adult 
vestiges of such). Abdominal flange development in Siphlonisca and the loss of 
the fork of the MP vein in the hindwings of Parameletus are autapomorphies 
that allow us to continue to regard the two lineages as separate genera. 

The claim that S. aerodromia disappeared for 50 years is not correct. It is 
true that 55 years occurred between the time Clemens (1915) discussed the 
species and the next publication reporting records of the species (Magnin and 
Harper 1970). However, the species was collected in New York up until the 
1930’s, when the type locality was altered due to the construction of the 
Sacandaga Reservoir. Fiance (1978) reported material that had been collected 
in 1952 in Canada. Lapses in records are most logically explained by the fact 
that there were few if any ephemeropterists collecting the habitats of S. aero- 
dromia in the Northeast between the time when Alexander and Crampton were 
collecting the type locality in New York and the time when intensive local re- 
search was begun more recently in Maine by Gibbs and her students. 

Finally, S. aerodromia does not stand alone among the North American 
mayflies in being monotypic. At the current time, 10 other North American 


146 ENTOMOLOGICAL NEWS 


genera in various families are known from only one species (see McCafferty 
1996). All of these can be considered just as unique in this respect as Siphlo- 
nisca. Certainly, S. aerodromia is a striking species from its size and some 
details of its morphology. Many other mayflies in North America, however, are 
equally extraordinary, and others from around the world possess more highly 
unusual morphological attributes. 

Local fascination with a particular species of aquatic insects can have very 
positive effects because it often leads to productive biological studies. The 
recent studies of Siphlonisca are just one case in point. Some examples of other 
species that are fascinating in their own respect and that have received concen- 
trated local attention in North America have included the behningiid Dolania 
americana Edmunds and Traver in Florida (e.g., Peters and Peters 1977, Peters 
et al. 1987), the polymitarcyid Ephoron album (Say) in Utah (e.g., Edmunds et 
al. 1956), and the heptageniid Stenacron interpunctatum (Say) in Indiana (e.g., 
McCafferty and Huff 1978, McCafferty and Pereira 1984, McShaffrey and 
McCafferty 1986). The only possible down side to such concentrated attention 
is that in interpreting and popularizing such data for general readership, distor- 
tions and exaggerations of information can and do occur to affect a degree of 
sensationalism, as has obviously been the case with Siphlonisca. Our intent 
here has been simply to set the record straight. 


LITERATURE CITED 


Anonymous. 1995. Carnivorous mayfly? Fly Fisherman Mag. 26 (Sept.): 30. 

Bae, Y. J. and W. P. McCafferty. 1991. Phylogenetic systematics of the Potamanthidae (Ephe- 
meroptera). Trans. Entomol. Soc. Am. 117: 1-143. 

Burian, S. K. and K. E. Gibbs. 1988. A redescription of Siphlonisca aerodromia Needham (Ephe- 
meroptera: Siphlonuridae). Aquat. Insects 10: 237-248. 

Burian, S. K. and K. E. Gibbs. 1991. Mayflies of Maine: an annotated faunal list. Maine Agr. Exp. 
Stat. Tech. Bull. 142: 1-109. 

Clemens, W. A. 1915. Mayflies of the Siphlonurus group. Can. Entomol. 47: 245-260. 

Edmunds, G. F. Jr. 1957. On the life history of Parameletus columbiae McDunnough (Ephe- 
meroptera). Proc. Utah Acad. Sci. arts lettr. 34: 24-25. 

Edmunds, G. F. Jr. 1962. The type localities of the Ephemeroptera of North America north of 
Mexico. Univ. Utah Biol. Ser. 12: viii + 1-39. 

Edmunds, G. F. Jr. and R. K. Allen. 1957. A checklist of the Ephemeroptera of North America 
north of Mexico. Ann. Entomol. Soc. Am. 50: 317-324. 

Edmunds, G. F. Jr., S. L. Jensen and L. Berner. 1976. The mayflies of North and Central America. 
Univ. Minn. Press, Minneapolis. 

Edmunds, G. F. Jr., L. T. Nielsen and J. R. Larsen. 1956. The life history of Ephoron album 
(Say) (Ephemeroptera: Polymitarcidae). Wasmann J. Biol. 14: 145-153. 

Fiance, S. B. 1978. A new locality for Siphlonisca aerodromia (Ephemeroptera: Siphlonuridae). 
Entomol. News 89: 208. 

Gibbs, K. E. 1980. The occurrence and biology of Siphlonisca aerodromia Needham (Ephe- 
meroptera: Siphlonuridae) in Maine, U.S.A. pp. 167-168. Jn: J. F. Flannagan and K. E. Mar- 
shall [Eds.], Advances in Ephemeroptera biology. Plenum, New York. 


Vol. 108, No. 2, March & April, 1997 147 


Gibbs, K. E. and T. M. Mingo. 1986. The life history, nymphal growth rates, and feeding habits of 
Siphlonisca aerodromia Needham (Ephemeroptera: Siphlonuridae) in Maine. Can. J. Zool. 64: 
427-430. 

Gibbs, K. E. and M. Siebenmann. 1996. Life history attributes of the rare mayfly Siphlonisca 
aerodromia Needham (Ephemeroptera: Siphlonuridae). J. N. Am. Benthol. Soc. 15: 95-105. 

Hubbard, M. D. 1987. Ephemeroptera. Fossilium catalogus, I: Animalia, pars 129. Kluger Publ., 
Amstelveen, The Netherlands. 

Hutchinson, R. 1989. Premiére mention de Siphlonisca aerodromia Needham (Ephemeroptera: 
Siphlonuridae) au Québec et répartition en Amérique du Nord. Faberies 14: 25-27. 

Kluge, N., D. Studemann, P. Landolt and T. Gosner. 1995. A reclassification of Siphlonuroidea 
(Ephemeroptera). Bull. Soc. Entomol. Suisse 68: 103-132. 

Magnin, E. and P. P. Harper. 1970. La nourriture des esrugeions Acipenser fulvescens de la Baie 
James. Natur. Can. 97: 73-85. 

McCafferty, W. P. 1990. Ephemeroptera, pp. 20-50. In: D. A. Grimaldi [Ed.], Insects from the 
Santana Formation, Lower Cretaceous, of Brazil. Bull. Am. Mus. Nat. Hist. 195, New York. 

McCafferty, W. P. 1991. Toward a phylogenetic classification of the Ephemeroptera (Insecta): a 
commentary on systematics. Ann. Entomol. Soc. Am. 84: 343-360. 

McCafferty, W. P. 1996. The Ephemeroptera species of North America and Index to their com- 
plete nomenclature. Trans. Am. Entomol. Soc. 122: 1-54. 

McCafferty, W. P. and G. F. Edmunds, Jr. 1979. The higher classification of the Ephemeroptera 
and its evolutionary basis. Ann. Entomol. Soc. Am. 75: 5-12. 

McCafferty, W. P. and B. L. Huff. 1978. The life cycle of the mayfly Stenacron interpunctatum . 
Great Lakes Entomol. 11: 209-216. 

McCafferty, W. P. and C. Pereira. 1984. Effects of developmental thermal regimes on two mayfly 
species and their taxonomic interpretation. Ann. Entomol. Soc. Am. 77: 69-87. 

McCafferty, W. P. and T.-Q. Wang. 1994a. Relationships of the genera Acanthametropus, Anale- 
tris, and Siphluriscus, and re-evaluation of their higher classification (Ephemeroptera: Pisci- 
forma). Great Lakes Entomol. 27: 209-215. 

McCafferty, W. P. and T.-Q. Wang. 1994b. Phylogenetics and the higher classification of the 
Timpanoga complex (Ephemeroptera: Ephemerellidae). J. N. Am. Benthol. Soc. 13: 569-579. 

McShaffrey, D. and W. P. McCafferty. 1986. Feeding behavior of Stenacron interpunctatum (Ephe- 
meroptera: Heptageniidae). J. N. Am. Benthol. Soc. 5: 200-21 0. 

Needham, J. G. 1908. A peculiar mayfly from Sacandaga Park, New York State Mus. Bull. 134: 
71-75. 

Peters, W. L. and J. G. Peters. 1977. Adult life and emergence of Dolania americana in north- 
western Florida (Ephemeroptera: Behningiidae). Int. Rev. Hydrobiol. 62: 409-438. 

Peters, W. L., J. G. Peters and T. J. Fink. 1987. Seasonal synchronization of emergence in Dolania 
americana (Ephemeroptera: Behningiidae). Can. J. Zool. 65: 3177-3185. 

Traver, J. R. 1935. Part II. Systematic, pp. 239-739 In: J. G. Needham, J. R. Traver and Y-C. Hsu 
[eds.], The biology of mayflies with a systematic account of North American species. Com- 
stock Publ. Co., Ithaca, New York. 


148 ENTOMOLOGICAL NEWS 


ESTABLISHMENT OF ALEIODES (= ROGAS) 
INDISCRETUS (HYMENOPTERA: 
BRACONIDAE), AN INTRODUCED PARASITE OF 
THE GYPSY MOTH IN NORTH AMERICA! 


Robert F. W. Schroder, Anne M. Sidor2 


ABSTRACT: Aleiodes indiscretus (Reardon);, an introduced braconid endoparasite released as a 
part of the gypsy moth biological control program, was recovered from Dasychira basiflava larvae 
collected in 1994 at Brandywine, Prince Georges County, Maryland. This confirms establishment 
of the parasite for the first time in North America. 


Aleiodes (= Rogas) indiscretus (Reardon), a solitary braconid endoparasite 
of Lymantria obfuscata Walker (Lepidoptera: Lymantriidae), was initially col- 
lected in Kotgarh, Kashmir, India. It was imported from India to the United 
States for rearing and release as a part of a biological control program against 
the gypsy moth, Lymantria dispar (L.) by the USDA, APHIS, Otis Air National 
Guard Base (ANGB), Massachusetts (Reardon, 1970). The parasite was first 
reared in 1967, at Otis ANGB and then released in 1968 at Ledyard, CT and 
Falmouth, MA. A total of 30,847 adults were released from 1968 to 1977 in 
New Jersey, Massachusetts, Connecticut and southeastern Pennsylvania (Met- 
terhouse, 1981). According to Anderson et al. (1977), extensive efforts were 
made to establish A. indiscretus and other new species for control of gypsy 
moth in North America, but no success was achieved in recent years except for 
the recovery in 1978 by Schaefer et al. (1989) of the pupal parasite Coccygo- 
mimus disparis (Viereck) (Hymenoptera: Ichneumonidae) on alternate hosts. 
According to Schaefer the parasite is well established in North America. 

One of the reasons given for the failures of A. indiscretus and some other 
exotic parasites was the lack of suitable alternate hosts (Hedlund and Schroder, 
1981). The gypsy moth is a univoltine species that is normally active from late 
April through July. The egg stage is present during the remainder of the year. 
Therefore, larval parasites including A. indiscretus and other pupal parasites 
must adapt to this short period of activity or find other alternate hosts. In 1974, 
studies were initiated and continued through 1994 in designated Maryland wood- 
lots to identify potential alternate hosts available prior to the arrival of intro- 
duced gypsy moth parasites, and potential native parasites that may select the 
gypsy moth upon its arrival at the study sites. We report here on the first estab- 
lishment of A. indiscretus in North America. 


1 Received January 22, 1996. Accepted October 9, 1996. 
2 USDA, ARS, PSI, Insect Biocontrol Laboratory, Beltsville, Md 20705. 


ENT. NEWS 108(2): 148-150, March & April, 1997 


Vol. 108, No. 2, March & April, 1997 149 


MATERIAL AND METHODS 


Twenty woodlot sites were selected in central and western Maryland in 1974 
(Fig. 1). At each site, 10 hardwood trees were each banded with burlap. During 
the growing season, we periodically examined the burlap bands and collected 
all the lepidopteran insects and any associated parasitic cocoons present under 
burlap. This study was conducted from 1974 to 1994. In addition to this study, 
we banded trees in southern Maryland during the 1993 - 94 growing season. We 
were particularly interested in the native lymantriids and associated parasites 
present at the sites. The insects collected were fed on leaves in our laboratory at 
20° C, 50% RH and under natural light for emergence of parasites. Specimens 
of each species were submitted for identification to the USDA, Systematic 
Entomology Laboratory (SEL). 


RESULTS AND DISCUSSION 


During the course of our alternate host studies over the past 19 years, it 
became apparent that Dasychira (Lepidoptera: Lymantriidae) was one of the 


Pennsylvania 


Virginia 


E-2 


Chesapeake Bay 


Fig 1. Area of woodlot sites in Maryland selected for survey of alternate hosts of imported gypsy 
moth parasites, including established site of Aleiodes indiscretus. 


Shaded portion was area where woodlot sites were located. Star represents site in Prince Georges 
County where A. indiscretus was discovered. 


150 ENTOMOLOGICAL NEWS 


predominant species present (R.F.W.S., unpublished data). According to Fergu- 
son (1978), Dasychira species all overwinter as partly grown, diapausing lar- 
vae. We therefore considered them as potential alternate hosts for introduced 
parasites of the gypsy moth. 

As one sample among hundreds collected in the ongoing study in 1994, we 
recovered 9 braconid parasites from 38 Dasychira basiflava (Packard) larvae 
collected 5/8 - 5/16 at Brandywine, Prince Georges County, Maryland (Fig. 1). 
The parasites were identified as Aleiodes indiscretus by P. M. Marsh, Research 
Entomologist (retired), SEL. The genus Aleiodes was resurrected out of syn- 
onymy with the genus Rogas by van Achterberg (1982). According to Marsh 
(personal communication), the genus Rogas is restricted to tropical areas, and 
all North American species listed in catalogues as Rogas should be called 
Aleiodes. The closest release site of this parasite was near Philadelphia in Schuyl- 
kill County, Pennsylvania, where it was released in 1971. This parasite was 
never recovered from gypsy moth larvae or other Dasychira collected in the 
study. Perhaps more extensive surveys like this one should have been conducted. 
The fact that this parasite was never recovered from gypsy moth larvae col- 
lected at our study sites, would suggest that it has chosen D. basiflava as its 
primary host. Ferguson (1978) states that D. basiflava is one of the closely knit 
complex of 16 North American species limited to the United States and Canada. 
Further studies can only verify if A. indiscretus has made the permanent transi- 
tion to Dasychira or if its primary host is still the gypsy moth. After nearly 20 
years of searching for alternate hosts of introduced gypsy moth parasites, we 
can now confirm the establishment of one more parasite, the impact of which 
must now be determined. 


LITERATURE CITED 


Anderson, J.F., M.A. Hoy, and R.M. Weseloh. 1977. Field cage assessment of the potential for 
establishment of Rogas indiscretus against the gypsy moth. Environ. Entomol. 6: 375-80. 
Ferguson, D.C. 1978. The moths of America north of Mexico. Fascicle 22.2. Noctuoidea. Lyman- 
triidae. E.W. Classey Limited and The Wedge Entomological Research Foundation, London. 

110 pp. 

Hedlund, R.C. and R.F.W. Schroder. 1981. Alternate host studies: pp. 372-75. In C.C. Doane & 
M.L. MacManus (eds.), The gypsy moth: research toward integrated pest management. USDA 
Tech. Bull. 1584, 757 pp. 

Metterhouse, W. 1981. Rogas indiscretus Reardon, pp. 368-69. Jn C.C. Doane & M.L. MacManus 
(eds.), The gypsy moth: research toward integrated pest management. USDA Tech. Bull. 1584, 
757 pp. 

Reardon, R.C. 1970. A new species of Rogas parasitic on the gypsy moth, Porthetria dispar 
(Hymenoptera: Braconidae). Proc. Entomol. Soc. Wash. 72: 473-5. 

Schaefer, P.W., R.W. Fuester, R.J. Chianese, L.D. Rhoads and R.B. Tichenor, Jr. 1989. Intro- 
duction and North American establishment of Coccygomimus disparis (Hymenoptera: Ichneu- 
monidae), a polyphagous pupal parasite of Lepidoptera, including gypsy moth. Environ. Entomol. 
18: 1117-1125. 

van Achterberg, C. 1982. Notes on some type species described by Fabricius of the subfamily 
Braconinae, Rogadinae, Microgastrinae and Agathidinae. Entomol. 42: 133-39. 


Vol. 108, No. 2, March & April, 1997 151 


A NEW NORTH AMERICAN SPECIES OF THE GENUS 
CLUZOBRA (DIPTERA: MYCETOPHILIDAE)! 


Edward I. Coher2 


ABSTRACT: The first North American species of the principally Neotropical genus Cluzobra is 
described from a short series of males and females taken in Ohio. 


Twenty-nine species of the sciophiline genus Cluzobra have been described, 
all from the Neotropical Region. Matile (1996) described twenty new species 
and reviewed nine species described by Lane (1948, 1956, 1959, 1960) and 
Edwards (1934, 1940, 1941). Excluding two questionable records of females, 
nineteen species are known from a single country and ten from a single 
collection. 

Only 2 species, C. aitkeni and C. spinulifera are reported as being widely 
distributed, although C. binocellaris and C. lanei would also fall into this cat- 
egory if collections from Mexico for the former and Nicaragua for the latter are 
confirmed by capture of males. Except for C. /anei, the other three species have 
been collected virtually year-round. 


Cluzobra Edwards, 1940 
Type species: Acnemia binocellaris Edwards, 1934 


Edwards , 1940. Rev. Entomol. 11:463. 
Vockeroth, 1981. Agric. Canada, Monogr. No. 27:231,237. 
Matile, 1996. Ann. Soc. Entomol. France (N.S.) 32(1):3-57. 


Cluzobra has been characterized in great detail by Matile (1996). Briefly, 
these sciophiline flies are distinguished by wings with Sc» absent, Sc ending in 
C, and Cu simple. 


Cluzobra antennulata Coher, NEW SPECIES 


Habitus: a moderately slim, orange-yellow fly. 


Male. Head (Fig. 1) with vertex and occiput light red-brown, vertex light between antennae: each 
ocellus about its diameter from the eye with an irregular row of short setae between them; frons 
yellowish, narrow, 5:2 and setose: clypeus yellowish, setose; palpus brown, length of apical seg- 
ment subequal to basal three segments; antenna (2+14) with scape and pedicel and first two flagel- 
lar segments cream-colored with base of darker distal segments narrowly infuscated giving antenna 
a ringed appearance; scape and pedicel with strong dorsal apical setae; ventral apical setae of pedicel 
short and stout. Thorax with mesonotum yellowish with brown acrostichal stripes from midway 


! Received February 10, 1995. Accepted August 10, 1996. 
2 Division of Natural Sciences, Southampton College, Southampton, N.Y. 11968. 


ENT. NEWS 108(2): 151-154, March & April, 1997 


152 ENTOMOLOGICAL NEWS 


that are contiguous anteriorly with broad humeral stripes, dorsocentral stripe narrow or absent; 
pleura variably pigmented yellowish or brownish except katepisternite that is always darker but 
light ventrally; pleurotergite with a pair of long central setae; scutellum and postnotum concolorous 
with pleura; long scutellar setae in a row; posterior postnotum with an irregular row of large, long 
setae. Wing: (3.0-3.2mm); patterned, (Fig. 2); 1stA barely divergent from Cu and ending beyond 
fM as a line of setae; 2dA nearly obsolete, represented by a row of setae and paralleling basal 
posterior wing margin; fully trichiate. Halter with stem yellow, knob variously pigmented but at 
least partially darkened. Legs with coxae brownish, anterior forecoxa, distal 2/3 of midcoxa, lateral 
apical fourth of hindcoxa setose, hindcoxa with a single long basal posterior seta; femora lighter 
with tibiae and tarsi yellowish; tibial spurs long, foretibia with an apical comb; foretibia/ 
forebasitarsus/ tarsus 2, 6/8/5. Abdomen orange yellow, very setiferous; TVIII (Fig. 3): SVIII (Fig. 
4). Terminalia: (Fig. 5, dorsal). 

Female. Description as for male. Terminalia: The paired cerci (fig. 6,c, lateral, setae omitted), are 
bore on tergite X (fig. 6, TX, dorsal,) which is bare and lightly pigmented. 


Material examined: Holotype, male, Ohio: Pike Co., Benton Twp., 4/8/79, GA Dahlem. In the 
collection of Michigan St. University. 

Allotype. Female. Same data as the holotype. 

Paratopotypes: 1 male and | female. 


DISCUSSION 


C. antennulata is most closely related to the Brazilian brunneicauda Matile, 
1996 from which the male of the new species may be easily distinguished by 
the form of its dark broadly bifid internal style and lack of median dorsal styles. 
The shape of the female cercus is closest to that of C. annulicornis as figured by 
Matile (loc. cit.), but with the apex of the basal cercal segment distinctly nar- 
rowed in C. antennulata. 

Vockeroth (1981), ina key to genera of the Mycetophilidae, noted a Cluzobra 
from Louisiana. Inquiries concerning this material have been to no avail; its 
identity cannot presently be determined. Thus, antennulata is the first Nearctic 
species of Cluzobra to be described. 


Biology: Limited capture data suggest that October to February may be the 
peak activity period for many of the continental South American species 
although aitkeni, binocellaris and spinulifera have been collected virtually 
throughout the year. Nothing is known of the bionomics. 


ACKNOWLEDGMENTS 


R.L. Fischer of Michigan State University made this material available for study. Annette 
Vollers prepared the drawings. 


LITERATURE CITED 


Edwards, F.W. 1934. New Neotropical Mycetophilidae (III). (Diptera). Rev. Entomol. 4(3):354- 
372. Figs. 1-5. 

Edwards, F.W. 1940. New Neotropical Mycetophilidae (1V). (Diptera). Rev. Entomol. 11 (1-2):440- 
465, Pls. 18-19. 


Vol. 108, No. 2, March & April, 1997 153 


Cluzobra 


SVIII TVIII 


Head. Lateral do terminalia. Dorsal view 


Plate. Cluzobra antennulata n.sp. Fig. |. Head, lateral view. Fig. 2. Wing. Fig. 3. Tergite VIII, male. 
Fig. 4. SVUI, male. Fig. 5. Male terminalia, dorsal aspect. Fig. 6. Cercus (c, lateral) and tergite X 


(TX, dorsal), female. 


154 ENTOMOLOGICAL NEWS 


Edwards, F.W. 1941. Mycetophilidae (Diptera) collected by the expedition to Matto Grosso of the 
Brazilian Zoological Club, in July 1939. Rev. Entomol. 12(1-2):303-314, Pl. 15. 

Lane J. 1948. “Mycetophilidae” do Brasil (Diptera, Nemocera). Rev. Brasil. Biol. 8(2):247-254, 
Figs. 1-15. 

Lane, J. 1956. New Neotropical Sciophilinae (Diptera: Mycetophilidae). Dusenia 7(3): 119-124, 
Figs. 1-8. 

Lane, J. 1959. Insecta Amapaensia. -Diptera: Mycetophilidae. (Second Contribution). Studia Ent. 
2(1-4): 105-118, Figs. 1-7. 

Lane, J. 1960. Mycetophilidae from Trinidad, B.W.I. (Diptera, Nematocera). Studia Entomol. 
3(1-4):375-384, Figs. 1-9. 

Lane, J. 1961. Further New Neotropical Mycetophilidae. (Diptera, Nematocera). Rev. Brasil. Ento- 
mol. 10:1-15, Figs. 1-5. 

Matile, L. 1996. Révision des Cluzobra Néotropicaux (Diptera: Mycetophilidae). Ann. Soc. Ento- 
mol. France (N.S.) 32:3-57, Figs. 1-75, 1 Tbl. 

Vockeroth, J.R. 1981. /N Manual of Nearctic Diptera Vol. 1. Research Branch Agriculture Canada, 
Monograph No. 27:223-246, Figs. 1-103, 


Vol. 108, No. 2, March & April, 1997 155 


FIRST RECORD OF ANTHOPOTAMUS VERTICIS 
AND NEW RECORDS OF A. DISTINCTUS 
(EPHEMEROPTERA: POTAMANTHIDAE) 

IN NEW ENGLAND! 


Steven K. Burian2 


ABSTRACT: The species Anthopotamus verticis is recorded for the first time in New England from 
the Housatonic River in western Connecticut. Notes on the larval habitat and observations on the 
sympatric occurrence of this species with A. distinctus are presented. New distribution records for 
A. distinctus are presented for Massachusetts and Vermont. 


In North America the family Potamanthidae is represented by a single ge- 
nus, Anthopotamus McCafferty & Bae, which contains four species and is broadly 
distributed over central and eastern North America (Bae & McCafferty 1991). 
Currently only one species, A. distinctus (Traver), has been reported from New 
England (Bae & McCafferty 1991, Burian and Gibbs 1991). Throughout New 
England A. distinctus occurs unpredictably in medium to large streams with 
loose cobble bottoms. In October of 1994, routine sampling of a river in west- 
ern Connecticut produced specimens of Anthopotamus larvae that were clearly 
not those of A. distinctus. In June of 1995, a series of specimens was reared 
from this site and determined to be A. verticis (Say). This is the first record of 
this species in New England. The new record is presented here, along with 
notes on the larval habitat and its co-occurrence with A. distinctus. New distri- 
bution records are presented for A. distinctus in Massachusetts and Vermont. 


METHODS 


Benthic samples were obtained with a D-frame aquatic net and standard 
kick technique. Live material for rearing was placed in foam cups and chilled 
for transport to the laboratory. Other specimens were kill-fixed by placing them 
directly into a modified Carnoy’s fluid (Edmunds et al. 1976) and later trans- 
ferred to 80% ethanol. Reared adults and larvae were identified using the most 
recent taxonomic keys (Bae & McCafferty 1991). Specimens of A. verticis were 
deposited in the Ephemeroptera Research Collection at Southern Connecticut 
State University. New locality records for A. distinctus were based on speci- 
mens recorded by Betsy Colburn of the Massachusetts Audubon Society, Steve 
Fiske of the Vermont Department of Environmental Conservation (DEC), and 


1 Received May 16, 1996; Accepted September 17, 1996. 


2 Dept. of Biology, Southern Connecticut State University, 501 Crescent Str., New Haven, CT 
06515. 


ENT. NEWS 108(2): 155-158, March & April, 1997 


156 ENTOMOLOGICAL NEWS 


specimens in the insect collection of the Peabody Museum, Yale University. 
Voucher specimens supporting new localities of A. distinctus were deposited 
with each of the three collections. For specimens from the Vermont DEC, refer- 
ence collection code numbers are cited as part of the new locality records. Lati- 
tude and longitude coordinates were not available for new Massachusetts records. 
Coordinates for Vermont sites were taken from U.S.G.S. 7.5' Quadrangle Maps. 
The Connecticut coordinates were determined using a Trimble Geoexplorer® 
G.P.S. receiver and checked using a U.S.G.S. 7.5' Quadrangle Map. 


New Potamanthidae Record 


Anthopotamus verticis (Say) 1839 


Specimens: 7 Imagos (3 Males, 4 Females - Reared); 2 Subimagos (Male - Reared); and 6 Larvae, 
S.K.Burian 16-VI-1995; 7 Larvae, S.K.Burian 14-X-1994. 


CONNECTICUT: Litchfield Co., Housatonic River at Housatonic Meadows State Park (Fly fish- 
ing Area) off Route 7[41°49'25"N/73°22'39"W]. 


Habitat: The channel of the Housatonic River at the site of the new record is 
about 40 to 45 m wide. Depth was variable because flow was regulated by a 
hydroelectric dam upstream of the collection site. Larvae of A. verticis were 
collected from side-channel areas in water about 0.5 m deep during low-flow 
conditions. Substrates at these locations were dominated by a mixture of me- 
dium to large cobbles. Larger particles were covered by a thick periphyton layer. 
Deposits of poorly-sorted medium gravel and sand occurred on the downstream 
sides of piles of cobbles and small boulders. Packs of coarse particulate organic 
matter were lodged among the cobbles in places. Filamentous green algae was 
present in tufts on the tops of some of the larger particles. Beneath the deposits 
of gravel and sand were thick deposits of fine sediments composed of gray- 
brown fine sand mixed with silt. 

Most larvae were obtained from piles of mixed cobbles overlaying deposits 
of poorly sorted gravel and sand in areas out of the main flow. No specimens 
were obtained from similar substrates in the deeper and swifter mid-channel 
area. 

At this locality, larvae of A. verticis share the river with larvae of A. distinctus. 
Larvae of both species were obtained simultaneously in kick samples. Field 
counts of larvae in samples obtained in October 1994 and June 1995 indicated 
that A. distinctus was more abundant on both dates than A. verticis. In fact, very 
few larvae of A. verticis were obtained in June of 1995 after repeated sampling, 
whereas numerous specimens of A. distinctus occurred in almost every sample. 
Although the life history and microhabitat preference of A. verticis have been 
studied (McCafferty & Bae 1994, Bae & McCafferty 1994) little is known about 
multiple species interactions. The co-occurrence of A. verticis and A. distinctus 


Vol. 108, No. 2, March & April, 1997 157 


at this site provides a unique opportunity to investigate their interactions and 
microhabitat distribution patterns. 


New Localities for A. distinctus 


A. distinctus has a range that spans the New England States; however, this is 
based on only a few published locations. Many suitable habitats between known 
localities are unoccupied, resulting in an erratic distribution pattern. The fol- 
lowing new records provide a clarification of the actual distribution of A. 
distinctus in New England: 


MASSACHUSETTS: Franklin Co., (Larvae, Female Imago) East Branch of the North River, 
Colrain, B. Colburn and F. Garretson 9-VII-1992, 15-XI-1994, 23-VI-1995; (Larvae) Burrington 
Brook (headwaters of the West Branch of the North River), B. Colburn and F. Garretson 15-XI- 
1994; (Larvae) West Branch of the North River, B. Colburm and F. Garretson 17-XI 1994; (Larvae) 
Green River, B. Colburn and F. Garretson 17-XI-1994. 

VERMONT: Addison Co., (Larvae) Lewis Creek, code 91.046a [44° 14'55"N/73°14'01"], 29-IX- 
1991; Bennington Co., (Larvae) North Branch of the Deerfield River, code 89.075a&b [42°54' 
50"'N/72"S0'16'""W], 4-X-1989; (Larvae) Walloomsac River, code 89.077a [42°55'08"'N/73° 16'08''W], 
5-X-1989; Caledonia Co., (Larvae) Passumpsic River, code 90.041 a [44°23’37"'N/72°01'23"'W],2- 
VIII-1990; Chittenden Co., (Larvae) Lamoille River, code 92.055a&b streams L44°41'00"N/ 
73°03'S7"W)], 29-IX-1992; (Larvae) Winooski River, code 91.023a [44°20'35"N/72°46'20"'W], 5- 
IX-1991; Franklin Co., (Larvae) Missisquoi River, code 91.062b [44°49'10''N/72°39'50"'W], 4-X- 
1991; Lamoille Co., (Larvae) Lamoille River, Hyde Park, [44°35'30"'N/72°38'30"'W], W.G. Downs 
20-VI-1975; Orange Co., (Larvae) First Branch of the White River, code 92.011b [43°58'34"N/ 
72°27'24""W], 1-I[X-1992; (Larvae) Connecticut River, code 92.034b [44°09'05'/72°02'32"'W], 17- 
IX-1992; Windam Co., (Larvae) Saxtons River, code 92.112b [43°08'27"'N/72°30'02"W], 4-X- 
1993; (Larvae) Williams River, code 92.043a&b [43°11'30"N/72°29'17""W], 18-I[X-1992; Wind- 
sor Co., (Larvae) White River, code 88.083a&b [43°49'30"N/72°34'00"W], 26-IX-1988. 


New locality records for Massachusetts and Vermont all occur in streams 
between the Connecticut River and the Hudson River. Records for Vermont are 
the most extensive for any of the New England States and show A. distinctus to 
be broadly distributed with populations in all major physiographic regions. In 
southern New England the only records east of the Connecticut River were 
those reported by Burian and Bednarik (1994). In Maine A. distinctus has only 
been recorded from streams in Aroostook County (Burian and Gibbs 1991). 
Currently, no records are available for New Hampshire and Rhode Island. 


ACKNOWLEDGMENTS 


I would like to thank Betsy Colburn and Fran Garretson of the Massachusetts Audubon 
Society for providing the data on A. distinctus in the Deerfield River System, Steve Fiske 
and his staff for generously providing specimens and site data from Vermont, and Leonard 
E. Munstermann of the Peabody Museum, Yale University, for the loan of specimens from 
the entomology collection. 


158 ENTOMOLOGICAL NEWS 


LITERATURE CITED 


Bae, Y.J. and W.P. McCafferty. 1991. Phylogenetic systematics of the Potamanthidae (Ephe- 
meroptera). Trans. Am. Entomol. Soc. 117: 1-143. 

Bae, Y.J. and W.P. McCafferty. 1994. Microhabitat of Anthopotamus verticis (Ephemeroptera: 
Potamanthidae). Hydrobiol. 288: 65-78. 

Burian, S.K. and A.F. Bednarik. 1994. The mayflies (Ephemeroptera) of Connecticut: An initial 
faunal survey. Entomol. News 105: 204-216. 

Burian, S.K. and K.E. Gibbs. 1991. Mayflies of Maine: an annotated faunal list. Maine Agricul- 
tural Exp. Sta. Tech. Bull. No. 142, 109 p. 

Edmunds, G.F., Jr., S.L. Jensen, and L. Berner. 1976. The mayflies of North and Central America. 
Univ. Minnesota Press, Minneapolis, 330 p. 

McCafferty, W.P. and Y.J. Bae. 1994. Life history of Anthopotamus verticis (Ephemeroptera: 
Potamanthidae). Great Lakes Entomol. 27: 57-67. 

Say, T. 1839. Description of new North American neuropterous insects, and observations on some 
already described. J. Acad. Nat. Sci. Phil. 8: 9-46. 


Vol. 108, No. 2, March & April, 1997 159 


FIRST ALABAMA RECORD OF THE PALE- 
BORDERED COCKROACH, PSEUDOMOPS 
SEPTENTRIONALIS (DICTYOPTERA: 
BLATTELLIDAE)! 


T’ai H. Roulston, Arthur G. Appel2 


ABSTRACT: Pseudomops septentrionalis was collected at two sites near Auburn, Alabama during 
the spring and summer of 1996, and appears well established locally. These sites are approximately 
560 kilometers east of the published range for the species and probably reflect accidental human 
introduction. 


Pseudomops septentrionalis Hebard is an outdoor cockroach most com- 
monly found on foliage and in ground litter (Helfer 1972). In May, 1996 one 
male P. septentrionalis was captured during the day at the Auburn University 
Arboretum (Auburn, Lee County, Alabama). The individual was perched 1 m 
off the ground on the foliage of privet (Ligustrum sp.), which grew along a 
small stream. In June, 1996 we returned to the same site and caught 13 P. sep- 
tentrionalis (two females, 11 males) in less than two hours by scanning the 
vegetation at night with a flashlight. All individuals collected were perched 
0.5 - 2 m above the ground on the foliage of various herbs and shrubs. No P. 
septentrionalis came to two black lights set up adjacent to the same area during 
the same evening. Also, no P. septentrionalis were caught in five, adjacent boll 
weevil pheromone traps, as they have been in Louisiana (Chapin 1980). 

Another specimen was taken in June, 1996 in ground litter at an Auburn 
University Fish Hatchery pond approximately eight kilometers from the arbo- 
retum. A specimen from Lee County was also turned in to an Auburn Univer- 
sity entomology course during the 1996 spring quarter (J.T. Vogt, pers. comm.). 
All specimens were adults and agreed with Helfer’s (1972) description. Speci- 
mens are deposited in the Auburn University Entomological Museum. 

The genus Pseudomops contains 40 species (Princis 1969), all of which are 
New World and most of which are tropical. Only P. septentrionalis reaches the 
United States. It has also been collected in Costa Rica (Princis 1969), and at 
least three states of northern Mexico (Atkinson et al. 1991). 

Hebard (1917) initially described P. septentrionalis from Texas, but later 
also recorded specimens from southern Oklahoma (Hebard 1938). Chapin (1980) 
published a range extension into much of Louisiana and Atkinson et al. (1991) 
extended the range into the southwestern corner of Arkansas. Since these are 


! Received July 29, 1996. Accepted August 28, 1996. 
2 Department of Entomology, Auburn University, Auburn, Alabama 36849-5413. 


ENT. NEWS 108(2): 159-160, March & April, 1997 


160 ENTOMOLOGICAL NEWS 


the first collections reported from Alabama and since none has been reported 
from Mississippi, which separates Alabama from the rest of the range of P. 
septentrionalis, we consider local collections of the species to reflect recent 
human activity rather than a natural range extension. 

The adventive introduction and establishment of cockroaches by human 
transport is common among distant exotic species but apparently rare among 
native American species into new regions of the United States. Atkinson et al. 
(1991) consider 69 species of cockroaches to have breeding populations within 
the continental United States and Canada. Of those species, 24 are considered 
exotic species introduced through human transport. The authors state that there 
is no evidence of native species having extended their range in that way. If P. 
septentrionalis remains established in Auburn and spreads further into Alabama, 
and if we are correct in treating their occurrence as adventive rather than as an 
overlooked, eastern arm of their range, then this could be the first case of a 
native American cockroach spreading in the United States by means of human 
transport. 


ACKNOWLEDGMENTS 


The authors wish to thank Eddie Watkins for providing the specimen from the Fish Hatch- 
ery. We also wish to thank Robert Rush for permitting us access to the Auburn Arboretum for 
research. This is Alabama Agricultural Experiment Station Journal No. 17-965251. 


LITERATURE CITED 


Atkinson, T.H., P.C. Koehler, and R.S. Patterson. 1991. Catalog and atlas of the cockroaches 
(Dictyoptera) of North America North of Mexico. Misc. Publ. Entomol. Soc. Amer. 78:1-85. 

Chapin, J.B. 1980. New distribution records for Pseudomops septentrionalis Hebard and Panchlora 
nivea (L.) (Dictyoptera: Blatellidae, Blaberidae). Proc. Entomol. Soc. Wash. 82:335-336. 

Hebard, M. 1917. The Blattidae of North America north of the Mexican boundary. Mem. Am. 
Entomol. Soc 2:1-284 I-vi 10 pl. 

Hebard, M. 1938. An ecological survey of the Orthoptera of Oklahoma. Okla. Agric. Exp. Stn. 
Tech. Bull. 5:1-31. 

Helfer, J.R. 1972. How to know the grasshoppers, cockroaches, and their allies, 2nd ed. Wm. C. 
Brown, Dubuque, Iowa. 

Princis, K. 1969. Blattaria: subord. Epilamproidea: fam. Blattellidae, pars 13: pp. 1041-1224. In: 
M. Beier [ed.], Orthopterorum catalogus. Junk, The Hague. 


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QL 
46 | 


EX 2 
7 3 / species of Atopsyche (Trichoptera: Hydro- 
ENT  piosidae) from Costa Rica R.J. Blahnik, R.M. Gottschalk 161 
Neoneides muticus (Heteroptera: Berytidae): host 
plants & seasonality in mid-Appalachian shale 
barrens A.G. Wheeler, Jr. 175 
Paddlefish, Polyodon spathula, as samplers of riffle 
beetles (Coleoptera: Elmidae) 
S.G. George, T.J. Hoover, H.P. Brown 179 
Notes on ant Eurhopalothrix floridana, with descrip- 
tion of male (Hymenoptera: Formicidae) 
M. Deyrup, C. Johnson, L. Davis 183 
Distribution & abundance of Eulonchus marialiciae 
(Diptera: Acroceridae) P. H. Adler, S.R. Reitz, C.N. Watson 190 
Ephemeroptera of Spring Creek, Oklahoma, with 
remarks on notable records 
W.P. McCafferty, R.K. Heth, R.D. Waltz 193 
Southward range extension of common ringlet, 
Coenonympha tullia inornata (Lepidoptera: 


Satyridae) David C. Iftner 201 
Efficiency of arthropod extraction from soil 
cores R.M. Snider, R.J. Snider 203 


A new Spiroplasma (Entomoplasmatales: Spiroplasmata- 
ceae) record for Georgia, & attempted horizontal 
transmission via predation 
J. Wedincamp, Jr., F.E. French, R.F. Whitcomb 209 
Preliminary list of Bruchidae (Coleoptera) of 
Cuba D.A. Marin, J.M. Kingsolver 215 
Cercopoidea types of species described by E. Schmidt in 
U.S. National Museum Natural History) with lecto- 
type designations (Homoptera: Cercopoidea) Ai-Ping Liang 222 
Amblycerus schwarzi (Coleoptora: Bruchidae) attacking 
seeds of tropical-almond terminalia in Cuba 
J. Genaro, J.M. Kingsolver 229° 
Seasonal distribution of Embolemidae (Hymenoptera) in 


central & northern Arkansas C.N. Lewis, J.B. Whitfield 231 
Ergatandromorphism in Odontomachus clarus (Hymen- 
optera: Formicidae) Juan A. Rodriguez-Garza 236 


Abundance & seasonal activity of Cantharidae, Lam- 

pyridae & Lycidae (Coleoptera) in a raspberry 

plantation in southern Quebec C. Levesque, G.-Y. Levesque 239 
SCIENTIFIC NOTES: 
Chamaemyiids as predators of Diuraphis noxia (Homop- 

tera: Aphididae) in Konya Province, Turkey Meryem Elmali 174 
Anthicus unicolor (Coleoptera: Anthicidae) a new preda- 


tor of Diuraphis noxia from Turkey Meryem Elmali 208 
Female Phengodes (Coleoptera: Phengodidae) feeding 

and an associated risk Richard S. Miller 215 
OBITUARY: Marvin H. Brunson 178 
BOOK REVIEWS 189 
BOOKS RECEIVED & BRIEFLY NOTED 221, 228,238 


SOCIETY MEETING OF FEBRUARY 19, 1997 182 


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Vol. 108, No. 3, May & June, 1997 161 


NEW SPECIES OF ATOPSYCHE FROM COSTA RICA 
(TRICHOPTERA: HYDROBIOSIDAE)! 


Roger J. Blahnik, Robert M. Gottschalk 


ABSTRACT: Three new species in the caddisfly genus (Trichoptera: Hydrobiosidae) from Costa 
Rica are described and illustrated: A. jaba, A. minimajada, and A. tapanti. Eleven additional 
species are known from the country. Illustrations of male genitalia and wing venation are pro- 
vided for the new species. A correction is made in the distribution of A. trifida, mistakenly re- 
ported by Schmid (1989) to occur in Costa Rica. 


RESUMEN: Tres nuevas especies de (Trichoptera: Hydrobiosidae) de Costa Rica son descritas e 
ilustradas: A. jaba, A. minimajada y A. tapanti. Ilustraciones de la venacion de las alas y de los 
genitales de los machos de estas nuevas especies son incluidas. Once especies mas son reportadas 
pare el pais. Finalmente, se presenta una correccion a la distribucion de A. trifida, erroneamente 
registrada por Schmid (1989) en Costa Rica. 


The caddisfly genus Atopsyche is very species diverse and predominantly 
Neotropical in distribution, ranging from northern Argentina to Texas and Ari- 
zona (Ross and King 1952). Larvae are typically found in cool streams and are 
free-living and predacious, characterized by unusual chelate front legs, un- 
doubtedly adapted for securing prey (Wiggins 1996). Larval morphology is 
illustrated by Wiggins (1996). The genus was first revised by Ross and King 
(1952) when less than 30 species were known. Biogeographical and evolution- 
ary relationships of the species were also discussed by Ross and King and 
expanded on by Ross (1953), who also described several additional species. 
Their biogeographical conclusions require reassessment in light of more re- 
cent developments in the field. Schmid (1989) presented a complete treatment 
of the family Hydrobiosidae, including a review of the genus and also de- 
scribed many new species. He criticized the phylogenetic framework erected 
by Ross and King, but chose to retain it because a phylogenetic analysis of the 
genus was outside the scope of his work. Acomprehensive phylogenetic analysis 
of the genus is needed before relationships between species can be stated with 
confidence. Although the diversity in the genus is much better understood than 
when Ross and King did their revision, additional species continue to be de- 
scribed and undoubtedly many more remain to be discovered. Schmid (1989) 
commented that a more thorough knowledge of the genus may be necessary 


! Received August 29, 1996. Accepted October 19, 1996. 


2 University of Minnesota, Department of Entomology, 219 Hodson Hall, 1980 Folwell Avenue, 
St. Paul. Minnesota 55108-6125 USA. 


3 Louisiana State University, Department of Plant Biology, 502 Life Sciences Building, Baton 
Rouge, Louisiana 70803-1705 USA. 


ENT. NEWS 108(3): 161-174, May & June, 1997 - 


\ 


AUG 18 1997 


LIBRARIES 


162 ENTOMOLOGICAL NEWS 


for a meaningful phylogenetic treatment. Descriptions of new species contrib- 
ute to that base of knowledge and therefore serve as an important prelude to a 
new phylogenetic analysis of the genus. Since the treatment of the genus by 
Schmid, additional new species from Ecuador were described by Sykora (1991) 
and a new species from Haiti by Botosaneanu (1991). With the inclusion of the 
three species described here, 116 species have been described to date. The 
species descriptions presented here represent part of an ongoing research project 
by Ralph W. Holzenthal, University of Minnesota, and his colleagues to fully 
document and describe the caddisfly fauna of Costa Rica. 

Including the three new species described below, 14 species are now known 
to occur in Costa Rica. The 11 previously described Costa Rican species are: 
A. callosa (Navas, 1924), A. chimpuocllo Schmid 1989, A. cira (Mosely, 1949), 
A. dampfi Ross and King 1952, A. erigia Ross 1947, A. huainacapac Schmid 
1989, A. implexa (Navas. 1924), A. majada Ross 1947, A. pachacamac Schmid 
1989, A. paucartampu Schmid 1989, and A. talamanca Flint 1974. Species of 
Atopsyche in Costa Rica have been collected between 30 m and 3120 m above 
sea level. However, most of the Costa Rican species were collected at middle 
elevations between 1400 m and 1650 m. 

Schmid (1989) listed A. trifida Denning as occurring in Costa Rica, prob- 
ably as a lapsus. The holotype locality was given as Puerto Rico by Denning 
(1948). Despite intensive collecting by Holzenthal, A. trifida has not been found 
in Costa Rica, and we have concluded that this species does not occur in Costa 
Rica. 

Terminology used to describe the male genitalia follows Schmid (1989). A 
summary of genitalic terminology can be found in Fig. 1. Holotypes for the 
three species are deposited in the collection at the National Museum of Natu- 
ral History [NMNH], Smithsonian Institution, Washington DC. Paratypes for 
A. jaba and A. minimajada are deposited in the University of Minnesota Insect 
Collection, St. Paul, Minnesota [UMSP], and at Costa Rica’s National 
Biodiversity Institute [INBIO], as stated in the species descriptions. Unless 
otherwise stated, all specimens are represented by pinned material. 


DESCRIPTION OF NEW SPECIES 


Atopsyche jaba, NEW SPECIES 
Figs. IA-D, 4A-B, 7 


Atopsyche jaba most closely resembles A. boneti Ross and King (1952), A. 
cordoba Denning (1968), and A. dampfi Ross and King (1952), especially in 
the armature of the parapods. However, the new species can be distinguished 
from the others by the shape of the first article of the inferior appendage, which 
is narrower, projects dorsally, and lacks any trace of an apicoventral projec- 
tion. There are also differences in the structure of the apical appendages of the 


Vol. 108, No. 3, May & June, 1997 163 


al 
sy 


SANs 
\ 


5 
sue 


\ 
WAS A 


a\ 
AAA 
VA 

RNey 

SA 


! 


“inf app 


Fig. 1. Atopsyche jaba n. sp., male genitalia: A, segments IX, X, and proctiger, lateral; B, phallic 
apparatus, lateral; C, phallic apparatus, ventral; D, inferior appendage, ventral. Terminology: IX 
= segment IX, proc = proctiger, par = parapod, fil = filipod, pr app = preanal appendage, inf app 
= inferior appendage, phal = phallotheca, end = endotheca, aed = aedeagus, car = anterior carina 
of phallotheca. 


164 ENTOMOLOGICAL NEWS 


phallotheca. Unlike A. cordoba and A. dampfi, A. jaba lacks distinctive black 
scales associated with a thickened base of Cu2 on the hindwing, but like those 
species has elongate, curled setae emerging from the first anal cell of the 
hindwing (Fig. 4B). 


ADULT: Forewing length 5.5-6.0 mm (male). Overall body color yellowish brown to 
light brown; antennae fuscous, yellowish basally; setae of palps fuscous. Wings light brown, 
erect setae of veins forming mottled pattern of alternate dark brown and amber colored se- 
tae, apex of wing fringed with light brown setae at apices of veins and dark brown setae 
between. Forewing venation (Fig. 4A) complete; Rj apparently unbranched. apically bor- 
dered by dense brown setae in region of pterostigma; fork I near apex of wing, fork II sessile; 
stem of M distinctly curved between m-cu crossvein and first fork of M; Cuz converging 
near fused anal veins (1A + 2A + 3A), with crossvein near apex forming small cell on poste- 
rior margin of wing (Fig. 4A). Hindwing with Rj incomplete, forked apically to Sc and stem 
of R2 + R3; forks II and IV absent; anal veins very strongly arched toward anal margin; male 
with first anal cell possessing elongate setae (Fig. 4B). Nygmas apparently absent from both 
wings. Terga III and IV of male (Fig. 7) each with pair of prominent, concavely rounded 
glands, lined internally with numerous short setae, located at anterolateral margins of re- 
spective terga. Sternum V of male with pair of small, convexly rounded glands, located at 
anterolateral margin of sclerite. Sterna VI and VII of male each with prominent spinelike 
ventral process on posteromesal margin; process on VII nearly straight, process on VI longer 
and curved posteriad, with row of short setae along anterior margin and terminating with 
large, blunt, spinelike seta. 

Male genitalia: Segment IX, in lateral view, narrow (Fig. 1A). Parapod with two promi- 
nent spines on dorsal edge, both slightly curved anteriad, the posterior spine smaller than 
anterior spine (Fig. 1A). Filipod long, slender, sparsely covered with elongate setae. Preanal 
appendage small, irregularly rounded, setose. Proctiger, in lateral view, broadly widened 
apically, covered externally with numerous minute setae; apicodorsal and lateral margins 
with elongate, coarse setae, sparse on lateral margin. Inferior appendage with first article 
constricted basally, otherwise relatively narrow and of uniform width, slightly curved mesad, 
angularly projecting apicodorsally, mesal surface with small preapical appendage; second 
article obliquely joined to first, slightly tapering and curved at apex. Phallotheca (Figs. 1B 
and 1C) with keel-like carina at anterior end; ventrally with short curved process articulat- 
ing with inferior appendages; apex deeply divided mesally, forming paired lobes, each rather 
broadly rounded at apex and associated externally with pair of elongate, narrow dorsal and 
ventral processes, dorsal ones distinctly sinuous, with apices acute and curved ventrad. 
Aedeagus an elongate, stout, spine-like structure, with distinct curvature. 

Type material: Holotype: OC, COSTA RICA: Puntarenas: Rio Jaba at rock quarry, 1.4 
km (air) W Las Cruces, 8.79°N, 82.97°W, 15.i11.1991, 1150 m, Holzenthal, Munoz, Huisman 
(NMNH). Paratypes: COSTA RICA: same location as holotype, 9.viii. 1990, 2 CO (UMSP), 
1 O (INBIO). 

Etymology: Named for the Jaba River, near the Wilson Botanical Gar- 


den, Las Cruces, Costa Rica. 
Atopsyche minimajada, NEW SPECIES 
Figs. 2A-D, 5A-B, 8A-B 


The wing and body color pattern and structure of the male genitalia of 
Atopsyche minimajada are very similar to Atopsyche majada Ross (1947). 
Specimens of the two species were initially confused by us. However, male 
genitalia of the two species have several characters which readily serve to sepa- 
rate them. Atopsyche minimajada, can be distinguished from A. majada by 
having the apicoventral lobe of the first article of the inferior appendage subequal 


Vol. 108, No. 3, May & June, 1997 165 


= 5) 
LDPE 1777) a 


Fig. 2. Atopsyche minimajada n. sp., male genitalia: A, segments IX, X, and proctiger, lateral; B, 
phallic apparatus, lateral; C, phallic apparatus, ventral; D, inferior appendage, ventral. 


166 ENTOMOLOGICAL NEWS 


in length to the second article, as opposed to being distinctly shorter in A. 
majada, and also by the shape of the parapods, which in A. minimajada have 
more prominent apices and are less constricted preapically (Fig. 2A). Addi- 
tionally, the elongate, recurved dorsal spine, associated with the base of the 
phallotheca in both species is differently formed; in A. minimajada the spine 
widens preapically and possesses numerous minute setae (Fig. 2B), whereas 
in A. majada the spine is narrow throughout its length and lacks any associated 
setae. The name for the new species derives from the fact that specimens we 
examined are smaller on average than those of A. majada. The size of A. 
minimajada, however, falls within the range of variability for A. majada and 
size alone is not sufficient to distinguish the two species. 


ADULT: Forewing length 5.1-5.5 mm (male), 5.8-6.7 mm. (female). Head, body, and legs 
pale yellow to light brown, except terga of abdomen fuscous. Setae of head whitish. Palps pale 
yellow with fuscous setae; antennae yellowish at base, fuscous apically. Wings light brown, with 
dark brown setae along costal margin, dense in region of pterostigma, and with patch of dark 
brown along anal margin at midlength; veins with erect setae, alternating in short segments be- 
tween whitish and dark brown. Forewing venation complete (Fig. 5A); Rj apparently unbranched 
apically, bordered by dense brown setae in region of pterostigma; fork I near apex of wing, fork 
II sessile; stem of M distinctly curved between m-cu and first fork of M; Cu? converging near 
fused anal veins (1A + 2A + 3A), with crossvein near apex forming small cell on posterior 
margin of wing. Hindwing (Fig. 5B) with Ry incomplete, forked apically to Sc and stem of Rj+ 
R3; forks II and IV absent. Male with Cup possessing elongate, thickened region in proximal 
part of vein; anal veins very strongly arched toward anal margin, base of 2A with brush of elon- 
gate setae extending over thickening of Cuz. Forewing with nygma near base of fork II, nygma 
apparently absent from hindwing. Tergum III of male (Fig. 8A) with pair of concavely rounded 
glands laterally on anteroventral margin. Sternum V of male (Fig. 8B) with pair of short, project- 
ing, saccate glands laterally on anterodorsal margin. Sterna VI and VII of male each with promi- 
nent spinelike ventral process on posteromesal margin; process on VII nearly straight, process on 
VI longer and curved posteriad, with row of short setae along anterior margin and terminating 
with large, blunt, spinelike seta. 

Male genitalia:: Segment IX, in lateral view, relatively narrow. Parapod nearly parallel sided, 
moderately inflected at middle, apicodorsally with subtriangular, spinelike process (more promi- 
nent than in A. majada . Preanal appendage small, rounded, with few setae. Filipod elongate, 
narrow, gradually widened apically, sparsely clothed with elongate setae. Proctiger, as viewed 
laterally, broadly widened apically, covered externally with numerous minute setae; dorsal, lat- 
eral and apical margins without elongate setae. Inferior appendage, as viewed laterally, with first 
article moderately wide, slightly bulging at middle; posteriorly, with narrow, projecting, 
apicoventral lobe, nearly equaling second article in length; in ventral view (Fig. 2D), with mesal 
curvature; second article narrow, emerging from apicodorsal margin of first article, narrowly 
separated from ventral lobe of first article and only slightly wider and longer. Phallic apparatus 
relatively simple, similar to A. majada (Figs. 2B, 2C); apex divided mesally; in lateral view, 
apical lobes slightly projecting ventrad and with numerous minute setae; phallotheca with elon- 
gate, recurved basal spine, branched at base and membranously attached to phallotheca, widen- 
ing preapically and covered with numerous minute setae; aedeagus forming elongate curved 
spine with basal enlargement. 

Type Material: Holotype: O, COSTA RICA: Guanacaste: Estacion Pitilla, Rio Orosi, 
10.931°N’ 85.428°W, 700 m, 22-25.v.1990, Holzenthal and Blahnik (NMNH). 

Paratypes: COSTA RICA: Alajuela: Cerro Campana, Rio Bochinche trib., 6 km (air) NW 
Dos Rios, 10.945°N, 85.413°W, 600 m, 22-23.vii. 1987, Holzenthal, Morse, Clausen, | O (UMSP); 
Rio Pizote, ca. 5 km (air) S Brasilia, 10.972°N, 85.345°W, 390 m, 12.11.1986, Holzenthal and 


Vol. 108, No. 3, May & June, 1997 167 


Fasth, 4 0’, 14 9 (pinned), 12 C (in alcohol) (UMSP); Rio Pizote, ca. 5 km N Dos Rios, 10.948°N, 
85.291°W, 470 m, 9.iii.1986, Holzenthal and Fasth, I CO’ (in alcohol) (UMSP); Guanacaste: Parque 
Nacional Guanacaste, El Hacha, Quebrada Pedregal, 10.983°N, 85.539°W, 300 m, 27.vii.1987, 
Holzenthal, Morse, Clausen, 1 CO (UMSP); Parque Nacional Guanacaste, Estacion Maritza, Rio 
Tempisquito, 10.958°N, 85.497°W, 550 m, 30-31.vii.1990, Huisman, Blahnik, Quesada, I CO (in 
alcohol) (INBIO); Rio Gongora (sulfur mine), 4 km (air) NE Quebrada Grande, 10.887°N, 
85.470°W, 590 m, 21.vii.1987, Holzenthal, Morse, Clausen, | O (UMSP); Rio Los Ahogados, 
11.3 km ENE Quebrada Grande, 10.865°N, 85.423°W, 470 m, 7.iii.1986, Holzenthal and Fasth, 
40,5 Q (inalcohol) (UMSP); Heredia: Rio Bijagual on road to Magsasay, 10.408°N, 84.076°W, 
140 m, Holzenthal, Morse, Fasth, 1 CO, | Q (UMSP); Rio Sarapiqui, 7 km W Puerto Viejo, 
10.452°N, 84.067°W, 50 m 11.ii.1986 Morse and Fasth, 5 C (in alcohol) (UMSP); Puntarenas: 
Rio Ceibo, route 2, ca. 6 km W road to Buenos Aires, 9.149°N, 83.377°W, 250 m, 20.ii.1986, 
Holzenthal, Morse, Fasth, 2 O', 6 9 (pinned), 6 CO’, 115 Q (in alcohol) (UMSP); San José: Res. 
Biol. Carara, Rio del Sur, 1.5 km (rd) S of Carara, 9.769°N, 84.531°W, 160 m, 13.iii.1991, 
Holzenthal, Mufioz, Huisman, 1 O' (UMSP). 

Etymology: Named _ because of the relatively small size of specimens of 
this species when compared to those A. majada, and because it was the size of 


the specimens that first drew our attention to differences between the species. 


Atopsyche tapanti, NEW SPECIES 
Fig. 3A-D 


This species is most similar to Atopsyche paucartampu Schmid (1989), but 
can be distinguished by several subtle differences. The inferior appendage of 
A. tapanti, in ventral view, has the first article less sinuously bulging at the 
middle and the second article is much narrower than in A. paucartampu. Addi- 
tionally, the filipods and parapods of A. tapanti are shorter. 


ADULT: Forewing length 7.4 mm (male). Overall body color yellowish brown, slightly darker 
dorsally, terga of abdomen fuscous; palps yellowish, with scant brownish setae; antennae yel- 
lowish basally, brown apically. Overall color of wings brownish; erect setae of veins mostly 
black or blackish with interspersed segments of pale yellow setae. Forewing venation complete 
(Fig. 6A); Rj branched apically, covered with dense brown setae in region of pterostigma; fork I 
near apex of wing, fork II sessile, Cuz converging near fused anal veins (1A + 2A + 3A), with 
crossvein near apex forming small cell on posterior margin of wing. Hindwing (Fig. 6B) with Rj 
incomplete, fork at apex intersecting Sc and stem of R2 + R3; forks Il and IV absent; anal veins 
distinctly arched toward anal margin; anal margin of wing, 2A (sparsely), and base of anal veins 
with elongate setae. Nygmas apparently absent from both wings. Tergum III of male with pair of 
concavely rounded glands on anterolateral margin (as in Fig. 8A). Sternum V of male with pair of 
elongate, projecting glands on anterolateral margin (Fig. 9). Sterna VI and VII of male each with 
prominent spinelike ventral process on posteromesal margin, nearly equal in size and shape, 
process on VI without stout apical seta; 

Male genitalia: Segment IX, in lateral view (Fig. 3A), relatively wide basally, anterior mar- 
gin broadly rounded. Parapod short, apex rounded, with one small, blunt tooth on posterodorsal 
margin. Filipod very short, fingerlike. Preanal appendage small, rounded, setose. Proctiger, as 
viewed laterally, broadly widened apically, covered externally with numerous minute setae; lat- 
eral margin with several elongate setae, apex without setae. Inferior appendage, in lateral view, 
with first article broadly, evenly widened at middle, second article wide basally, apically narrow 
and mesally curved; in ventral view, first article mesally curved, with conspicuous, setose, 
thumblike projection near base on mesal surface, second article narrowed and slightly hooked 
apically. Phallic apparatus (Fig. 3B) simple, mesally bifurcated, forming paired apical lobes, 


168 ENTOMOLOGICAL NEWS 


os 


HUES 


ee eae 


COSY SSS 


Fig. 3. Atopsyche tapanti n. sp., male genitalia: A, segments IX, X, and proctiger, lateral; B, 
phallic apparatus, lateral; C, phallic apparatus, ventral; D, inferior appendage, ventral. 


Vol. 108, No. 3, May & June, 1997 169 


Figs. 4-6. Atopsyche n. spp., wings: A, forewing; B hindwing. 4. A. jaba; 5. A. minimajada; 6. A. 
tapanti. 


170 ENTOMOLOGICAL NEWS 


tergum IV 


8B sternum V 


tergum Ill 


_sternum V gland 


Figs. 7-9 Atopsyche n. spp., abdominal glands. 7. A. jaba, dorsal view of terga III and IV and 
associated glands; 8. A. minimajada, A, dorsal view of tergum III and associated glands, B, 
ventral view of sternum V and associated glands; 9. A. tapanti, new species, lateral view of 
segment V and associated gland. 


Vol. 108, No. 3, May & June, 1997 171 


broadly rounded at apex, as viewed laterally. Aedeagus a stout, dorsally curved spine with basal 
enlargement. 

Type MateriaL: Holotype: Male, COSTA RICA: Cartago: Reserva Tapanti, Rio Grande de 
Orosi, 9.686 N., 83.756 W, 1650m, 8-9.vii. 1986, Holzenthal, Heyn, Armitage (NMNH). 


Etymology: Referring to the type locality, Tapanti, National Park, Costa 
Rica. : 


RELATIONSHIPS OF THE COSTA RICAN SPECIES 


A foundation for the phylogenetic placement of species of Atopsyche was 
erected by Ross and King (1952) and Ross (1953), who used character simi- 
larities to define a number of species groups, all of which were placed into two 
subgenera, Atopsyche and Atopsaura. Schmid (1989) placed his newly described 
species, and also those described subsequently to Ross (1953), within the frame- 
work erected by Ross and King, based on the characters they used. However, 
as discussed in the introduction, Schmid noted that these were probably not 
natural groupings. He also reduced the genus Dolochorema to subgeneric sta- 
tus within Atopsyche, thereby creating a third subgenus. A newly recognized 
species group and several additional species were left incertae sedis within 
Schmid’s taxonomic outline. 

It is possible to place the species described here within the existing taxo- 
nomic framework, based on the characters used to define the groups, and we 
have indicated these relationships below. However, in agreement with Schmid’s 
critique of the existing taxonomic system, we have refrained from formally 
placing the newly described species within the current taxonomic hierarchy. 
Instead, we have limited ourselves to a discussion of characters shared by spe- 
cies. This discussion is not intended as a justification for the phylogenetic 
placement of the species; rather it is intended to provide information that may 
be of use in an eventual cladistic revision. Despite our reservations about the 
existing taxonomic system, it is undoubtedly true that a number of species 
currently placed together are closely related. This statement is based on the 
fact that some species share a number of ostensible character similarities, at 
least some of which are likely apomorphic. We discuss below character affini- 
ties of members of the Costa Rican fauna, and place the discussion within the 
contextual reference of the characters used by Ross and King (1952) to define 
species groups. 

Setose glandular structures located on both terga III and IV of males is a 
character used by Ross (1953) to characterize a subgroup of species in the 
subgenus Atopsyche, including A. implexa, A. banksi, and A. vatucra. He placed 
this subgroup within his kingi group or complex, a new name designation for 
the tkonnikovi group of Ross and King (1952). Schmid (1989) merged the 
kingi group with the bolivari group of Ross and King (1952). One of our new 
species, Atopsyche jaba, has males with setose glands located on terga III and 
IV, and on this basis would be included within the subgroup of species recog- 


172 ENTOMOLOGICAL NEWS 


nized by Ross. In addition to A. jaba and A. implexa, other species from Costa 
Rica that possess setose glands on both terga III and IV include A. pachacamac 
and A. huainacapac. Atopsyche pachacamac is similar to A. jaba and A. implexa 
in a number of respects, including overall coloration and the structure of the 
apex of the phallotheca. However, A. huainacapac is distinctly different. It 
was placed by Schmid (1989) in the /ongipennis group of Ross and King (1952), 
and in the subgenus Afopsaura, indicating a very distant relationship to these 
other species. It is possible that the glandular character is homoplasic in this 
species, or perhaps the character is primitive and not phylogenetically infor- 
mative. This is difficult to assess because the possession of glandular struc- 
tures is not often mentioned in species descriptions, despite the fact that all of 
the species of Atopsyche from Costa Rica have glands present on sternum V 
and also on tergum III, or on both terga III and IV, with the sole exception of A. 
chimpuocllo, which lacks prominent glands on both terga III and IV. However, 
in none of the other species from Costa Rica are the glands internally setose. 
Atopsyche dampfi, A. boneti, and A. cordoba, are all structurally very similar 
to A. jaba, A. implexa, and A. pachacamac. However, in the descriptions of 
none of these species is the presence of setose glandular structures mentioned. 
In A. dampfi, which we had the opportunity to examine, glands are present on 
both segments, but are much reduced in size and lack setation. Clearly a phy- 
logenetic analysis is required before these glandular structures, either by their 
morphology or by their presence or absence, can be confidently used to define 
groups. 

Atopsyche minimajada is undoubtedly very closely related to A. majada, 
which was placed by Ross and King (1952) in the batesi group of Atopsyche 
and in the subgenus Atopsaura by Ross (1953). The defining character of this 
group is the possession of an unpaired dorsal spine emerging from the base of 
the phallotheca. Species with this character, and in which the spine is bifurcate 
at the base and attached membranously to the phallotheca, probably form a 
monophyletic assemblage, since the character is distinctive enough to make 
parallel evolution unlikely. The batesi group includes all of the species de- 
scribed from the Greater Antilles and also several additional species with dis- 
tributions in Central America or the northern part of South America, extending 
as far south as Bolivia. Atopsyche callosa is the only additional species from 
Costa Rica included in this group. 

The last new species described in this paper, Atopsyche tapanti, is very 
similar to A. paucartampu Schmid, and the two undoubtedly form a closely 
related species pair. Schmid (1989) placed A. paucartampu in the tripunctata 
group of Ross and King (1952). Other species from Costa Rica with a general 
similarity to these two species include A. cira, A. talamanca, and A. chimpuocllo. 
The latter species was placed by Schmid (1989) in the bolivari group of Ross 
and King. However, characters which A. chimpuocllo share with the other four 
species listed above include the possession of a mesally-directed, thumblike 


Vol. 108, No. 3, May & June, 1997 173 


protrusion on the first article of the inferior appendage at midlength (Fig. 3D) 
and elongate glandular structures associated with sternum V (Fig. 9). There is 
also a general color similarity between the species mentioned above; all have a 
rounded, blackish spot extending from the costal margin of the wing, bordered 
by golden setae. As mentioned above, A. chimpuocllo lacks noticeable glandu- 
lar structures on either terga III or IV, whereas all of the other four species 
have rounded glandular structures without internal setation located on tergum 
III only. This last character is also found in the members of the batesi group 
found in Costa Rica (Fig. 8A), and is probably a plesiomorphic character. 

Atopsyche erigia, also placed by Ross and King in the tripunctata group, is 
different in a number of respects from the other Costa Rican species placed in 
this group. The character used by Ross and King to define the group is the 
possession of a second article of the inferior appendage that is simple in struc- 
ture and broad at the base. Atopsyche erigia was used as an archetypal ex- 
ample for the group. This character is rather superficial and species currently 
placed together on this basis may have only a spurious relationship. It seems 
likely that the tripunctata group is not a natural, or at least not a closely related 
assemblage of species. Unlike the other species from Costa Rica placed in the 
group, A. erigia has glandular structures on sternum V that are short, and 
rounded glandular structures without internal setation on both segments III 
and IV. Also, unlike the other species of the tripunctata group discussed above, 
the first article of the inferior appendage lacks a thumblike process on the 
mesal surface. On the other hand, in coloration A. erigia closely resembles A. 
majada, although it lacks the distinctive phallic spine that characterizes mem- 
bers of the batesi group. It is also now placed in a different subgenus. A close 
relationship of A. erigia to other species of the tripunctata group from Costa 
Rica is not well supported by character evidence. Hopefully, a cladistic analy- 
sis of the genus at some future date will sort out the seemingly contradictory 
character relationships existing among the species in the various subgenera 
and species groups. — 


ACKNOWLEDGMENTS 


This research was supported by National Science Foundation grants BSR-8917684 and DEB 
9400632, awarded to Ralph W. Holzenthal, and also by the University of Minnesota Under- 
graduate Research Opportunities Program and the College of Biological Sciences Honors Pro- 
gram at the University of Minnesota. A special thanks is due Ralph W. Holzenthal, who provided 
the illustrations of the wings of the new species and served as faculty advisor to the second 
author for the College of Biological Sciences Honors Program. The advice of Atilano Contreras, 
Victoria Nations, and Fernando Munoz, as well as the editing of Susan Weller and Frank Barnwell 
were greatly appreciated. Paper number 22,541 Scientific Journal Series, Minnesota Agricultural 
Experiment Station, St. Paul, Minnesota. 


174 ENTOMOLOGICAL NEWS 


LITERATURE CITED 


Botosaneanu, L. 1991. Trichoptéres d’ Haiti. Bulletin de L’ Institut Royal des Sciences Naturalles 
de Belgique 61: 113-134. 

Denning, D. G. 1948. A Review of the Rhyacophilidae (Trichoptera). Can. Entomol. 80: 97-115. 

Denning, D. G. 1968. New and interesting North American Trichoptera. Pan-Pacific Entomol. 
44: 17-26. 

Ross, H. H. 1947. Descriptions and records of North American Trichoptera, with synoptic notes. 
Trans. Amer. Entomol. Soc. 73: 125-168. 

Ross, H. H. 1953. Additional material on the phylogeny and dispersal of Atopsyche (Trichoptera: 
Rhyacophilidae). Journ Wash. Acad. Sci. 43: 287-293. 

Ross, H. H., and King, E. W. 1952. Biogeographic and taxonomic studies in Atopsyche 
(Trichoptera, Rhyacophilidae). Ann. Entomol. Soc. Amer. 45: 177-204. 

Schmid, F.N. 1989. Les Hydrobiosides (Trichoptera, Annulipalpia). Bulletin de L’ Institut Royal 
des Sciences Naturalles de Belgique 59 (supplement): 1-154. 

Sykora, J. L. 1991. New species of Hydrobiosidae from Ecuador (Insecta: Trichoptera: 
Annulipalpia). Ann. Carnegie Museum 60: 243-251. 

Wiggins, G. B. 1996. Larvae of the North American Caddisfly Genera (Trichoptera), Second 
Edition. Univ. Toronto Press, Toronto. 


SCIENTIFIC NOTE 


CHAMAEMYIIDS AS PREDATORS OF DIURAPHIS NOXIA 
(HOMOPTERA: APHIDIDAE) IN KONYA PROVINCE, TURKEY! 


Meryem Elmali2 


Three species in two genera of Chamaemyiidae (Diptera) have been found to be predators 
on Diuraphis noxia (Kurdjumov) colonies on wheat and other alternate host plants during four 
years (1989-1992) in Konya province, Turkey. Leucopis (Leucopis) pallidolineata Tanasijtshuk 
was previously reported from Turkey (Diizgiines et al. 1982) and as a predator of D. noxia 
(Tanasijtshuk 1984, Kovalev et al. 1991). Leucopis (Leucopis) kerzhneri Tanasijtshuk and 
Parochthiphila sp. are the first records in the world as predators of D. noxia. These two are also 
first records for Turkish fauna. 


LITERATURE CITED 


Diizgiines, Z., S. Toros, N. Kilincger, and B. Kovanci. 1982. Ankara ilinde saptanan afit predatorii 
Leucopis tiirleri (Dip.: Chamaemyiidae). Turk. Bit. Kor. Derg., 6: 91-96. 

Kovalev, O.V., T.J. Poprawski, A.V. Stekolshchikov, A.B. Vereshchagina, and S.A.Gandrabur. 
1991. Diuraphis Aizenberg (Hom., Aphididae): key to apterous viviparous females and re- 
view of Russian language literature on the natural history of Diuraphis noxia (Kurdjumov, 
1913). J. Appl. Entomol. 112: 425-436. 

Tanasijtshuk, V.V. 1986. [Silver flies (Chamaemyiidae)]. Keys to the fauna of the USSR. Diptera 
14 (7). Leningrad: Nauka. 


! Received September 16, 1996. Accepted December 10, 1996. 
2 Plant Protection Department, Faculty of Agriculture, Selcuk University, 42031 Konya, Turkey. 


Vol. 108, No. 3, May & June, 1997 N75 


NEONEIDES MUTICUS (HETEROPTERA: 
BERYTIDAE): HOST PLANTS AND SEASONALITY IN 
MID-APPALACHIAN SHALE BARRENS! 


A. G. Wheeler, Jr.2.3 


ABSTRACT: The stilt bug Neoneides muticus, a characteristic insect of shale barrens, was ob- 
served on moss phlox, Phlox subulata (Polemoniaceae), in 31 mid-Appalachian shale barrens 
and outcrops in Maryland, Pennsylvania, Virginia, and West Virginia. Overwintered adults mated 
and oviposited as early as mid-April, and first-generation adults appeared by late June. A small 
second generation was found on moss phlox at only two sites. Other hosts in shale barrens were 
a fern, Cheilanthes lanosa (Adiantaceae); the composite Aster oblongifolius (Asteraceae); a mint, 
Scutellaria ovata (Lamiaceae); and a beardtongue, Penstemon canescens (Scrophulariaceae). 
The record of N. muticus from moss phlox is the first for a berytid from the Polemoniaceae, and 
its association with woolly lipfern, C. /anosa, is the second for this stilt bug on a fern species. 


Neoneides muticus (Say), formerly placed in the Old World genus Neides 
Latreille (see Henry 1997a), is one of the most widely distributed North Ameri- 
can berytids. Its U.S. distribution includes all 48 conterminous states, and it 
occurs across much of southern Canada and ranges into northern Mexico (Henry 
1997b). Little biological information on this commonly collected stilt bug was 
available until its seasonality and habits were studied in New York and Penn- 
sylvania (Wheeler 1978). This bivoltine species develops on common mullein 
(Verbascum thapsus L.), adults overwintering in the basal rosettes. Mating and 
Oviposition in New York take place from late April to early May, and first- 
generation adults appear in late June to early July. A small second generation is 
produced during midsummer to early fall. Both phytophagous and zoophagous, 
N. muticus feeds on leaves and flower buds of common mullein and on the 
mullein thrips (Haplothrips verbasci (Osborn)). Other scrophulariaceous hosts 
are moth mullein (V. blattaria L.) and hairy beardtongue (Penstemon hirsutus 
(L.) Willd.) (Wheeler 1978, Wheeler and Henry 1981). Its hosts also include 
members of the Asteraceae (Hieracium aurantiacum L. and H. pratense Tausch) 
and Rosaceae (Potentilla norvegicus L. and Rubus sp.), as well as hayscented 
fern (Dennstaedtia punctilobula (Michx.) Moore) (Wheeler 1978). 

Here I report the use of several additional host plants by N. muticus, includ- 
ing another fern species and the first polemoniaceous plant, moss phlox (Phlox 
subulata L.), recorded as a stilt bug host. Its seasonal history on moss phlox in 
mid-Appalachian shale barrens and shale outcrops is summarized. 


| Received October 3, 1996. Accepted November 5, 1996. 
2 Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, PA 17110. 


3 Present address: Department of Entomology, Clemson University, Clemson, SC 29634. 


ENT. NEWS 108(3): 175-178, May & June, 1997 


176 ENTOMOLOGICAL NEWS 


STUDY SITES AND METHODS 


The seasonality of N. muticus was followed during a study of plant bugs 
(Heteroptera: Miridae) associated with moss phlox in shale barrens occurring 
from south-central Pennsylvania to southwestern Virginia (see Wheeler 1995). 
The sample sites included several classic shale barrens: Eagle Rock, Head Wa- 
ters, Millboro, and Short Mountain in Virginia, and Kates Mountain in West 
Virginia (see Wheeler 1995 and references therein). Sampling was conducted 
at irregular intervals from early April to early August 1989 to 1995. All stages 
of N. muticus that were shaken from mats of moss phlox were recorded. Obser- 
vations on additional host plants were made in several shale barrens that har- 
bored populations of N. muticus on moss phlox. Voucher specimens of the berytid 
are deposited in the collection of the National Museum of Natural History, 
Washington, D.C. 


RESULTS 


Phlox subulata 

Populations of N. muticus were observed on this plant in 31 shale barrens 
and outcrops in the mid-Appalachians, including 13 in West Virginia (Grant, 
Greenbrier, Hampshire, and Pendleton counties), 10 in Virginia (Alleghany, 
Bath, Botetourt, Highland, Roanoke, Rockbridge, and Shenandoah counties), 
6 in Maryland (Allegany and Washington counties), and 2 in Pennsylvania 
(Bedford Co.). 

Adults, which apparently overwintered under prostrate mats of moss phlox, 
began mating by mid-April. Oviposition on the sepals occurred as early as 
mid-April. Mating and oviposition continued in shale barrens until early May. 
First instars were found by late April and were present until late May. Over- 
wintered adults were not seen after mid-May. Mid- to late instars were often 
seen on moss phlox during June and July, adults of the first generation appear- 
ing by the last week of June. 

A small second generation was observed on moss phlox only at Smoke 
Hole in Pendleton Co., West Virginia, and in the George Washington National 
Forest in Rockingham Co., Virginia. Moss phlox colonies often brown during 
July and August, and in shale barrens N. muticus was always more numerous 
on more succulent hosts during midsummer. 


Other Hosts in Shale Barrens 

In addition to moss phlox, common first-generation hosts of N. muticus in 
Maryland, Virginia, and West Virginia were Appalachian beardtongue, Penste- 
mon canescens (Britt.) Britt. (Scrophulariaceae), and aromatic aster, Aster 
oblongifolius Nutt. (Asteraceae). Early instars of a second generation were 
present on P. canescens in Pendleton Co., West Virginia, in early July. In Bath 


Vol. 108, No. 3, May & June, 1997 177 


Co., Virginia, second-generation nymphs and adults were found on forest skull- 
cap, Scutellaria ovata L. (Lamiaceae), in mid-July. 

Woolly lipfern, Cheilanthes lanosa D.C. Eat., served as a host at the Romney- 
Oldtown shale barren in Allegany Co., Maryland. In early July 1994, fourth- 
and fifth instar nymphs and adults were observed on this fern in one area of the 
barren. The following year, overwintered adults and first through third instars 
were collected on woolly lipfern colonies throughout the barren in late May. 


DISCUSSION 


Although a common, polyphagous stilt bug species, NV. muticus should be 
considered a characteristic shade barren insect, occurring on moss phlox with 
specialists such as the mirids Polymerus tinctipes Knight and P. wheeleri Henry 
(Wheeler 1995) and the psyllid Craspedolepta eas (McAtee) (Wheeler 1994). 
The pattern of host use by N. muticus in mid-Appalachian shale barrens con- 
forms to that previously recorded (Wheeler 1978) and is typical of most other 
Berytidae—that is, glandular-hairy plants, mostly dicots. Moss phlox is the 
first member of the Polemoniaceae reported as a berytid host (see Wheeler and 
Schaefer 1982). In shale barrens, N. muticus develops on the glandular subspe- 
cies of this plant, Phlox subulata brittonii (Small) Wherry. Other shale barren 
hosts are similarly glandular, pubescent, or both, including the fern Cheilanthes 
lanosa, whose leaves are hirsute. This association, the second for N. muticus 
with a fern species (Wheeler 1978), is atypical within the Berytidae er 
and Schaefer 1982). 

Neoneides muticus does not often produce a second generation on moss 
phlox, which becomes essentially dormant in shale barrens during midsum- 
mer. Instead, this stilt bug develops on more succulent hosts, such as the late- 
blooming Aster oblongifolius. 


ACKNOWLEDGMENTS 


I thank T. J. Henry (Systematic Entomology Lab., USDA, ARS, PSI, c/o National Museum 
of Natural History, Washington, DC) for providing prepublication copies of his berytid studies, 
E. R. Hoebeke (Department of Entomology, Cornell University, Ithaca, NY) and G. L. Miller 
(SEL, USDA, ARS, PSI, c/o NMNH, Beltsville, MD) for useful comments on an early draft of 
the manuscript, and M. A. Berdine and D. A. Samson (The Nature Conservancy) for permitting 
access to Maryland’s Oldtown Shale Barren Preserve. 


LITERATURE CITED 


Henry, T. J. 1997a. Cladistic analysis and revision of the stilt bug genera (Heteroptera: Berytidae) 
of the world. Contr. Am. Entomol. Inst. 30(1): 1-100. 

Henry, T. J. 1997b. Monograph of the Berytidae, or stilt bugs (Heteroptera), of the Western 
Hemisphere. Mem. Entomol. Soc. Wash. in press. 


178 ENTOMOLOGICAL NEWS 


Wheeler, A. G., Jr. 1978. Neides muticus (Hemiptera: Berytidae): life history and description of 
fifth instar. Ann. Entomol. Soc. Am. 71:733-736. 

Wheeler, A. G., Jr. 1994. Craspedolepta eas: distribution, hosts, and habits of a phlox specialist 
(Homoptera: Psylloidea: Aphalaridae). Proc. Entomol. Soc. Wash. 96:91-97. 

Wheeler, A. G., Jr. 1995. Plant bugs (Heteroptera: Miridae) of Phlox subulata and other narrow- 
leaved phloxes in eastern United States. Proc. Entomol. Soc. Wash. 97:435-451. 

Wheeler, A. G., Jr. and T. J. Henry. 1981. Jalysus spinosus and J. wickhami: taxonomic clarifi- 
cation, review of host plants and distribution, and keys to adults and 5th instars. Ann. Entomol. 
Soc. Am. 74:606-615. 

Wheeler, A. G., Jr. and C. W. Schaefer. 1982. Review of stilt bug (Hemiptera: Berytidae) host 
plants. Ann. Entomol. Soc. Am. 75:498-506. 


Marvin H. Brunson 


Marvin H. Brunson died at age 94 on December 23, 1996 in a nursing home in Moorestown, 
NJ. He was a former President (1967 and 1968) and Vice-President (1965 and 1966) of the Ameri- 
can Entomological Society. 

One of 15 children, Marvin was born on a Meridian, Mis- 
sissippi farm on December 7, 1902. He graduated from Missis- 
sippi State College in 1926, and earned his M.S. in entomology 
from Iowa State College in 1928. Marvin was an extension en- 
tomologist at Clemson for two years, before joining the USDA’s 
Oriental Fruit Moth research unit at Moorestown, NJ in 1931. 

At Moorestown, Marvin performed research on the chemi- 
cal and biological control of fruit insects for 27 years, until trans- 
ferring to Washington State. There, as Research Leader and Sta- 
tion Head, he reorganized and led chemical and biological con- 
trol research on fruit tree pests at the USDA’s Yakima and Wenat- 
chee laboratories from 1959-1963. In 1963, he returned to 
Moorestown to become Investigations Leader of the USDA In- 
troduced Beneficial Insects Laboratory. During his leadership, 
and with the assistance of R. I. Sailer in Beltsville, landmark 
classical biological control successes were developed by this laboratory, in cooperation with one 
or two other USDA units, to combat the alfalfa weevil (present savings to agriculture of $90 
million per year) and the cereal leaf beetle (present savings $20 million/year). This work estab- 
lished the Moorestown laboratory as one of the few to successfully develop permanent biological 
controls for modern agriculture, and eliminate the need for insecticides over large areas of the 
United States, and was accomplished despite a limited budget. 

Marvin was a forthright leader, and was not reluctant to differ with administrators who had 
little understanding of how classical biological control or practical agricultural research actually 
“worked”. This protected his researchers from frequent changes in direction or inappropriate 
goals, and contributed significantly to their successful results. After he retired from the USDA in 
1970, he continued to work in the laboratory’s library until the unit moved to Newark, Delaware 
in the fall of 1973. 

Marvin remained in Moorestown, enjoying golf and retirement. He was a member of the 
YMCA’s Men’s Club and Trinity Episcopal Church in Moorestown. He also served on the 
Moorestown School Board in the 1950’s. His wife Martha (“Matt”) (Strickler) died in 1991. He 
is survived by a daughter, Lois (Mrs. Charles Y. Murphey III) of Cherry Hill, NJ and two 
grandchildren. 


W. H. Day J. J. Drea, Jr. D.O. Hathaway B.A. Butt 
Newark, DE. Oceanside, CA Wapato, WA McMinnville, TN 


Vol. 108, No. 3, May & June, 1997 179 


PADDLEFISH (POLYODON SPATHULA) AS 
SAMPLERS OF RIFFLE BEETLES 
(COLEOPTERA: ELMIDAE)! 


Steven G. George2, Jan Jeffrey Hoover, Harley P. Brown4 


ABSTRACT: Stomach contents of 17 paddlefish, collected in the Mississippi delta, yielded 41 
specimens of riffle beetle, Stenelmis spp. The specimens provide documentation of riffle beetles 
in the Big Sunflower River and the first record of Stenelmis parva in Mississippi. 


Riffle beetles (Coleoptera: Elmidae) are sensitive indicators of aquatic en- 
vironments, with most species occurring in erosional habitats of high water 
quality (Leech and Chandler, 1956; Hilsenhoff, 1982; Brown, 1987; White 
and Brighan, 1996). Techniques for their collection include drift nets, light 
traps, dip nets, and hand-picking from submersed rocks, wood, vegetation, 
and debris (Cowell and Carew, 1976; Brown 1976). These are effective in most 
streams, littoral zones, and small water bodies, but occurrences of riffle beetles 
in some habitats are insufficiently documented. For sampling large, turbid riv- 
ers unsuitable for traditional collecting techniques, we recently stumbled upon 
an unthought of assistant — the paddlefish (Polyodontidae: Polyodon spathula), 
a riverine filter-feeder. 

With no thought of riffle beetles, paddlefish diets were studied 1 Mar 94 - 
28 Feb 95 in the Big Sunflower River, a turbid, soft-bottomed, frequently hy- 
poxic stream of the Mississippi delta. River stages during this period ranged 
from 5.2 to 9.6 m National Geodetic Vertical Datum (NGVD), water surface 
elevation above mean sea level. Adult and sub-adult fish were obtained as 
bycatch from a commercial fisherman working a 60 km reach daily from Round 
Lake, Sunflower County, MS, downstream to Osceola, Washington County, 
MS. The upper boundary of this reach occurs near the mouth of the Quiver 
River, the lower boundary near the mouth of the Bogue Phalia. Stomachs of 99 
fish 411-1161 mm eye-to-fork length were examined. Almost 99% of the food 
organisms taken from the paddlefish stomachs consisted of copepods and cla- 
docerans, and most of the remaining percentage consisted of ostracods and 
assorted aquatic insects, but 17 stomachs contained adult riffle beetles. 

The mean number of beetles, in those 17 fish, was 2.4 beetles/fish (SD = 


1 Received October 7, 1996. Accepted October 30, 1996. 
2 Dyntel CZP, CEWES-ER-A, 3909 Halls Ferry Road, Vicksburg, MS 39180-6199. 


3 U.S. Army Engineer Waterways Experiment Station ER-A, 3909 Halls Ferry Road, Vicksburg, 
MS 39180-6199. 


4 The University of Oklahoma, Department of Zoology, Norman, OK 73019-0235. 


ENT. NEWS 108(3): 179-182, May & June, 1997 


180 ENTOMOLOGICAL NEWS 


1.17), and the maximum number was 5 beetles/fish. Of the 41 beetles, 33 were 
retained as voucher specimens and deposited in the Oklahoma Museum of 
Natural History. The species (and numbers of each) were Stenelmis decorata 
Sanderson 1938 (26), S. grossa Sanderson 1938 (6), S. parva Sanderson 1938 
(1). Beetles were obtained from paddlefish collected in late spring (9 May 94- 
7 Jun 94) at river stages 5.5-7.9 m NGVD, and in late summer/early autumn (2 
Sep-10 Oct 94) at river stage 5.2-5.6 m NGVD, during the hours 0500-0930. 
Fish that contained riffle beetles were 411-724 mm eye-to-fork length. Be- 
cause the Quiver River enters the Sunflower River not far above the collection 
sites, it is possible that some — perhaps most — of the beetles taken were de- 
rived from that stream. The Quiver River is a smaller, soft-bottomed, turbid 
stream; length and channel capacity are approximately one fourth that of the 
Big Sunflower River (USACE, 1955). Considering the paddlefish’s feeding 
mode, all beetles were probably drifting in the water column when ingested. 

A previous, unpublished invertebrate survey did not indicate riffle beetles 
in the Big Sunflower River (Parker and Robinson, 1972), but temporal occur- 
rence and taxonomic composition are consistent with previous descriptions of 
elmid biology. Because only adults were observed, beetles may have been in- 
tercepted during post-emergence migrations. Beetles may also have been in- 
tercepted while passively drifting or while surfacing. Drifting is common among 
elmids, especially at night (Waters, 1972; Brown, 1987), and the periods when 
riffle beetles were observed in the Big Sunflower River correspond to June and 
October peak abundances of drifting adults in other southern streams (Cowell 
and Carew, 1976; Reisen, 1977). Riffle beetles in well-aerated streams seldom 
ascend to the surface, but may do so when oxygen becomes limiting (Leech 
and Chandler, 1956). Hydrofuge hairs on the insect’s body allow retention of 
an air bubble that functions as an air store and/or a gill (Chapman, 1969). In 
the Big Sunflower and Quiver Rivers, mid-day dissolved oxygen is high (> 7.0 
mg/l) in late summer and early autumn but low (< 3.0 mg/l) in late spring. 
Beetles may respond to hypoxia in spring by drifting or surfacing, either of 
which would increase their risk of ingestion by paddlefish. 

S. decorata, the most abundant species, is tolerant of organic pollutants 
(Brown, 1976) which are high in the Big Sunflower River due to intensive 
agriculture in the delta (Johnson et al. 1994). S. grossa, of lesser abundance, is 
characteristic of sandy streams under logs (Brown, 1976), a habitat that is lim- 
ited in this drainage by the high rates of silt deposition and removal of sub- 
merged wood due to agriculture and flood control activities. S. grossa is previ- 
ously documented from Mississippi (Brown, 1976). S. decorata and S. parva 
are not listed for Mississippi by Brown (1983), but Schmude (1992) includes 
records of S. decorata from six counties in Mississippi. The finding of S. parva 
is most surprising because it heretofore has been known only from southeast- 
ern Oklahoma, southeastern Texas (Edna, between Houston and Corpus Christi), 


Vol. 108, No. 3, May & June, 1997 181 


and western Arkansas, and occurs chiefly on submerged wood. 

Low numbers of beetles, and the precarious nature of paddlefish popula- 
tions, make exclusive sampling of paddlefish for elmids inefficient and ex- 
travagant. Because paddlefish are commercially exploited, however, and be- 
cause numerous diet studies have been and are being conducted, existing 
samples may be utilized by curious coleopterists. Previously, only crawling 
water beetles (Haliplidae) have been recorded from paddlefish stomachs 
(Gannon and Howmiller, 1973), but “Insecta” are frequent and sometimes abun- 
dant foods (Ruelle and Hudson, 1977; Russell, 1983). Riffle-beetles ingested 
by paddlefish are mostly in an excellent state of preservation (i.e., limbs and 
heads intact). Thus, paddlefish can provide interesting insights into the biol- 
ogy of riffle beetles that would be otherwise unobtainable. 


ACKNOWLEDGMENTS 


We thank W.E. Lancaster for providing the fish for this study; K.J. Killgore reviewed the 
manuscript and F. Griggs provided river stage data. Permission was granted by the Chief of 
Engineers to publish this information. Funding was provided by the U.S. Army Engineer, Vicksburg 
District. 


LITERATURE CITED 


Brown, H. P. 1976. Aquatic Dryopoid Beetles (Coleoptera) of the United States. Water Pollution 
Control Research Series 18050 ELDO4/72. U.S. Environ. Protec. Agency, Cincinnati, OH. 
81 pp. 

Brown, H.P. 1983. A catalog of Coleoptera of America north of Mexico. Family:Elmidae. U.S. 
Dept. Agric., Handbook No. 529-50, x+ 33 pp. 

Brown, H.P. 1987. Biology of riffle beetles. Ann Rev. Entomol. 32: 253-273. 

Chapman, R.F. 1969. The insects - structure and function. American Elsevier Publishing Co., 
New York, NY, 819 pp. 

Cowell, B.C. and W.C. Carew. 1976. Seasonal and diel periodicity in the drift of aquatic insects 
in a subtropical Florida stream. Freshwat. Biol. 6: 587-594. 

Gannon, J.E. and R.P. Howmiller. 1973. Ecological notes on paddlefish (Polyodon spathula) 
with short rostrums. Mich. Acad. 6: 217-222. 

Hilsenhoff, W.L. 1982. Using a biotic index to evaluate water quality in streams. Wis. Dept. Nat. 
Resadech: Bull532-31-22° 

Johnson D.R., B.A. Sullivan, and D.L. Wallace. 1994. Water quality appendix. Appendix L, 
pp. 1-81. Jn: Project report and supplement No. 2 Appendices E-M, to the final environmen- 
tal impact statement — flood control, Misssissippi River and tributaries, Yazoo Basin, Missis- 
sippi — Big Sunflower River Maintenance Project. U.S. Army Corps of Engineers, Vicksburg 
District, Vicksburg, MS. 

Leech, H.B. and H.P. Chandler. 1956. Aquatic Coleoptera. pp. 293-371. Jn: R.L. Usinger, [ed.], 
Aquatic insects of California, Univ. Calif. Press, Berkeley, CA. 508 pp. 

Parker, W. and D. Robinson. 1972. Survey of sources of stream pollution and its effect upon 
bottom organisms, fishes and chemical qualities. A report to the Mississippi Game and Fish 
Commission, Project No. F-9-14. Jackson, MS. 61 pp. 


182 ENTOMOLOGICAL NEWS 


Reisen, W.K. 1977. The ecology of Honey Creek, Oklahoma: the downstream drift of three 
species of aquatic dryopoid beetles (Coleoptera: Dryopoidea). Entomol. News 88: 185-191. 

Ruelle, R., and P.L. Hudson. 1977. Paddlefish (Polyodon spathula): growth and food of young 
of the year and a suggested method for measuring length. Trans. Am. Fish. Soc. 106: 609- 
613. 

Russell, T.R. 1983. Biology and Life History of the Paddlefish A Review. pp. 2-21. In: J.G 
Dillard, L.K. Graham, and T.R. Russell. [eds.], The paddlefish: status, management, and propa- 
gation. Amer. Fish. Soc. Spec. Pub. No. 7: vit 159. 

Schmude, K.L. 1992. Revision of the riffle beetle genus Stenelmis (Coleoptera: Elmidae) in 
North America, with notes on bionomics. Doctoral thesis, University of Wisconsin, Madison, 
WI. 388 pp. 

USACE. 1955. General design memorandum No. |. Big Sunflower, Little Sunflower, Hushpuckena 
and Quiver Rivers and their tributaries, and Deer Creek, Steele Bayou, and Bogue Phalia, 
Mississippi. U.S. Army Engineer District, Vicksburg, MS. 33 pp. + 16 plates. 

Waters, T.F. 1972. The drift of stream insects. Ann. Rev. Entomol. 17: 253-272. 

White, D.S. and W.U. Brighan. 1996. Aquatic Coleoptera. pp. 399-473. In: R.W. Merritt and 
K.W. Cummings [eds.], An introduction to the aquatic insects of North America. Third Edi- 
tion. Kendall/Hunt Publishing. Company. Dubuque, IA. xili+ 862 pp. 


SOCIETY MEETING OF FEBRUARY 19, 1997 


Dr. Dale F. Schweitzer 
The Nature Conservancy, Port Norris, New Jersey 


LEPIDOPTERA OF THE NEW JERSEY PINE BARRENS 


Dr. Schweitzer presented a richly illustrated overview of the diverse butterflies and moths 
found in this region. The distinctive character of its fauna was recognized at least as early as 
1910 in J.B. Smith’s insect list and by mid-twentieth century lepidopterists like Fredrick Lemmer, 
Otto Bucholz, and Joseph Muller. Many species common in the Pine Barrens are rare elsewhere 
and there are a number of endemics or near-endemics as well as undescribed species. Groups like 
saturniid moths that have declined drastically in the Northeast are doing well in the Barrens, and 
he suggested that overall, the Pinelands Protection Act may have done more for biodiversity than 
the Alaska Lands Act which was signed into law about the same time. 

Dr. Schweitzer discussed the southern affinities of much of the Pine Barrens fauna, particu- 
larly with pine-dominated habitats in Florida and the Carolinas. 

He went on to describe the key role that fire has played in creating and maintaining habitat 
for these unusual species. Much of this habitat has been altered or eliminated by fire suppression, 
and even the dwarf pine plains are changing as a result of fewer fires. Although presumably once 
more widespread, Pine Barrens reed grass savannas are now found almost exclusively on the 
Fort Dix impact area, where fires are frequently started by stray munitions and are often allowed 
to burn themselves out. That many of these fires occur during the growing season may be par- 
ticularly important, he said. Two out of less than ten known remaining populations of the imper- 
iled Arogos Skipper (Atrytone arogos arogos) are found there, along with a number of other rare 
species, in much greater abundance than elsewhere. 


(continued on page 202) 


Vol. 108, No. 3, May & June, 1997 183 


NOTES ON THE ANT EURHOPALOTHRIX 
FLORIDANA, WITH A DESCRIPTION OF THE MALE 
(HYMENOPTERA: FORMICIDAE)! 


Mark Deyrup2, Clifford Johnson, Lloyd Davis4 


ABSTRACT: The basicerotine ant Eurhopalothrix floridana is a relatively common woodland 
species in peninsular Florida. In small (2 liter) litter samples, FE. floridana usually co-occurs with 
one or more of at least 32 other species of ants. There is some evidence that F. floridana may 
have been introduced into Florida. We provide a diagnosis and habitus illustration of the previ- 
ously unknown male 


The ant Eurhopalothrix floridana Brown and Kempf (Fig. 1) is the only 
U.S. representative of the tribe Basicerotini, a group of cryptic, slow-moving, 
pantropical myrmicines. Eurhopalothrix species are predatory, and observa- 
tions of one species suggest that these ants may be somewhat specialized preda- 
tors of termites (Wilson and Brown 1984). Eurhopalothrix floridana was de- 
scribed in 1960 (Brown and Kempf) on the basis of a single specimen, but the 
species is not nearly as rare as one might expect from the late date of its dis- 
covery. In this note we update the distribution and habitat information on E. 
floridana, consider the possibility that it may be an exotic species, and de- 
scribe the previously unknown male. 


DISTRIBUTION AND HABITAT 


Eurhopalothrix floridana occurs throughout peninsular Florida (Fig. 2), 
based on about 130 collections from over 70 sites. Although we sampled leaf 
litter and rotten wood from all over Florida, we made no effort to sample evenly, 
and the distribution pattern in Fig. 2 reflects biases based on convenience and 
access. The total sampling effort, however, involved many thousands of Tulgren 
funnel extractions, collected in all regions of the state. We collected, for ex- 
ample, hundreds of samples from the western panhandle, without finding any 
E. floridana. The pattern of records reflects the distribution of woodlands in 
peninsular Florida, including the dry tropical hammocks of the Florida Keys, 
the Atlantic coastal ridge hammocks, the scrub forests and ecotonal hardwood 
forests of the southern and central ridges, and the mesic and xeric forests of the 
northern peninsula. 

In a study of E. floridana using standardized, unsifted, approximately 


! Received October 15, 1996. Accepted November 13, 1996. 


2 Archbold Biological Station, P.O. Box 2057, Lake Placid, FL 33862. 
3 Department of Zoology, 223 Bartram Hall, University of Florida, Gainesville, FL 32611. 
4 3920 NW 36th Place, Gainesville, FL 32606. 


ENT. NEWS 108(3): 183-189, May & June, 1997 


184 ENTOMOLOGICAL NEWS 


Figure 1. Eurhopaalothrix floridana, worker. 


2-liter samples of litter, we found EF. floridana in 65 samples on 32 occasions. 
The 32 habitat records include: 8 from xeric forest (old growth sand pine scrub, 
sandhill invaded by large oaks), 11 from mesic forest (usually mixed oaks and 
pines), 5 from wet forest (mixed pine and hardwoods, usually including oaks 
and magnolia); and 8 from coastal tropical hardwood forest. This is apparently 
a woodland species that is not particular about drainage. 

The 2-liter litter samples used in this analysis were indexed by site and 
date, with an ant species list for each sample. This allows us to say that, out of 
346 samples collected at a site and date where E. floridana was found, 65 
(19%) contained in E. floridana. At a number of sites less than one out of 10 
samples had E. floridana, so it is evident that in many cases at least 5, and 
often more, samples must be extracted to demonstrate the presence on this 
species at a particular site. This need for large numbers of samples to show 


Vol. 108, No. 3, May & June, 1997 185 


presence or absence of a species is acommon problem in surveys of litter ants; 
many species are much rarer. 

In our small litter samples, FE. floridana is usually found with other species 
of ants, with which it must be somewhat compatible and with which it must 
share microhabitat requirements. The numbers of co-occurring species are as 
follows: 6 samples had no other ant species, 12 samples had 1 other species; 


Figure 2. Known distribution of Eurhopalothrix floridana in Florida. 


186 ENTOMOLOGICAL NEWS 


15 samples had 2 other species; 13 samples had 3 other species, 11 samples 
had 4 other species, 6 samples had 5 other species, and 2 samples had 6 other 
species.The list of 32 co-occurring species and their frequency of occurrence 
appears in Table 1. A statistically different frequency of species in the samples 
that did not have E. floridana (samples from the same place and date as those 
that did have E. floridana) might hint at another level of ecological relation- 
ships. Unfortunately, there were no sites where there were many samples with 
E. floridana, and comparisons between sites brings in larger scale biogeo- 
graphic factors. 


NATIVE OR EXOTIC? 


Brown and Kempf (1960) suggest that EF. floridana could be a recent intro- 
duction from the Neotropics. There is additional circumstantial evidence that 
supports this suggestion. Expanding ranges are typical of exotic species; E. 
floridana is now easily obtained in Alachua and Putnam counties (Fig. 2), 
where Van Pelt (1958) sampled extensively for dacetine ants without finding 
E. floridana. Eurhopalothrix. floridana has not been found in the West Indies, 
the source of almost all tropical ants that have, we assume, dispersed naturally 
to Florida. It is unlikely to be a native from the forests of the southern Appala- 
chians, because such species usually have a large range to the north and west 
of Florida. E. floridana occurs in Mexico (W. L. Brown, Jr., 1985, pers. comm.), 
so there is a possible source of introduction. There are species of insects, in- 
cluding ants, that occur in southwestern North America, with a disjunct popu- 
lation or closely related congeners in Florida, but these are species of open, 
desert or savannah habitats, not woodlands (Deyrup 1990). 

On the other hand, it could be native to Florida. The oldest record is a 
damaged specimen found recently by David Smith of the U. S. National Mu- 
seum of Natural History. This specimen is from Key West (once a major com- 
mercial port), dated 1887, and is in the Pergande collection. A record of this 
antiquity coming from a less settled part of Florida might suggest that E. 
floridana is a native species, but in tropical Florida a great amount of entrepre- 
neurial horticulture occurred well before 1887. Henry Perrine, for example, 
was in a good position in the 1830’s to be an unintentional purveyor of 
cryptobiotic ants to the Miami area. Perrine displayed a missionary zeal in the 
importation of plants from southern Mexico, many in boxes or tubs of soil. 
Marjory Douglas (1978) cites a report of “more than 100 boxes of plants shipped 
from the Yucatan” by Perrine outside one building in the Miami area. 
Eurhopalothrix. floridana is a cryptic species that could have escaped being 
noticed by some collectors; it does not show any association with highly dis- 
turbed habitats, its ability to live in xeric woodlands might have allowed it to 


Vol. 108, No. 3, May & June, 1997 187 


move to Florida from Mexico along with other upland species at the end of the 
Pliocene or in the early Pleistocene. We conclude that this species is probably 
introduced into Florida, but would like to see better documentation of the spe- 
cies in Mexico, especially southern Mexico. 


Male of E. floridana 


Diagnosis: the male (Fig. 3) generally resembles a male dacetine in its dense, reticulate 
sculpture, the facial projection trom which the antennae emerge, reduced venation, and long 
petiole. It is distinguished from Florida dacetines by the following features: 1.) The post petiole 
is twice as wide as the petiole, with a strongly concave anterior border and a conspicuous spatu- 
late hair on each side. 2.) The first submarginal cell is clearly developed. 3.) The antennal scape 
has a strong bristle on the proximal corner of the large bulge on the inner side. This may be a 
character found throughout the genus (Brown and Kempf 1960) 4.) The size is relatively large. 


Figure 3. Eurhopalothrix floridana, male. 


188 ENTOMOLOGICAL NEWS 


Total length: 2.02-2. 15 mm Length of forewing: 2.32-2.45 mm. 

There are only 2 additional species of Eurhopalothrix (both Old World species) whose males 
have been described (Brown and Kempf 1960). E. floridana differs from these, according to the 
descriptions, in the smooth, shining areas on the mesopleuron (Fig. 3). 

Seasonality of males: males were collected with workers on 12 July and 19 August. Fifty 
males were collected in Townes traps at several sites between 15 July and 5 December. Deposi- 
tion of males: males, along with workers, have been deposited in the following collections: Harvard 
Museum of Comparative Zoology (Cambridge, MA), U. S. Museum of Natural History (Wash- 
ington, D.C ), Los Angeles County Museum, British Museum of Natural History (London), Florida 
State Collection of Arthropods (Gainesville), Canadian National Collection (Ottawa), Collection 
of Mark Dubois (Washington, IL), Collection of William MacKay (El Paso, TX), Archbold Bio- 
logical Station Collection of Arthropods (Lake Placid, FL). 


Table 1. Species of ants found together with E. floridana in small litter samples. 


No. of Species 
Co-occurrences 
28 Hypoponera opacior 
22 Solenopsis abdita 
19 Pheidole dentigula 
11 Strumigenys eggersi 
8 Solenopsis tennesseensis 
7 Strumigenys louisianae 
6 Pheidole moerens, Smithistruma talpa 
5 Brachymyrmex depilis, Pheidole dentata, P. floridana, 


Smithistruma ornata, Wasmannia auropunctata 


4 Aphaenogaster miamiana or carolinensis, Paratrechina 
faisonensis, Quadristruma emmae 

3 Solenopsis carolinensis 

2 Myrmecina americana, Odontomachus brunneus, Pachycondyla 
Stigma, Strumigenys rogeri 

1 Aphaenogaster fulva, Cyphomyrmex minutus, C. rimosus, 


Hypoponera inexorata, H. punctatissima, Odontomachus 
ruginodis, Paratrechina guatemalensis, P. wojciki, Smithistruma 
brevisetosa, S. dietrichi, Trichoscapa membranifera. 


ACKNOWLEDGMENTS 


We are especially grateful to Walter Suter (Carthage College, Kenosha, WI) for allowing us 
to look through a large number of Tulgren funnel residues from all over Florida, and to Virendra 
Gupta (Florida State Collection of Arthropods) for sending us ants (pre-sorted!) from Townes 
trap samples collected in Gainesville. Zachary Prusak (graduate student at the University of 
Central Florida, Winter Park) also contributed specimens from several sites. David Wang (senior 
at Lake Placid High School) compiled all the data from litter samples collected in the Archbold 
study of litter ants. 


LITERATURE CITED 


Brown, W. L. Jr., and W. W. Kempf. 1960. A world revision of the ant tribe Basicerotini. Studia 
Entomol. 3: 161-250. 


Vol. 108, No. 3, May & June, 1997 189 


Deyrup, M. 1990. Arthropod footprints in the sands of time. Florida Entomol. 73: 529-538. 

Douglas, M.S. 1978. The Everglades: River of Grass (revised edition). Banyan Books Inc. Mi- 
ami, 447 pp. 

Van Pelt, A. F., Jr. 1958. The ecology of the ants of the Welaka Reserve, Florida, (Hymen- 
optera: Formicidae). 2. Annotated list. Amer. Midland Natur. 59: 1-57. 

Wilson, E. O., and W L. Brown, Jr. 1984. Behavior of the cryptobiotic predaceous ant 
Eurhopalothrix heliscata, n. sp. (Hymenoptera. Formicidae: Basicerotini). Insectes Sociaux, 
Paris 31: 408-428. 


BOOK REVIEW 


FIELD GUIDE TO NORTHEASTERN LONGHORNED BEETLES (CO- 
LEOPTERA: CERAMBYCIDAE). Douglas Yanega. 1996. Illinois Natural 
History Survey Manual 6, Champaign, Illinois. 


This field guide has an ambitious goal — to assist nonspecialists in making species deter- 
minations of cerambycids. What makes this manual so successful is that it is focused on this 
goal. It does not attempt to be more than an identification manual. But that is not a criticism. 
Rather, this book succeeds in its task so well that it may change the format of future field 
guides for nonspecialists. 

The book starts with a brief introduction to beetle terminology using the unique mor- 
phological details of the cerambycids. It does not go into any detail that is not easily seen 
with the naked eye or a 10X hand lens, as that would be of little-help with species identifica- 
tion. 

Next follows a brief section on the natural history of the Cerambycidae including infor- 
mation on biogeography, climate, life histories, phenology, reproduction, interactions with 
other species, and variation. There is also a short section on collecting techniques and the 
curation of specimens. All of this information is presented in the first 23 pages! 

The field guide covers 342 species of longhorned beetles which includes all the north- 
eastern species save one which is undescribed. The species identifications are presented in 
an unusual manner compared to other field guides. Yanega does not use keys in the formal 
sense. Keys in technical books tend to be of little use to the nonspecialist because they do 
not convey a sense of confidence for the user. Rather, Yanega presents a system that assumes 
that the user will scan photographs to compare with the specimen. This is where the manual 
is exceptional. The author has taken considerable care in finding the best specimens to pho- 
tograph. Photographs of both sexes are presented and small arrows point to diagnostic fea- 
tures. I have tested the manual with undergraduate students and they had no difficulty in 
determining the species and the sex of beetles they had collected. 

In addition to the photographs, Yanega provides a synopsis of each species giving its 
flight period, larval feeding habits, size, and general notes. The information is brief. If the 
user needs more detailed information, then s(he) is referred to the original description and 
other works. 

This field guide is a successful manual in that it is accessible to the nonspecialist and 
recognizes how nonspecialists go about the process of determining an identification. Any- 
one interested in cerambycids will benefit from this book and I am sure that it will be the 
first field guide to be consulted to determine the identification of an unknown cerambycid. 


Gene Kritsky, Department of Biology 
College of Mount St. Joseph, Cincinnati, OH 45233 


190 ENTOMOLOGICAL NEWS 


DISTRIBUTION AND ABUNDANCE OF EULONCHUS 
MARIALICIAE (DIPTERA: ACROCERIDAE)! 


Peter H. Adler2, Stuart R. Reitz>, Charles N. Watson4 


ABSTRACT: Eulonchus marialiciae, a rare parasitoid of the folding trapdoor spider Antrodiaetus 
unicolor, was recorded from cool, humid, mixed forests at elevations above 1200 m in five coun- 
ties of western North Carolina. Adults were on the wing during June and August. Smooth black- 
berry (Rubus canadensis) served as a nectar source. Host spiders and nectar sources of this fly 
are abundant, and most of the habitat in which it has been found is federally protected. 


The larvae of all known members of the family Acroceridae are internal 
parasitoids of spiders; however, the biologies of many of the species are poorly 
known (Schlinger 1981). Some of the more colorful species are in the genus 
Eulonchus which consists of seven described species and at least 17 species 
that are undescribed; all but one species in the genus are restricted to western 
North America (Schlinger 1966, 1981). 

Eulonchus marialiciae Brimley, informally known as Mary Alice’s small- 
headed fly (Fig. 1), was described from a single male captured 23 June 1923 on 
Andrews Bald Mt., Swain Co., NC (Brimley 1925). The only other report of 
this fly comes from a study of the folding trapdoor spider Antrodiaetus unicolor 


Fig. 1. Male of Eulonchus marialiciae collected 8 June 1965 at Heintooga Overlook, Blue Ridge 
Parkway, Swain Co., NC (specimen housed in Canadian National Collection). 


! Received September 11, 1996. Accepted October 30, 1996. 

2 Department of Entomology, Clemson University, Clemson, SC 29634-0365. 
3 Department of Entomology, University of California, Riverside, CA 92521. 
4 Aquatic Resources Center, P. O. Box 680818, Franklin, TN 37068-0818. 


ENT. NEWS 108(3): 190-192, May & June, 1997 


Vol. 108, No. 3, May & June, 1997 191 


(Hertz) (Araneae: Antrodiaetidae) by Coyle (1971). On 1 and 2 August 1966, 
Coyle (1971) recovered one pupa that produced an adult, one freshly emerged 
adult, and an unspecified number of pupal exuviae from “several” burrows of 
A. unicolor along Rhododendron Trail at the Highlands Biological Station (1250 
m elevation), Macon Co., NC. He also observed 18 adults hovering near bur- 
rows, and noted that adults often aggregated around burrows that he was exca- 
vating during the day, leading him to suggest that the flies were responding to a 
chemical released during the excavations. Coyle’s original field notes state that 
of five flies taken from these aggregations, all were males; this observation 
suggests to us that mating might take place near the host burrows. Although 
Coyle (1971) unearthed many burrows of A. unicolor from Pennsylvania to 
Alabama, he found no other specimens of E. marialiciae. 

Eulonchus marialiciae was one of 15 Diptera in the United States formerly 
placed in Category 2 of the Federal Register (United States Fish and Wildlife 
Service 1994). This designation denotes taxa for which endangered or threat- 
ened status is possibly appropriate but for which conclusive data are lacking. 
To obtain additional information on the distribution and status of this fly, we 
spent 42 days from June through mid-September, 1993 and 1994, prospecting 
at 43 sites in the mountains of western North Carolina and South Carolina. We 
ran Malaise traps, excavated burrows of A. unicolor, and inspected potential 
nectar sources; we also examined museum collections. Five males from our 
study have been deposited as voucher specimens in the Clemson University 
Arthropod Collection. 

Other than the holotype (U. S. National Museum), we located only one 
other museum specimen. The Canadian National Collection in Ottawa con- 
tains a male taken on Rubus by J. G. Chillcott, 8 June 1965, at Heintooga 
Overlook along the Blue Ridge Parkway, Swain Co., NC. 

During our field work, we found 10 flies (80, 2 sex unknown [not cap- 
tured]) feeding on the floral nectar of smooth blackberry (Rubus canadensis 
L.) along the Blue Ridge Parkway in North Carolina: 


Haywood Co., Beartrail Ridge Parking Area, 1790 m, 16 June 1994 (1 sex unknown); Haywood 
Co., Spot Knob Overlook, 1723 m, 16 June 1994 (10); Jackson Co., 1.0 km N of Doubletop 
Mt. Overlook, 1635 m, 16 June 1994 (1C’); Jackson Co., 1.1 km N of Doubletop Mt. Overlook, 
1634 m, 16 June 1994 (1C); Swain Co., Heintooga Overlook, 1626 m, 15 June 1994 (1 CO, i sex 
unknown); Swain Co., Mollies Gap, 1631 m, 13 June 1994 (2C’); Transylvania Co., Mt. Hardy 
Viewing Area, 1650 m, 17 June 1994 (20). 


We observed feeding from 1200 to 1600 h on sunny and overcast days. 
Flies could be approached within less than a meter as they probed individual 
blooms while stationed on the flowers. One male fed for 3-100 sec (mean + SE 
= 25.5 + 6.8 sec, 24 °C) per flower. Flies carried a light dusting of pollen on 
their bodies. Three males that were brought to the laboratory fed on a 10% 


192 ENTOMOLOGICAL NEWS 


solution of honey. Dissections 24 h later revealed a lack of stored nutrient in 
the abdomen, suggesting a dependence on sugar sources such as nectar. Our 
small sample suggests that males are more likely to be found nectaring than 
are females; in fact, we have no evidence that females take nectar, although we 
suspect that they do. Other described species in the genus Eulonchus are usu- 
ally found probing flowers for nectar and are considered important pollinators 
(Schlinger 1981). 

In total, we account for 32 adult flies that have been seen or collected since 
the type specimen was taken in 1923. All were found at elevations of 1250 m or 
higher in five mountainous counties of western North Carolina. This region is 
characterized by cool, humid woodlands with a liberal humic layer and domi- 
nated by hemlock (Tsuga canadensis (L.) Carr.), oaks (Quercus spp.), and rhodo- 
dendron and flame azalea (Rhododendron spp.). Our records, coupled with those 
of Coyle (1971), suggest either that two generations of flies are produced yearly, 
with adults on the wing during June and August, or that a single brood with 
prolonged emergence occurs. Flies emerging after June would require a nectar 
source other than R. canadensis, which does not bloom beyond that time. 

We conclude that E. marialiciae is rare but probably not threatened or en- 
dangered. Its known spider host and nectar source, as well as habitat, are abun- 
dant over much of western North Carolina, southwestern Virginia, and eastern 
Tennessee, suggesting that it exists over an area greater than our records indi- 
cate. Most sites where this fly has been recorded are within federally protected 
areas. 


ACKNOWLEDGMENTS 


This study was supported by a grant from the U. S. Fish & Wildlife Service. We thank J. A. 
Ratzlaff (U. S. Fish & Wildlife Service, Asheville, NC) for facilitating our study, F. A. Coyle 
(Western Carolina University) for sharing his original field notes, J. D. Culin (Clemson Univer- 
sity) for photographing the fly, J. M. Cumming (Eastern Cereal and Oilseed Research Centre, 
Ottawa) for loaning the specimen in the Canadian National Collection, and A. G. Wheeler, Jr. 
(Clemson University) for commenting on the manuscript. This is Technical Contribution No. 
4232 of the South Carolina Agricultural Experiment Station, Clemson University. 


LITERATURE CITED 


Brimley, C. S. 1925. New species of Diptera from North Carolina. Entomol. News 36: 73-77. 

Coyle, F. A. 1971. Systematics and natural history of the mygalomorph spider genus Antrodiaetus 
and related genera (Araneae: Antrodiaetidae). Bull. Mus. Comp. Zool. 141: 269 402. 

Schlinger, E. I. 1966. An analysis of the distribution of Eulonchus Gerstaecker (Diptera: 
Acroceridae). Bull. Entomol. Soc. Am. 12: 112-113. 

Schlinger, E. I. 1981. Acroceridae. Pp. 575-584. In J. F. McAlpine, B. V. Peterson, G. E. Shewell, 
H. J. Teskey, J. R. Vockeroth and D. M. Wood (coordinators). Manual of Nearctic Diptera. 
Vol. 1. Res. Branch Agric. Can. Monogr. 27. 

U.S. Fish and Wildlife Service. 1994. Endangered and threatened wildlife and plants; animal 
candidate review for listing as endangered or threatened species: proposed rule. Federal Reg- 
ister, part IV. U. S. Depart. Interior SO CFR part 17, 59 (219): 58982-59028. 


Vol. 108, No. 3, May & June, 1997 193 


THE EPHEMEROPTERA OF SPRING CREEK, 
OKLAHOMA, WITH REMARKS 
ON NOTABLE RECORDS! 


W. P. McCafferty2, R. K. Heth3, R. D. Waltz4 


ABSTRACT: A total of 40 nominal species of mayflies are reported from nine sites on Spring 
Creek, a cold-water, western Ozarkian stream in Cherokee, Delaware, and Mayes Counties, Okla- 
homa. Of these species, 28 represent new state records, bringing the published total for Okla- 
homa to 59. The essential geographic affinity of the Spring Creek fauna is eastern North America. 
Many of the species were predictably found, being part of a typical pattern that includes the 
Ozarks and Ouachita Mountains of Arkansas, the southern Appalachians and the Cumberland 
Plateau. A large number are also known from eastern and midwestern North America in general. 
A few are transcontinental, or previously known only from eastern Canada and northeastern and 
north-central USA. Most new state records also represent westernmost range extensions. The 
notable geographic records of Ephemerella rotunda, Nixe flowersi, Paraleptophlebia jeanae, P. 
moerens, Rhithrogena impersonata, R. jejuna, and Stenonema vicarium are discussed. 


Oklahoma is one of several states in the USA that have been very poorly 
documented with respect to their mayfly fauna (McCafferty et al. 1990). The 
previous first published reports of 31 nominal Ephemeroptera species in the 
state are as follows: Traver (1934): Choroterpes oklahoma Traver [ = 
Neochoroterpes oklahoma (Traver)] and Habrophlebiodes annulata Traver; 
Traver (1935): Caenis delicata Traver [ = C. latipennis Banks] and Tricorythodes 
fictus Traver; Spieth (1938): Ephemera traverae Spieth; Spieth (1941): 
Hexagenia bilineata (Say), H. limbata (Serville), and H. rigida McDunnough; 
Allen and Edmunds (1965): Ephemerella excrucians Walsh; McKinley et al. 
(1972): Caenis simulans (McDunnough) [ = C. amica Hagen] and Stenonema 
tripunctatum (Banks) [ = S. femoratum (Say)]; Lewis (1974): Stenonema 
interpunctatum (Say) [ = Stenacron interpunctatum (Say)]; Reisen (1975): 
Baetis bicaudatus Dodds and Dactylobaetis mexicanus Traver and Edmunds 
[ = Camelobaetidius mexicanus (Traver and Edmunds)]; Wilhm et a/. (1978): 
Heptagenia diabasia Burks, H. maculipennis Walsh [| = Leucrocuta maculi- 
pennis (Walsh)], and Stenonema ares Burks [ = Stenonema terminatum 
terminatum (Walsh)]; Morihara and McCafferty (1979): Baetis quilleri Dodds 
[ = Fallceon quilleri (Dodds)]; Bednarik and McCafferty (1979): Stenonema 
exiguum Traver and S. mediopunctatum arwini Bednarik and McCafferty; 
Wilhm et al. (1979): Stenonema luteum (Clemens); Magdych (1979): Baetis 


! Received October 18, 1996. Revision received December 7, 1996. Accepted December 9, 1996. 
2 Department of Entomology, Purdue University, West Lafayette, IN 47907. 
3 2136 Faculty Row, Bacone College, Muskogee, OK 74403. 


4 IDNR, Division of Entomology and Plant Pathology, 402 West Washington, Rm W-290, 
Indianapolis, IN 46204. 


ENT. NEWS 108(3): 193-200, May & June, 1997 


194 ENTOMOLOGICAL NEWS 


flavistriga McDunnough; Pescador and Berner (1981): Baetisca lacustris 
McDunnough; Henry and Kondratieff (1982): Leptophlebia bradleyi (Need- 
ham); Kondratieff and Voshell (1984): Isonychia rufa McDunnough; Provonsha 
(1990): Caenis anceps Traver, C. hilaris (Say), and C. punctata McDunnough; 
Bae and McCafferty (1991): Anthopotamus neglectus disjunctus (Traver); 
McCafferty (1994): Ephemera simulans Walker; and Waltz et al. (1996): Baetis 
intercalaris McDunnough. Of these published records, only the report of the 
western species B. bicaudatus from south-central Oklahoma by Reisen (1975) 
is improbable. 

We can account for an additional 28 nominal species in Oklahoma based 
on identification of larval and adult Ephemeroptera collected during a study of 
Spring Creek in Cherokee, Delaware, and Mayes Counties in the western Ozarks 
(Table 1, Fig. 1). Many of these species were predictably found in Oklahoma 
based on their known continental ranges, and particularly if previously known 
from the Ozark and Ouachita Mountains of Arkansas (McCafferty and Pro- 
vonsha 1978). Moreover, several of these predictable species have also been 
collected in southeastern Oklahoma by D. E. Baumgardner (pers. comm.). Some 
of the new records from Spring Creek, however, constitute significant species 
range extensions, disjunctions, or reports of poorly known species. The may- 
flies of Spring Creek are mainly northeastern or southeastern species or both, 
in terms of the North American geographic regional affinities of McCafferty 
and Waltz (1990). 

As pointed out by Jester et a/. (1988) in their study of the ecology of 
Spring Creek, the stream is representative of large creeks in the western Ozarks 
that have diverse communities of unique cool-water fish species and ubiqui- 
tous warm-water fish species. We have found that the mayflies of Spring Creek 
(Table 1) also constitute a mixture of faunal elements, being composed of 1) 
several species that are found in the southern Appalachians, Cumberland Plateau 
and the Ozark-Ouachita Mountains; 2) some species that are basically 
midwestern fauna typical of low gradient, warm-water habitats; 3) a few species 
that were previously only known from cold-water trout streams of eastern 
Canada and northeastern and north-central USA; and 4) some species that are 
ubiquitous and transcontinental. None of the species were indigenous to western 
North America. 

Sites at which taxa were collected are described below, and their locations 
along Spring Creek are shown in Figure 1. Additional physical and chemical 
data associated with Spring Creek and many of our sites may be found in 
Jester et al. (1988). Table 1 gives the site distribution of all species taken at 
Spring Creek. Following the description of sites, we present a distributional 
analysis of those species representing significant range extensions or disjunc- 
tions based on their discovery in Oklahoma. Voucher specimens are deposited 
in the Purdue Entomological Research Collection, West Lafayette, Indiana, 
and at Northeastern State University, Tahlequah, Oklahoma. 


Vol. 108, No. 3, May & June, 1997 195 


DESCRIPTION OF SAMPLE SITES (FIG. 1) 


Site A— Delaware County, Coppage Farm, 322m elevation. Spring Creek is a first order stream 
at this site; it is a ford approximately 2-3m in width; substrate consists of cobble, gravel, and 
sand; there is partial shading; and there are no backwater areas. 

Site B — Delaware County, Oaks Mission, 305m elevation. Spring Creek is a first order stream 
at this site; it is approximately 3m wide; substrate consists of cobble and gravel; there is com- 
plete shade cover; and no backwater areas are present. 

Site C — Cherokee County, Rocky Ford, 277m elevation. Spring Creek is a second order stream 
at this site; it is approximately 9 m wide with bedrock and mixed sand and gravel; there is partial 
shade present. 

Site D — Cherokee County, Teresita at bridge, 256m elevation. Spring Creek is a second order 
stream at this site; it is very diverse with various currents and backwater areas; it ranges from 
approximately 6-8m in width; substrate consists of cobble, gravel, and sand; and there is partial 
shade. 

Site E — Cherokee County, Cherokee Cattle Company, 232m elevation. Spring Creek is a third 
order stream at this site; it ranges from approximately 5-8m in width; substrate consists of small 
cobble, gravel, and sand; there was partial shade present; some backwater areas are present; and 
cattle are often present. 


x 
Cl 
———S— gov UZ Fe 
3 { Kansas 
SCALE OF MILES Pee e: 
jo} 
1S) 
gig ® 
als 
=15 
=/4 @ Oaks DELAWARE CO. 


gq 


&, 

oe creek ™~ 
Cedar ee eal vf 
Crest Ne ‘ a7 


Fort 
Gibson P 
Reservoir 


(C) Rocky Ford 
State Park 


@ Luck Spring 


MAYES CO. 
CHEROKEE CO. 


LOCATION MAP 


Fig. 1. Spring Creek, Oklahoma study area, with lettered collection sites indicated (described in 
List of Sites in the text and referred to in Table 1). Location map with Cherokee, Delaware, and 
Mayes Counties, Oklahoma darkened. 


196 ENTOMOLOGICAL NEWS 


Site F — Cherokee County, Timbercreek Ranch, 213m elevation. Spring Creek is a third order 
stream at this site; it is at a ford approximately 10m wide; substrate consists of coarse gravel; 
there is no shade; and backwater areas are present. 

Site G— Mayes County, Camp Garland, 194m elevation. Spring Creek is a third order stream at 
this site; it is approximately 3-4m wide; substrate consists of gravel; it is partially shaded; and 
there are backwater areas present. 

Site H — Mayes County, Twin Bridges, 183m, elevation. Spring Creek is a third order stream at 
this site; it is approximately 10m wide; substrate consists of gravel; it has full exposure to sun; 
and there are backwater areas present. 


SIGNIFICANT RECORDS 


Ephemerella rotunda Morgan — This species is known from eastern Canada, 
eastern USA along the Appalachians from Maine to Georgia, but also extending 
into Florida (Berner 1958, 1977). It has also been reported from Kentucky, 
Michigan, and Wisconsin (Allen and Edmunds 1965). It has not been known 
west of Wisconsin and Kentucky. Its presence in the Oklahoma Ozarks 
represents a considerable range extension westward, but as for so many other 
Ozark species, it is acommon Appalachian species also occurring in Kentucky. 
The larvae of this species cannot consistently be told from those described as 
E. inconstans Traver. Thus, there remains the possibility that the Oklahoma 
record of E. rotunda is attributable to E. inconstans, assuming the latter name 
is not a junior synonym of the former. Ephemerella inconstans has been recorded 
from Georgia, Kentucky, North Carolina, Tennessee, and Virginia (see Allen 
and Edmunds 1965, Berner 1977). Larvae of E. rotunda were collected on III- 
14,18-1995 and V-18-1996. 


Nixe flowersi McCafferty — The discovery of N. flowersi in Oklahoma is 
significant because the species was previously known only from the type locality 
in Indiana. The species was described from adults (McCafferty 1982) and reared 
larvae that had previously been described as the larvae of Heptagenia persimplex 
McDunnough (McCafferty 1977). In Indiana, the species occurs in the far south 
central unglaciated area of the Ohio River Basin. Several mayflies from this 
latter area appear to be more typical of other southern unglaciated areas of 
North America, including the Ozarks and Smoky Mountains. Adults of N. 
flowersi were taken on VI-29-1996. 


Paraleptophlebia jeanae Berner — Berner (1955, 1975) reported this species 
from Alabama, South Carolina, and Virginia. It was recently discovered in far 
southern Indiana (Randolph and McCafferty 1996) in the unglaciated area of 
the Ohio River Basin, and therefore may occur in Kentucky. Its presence in the 
Oklahoma Ozarks reflects a common distributional pattern (i.e., western Ozark- 
Oauchita and eastern Cumberland Plateau-Appalachian pattern) pointed out 
by McCafferty and Provonsha (1978) for much of the Ozark-Oauchita mayfly 
fauna. Harris (1990) considered this species among rare and possibly 


Vol. 108, No. 3, May & June, 1997 197 


endangered mayflies in Alabama, as did Kondratieff and Kirchner (1991) for 
Virginia, and Morse et al. (1996) generally. The fact that its larvae were not 
identifiable prior to the time of the larval description by Randolph and 
McCafferty (1996) may account in part for its not having been reported from 
mountainous Arkansas and the fact that it has sometimes been considered rare. 
At least one species of Paraleptophlebia, P. calcarita Robotham and Allen, 
known from the Ozarks (Robotham and Allen 1988), remains undescribed as 
larvae. Adults of P. jeanae were taken on V-21-1996. 


Paraleptophlebia moerens (McDunnough) — This is an eastern and midwestern 
North American species that has not been previously reported from the Ozarks 
or any areas adjacent to Oklahoma. Oklahoma represents the westernmost 
known range of the species. This species apparently fits the second element of 
the Ozark mayflies identified by McCafferty and Provonsha (1978). That is 
those that range generally into the Midwest, are widely adapted, and common 
in the East and states such as Indiana and Illinois. It has been rarely reported 
from the Southeast (Berner 1975, Harris et a/. 1996). Larvae of P. moerens 
were taken on I-25-1996 and V-21-1996. 


Rhithrogena impersonata (McDunnough) — This species has been reported 
from northeastern North America and north-central USA, including Michigan 
and Wisconsin, where it is relatively common in northern counties (see Leonard 
and Leonard 1962, Flowers and Hilsenhoff 1975, Yanoviak and McCafferty 
1996). Its presence in western Oklahoma evidently represents a significant 
disjunct range extension, as does its previously unreported presence in Kentucky 
(Randolph and McCafferty, unpublished). It has not been reported from other 
areas of the Midwest, including adjacent mountainous areas of Missouri and 
Arkansas. Larvae of R. impersonata were taken on V-18-1996. 


Rhithrogena jejuna Eaton — The distribution of this species is essentially the 
same as that of R. impersonata. Moreover, Leonard and Leonard (1962) found 
the two species cohabiting in Michigan. Its presence in western Oklahoma 
also represents a significant disjunct range extension. It has not been reported 
from adjacent areas of Missouri and Arkansas. Larvae of R. jejuna were taken 
on III-21-1995. 


Stenonema vicarium (Walker) — This is mainly a northeastern and midwestern 
species in North America (Bednarik and McCafferty 1979), although it has 
been reported recently from Alabama by Harris et al. (1996). Its presence in 
Oklahoma represents a notable southwestern disjunction, having not been found 
in Arkansas, or confirmed from Missouri. Larvae of S. vicarium were taken on 
III-13-14, 18, 21-1995, VH-14-15-1995, X-17-1995, and I-16-17-1996. 


198 ENTOMOLOGICAL NEWS 


Table 1. Alphabetical listing of Ephemeroptera collected at Spring Creek sites. Sites are 
described in the text and shown in Figure 1. Asterisks indicate new Oklahoma records. 


SPECIES COLLECTING SITES 


~Acentrellaitunbidauer: aereietietrernie os) cree 1 <= AC) JB 
aI Ne qaniriepuntlelona soeccbbaghoucdcaoc ALB 
Baetisiflavistri la ecy2d ayscerivale terse o,0.0 15s A.B 
BaetismitercalanlSe epee ran sean eyeiste aaa A 
*Callibactis flordanuS.ac.te ile 2 ass « aieis «le 
Gaenistancepsmem. ote ose selec eteere ner A 
(CAEMSIETITSIMNS oeegoeccdocsaseonab one A 
*Ghoroterpes|basalisien)ei ee -eieiae tet) foe B E 
SDiphetonmbarenler cesses era eae A B DE 
EphemetaltnaVerae rer. ttre fla chaperone A B 
SEphemerciiarommdar erie « letecws oe erors 
*Eurylopnellainicolonees tori. oto lela ott B 
*Eurylophellaienoensis 71s ec. scree ks I 
*Eurylophella macdunnoughi ............. F 
ISU AAG So cee suuploepsoneo oases Blac B D FoaG 
HexagenialDilaneatats.ac sere «/<\e ie olsrasei oversee G 
Hexacentallimbatalsee sar tateetstarae sts cto) isle 
+lsonychialbicolor certs) te ole i rl 
-4) Koy NTE NEA SGC Ve rg dan oe cm omntbidc Soap oF 
+Leptophiebramebulosa rie «lane - <li. 9 <1 DEVE 
SWeucrocutaiheDemrr sae cirayee ss cele ol 
PIVSUCKOCULANMINERVal >, ae prr-te)-frctcucfes het cena cn ars 
ENIXeMIOWENSI act meric sotto stat oior dans chet E 
+ Nix INCONSPlCUAies stint ois aueicie sists 
BNIXE) PEMlIalgerareopcee reseyo ysis ichexctene Nousieyer ers E G 
*Paraleptophlebia guttata ................ Is 
*Paraleptophlebiatjeanae’ 7-)4-)-1)< 3). | ot 
+Paraleptophlebialmoerensy= o.-.-@ 2.2) 320-8 = 2) 
*Paraleptophlebiaymollisiaor -)-t121)-)-- oer Cc 
Paraleptophiebia sp»... 1. -saees<ie) = =o) 
+Procloecon rubropictum: . 45.4.5 2s. 
*Riithrosena impersonatary irs cre cee selects ie 
*Rhithrogena jejuna:. © 20/520. Sve ae a trees I 
* Sipnlonurus marshall eyscue seperti ostet Div E 
Stenacron interpunctatum\y-/)5/)5 2 os rye Aa) WB E 
*Stenonetia bednavikiee. ssc. ert ee 
Stenonemaifemoratume si. > cies oie = eis erates DE: 
Stenonemailuteum/s. 4-4 eels re B 
Sténonemaims anwiniv ace cm snccr nectar ok G 
#Stenonema modestuimise yee eee te eee E 
Stenonemait.tenminatum scien. cransene ole B H 
+Stenonema! Vicanum\ ern es roe eer DE: 
‘Iricorythodesispy.-)..08.0 23-1 ae ee B E Gi Shiite 


Aer La Aoe 
Ss) As) eos, 
™ ™ 

7 

QQQQ 

xx xx 


(ele) 


ao Lk 


—— 


Giofy eh\9) 


Vol. 108, No. 3, May & June, 1997 199 


ACKNOWLEDGMENTS 


We thank members of the Spring Creek Coalition and their president, Jennifer Owens (Peggs, 
OK), for support and access to study sites. We also thank A. V. Provonsha (Purdue University) for 
preparation of Figure 1, and R. P. Randolph (Purdue University) for his technical assistance. 
Boris Kondratieff kindly drew our attention to some of the previous Oklahoma mayfly records. 
This paper has been published as Purdue University Agricultural Research Program Journal No. 
15233. 


LITERATURE CITED 


Allen, R. K. and G. F. Edmunds, Jr. 1965. A revision of the genus Ephemerella (Ephemeroptera, 
Ephemerellidae) VIII. The subgenus Ephemerella in North America. Misc. Publ. Entomol. 
Soc. Am. 4: 243-282. 

Bae, Y. J. and W. P. McCafferty. 1991. Phylogenetic systematics of the Potamanthidae 
(Ephemeroptera). Trans. Am. Entomol. Soc. 117: 1-143. 

Bednarik, A. F. and W. P. McCafferty. 1979. Biosystematic revision of the genus Stenonema 
(Ephemeroptera: Heptageniidae). Can. Bull. Fish. Aquat. Sci. 201: 1-73. 

Berner, L. 1958. A list of the mayflies from the lower Apalachicola River drainage. Quart. J. Fla. 
Acad. Sci. 21: 25-31. 

Berner, L. 1975. The mayfly family Leptophlebiidae in the southeastern United States. Fla. 
Entomol. 58: 137-156. 

Berner, L. 1977. Distributional patterns of southeastern mayflies (Ephemeroptera). Bull. Fla. St. 
Mus. Biol. Sci. 22:1-55. 

Flowers, R. W. and W. L. Hilsenhoff. 1975. Heptageniidae (Ephemeroptera) of Wisconsin. Gr. 
Lakes Entomol. 8: 201-218. 

Harris, S. C.1990. Preliminary considerations on rare and endangered invertebrates in Alabama. 
J. Alabama Acad. Sci. 61: 64-92. 

Harris, S. C., B. C. Kondratieff, and B. P. Stark. 1996. New records of Ephemeroptera, 
Plecoptera and Trichoptera from Alabama. Entomol. News 107: 237-242. 

Henry, B. C. and B. C. Kondratieff. 1982. New state records of the mayfly Leptophlebia bradleyi 
Needham. Entomol. News 93:125-126. 

Jester, D. B., E. M. Grigsby, P. D. Powell, B. Carroll, J. Fowlkes, L. Lehman, L. Lindley, W. 
Meads, and B. Stone. 1988. Ecology of Spring Creek, a large Ozark creek in Oklahoma. 
Okla. Wat. Res. Brd. Publ. No. TR88-3: 1-116. 

Kondratieff, B. C. and R. F. Kirchner. 1991. Mayflies. pp. 194-197. Jn: K. Terwilliger [ed.], 
Virginia’s endangered species. McDonald and Woodward Publ. Co., Blacksburg, Va. 

Kondratieff, B. C. and J: R. Voshell, Jr. 1984. The North and Central American species of 
Isonychia (Ephemeroptera: Oligoneuriidae). Trans. Am. Entomol. Soc. 110: 129-244. 

Leonard, J. W. and F. A. Leonard. 1962. Mayflies of Michigan trout streams. Cranbrook Inst. 
Sci. Bloomfield Hills, Mich. 

Lewis, P. A. 1974. Taxonomy and ecology of Stenonema mayflies (Heptageniidae: Ephe- 
meroptera). U. S. E. P. A. Environ. Monit. Ser. Rep. EPA-670/4-74-006: 1-81. 

Magdych, W. P. 1979. The microdistribution of mayflies (Ephemeroptera) in Myriophyllum beds 
in Pennington Creek, Johnston County, Oklahoma. Hydrobiol. 66: 161-175. 

McCafferty, W. P. 1977. Newly associated larvae of three species of Heptagenia (Ephemeroptera: 
Heptageniidae). J. Georgia Entomol. Soc. 12: 350-358. 

McCafferty, W. P. 1982. A new species of Nixe from Indiana (Ephemeroptera: Heptageniidae). 
Gr. Lakes Entomol.15: 227-229. 

McCafferty, W. P. 1994. Distributional and classificatory supplement to the burrowing mayflies 
(Ephemeroptera: Ephemeroidea) of the United States. Entomol. News 105: 1-13. 


200 ENTOMOLOGICAL NEWS 


McCafferty, W. P. and A. V. Provonsha. 1978. The Ephemeroptera of mountainous Arkansas. J. 
Kans. Entomol. Soc. 51: 360-379. 

McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) 
of North and Middle America. Trans. Am. Entomol. Soc. 116; 769-799. 

McCafferty, W. P., B. P. Stark, and A. V. Provonsha. 1990. Ephemeroptera, Plecoptera, and 
Odonata. pp. 43-58. Jn: M. Kosztarab and C. Schaefer [eds.], Systematics of the North Ameri- 
can insects and arachnids: status and needs. Va. Agr. Exp. Stat. Inform. Ser. No. 90-1, Va. 
Polytech. Inst. St. Univ., Blacksburg. 

McKinley, R. E., R. Prins, and L. E. Jech. 1972. Occurrence and distribution of arthropods in 
Travertine Creek, Platt National Park, Murray County, Oklahoma. Proc. Okla. Acad. Sci. 52: 
49-52. 

Morihara, D. K. and W. P. McCafferty. 1979. The Baetis larvae of North America (Ephe- 
meroptera: Baetidae). Trans. Am. Entomol. Soc. 105: 139-221. 

Morse, J. C., B. P. Stark, W. P. McCafferty, and K. J. Tennessen. 1996. Southern Appala- 
chian and other southeastern streams at risk: implications for mayflies, dragonflies and dam- 
selflies, stoneflies, and caddisflies. Jn: G. W. Benz and D. E. Collins [eds.], Aquatic fauna in 
peril: the southeastern perspective. Special Publ. No. 1, Southeast Aquat. Res. Inst., Lenz 
Design & Communications, Decatur, Georgia, in press. 

Pescador, M. L. and L. Berner. 1981. The mayfly family Baetiscidae (Ephemeroptera). Part II 
biosystematics of the genus Baetisca. Trans. Am. Entomol. Soc. 107: 163-228. 

Provonsha, A. V. 1990. A revision of the genus Caenis in North America (Ephemeroptera: 
Caenidae). Trans. Am. Entomol. Soc. 116: 801-884. 

Randolph, R. P. and W. P. McCafferty. 1996. First larval descriptions of two species of 
Paraleptophlebia (Ephemeroptera: Leptophlebiidae). Entomol. News 107: 225-229. 

Reisen, W. K. 1975. The ecology of Honey Creek, Oklahoma: Spatial and temporal distribu- 
tions of the macroinvertebrates. Proc. Okla. Acad. Sci. 55: 25-31. 

Robotham, C. D. and R. K. Allen. 1988. Paraleptophlebia calcarica, n. sp. (Ephemeroptera: 
Leptophlebiidae) from western Arkansas. J. Kans. Entomol. Soc. 61: 317-320. 

Spieth, H. T. 1938. Taxonomic studies on Ephemerida, I: Description of new North American 
species. Am. Mus. Novitates 1002: 1-11. 

Spieth, H. T. 1941. Taxonomic studies on the Ephemeroptera II. The genus Hexagenia. Am. 
Midl. Natural. 26: 233-280. 

Traver, J. R. 1934. New North American species of mayflies (Ephemerida). J. Elisha Mitchell 
Sci. Soc. 50; 189-254, 

Traver, J. R. 1935. Part II. Systematic, pp. 239-739 /n: J. G. Needham, J. R. Traver and Y.-C. 
Hsu [eds.], The biology of mayflies with a systematic account of North American species. 
Comstock Publ. Co., Ithaca, New York. 

Waltz, R. D., D. E. Baumgardner, and J. H. Kennedy. 1996. An atypical larval color form of 
Baetis intercalaris (Ephemeroptera: Baetidae) from Pennsylvania and the Kiamichi River 
Basin of southeastern Oklahoma. Entomol. News 107: 83-87, 

Wilhm, J., J. Cooper, and S. Burks. 1979. Species composition of algae and benthic 
macroinvertebrates in the Blue and Kiamichi Rivers. Proc. Okla. Acad. Sci. 59: 85-88. 

Wilhm, J. H., H. Namminga, and C. Ferraris. 1978. Species composition and diversity of 
benthic macroinvertebrates in Greasy Creek, Red Rock Creek and the Arkansas River. Am. 
Midl. Natural. 99: 444-453. 

Yanoviak, S. P. and W. P. McCafferty. 1996. Comparison of macroinvertebrate assemblages 
inhabiting pristine streams in the Huron Mountains of Michigan, USA. Hydrobiol. 330: 195- 
ZAie 


Vol. 108, No. 2, May & June, 1997 201 


SOUTHWARD RANGE EXTENSION OF THE 
COMMON RINGLET, COENONYMPHA 
TULLIA INORNATA (LEPIDOPTERA: SATYRIDAE)! 


David C. Iftner2 


ABSTRACT: The occurrence of Coenonympha tullia inornata is recorded in New Jersey for the 
first time. Its present distribution in the northeast, and its potential for spread in New Jersey are 
also discussed. 


The common ringlet, Coenonympha tullia inornata (W. H. Edwards, 1861) 
is one of many described subspecies that make up the confusing “tullia” spe- 
cies complex. Some authors consider many or all of these subspecies to be 
distinct species (Miller and Brown, 1981; Opler and Malikul, 1992). This spe- 
cies is circumpolar in occurrence, and extremely variable geographically. In 
North America, this species is found from Alaska to southern California and 
Arizona in the west, throughout much of lower Canada, and in the eastern United 
States as far south as the northern parts of Minnesota, Wisconsin and Michigan, 
and central New England (Opler and Krizak, 1984). 

Since its first appearance in New England (Ferris, 1970; Shapiro, 1974), the 
southward expansion of this species into the northeastern United States has 
been truly remarkable. In a period of less than twenty five years, this species 
has moved steadily southward from where it first entered both Maine and New 
York from Canada to localities as far south as New York City, and Westchester 
County, New York (Glassberg, 1993). 

On 26 June 1994 while conducting survey work for The Nature Conser- 
vancy in the Mashipacong Bogs Preserve, Sussex County, New Jersey, I ob- 
served a small, unfamiliar orange brown butterfly flying across an old pasture 
south of the utility buildings. The individual was soon collected, and surpris- 
ingly turned out to be a worn female of C. t. inornata, a new state record for 
New Jersey. This specimen is housed in the collection of the author. 

It had been speculated that this species would soon be found in New Jersey, 
with most collectors thinking that it would first be located in the northeastern 
portion of the state, somewhere adjacent to Westchester County, New York where 
it was already established. The presence of this butterfly in northwestern New 
Jersey indicates that the spread of this species has probably already encom- 
passed all of extreme northern New Jersey. Subsequent collecting by James J. 
Popelka and the author in 1995 supports this notion, yielding eight additional 


| Received January 26, 1996. Accepted March 1, 1996. 
2 8 Alpine Trail, Sparta, New Jersey 07871. 


ENT. NEWS 108(3): 201-202, May & June, 1997 


202 ENTOMOLOGICAL NEWS 


locality records for this butterfly, including new county records for Passaic 
and Morris counties. The presence of a second brood was also confirmed. 
While it is unknown how far south and east C. ft. inornata will eventually 
spread, suitable habitat exists throughout much of the northern half of New 
Jersey, and northeastern Pennsylvania. Individuals hoping to find this butterfly 
should search open grassy areas (meadows, pastures, old fields, etc.) that have 
a profusion of nectar sources during the months of June and August when the 
two broods are on the wing. Various grasses (Poaceae) serve as hostplants in 
North America, while sedges (Cyperaceae) are utilized in Eurasia (Scott, 1986). 


ACKNOWLEDGMENTS 


I kindly thank John V. Calhoun, John W. Peacock, and two anonymous reviewers for 
critically reviewing the draft of this manuscript. I also wish to thank James J. Popelka for his 
assistance and companionship in the field. 


LITERATURE CITED 


Ferris, C. D. 1970. Occurrence of Coenonympha inornata (Satyridae) in Maine. J. Lepid Soc. 
24(3): 202. 

Glassberg, J. 1993. Butterflies through binoculars: a field guide to butterflies in the Boston-New 
York Washington region. Oxford Univ. Press. New York. 160 pp., 40 pls. 

Miller, L. D. and F. M. Brown. 1981. A catalogue/checklist of the butterflies of America north 
of Mexico. Lepid. Soc. Mem. No. 2. 280 pp. 

Opler, P. A. and G. O. Krizek. 1984. Butterflies east of the Great Plains: an illustrated natural 
history. Johns Hopkins Univ. Press. Baltimore. xv + 294 pp., 54 pls. 

Opler, P. A. and V. Malikul. 1992. A field guide to eastern butterflies. Houghton Mifflin Com- 
pany. New York. xvii + 396 pp., 48 pls. 

Scott, J. A. 1986. The butterflies of North America, a natural history and field guide. Stanford 
Univ. Press. Stanford, California. xii + 583 pp., 64 pls. 

Shapiro, A. M. 1974. Butterflies and skippers of New York State. Search Agriculture 4(3): 1-60. 


(Continued from page 182) 


Some Pine Barrens species are no longer found in natural habitats. The Frosted Elfin 
(Callophrys irus) is nearly restricted to railroads, powerlines and airports, where its host, Bapti- 
sia, still occurs in abundance. Thus, while there may be a large enough protected area of land to 
permit their survival, present management practices appear to be causing declines of some of the 
rarer, highly fire-dependent species found in the Pine Barrens. 

In news of entomological interest, Dr. Schweitzer reported seeing two Question Marks 
(Polygonia interogationis) on February 19, although he had seen his first moths of the year on 
January 3 in Cumberland Co, NJ. Ichneumonids have also been seen at light. There was some 
discussion and speculation on the reasons for low yellowjacket (Vespula sp.) numbers the past 
year. Possible causes included heavy spring rains and increasing implementation of pack-in, 
pack-out rules for food waste in public parks. 

W.J. Cromartie, 
Corresponding Secretary 


Vol. 108, No. 3, May & June, 1997 203 


EFFICIENCY OF ARTHROPOD EXTRACTION 
FROM SOIL CORES! 


Renate M. Snider2, Richard J. Snider 


ABSTRACT: Soil cores (5 x 15 cm) were heat-extracted in Tullgren-type funnels, then floated 
three consecutive times in a saturated sugar solution. The organic supernatant was decanted and 
searched for Collembola and Acari. Heat-extraction efficiency varied greatly between arthropod 
taxa, years, dates within years, and single samples within sampling dates. Using Onychiuridae as 
an example, effects of sugar flotation on the accuracy of population density estimates were dis- 
cussed. 


Soon after the beginning of a long-term study whose goals included moni- 
toring of soil faunal dynamics in Michigan forests (“Project ELF”, Snider & 
Snider 1987), we realized that our data could be severely biased by the method 
used to extract fauna from soil cores. One of the objectives of the study con- 
cerned potential effects of Extremely Low Frequency electromagnetic fields 
on collembolan and acarine populations. An accurate estimate of population 
densities was of primary importance. We therefore sought to validate the effi- 
ciency of standard heat-extraction by post-extraction treatment of samples with 
saturated sugar solution (sugar flotation). Examples of results are presented 
below. 


METHODS 


Soil cores (5 cm diameter, 15 cm depth) were taken in two hardwood forest 
sites in northern Michigan (Test and Control sites of “Project ELF”, described 
in Snider & Snider 1987). Soils in both sites were well-drained Spodosols (Alfic 
Haplorthods, coarse-loamy, mixed, frigid), with approximately 60% sand, 23% 
silt, and 17% clay. Cores were transported in coolers and placed on sieves in 
Tullgren-type extractors (Tullgren 1918). Large, inverted funnels housing 40 
watt light bulbs completely covered those which housed the samples. Heat, 
controlled by rheostats, was increased gradually over a period of 7 days, or until 
samples were completely dry. Following heat-extraction of soil fauna into col- 
lection jars, the dry core samples were removed, bagged in plastic,and treated as 
follows: 1) Soil cores were gently crumbled while still in plastic bags, placed 
into wide-mouth jars (1 liter capacity), moistened with distilled water, and im- 
mersed in saturated sugar solution, leaving approximately 3 cm head space. 
2) Jars were covered with lids, gently shaken several times, and left standing for 
2 hours to allow organic matter to float to the surface. 3) The solution, including 


1 Received June 12, 1996. Accepted December 9, 1996. 
2 Department of Zoology, Michigan State University, East Lansing, Michigan 48824. 


ENT. NEWS 108(3): 203-208, May & June, 1997 


204 ENTOMOLOGICAL NEWS 


roots and other organic debris, was decanted through a 200 mesh sieve into a 
large bowl. Care was taken to not include silt which had settled in the bottom of 
the jar. 4) The sugar solution from the bowl was returned to the jar, and the 
organic matter was rinsed with water and then washed from the sieve into sample 
jars with 95% ethyl alcohol. 5) Steps 2 to 4 were repeated two more times and 
all organic matter from each core was combined in one sample jar. By the third 
decanting, very little organic matter was recovered. When the method was first 
developed, we floated and decanted organic supernatant four times, and sorted 
each “run” separately; results showed that the fourth iteration was not neces- 
sary, since it yielded < 1% of the total specimens extracted by flotation. 

Most Collembola and mites could be identified with a dissecting micro- 
scope. Small specimens, and particularly members of the family Onychiuridae, 
were mounted on slides after rehydration in an acetic acid/water solution for up 
to 24 hours. It was also necessary to train personnel in distinguishing exuviae 
(which were shriveled and twisted, clear to near-opaque, and tended to float in 
alcohol) from whole animals (which resembled live specimens and usually did 
not float in alcohol). 


RESULTS AND DISCUSSION 


From 1986 through 1992, we used sugar flotation for all soil samples taken 
in Test and Control sites of “Project ELF” (Snider & Snider 1987, 1995). In 
Table 1, annual extraction efficiency data are summarized for frequently en- 
countered taxa. Several taxa were always poorly extracted by heat (e.g., 
Onychiuridae in general, and the undescribed mesostigmatid mite “sp. A”). 
Others were consistently obtained with relatively high efficiency (Nanorchestes 
“sp. A”, Isotoma notabilis Schaeffer). Species-specific differences were also 
observed between members of the same family (e.g., Isotomidae: /sotomiella 
minor (Schaeffer) vs. lsotoma notabilis). 

The most commonly used methods for obtaining microarthropods from 
soil samples are Tullgren-type heat extractors and flotation procedures. Many 
variants of these techniques have been devised, compared and reviewed (e.g., 
Edwards & Fletcher 1971; Peterson 1978; Walter et al. 1987; Dunger & Fiedler 
1989; Edwards 1991). None of them are 100% efficient for any given taxon. 
For Collembola, funnel extraction efficiency may range from 16% (Tamura 
1976) to >90% (Petersen 1978; Lussenhop 1971). Edwards (1991) suggested 
that a correction factor should be applied to density estimates obtained by a 
particular technique in a given site. We had hoped to derive species-specific 
correction factors from first-year data (eliminating the need to use sugar flota- 
tion), but it soon became clear that this would not be possible. Not only did 
efficiency of heat extraction vary between dates (Table 2), but it also varied 
greatly between individual samples taken on the same date (Table 3). A poste- 
riori correction for extraction efficiency would have to be applied to individual 
samples, since statistical tests of density differences between dates, sites or 


Vol. 108, No. 3, May & June, 1997 


205 


Table 1. Efficiency of heat-extraction for selected arthropods, based on the total number of speci- 
mens obtained per year; values are given as percent [(N individuals obtained by heat-extraction) 
/ (N total obtained by heat plus flotation)] x 100; T and C = Test and Control sites. 


TAXON 
ACARINA 
Nanorchestes sp. A 


Mesostigmata sp. A 


COLLEMBOLA 
Tullbergia mala Christiansen 
& Bellinger 


Tullbergia granulata Mills 


Isotoma notabilis Schaeffer 


Isotomiella minor (Schaeffer) 


Total Onychiuridae 


Total Isotomidae 


PERCENT EXTRACTED BY HEAT 


SITE 1986 1987 1988 1989 1990 1991 1992 


100.0 
94.9 


36.9 
25.6 


(ey) ey | 


27.8 
23.4 


29.2 
28.5 


92.4 
94,2 


43.8 
50.7 


31.6 
24.5 


THA 
84.8 


OSes oO Ves @=) er 


80.1 
84.4 


23.9 
32.4 


78.3 
Fide 


92:3 
97.8 


38.9 
40.3 


80.3 
68.9 


11.1 
1957 


84.5 
90.1 


34,3 
48.3 


12.8 
12.6 


94.2 
94.1 


42.1 
34.8 


27.2 
9.1 


19.5 
12.8 


94.6 
91.6 


2907 
33.3 


20.3 
13.1 


78.7 
64.6 


87.2 100.0 
98.7 100.0 


20.9 18.9 
Pk ile 


122 ee 
S07 


16:25 4:8 
13°9) 7:8 


TR PET 
93.9 71.1 


NS)ef/ 374s) 
29.3 19.6 


16:59 5 56:6 
13e1. 644 


56.8 51.4 
66.1 48.9 


Table 2. Example of date-specific heat-extraction efficiencies for Tul/bergia mala (Onychiuridae) 
from the Control site, dates | through 13 (early May to late October), 1986; N = 10 samples / 
date. N Heat = number of individuals extracted by heat; N Float = number obtained by sugar 
flotation; % Effic. = percent of total number extracted by heat. 


TOTAL NUMBER OF INDIVIDUALS / DATE 


DATE 1 2 3 
INGieateenery: Ts, AWB" 7s) 
Ni Float 8. 228.25455209% 382 


Gos TBic weary 23.5 m-A5.3'16:4 


22 OT 83 


35 


3365) 281 PSO 410 203 


38 25-1 ~39:0 7-9 


15.8 


196 


10.1 


Ips 118) 


G7 on 823 


241 322 260 331 


36.4 13.4 20.5 6.5 


206 ENTOMOLOGICAL NEWS 


years are based on sample means and associated error estimates. If a general 
correction factor for heat extraction efficiency were uniformly applied to the 
data in Table 3, for example, the result would be a severe distortion rather than 
a correction of data. 

Population density estimates and analyses of community structure and com- 
position must depend to some degree on the methods used to obtain specimens. 
During the first 2 years of the study (1984-1985) we used heat-extraction alone. 
Species which were then thought to be rare or absent in either site (e.g., Willemia 
intermedia Mills and W. similis Mills: Hypogastruridae) were found to be com- 
mon in both communities once we initiated sugar flotation. Annual density es- 
timates, particularly for Onychiuridae (Table 4), also increased significantly 
beginning in 1986. 

Without doubt, standard procedures of handling, extraction, and subsequent 
sorting of soil faunal samples harbor numerous sources of error, variable with 
soil type, operator skills, extraction apparatus and available facilities. In the 
case of “Project ELF”, for instance, lack of temperature-controlled facilities 
precluded the use of high-gradient methods, which have been shown to in- 
crease efficiency (Merchant & Crossley 1970; Edwards & Fletcher 1971; Bieri 
et al. 1986; Crossley & Blair 1991; review in Edwards 1991), and may have 
contributed to variable extraction efficiencies over seasons or years. 

Some arthropods are extracted more efficiently by heat, others by flotation 
(Edwards & Fletcher 1971; Edwards 1991). We used both methods, and found 
it necessary to process all samples in this manner, due to the wide variation 
between individual samples (Table 3). Financial constraints usually preclude 
labor-intensive procedures such as flotation. We recorded sorting times of 10 
to 12 hours for a single sugar-floated forest soil sample, depending on the skill 
of individuals as well as on the structural complexities of samples (amount of 
root material or particulate organic matter). However, for any soil biological 


Table 3. Examples of sample-specific heat-extraction efficiencies for Tullbergia mala 
(Onychiuridae) from the Control site, May 5 and 19, 1986. Abbreviations as in Table 2. 


TOTAL NUMBER OF INDIVIDUALS / SAMPLE 


SAMPLE NO. 1 2 3 4 5 6 7 8 9 10 TOTAL 
May 5, 86: 

N Heat 9 0 0 0) 7 8 0 1 3 50 78 

N Float Nee AK 3 6 7) PRS AIG 2 80) S7/ 254 
% Effic. 3715" 00 O10) O10)" 29:2592221 0:0 1a. Stl Gar, 235 
May 19, 86: 

N Heat 9 4 34° I'S) Sie 9 3 1 34 «14 173 
N Float 2 0 5 20 3 11 Y a, iy. & 209 


% Effic. 81.8 100.0 87.2 47.4 92.5 63.3 30.0 16.7 18.8 60.9 45.3 


Vol. 108, No. 3, May & June, 1997 207 


study where the validity of conclusions rests on the accuracy of population 
estimates, we suggest that investing in a procedure such as the one described 
here may well be worthwhile. 


Table 4. Mean annual density /m2 of Onychiuridae (all species combined) in Test and Control 
sites, 1984 to 1992. 


DENSITY / M2 
METHOD Heat extraction Heat extraction plus flotation 
YEAR 1984 1985 1986 1987 1988 1989 1990 1991 1992 
TEST SITE 2854 3819 8538 13756 11893 12995 8505 10806 10085 


CONTROL SITE 7884 6442 25603 43425 28860 31739 28992 15847 21017 


ACKNOWLEDGMENTS 


We are grateful to Nancy J. Sferra, Frank Calandrino, Tina Schachterle, Thomas Harpstead 
and Chad Schaedig for their technical support and for training the numerous undergraduates who 
helped with sample processing. Support for this research was provided by the Naval Electronic 
Systems Command through a subcontract to IIT Research Institute under contract NO0039-81- 
C-0357. 

LITERATURE CITED 


Bieri, M., V. Delucchi and M. Stadler. 1986. Optimization of extraction conditions in the air 
conditioned Macfadyen funnel extractor for soil arthropods. Pedobiologia 30: 127-135. 
Crossley, D.A. Jr. and J.M. Blair. 1991. A high-efficiency, “low-technology” Tullgren-type 

extractor for soil microarthropods. Agric. Ecosystems Environ. 34: 187-192. 

Dunger, W. and H.J. Fiedler. 1989. Methoden der Bodenbiologie. Gustav Fischer Verlag, Jena. 

Edwards, C.A. 1991. The assessment of populations of soil-inhabiting invertebrates. Agric. Eco- 
systems Environ. 34: 145-176. 

Edwards, C.A. and K.E. Fletcher. 1971. A comparison of extraction methods for terrestrial 
arthropods. Jn: J. Phillipson (ed.), Methods of Study in Quantitative Soil Ecology: Popula- 
tion, Production and Energy Flow. IBP Handbook No. 18. Blackwell Scientific Publications, 
Oxford, pp. 150-185. 

Lussenhop, J. 1971. A simplified cannister-type soil arthropod extractor. Pedobiologia 11:40- 
45. 

Merchant, V.A. and D.A. Crossley, Jr. 1970. An inexpensive, high-efficiency Tullgren extrac- 
tor for soil microarthropods. J. Ga. Entomol. Soc. 5: 83-87. 

Petersen, H. 1978. Some properties of two high-gradient extractors for soil microarthropods, 
and an attempt to evaluate their extraction efficiencies. Nat. Jutl. 20: 95-122 

Snider, R.J. and R.M. Snider. 1987. ELF ecological monitoring in Michigan. I. Description of 
sites for soil biological studies. Pedobiologia 30: 241-250. 

Snider, R.J. and R.M. Snider. 1995. ELF Communications System Ecological Monitoring Pro- 
gram: Soil Arthropods and Earthworms. Final Report. IIT Research Institute Technical Re- 
port D06212-7, 286 pp. Available from National Technical Information Service, U.S. Dept. 
of Commerce. 


208 ENTOMOLOGICAL NEWS 


Tamura, H. 1976. Biases in extracting Collembola through Tullgren funnels. Rev. Ecol. Biol. 
Sol 13: 21-34. 

Tullgren, A. 1918. Ein sehr einfacher Ausleseapparat fiir terricole Tierfaunen. Z. angew. Entomol. 
4: 149-150. 

Walter, D.E., J. Kethley and J.C. Moore. 1987. A heptane flotation method for recovering 
microarthropods from semiarid soils, with comparison to the Merchant-Crossley high gradi- 
ent extraction method and estimates of microarthropod biomass. Pedobiologia 30: 221-232. 


SCIENTIFIC NOTE 


ANTHICUS UNICOLOR (COLEOPTERA: ANTHICIDAE), 
A NEW PREDATOR OF DIURAPHIS NOXIA 
(HOMOPTERA: APHIDIDAE) FROM TURKEY! 


Meryem Elmali2 


The natural enemies of Russian wheat aphid /Diuraphis noxia (Kurdjumoy)] are well docu- 
mented in South Africa (Aalbersberg et al. 1988), in USSR (Kovalev et al. 1991), in Eurasia 
(Gruber et al. 1991), in California (Bernal et al. 1993), in Morocco, Jordan, Syria (Miller et al. 
1992), and in Turkey (Miller et al. 1992, Elmali 1996). Although Anthicidae is commonly re- 
ferred to as a family containing predator species (McQuillan et al. 1982, Smith et al. 1987, Frank 
and Slosser 1991, Elmali 1996), Anthicus unicolor Schm. has not been previously reported in the 
literature as a predator of D. noxia. This anthicid was also a first record for Turkish fauna. It was 
locally seen in only one field infested with D. noxia at high density in Konya province. However, 
after A. unicolor adults suddenly appeared in large numbers, D. noxia declined rapidly. A. unicolor 
has a large brown egg which it deposits at the top of a short white fiber like a neuropteran egg. 
Eggs were where D. noxia colonies were present. A. unicolor was able to prey on D. noxia in 
rolled leaves and other concealed places. Although Sitobion avenae (F.), Rhopalosiphum maidis 
Fitch, R. padi (L.) (Homoptera: Aphididae), and Sipha (Rungsia) elegans Del Guercio (Homoptera: 
Chaitophoridae) were also prey of A. unicolor, D. noxia was preferred. 


LITERATURE CITED 


Aalbersberg, Y.K., M.C. Van Der Westhuizen, and P.H. Hewitt. 1988. Natural enemies and 
their impact on Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae) populations. Bull. Ent. 
Res. 78: 111-120. 

Bernal J. D. Gonzalez E.T. Tatwick J.G. Loya R Gonzalez and W.E. Bendixen. 1993. Natural 
enemies of Russian wheat aphid identified in California. California Agriculture 47: 24-28. 

Elmali, M. 1996. Russian wheat aphid /Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae)] 
in Konya province. 5" International Wheat Conference. Ankara. (in press). 

Frank, W.A. and J.E. Slosser. 1991. An illustrated guide to the predaceous insects of the North- 
ern Texas Rolling Plains. Misc. Publ. Texas Agric. Exp. Sta. No. MP-1718. 23 pp. 

Gruber F., T.J. Poprawski, and E. Rey. 1991. Survey for natural enemies of Diuraphis noxia 
(Mordvilko) in Eurasia. Bulletin-SROP. 14: 102-109. 

(Continued on page 212) 


! Received September 16, 1996. Accepted December 10, 1996. 
2 Plant Protection Department, Faculty of Agriculture, Selcuk University, 42031 Konya, Turkey. 


Vol. 108, No. 3, May & June, 1997 209 


A NEW SPIROPLASMA (ENTOMOPLASMATALES: 

SPIROPLASMATACEAE) RECORD FOR GEORGIA 

AND ATTEMPTED HORIZONTAL TRANSMISSION 
VIA PREDATION! 


Jim Wedincamp Jr.2, Frank E. French, Robert F. Whitcomb4 


ABSTRACT: Firefly larvae, Photuris lucicrescens and P. hebes, did not become infected with 
EC-1 or HYOS-1 Spiroplasma strains after feeding on injected Tenebrio molitor pupae. P. 
lucicrescens and P. hebes larvae had natural Spiroplasma infections but no mollicutes related to 
strains EC-1 or HYOS-1 were recovered. Injected 7: molitor pupae carried both the HYOS-1 (11/ 
13) and EC-1 (10/10) strains for up to five days. Isolations of naturally occurring spiroplasmas 
from P. hebes larvae and adults were identified as group XIX spiroplasmas. These isolations of 
spiroplasmas from Photuris spp. fireflies were the first isolates of group XIX from Georgia. 


Spiroplasmas are helical wall-less prokaryotes belonging in the Division 
Tenericutes, Class Mollicutes and Order Entomoplasmatales (Tully et al 1993; 
Whitcomb and Tully 1982). They are associated with arthropods (Whitcomb, 
1981), especially flies (Diptera) and beetles (Coleoptera) (Hackett, 1990). Little 
is known of the bionomics of Spiroplasma. The natural routes by which 
spiroplasmas colonize tabanids have never been demonstrated and limited sam- 
pling suggests that spiroplasmas do not occur or are rare in tabanid larvae 
(French et al 1992). If this is true we must look for alternate hosts that enable 
tabanid spiroplasmas to overwinter. Fireflies are rich sources of spiroplasmas 
including the group XIV strain EC-1 (Hackett et al 1992) which are also found 
in tabanids. The EC-1 strain was originally isolated from the gut and hemolymph 
of Ellychnia corrusca firefly beetles collected in Maryland (Hackett et al 1992). 
The HYOS-] strain was originally isolated from the tabanid, Hybomitra opaca, 
collected in Montana: 

Spiroplasmas isolated in other studies from fireflies include: the group XIV 
strain EC-1 isolated from Ellychnia corrusca, group XVI strain isolated from 
Photinus pyralis, group XIX strain PUP-1 isolated from Photuris lucicrescens, 
and group XIX strain PUP-1 isolated from Photuris spp. (Hackett et al 1992). 

To simulate natural acquisition of spiroplasmas via predation, tenebrionid 
pupae were injected with tabanid spiroplasmas and offered as prey to firefly 


! Received October 15, 1996. Accepted October 30, 1996. 


2 402 Life Sciences Building, Department of Entomology, Louisiana State University, Baton 
Rouge LA 70803. 


3 Biology Department and Institute of Arthropodology and Parasitology, Georgia Southern Uni- 
versity, Statesboro GA 30460. 


4 Vegetable Laboratory, U.S. Department of Agriculture, Beltsville MD 20705. 


ENT. NEWS 108(3): 209-212, May & June, 1997 


210 ENTOMOLOGICAL NEWS 


larvae, which are predaceous and are potential reservoirs for tabanid spiro- 
plasmas. The possibility that spiroplasmas could be acquired in the course of 
predation has been suggested (Hackett and Clark 1989). 


MATERIALS AND METHODS 


Spiroplasma were isolated from Photuris firefly larvae abdominal viscera 
after surface sterilization, abdominal snip, removal and mincing of viscera in 
MID broth (Whitcomb 1983) and filtration through 0.45um pores using the 
techniques of Markham et al (1983) with the exception that two drops of Photo- 
flo 200 was added to the hypochlorite solution to ensure wetting of the insect 
and surface sterilization. 

Serological deformation tests (Williamson and Whitcomb 1983) with a 
final antiserum dilution of 1:1280, were used to confirm the presence of 
Spiroplasma HY OS-1 and EC-1. 

Firefly larvae, 45 Photuris lucicrescens, were collected September 26-30, 
1993, in Mercer County, New Jersey. Photuris hebes (n = 61) were collected 
March 30-April 4, 1994, in Evans County, Georgia. The larvae were collected 
on the surface of the soil by observation of light flashes emitted from the abdo- 
men. Each larva was placed in a 60 mm disposable petri dish lined with 5.5 cm 
dia. #1 Whatman® filter paper moistened with distilled water and maintained 
at 16-25° C with a 12/12 hr photoperiod. 

Mealworm beetle pupae (Tenebrio molitor L., Coleoptera: Tenebrionidae) 
with lightly colored legs were selected for injection, since pupae with darker 
pigmented legs tended to develop into adults before the firefly larvae fed. Pu- 
pae were randomly assigned to | of 3 groups. Group | was injected with 5.0 wl 
of the HYOS-1 Spiroplasma strain grown in M1D broth using a 10 pl Osge® 
syringe. Group 2 was injected with 5.0 1 of the EC-1 Spiroplasma strain grown 
in MID broth. The titer of injected spiroplasmas, as estimated by serial dilu- 
tion, was 108-109 spiroplasmas per ml (strain HYOS-1) and 107-109 spiroplasmas 
per ml (strain EC-1). Group 3 was injected with 5.0 11 of MID broth as con- 
trols. Each 7: molitor pupa was injected 24 to 30 hrs before placement in a 60 
mm petri dish with a single firefly larva. A second injected pupa was placed 
with each firefly larva after evidence of feeding was observed. Feeding was 
indicated by evisceration or holes in the exoskeleton of the mealworm pupa. 
Viscera of 88 firefly larvae were cultured four days after feeding on a second 
mealworm pupa. Viscera of 11 adults reared from P. hebes larvae from the same 
field collection were also cultured for spiroplasmas three days after eclosion. 


RESULTS AND DISCUSSION 


The firefly larvae used in our test had natural infections of spiroplasmas and 


Vol. 108, No. 3, May & June, 1997 211 


Entomoplasma ellychnia, complicating the isolation of spiroplasmas from gut 
flora. P. Iucicrescens larvae had a natural Spiroplasma infection rate of 6.7% 
(3/45), but all spiroplasmas isolated were closely related to the group XIX PUP- 
1 strain (Whitcomb et al 1992). The P. hebes larvae had a natural infection rate 
of 21% (9/43) with Spiroplasma group XIX, but no mollicutes related to strains 
EC-1 or HYOS-1 were recovered. All nine isolates from 43 larvae of P. hebes 
and four isolates from 11 reared adults (same field collection) were identified 
serologically as group XIX spiroplasmas. Injected T. molitor control pupae car- 
ried both the HYOS-1 (11/13) and EC-1 (10/10) strains for up to five days. 

Firefly larvae, 45 P. lucicrescens and 43 P. hebes, did not become infected 
with EC-1 or HYOS-1 Spiroplasma strains after feeding on injected T. molitor 
pupae. 

The Evans County isolations of spiroplasmas from Photuris spp. firefly lar- 
vae and adults were the first isolates from Georgia. These isolations of group 
XIX from P. hebes firefly larvae and adults confirms the existence of this group 
in the southeastern USA. Shaikh et al (1987) reported the group XIX Spiroplasma 
from mosquitoes in Alabama. Adult P. hebes fireflies either carried the infec- 
tion through metamorphosis or became infected from the soil within one day 
after emergence from the pupal stage. 

Although our findings suggest that acquisition of spiroplasmas via preda- 
tion may not occur it should be noted that alternate routes for the transfer of 
spiroplasmas among fireflies and tabanids do occur. Wedincamp et al (1996) 
revealed evidence that spiroplasmas may be transmitted among adult fireflies 
and tabanids at shared carbohydrate feeding sites, ie. nectaries and honeydew 
deposits. 


ACKNOWLEDGMENTS 


We thank Anthony J. LeCroy (Statesboro, Georgia) for laboratory assistance and field col- 
lections of fireflies; Jonathan Copeland (Georgia Southern University, Statesboro, Georgia) for 
technical advice; Paul S. Johnson (Princeton, New Jersey) for obtaining firefly larvae and Frederic 
V. Vencl (State University of New York at Stony Brook, New York) for identification of fireflies. 


LITERATURE CITED 


French, F.E., R.F. Whitcomb, D.V. Hagan, J.A. Raftner, M. Konai and E.A. Clark. 1992. 
Dynamics of Spiroplasma infections in tabanid (Diptera: Tabanidae) flies, laboratory trans- 
mission, and in vitro tests. International Organization for Mycoplasmology Letter 2: 114. 

Hackett, K.J. and T.B. Clark. 1989. The Mycoplasmas. R.F. Whitcomb and J. G. Tully (eds), 
vol. 5, pp. 113-200. Academic Press, New York. 

Hackett, K.J. 1990. Adaptational biology and spiroplasmas. Zentral. Bacteriol. (Suppl) 20: 23- 
26. 

Hackett, K.J., R.F. Whitcomb, J.G. Tully, J.E. Lloyd, J.J. Anderson, T.B. Clark, R.B. Henegar, 

D.L. Rose, E.A. Clark and J.L. Vaughn. 1992. Lampyridae (Coleoptera): a plethora of mollicute 
associations. Microbial. Ecol. 23: 181-193. 


212 ENTOMOLOGICAL NEWS 


Markham, P., T.B. Clark and R.F. Whitcomb. 1983. Culture media for spiroplasmas from 
arthropods. J.G. Tully and S. Razin (eds) Methods in Mycoplasmology, vol 2. Academic 
Press, New York, pp 217-223. 

Shaikh, A.A., W.E. Johnson Jr., C. Stevens, and A.Y. Tang. 1987. The isolation of spiroplasmas 
from mosquitoes in Macon County, Alabama. J. Am. Mosq. Control Assoc. 3: 289-295. 
Tully, J.G., J.M. Bove, F. Laigret and R.F. Whitcomb. 1993. Revised taxonomy of the Class 
Mollicutes: Proposed elevation of a monophyletic cluster of arthropod-associated Mollicutes 
to ordinal rank (Entomoplasmatales ord. nov.), with provision for familial rank to separate 
species with nonhelical morphology (Entomoplasmataceae fam. nov.) from helical species 
(Spiroplasmataceae), and emended descriptions of the order Mycoplasmatales, family 

Mycoplasmataceae. Int. J. Syst. Bacteriol. 43: 378-385. 

Wedincamp Jr., J., FE. French, R.F. Whitcomb, and R.B. Henegar. 1996. Spiroplasmas and 
entomoplasmas (Procaryotae: Mollicutes) associated with tabanids (Diptera: Tabanidae) and 
fireflies (Coleoptera: Lampyridae). J. Invert. Pathol. 68: 183-186. 

Whitcomb, R.F. 1981. The biology of spiroplasmas. Ann. Rey. Entomol. 26: 397-425. 

Whitcomb, R.F. 1983. Culture media for spiroplasmas. S. Razin and J.G. Tully (eds) Methods in 
Mycoplasmology, vol 1. Academic Press, New York, pp 147-158. 

Whitcomb, R.F., F.E. French, J.G.Tully, G.E. Gasparich, J.M. Bové, P. Carle, E.A. Clark, 
and R. Henegar. 1992. Tabanid spiroplasma serovars. International Organization for 
Mycoplasmology Letter 2: 115. 

Whitcomb, R.F. and J.G. Tully. 1982. Taxonomy and identification of spiroplasmas. Rev. In- 
fect. Diseases 4:148-153. 

Williamson, D.L. and R.F. Whitcomb. 1983. Special serological tests for spiroplasma identifi- 
cation. J.G. Tully and S. Razin (eds) Methods in Mycoplasmology, vol 2. Academic Press, 
New York, pp 249-259. 


(Continued from page 208) 


Kovaley O.V., T.J. Poprawski, A.V. Stekolshchikov, A.B. Vereshchagina and S.A. Gandrabur. 
1991. Diuraphis Aizenberg (Horn., Aphididae): key to apterous viviparous females and re- 
view of Russian language literature on the natural history of Diuraphis noxia (Kurdjumov 
1913). J. Appl. Entomol. 112: 425-436. 

Mc Quillan P.B., J.E. Ireson and KE. Lee. 1982. Some aspects of the invertebrate fauna and its 
role in Tasmanian pastures. Proc. 3 Australasian Conf. on Grassland Invertebrate Ecology 
(edited by Lee K.E.) 101-106. Adelaide, Australia. South Australia Gov’t. Printer. 

Miller, R.H., KS. Pike, L.K. Tanigoshi and L.L. Buschman. 1992. The Russian wheat aphid. 
Diuraphis noxia (Mordvilko) (Homoptera: Aphididae) and natural enemies in Morocco, Jor- 
dan, Syria, and Turkey. Proc. 2™ Turkish Nat’l. Congress Entomol. January 28-31, 1992. 
Adana, 61-62. 

Smith, J.P., RD. Hall and G.D. Thomas. 1987. Arthropod predators and competitors of the 
stable fly, Stomoxys calcitrans (L.) (Diptera: Muscidae) in Central Missouri. J. Kansas Entomol. 
Soc. 60: 562-567. 


Vol. 108, No. 3, May & June, 1997 PN) 


SCIENTIFIC NOTE 


FEMALE PHENGODES FEEDING 
AND AN ASSOCIATED RISK 
(COLEOPTERA:PHENGODIDAE)! 


Richard Stuart Miller2 


Phengodes Illiger are remarkable beetles. Known females are difficult to distinguish exter- 
nally from larvae, except by their larger size and their ovipositional behavior. Although there are 
scattered notes published about Phengodes luminescence and the female’s resemblance to lar- 
vae, their biology remains comparatively unknown. No thorough study, such as that of Zarhipis 
LeConte by Tiemann (1967), has been published. This is surprising, given the interest that 
phengodids generate among amateur and professional entomologists alike. Several biological 
observations of Phengodes sp., which differ from published accounts of phengodid biology, are 
discussed here. 

In May 1986 two female Phengodes were discovered under the bark of a fallen hardwood 
tree 6-7 km west of Cloverdale, Putnam County, Indiana. The tree was supported above the 
ground by another tree, except at the roots. Its bark was loose, but the wood was still solid. Based 
on distribution (Wittmer 1975), these Phengodes were most likely one of three species - P. fusciceps 
intermedia Wittmer, P. plumosa Olivier, or P. laticollis LeConte. One was feeding on Narceus sp. 
(Diplopoda: Spirobolidae) about a meter distant from the other feeding on Polydesmis sp. 
(Diplopoda: Polydesmidae). Phengodids are known millipede specialists (Riley 1887, Balduf 
1935) and these observations corroborate Tiemann’s (1967) observations that prey selection is 
not species-specific. 

I brought both phengodids with additional living Narceus back to the lab and placed each 
phengodid separately in petri dishes with moist Whatman #1 filter paper. Since the Narceus prey 
in the field was approximately the same size as the phengodid female predator, I placed a similar- 
sized millipede sequentially in each petri dish. The following observations were made with the 
aid of a MS dissecting microscope. 

Both phengodids attacked the millipedes immediately after coming into contact with them. 
They encircled their prey, drawing them off the substrate while holding the millipedes’ anterior 
legs directed dorsally. The phengodids then reached around the millipedes in attempts to bite 
their ventral cervical membranes. The first female was successful and held the millipede for 
several minutes in this fashion, while leaving her mandibles embedded within the millipede, 
until the latter was quiescent. The phengodid then used her mandibles to remove the head from 
the rest of the body. She fed on the head first, and then inserted her head into the thorax and 
began to feed. Later, I noted that she had crawled further into the body as she continued to feed. 
This behavioral sequence was similar to that of larval Zarhipis integripennis (LeConte) reported 
by Tiemann (1967). The next morning the millipede was merely an empty exoskeleton with no 
observable liquid or soft contents. 

The second female did not seem able to keep herself free of the millipede’s legs. The con- 
stant motion of those legs ultimately exposed the phengodid’s cervical membranous region to the 
Narceus mandibles. Although the millipede did not pierce the membrane, it evidently crushed 


! Received November 2, 1996. Accepted December 28, 1996. 
2.618 South 15th Ave., Bozeman, MT 59715. 


ENT. NEWS 108(3): 213-214, May & June, 1997 


214 ENTOMOLOGICAL NEWS 


the subesophageal ganglion, since the phengodid immediately ceased attacking the millipede 
and her entire body began to quiver. The millipede moved away and ignored the beetle thereafter. 
Later, the phengodid lay quiescent on her side. The next morning she was able to right herself 
slightly, but she never regained mobility. Tiemann (1967) suggested that millipede defense from 
larval Zarhipis predation is limited to wriggling free and outrunning it. The results of the second 
encounter suggest that this is not the case, at least for Narceus. This may also explain Tiemann’s 
(1967) observation that Zarhipis invert their cervical membrane away from millipede mandibles 
to protect it when biting its prey. 

The first phengodid layed 2 eggs shortly after feeding on the millipede, but the second never 
oviposited in the lab. Barber (1906) reported 53 eggs from a single Phengodes female. There- 
fore, the first female either died before additional oviposition or she had previously oviposited. 
Alternatively, egg number is extremely variable in Phengodes. Both females died within days of 
the first’s oviposition. Soon thereafter, 2 first instar Phengodes eclosed. Tiemann (1967) hypoth- 
esized that female Zarhipis integripennis do not feed as adults, because they ignored several live 
millipedes offered as food. However, adult female Phengodes do feed after mating as exempli- 
fied in both the field and laboratory. 


ACKNOWLEDGMENT 


Thanks to Charles Withrow for determination of the millipedes and to Kevin O'Neil and 
Richard Hurley, two anonymous reviewers and the editor for manuscript review. 


LITERATURE CITED 


Balduf, W. V. 1935. The Bionomics of Entomophagous Coleoptera. John S. Swift, St. Louis. 220 


p- 

Barber, H. S. 1905 (1906). Note on Phengodes in the vicinity of Washington, D. C. Proc. Entomol. 
Soc. Washington 7(4): 196-197. 

Riley, C. V. 1887. [Comments on the discovery of the female of Phengodes]. Entomol. Ameri- 
cana 3(6): 107. 

Tiemann, D. L. 1967. Observations on the natural history of the Western banded glowworm 
Zarhipis integripennis (LeConte) (Coleoptera; Phengodidae). Proc. California Acad. Sci. 
Fourth Series: 35 (12): 235-264. 

Wittmer, W. 1975. The genus Phengodes in the United States (Coleoptera: Phengodidae). Coleopt. 
Bull. 29(4): 231-250. 


Vol. 108, No. 3, May & June, 1997 215 


A PRELIMINARY LIST OF THE BRUCHIDAE 
(COLEOPTERA) OF CUBA! 


Dania Alvarez Marin2, John M. Kingsolver? 


ABSTRACT: A preliminary list of 36 known species of Cuban bruchids with their synonyms is 
assembled as a basis for a faunal study of the island. Known distribution and plant host genera 
are provided. 


Thirty-six species of Bruchidae are herein recorded for the island of Cuba. 
Bruchus centrimaculatus Allard was listed by Blackwelder (1946) but it is a 
European species undoubtedly misidentified and should be stricken from the 
list. Both Suffrian (1870) and Gundlach (1891) listed Bruchus cinerifer Fahraeus 
as occurring in Cuba but this species, now in Mimosestes, has so far been found 
only in southeastern Mexico. Only a critical examination of the specimen in the 
Gundlach Collection will determine its correct assignment. We omit it from 
this list. 

Recent surveys of specimens in the Instituto de Ecologia y Sistematica, 
Havana, the Florida State Collection of Arthropods, Gainesville, and the Stewart 
B. Peck private collection, Ottawa, Ontario, have added seven species to the 
faunal list for the island. Each is designated in the list as NEW RECORD. 

Three species names in the list (Bruchus quadratus, Bruchus pantherinus, 
and Bruchus livens) were proposed by Suffrian in 1870, and although the types 
recently became available, no bruchid specialist has yet critically studied them. 

Faunal lists for other West Indian islands have not been compiled but 14 
species are known from the Dominican Republic, 12 from Jamaica, and 13 
from Puerto Rico. Florida now lists 48 species. 

Generic name(s) following each specific name indicate previous assign- 
ments of that name. Full bibliographic citations are given for valid names. 
Refer to the catalog by Udayagiri and Wadhi (1989) for citations for synonyms. 


PACHY MERINAE 


Caryobruchus gleditsiae (Johansson and Linné, 1767) (Dermestes, Bruchus, Pachymerus) 
= Bruchus fuscus Goeze 1777 
= Bruchus arthriticus Fabricius 1801 
Distribution: Bahamas, Bermuda, Cuba, Dominica, Dominican Republic, El Salvador, 
Guatemala, Honduras. Jamaica, Mexico, Panama, USA 
Host: Chamaedorea, Coccothrinax, Copernica, Hemithrinax, Livistona, Phoenix, Sabal, 
Serenoa, Thrinax, Washingtonia 


I Received May 15, 1996. Accepted November 7, 1996. 


2 Jefe, Departamento de Entomologia, Instituto de Ecologia y Sistematica, Academia de Ciencias 
de Cuba, Habana, Cuba. 


3 Research Associate, Florida State Collection of Arthropods, P.O. Box 147100, Gainesville, FL 
32614-7100 USA. 


ENT. NEWS 108(3): 215-221, May & June, 1997 


216 ENTOMOLOGICAL NEWS 


EL  ——————— 


Caryobruchus marieae Nilsson and Johnson, 1990 
Distribution: Cuba 
Host: Sabal palmetto ( Walt.) Lodd., Chamaeodorea elegans Mart. 


AMBLYCERINAE 
Amblycerini 


Amblycerus baracoensis Kingsolver, 1970 (ernended from baracoenis) 
Distribution: Costa Rica, Cuba, Mexico, Paraguay 
Host: Cordia gerascanthus L. 
Amblycerus cistelinus (Gy\\enhal, 1444) (Spermophagus) 
= Spermophagus centralis Sharp, 1685 
Distribution: Belize, Brazil, Costa Rica, Cuba, Guatemala, Mexico, Nicaragua, Panama 
Host: Guazuma ulmifolia Lam. 
Amblycerus eustrophoides (Schaefter, | 904) (Spermophagus) 
Distribution: Cuba, Costa Rica, Florida, Mexico 
Host: Drypetes laterifolia (Sw.) Krug & Urban 
Amblycerus pygidialis (Suffrian 1470) (Spermophagus) 
= Amblycerus chapini Kingsolver 1970 
Distribution: Cuba, Jamaica 
Host: Cordia gerascanthus L., 
Amblycerus sallei eke\, 1455) (Spermophagus), NEW RECORD 
« Amblycerus martorelli Bridwell 1944 
Distribution: Cuba, Dominican Republic, Haiti, Jamaica, Puerto Rico, Venezuela 
Host: Prosopis juliflora (Sw.) DC, 
Amblycerus schwarai Kingsolver, 1970, NEW RECORD 
Distribution: Cuba, Curagao, Dominican Republic, Grand Cayman, Jamaica, Puerto Rico, St. 
Croix, Virgin Is., Florida, 
Host: Mippomane mancinella L., Tectona grandis \., Ricinis communis L., Randia aculeata 
L, (flowers) 
Amblycerus simulator Jacque\in- Duval, 1457) (Spermophagus) 
Distribution: Costa Rica, Cuba 
Host: Luchea speciosa Willd, 
Amblycerus taeniatus (Suftrian, 1470) (Spermophagus) 
Distribution; Cuba 
Host: Caesalpinia bijuga Swarr 


Spermophagini 


ZLabrotes subfasciatus (Yoheman, |444) (Spermophagus) 

Spermophagus musculus Boheman 1444 

Spermophagus semifasciatus Boheman 1649 

Bruchus cingulatus Kunze in Suffrian 1470 

Bruchus leucogaster Kunze in Sharp 1665 

Spermophagus pectoralis Sharp 1665 

Spermophagus (Zabrotes) semicinctus Horn 1894 

Spermophagus basicornis Vie \926 

Spermophagus minusculus Vie \944 (nisspelling for musculus Hoheman) 
Distribution: Nearly worldwide, especially in tropical and subtropical regions 
Host: Many leguminous genera Cajanus, Cicer, Dolichos, Glycine, Phaseolus, Pisum, Vicia, 

Vigna 


Vol. 108, No. 3, May & June, 1997 217 


BRUCHINAE 
Bruchini 


Bruchus pisorum (Linné, 1758) 

= Laria salicis Scopoli 1763 

= Bruchus cruciger Geoffroy 1785 

= Bruchus sparsus Fabricius 1801 

= Bruchus intermedius Motschulsky 1854 

= Bruchus fabae Brulle 1864 

= Bruchus lunarius Rey 1893 

= Bruchus unifasciatus Rey 1893 

Distribution: Cosmopolitan 

Host: Prefers Pisum sp. but also reported in Cassia, Cytisus, Lathyrus, Phaseolus, Vicia 
Bruchus rufimanus (Boheman, 1833) 

= Bruchus fabae Motschulsky 1854 

= Bruchus granarius Marsham 1802 

Distribution: Cosmopolitan 

Host: Prefers Vicia but also reported in Cicer, Lathyrus, Lens, Lupinus, Phaseolus, Pisum, 

Vigna 


Bruchidiini 


Callosobruchus chinensis (Linné,1758) (Curculio, Mylabris, Bruchus, Pachymerus) 

= Bruchus pecticornis Fabricius 1775 

= Bruchus rufus De Geer 1775 

= Bruchus scutellaris Fabricius 1792 

= Bruchus barbicornis Fabricius 1801 

= Bruchus bistriatus Fabricius 1801 

= Bruchus biguttatus Fabricius 1801 

= Bruchus rufobrunneus Wollaston 1870 

= Bruchus elegans Sturm in Gemminger & Harold 1873 

Distribution: Cosmopolitan, mostly tropical and subtropical, major stored product pest 

Host: Many leguminous genera including Arachis, Cajanus, Cassia, Cicer, Cyamopsis, 
Dolichos, Glycine, Lathyrus, Lens, Phaseolus, Pisum, Vicia, Vigna. Nelumbo (Nelum- 
bonaceae) 

Callosobruchus maculatus (Fabricius, 1775) 

= Bruchus ornatus Boheman 1829 

= Bruchus vicinus Gyllenhal 1833 

= Bruchus ambiguus Gyllenhal 1839 

= Bruchus sinuatus Fahraeus 1839 

Distribution: Cosmopolitan, major stored product pest 

Host: Many leguminous genera including Acacia, Arachis, Cajanus, Cicer, Dolichos, Gly- 

cine, Lathyrus, Medicago, Phaseolus, Pisum, Vicia, Vigna 


Megacerini 


Megacerus ferruginosus Teran and Kingsolver, 1977 
Distribution: Cuba 
Host: Unknown 


218 ENTOMOLOGICAL NEWS 


Megacerus flabelliger (Fahraeus, 1839) (Bruchus), NEW RECORD 
= Bruchus luculentus Boheman 1859 
= Kytorhinus pygidialis Motschulsky 1874 
= Pachybruchus verticalis Pierce 1915 
Distribution: Argentina, Brazil, Costa Rica, Cuba, Ecuador, Mexico, Nicaragua, Venezuela 
Host: Species of Ipomoea and Merremia (Convolvulaceae) 
Megacerus porosus (Sharp) 1885 (Bruchus) 
= Bruchus obliquefasciatus Pic 1932 
= Bruchus notaticollis Pic 1932 
Distribution: Colombia, Cuba, Brazil, Mexico, Puerto Rico 
Host: Merremia aegyptia Dennst. 
Megacerus tricolor (Suffrian, 1870) (Bruchus, Acanthoscelides) 
Distribution: Cuba, Dominican Republic, Nicaragua 
Host: Ipomoea alba L. (= I. bonanox L.) 


Acanthoscelidini 


Acanthoscelides flavescens (Fahraeus, 1839) (Bruchus), NEW RECORD 
= Bruchus ochraceous Schaeffer 1907 (not Baudi 1886) 
= Bruchus ochraceicolor Pic 1913 
Distribution: Costa Rica, Cuba, Dominican Republic, Guatemala, Honduras, Nicaragua, 
Mexico, Panama, Puerto Rico, St. Vincent, USA 
Host: Eriosema, Galactia, Rhynchosia, Tephrosia 
Acanthoscelides livens (Suffrian, 1870) (Bruchus). Transferred by Blackwelder (1946) but 
correct generic placement depends on critical examination of the type in the Gundlach 
Collection. 
Distribution: Cuba 
Host: Unknown 
Acanthoscelides obtectus (Say, 1831) (Bruchus) 
= Acanthoscelides obsoletus of authors (misident.) 
= Bruchus irresectus Fahraeus 1839 
= Bruchus pallidipes Fahraeus 1839 
= Bruchus tetricus Gyllenhal 1839 
= Bruchus subellipticus Wollaston 1854 
= Bruchus incretus Walker 1859 
= Bruchus fabae Riley 1871 
= Bruchus varicornis Motschulsky 1874 
Distribution: Cosmopolitan, major stored product pest 
Host: Many leguminous genera including Cajanus, Cicer, Lathyrus, Lens, Phaseolus, Pisum, 
Sesbania, Vicia, Vigna 
Acanthoscelides pantherinus (Suffrian, 1870) (Bruchus). Transferred by Blackwelder (1946) 
but correct generic placement depends on critical examination of the type in the Gundlach 
Collection. 
Distribution: Cuba 
Host: Unknown 
Acanthoscelides quadratus (Suffrian, 1870) (Bruchus). Transferred by Blackwelder (1946) 
but correct generic placement depends on critical examination of type in the Gundlach 
Collection. 
Distribution: Cuba 
Host: Unknown 


Vol. 108, No. 3, May & June, 1997 219 


Acanthoscelides quadridentatus (Schaeffer, 1907) (Bruchus), NEW RECORD 
Distribution: Cuba, Mexico through Central America to Brazil, USA 
Host: Varieties of Mimosa pigra L., M. strigillosa Torr. & Gray, M. pigra var. berlandieri 
(Gray) Turner 
Ctenocolum podagricus (Fabricius, 1801) (Bruchus, Pseudopachymerus, Acanthoscelides, 
Caryedes), NEW RECORD 
= Bruchus crotonae Fahraeus 1839 
= Bruchus pictifemur Sharp \885 
Distribution: Brazil, Costa Rica, Cuba, Ecuador, El Salvador, Guyana, Mexico, Tobago, 
Venezuela 
Host: Lonchocarpus, Piscidia 
Meibomeus relictus (Suffrian, 1870) (Bruchus) 
Distribution: Cuba 
Host: unknown for Cuba; other species in genus attack seeds of Adesmia, Desmodium, Poiretia, 
and Zornia 
Merobruchus lysilomae Kingsolver, 1988 
Distribution: Bahamas, Cuba, Dominican Republic, Florida, Haiti 
Host: Acacia, Albizia, Lysiloma 
Mimosestes insularis Kingsolver and Johnson, 1978 
Distribution: Cuba, Dominican Republic, Puerto Rico, Hawaii 
Host: Acacia farnesiana (L.) Willd., Prosopis 
Mimosestes mimosae (Fabricius, 1781) (Bruchus) 
= Bruchus dominicanus Jekel 1855 
= Bruchus breweri Crotch 1867 
= Bruchus inornatus Hor 1873 
= Bruchus subrufus Motschulsky 1874 
= Bruchus strigatus Motschulsky 1874 
= Bruchus immunis Sharp 1885 
= Bruchus innotatus Pic 1912 
Distribution: Brazil, Colombia, Costa Rica, Cuba, Curagao, Dominican Republic, Guate- 
mala, Guyana, Haiti, Honduras, Jamaica, Mexico, Netherlands West Indies, Nicaragua, 
Puerto Rico, Trinidad, Venezuela 
Host: Acacia, Caesalpinia, Ceratonia, Parkinsonia 
Mimosestes nubigens (Motschulsky, 1874) (Bruchus, Acanthoscelides) 
= Bruchus sallaei Sharp 1885 
Distribution: Brazil, Costa Rica, Cuba, El Salvador, Guatemala, Hawaii, Honduras, Mexico, 
New Caledonia, Nicaragua, Panama, Philippines, Puerto Rico, USA 
Host: Acacia, Prosopis (occasionally) 
Sennius fallax (Boheman, 1839) (Bruchus, Acanthoscelides) 
= Bruchus californicus Boheman 1859 
= Bruchus xanthopus Suffrian, 1870 
= Bruchus probus Sharp 1885 
Distribution: Costa Rica, Cuba, Guatemala, Jamaica, Mexico, Panama, Puerto Rico, USA 
Host: Cassia 
Sennius morosus (Sharp, 1885) (Bruchus, Acanthoscelides), NEW RECORD 
Distribution: Costa Rica, Cuba, El Salvador, Guatemala, Honduran, Mexico, Nicaragua, 
Panama, USA 
Host: Cassia 
Sennius rufomaculatus (Motschulsky, 1874) (Bruchus, Acanthoscelides) 
= Bruchus instabilis Sharp 1885 
= Bruchus ricanus Pic 1929 


ENTOMOLOGICAL NEWS 


nN 
Nm 
i=) 


= Bruchus turrialbanus Pic 1930 
Distribution: Colombia, Costa Rica, Cuba, El Salvador, Grenada, Guadeloupe, Guatemala, 
Hispaniola, Honduras, Jamaica, Mexico, Nicaragua, Panama, Puerto Rico, St. Vincent, 
Tobago, Trinidad 
Host: Cassia 
Stator bottimeri Kingsolver, 1972 
Distribution: Bahamas, Cuba, USA 
Host: Acacia farnesiana (L.) Willd., Acacia pinetorum Hermann 
Stator rugulosus Kingsolver, 1972 
Distribution: Cuba 
Host: Pithecellobium discolor Britton 


LITERATURE CITED 


Blackwelder, R.E. 1946. Checklist of the coleopterous insects of Mexico, Central America, the 
West Indies, and South America. Bul. U.S. Nat. Mus. 185:551-763. 

Boheman, C.H. 1833. (Bruchidae) /n Schoenherr, C.J. Genera et species curculionidum, cum 
synonymia hujus familiae: species novae aut hactenus minus cognitae, descriptionibus a Dom. 
Leonardo Gyllenhal, C.H. Boheman, et entomologis aliis. Volume 1, Part 1:31-118. 

Boheman, C.H. 1839. (Bruchidae) /n Schoenherr, C. J. Genera et species curculionidum, cum 
synonymia hujus familiae: species novae aut hactenus minus cognitae, descriptionibus a Dom, 
L. Gyllenhal, C.H. Boheman, et entomologis aliis. Volume 5, Part 1:1-456. 

Fabricius, J.C. 1775. Systema entomologia, sistens insectorum classes, species, adiectus syn- 
onyms, locis, descriptionibus, observationibus. Lipsiae. 832 pp. 

Fabricius, J.C. 1781. Species insectorum, exhibentes eorum differential specificas, synonyma 
auctorum, loca natalia, metamorphosis, adjectis observationibus, descriptionibus. Vol. 1. 
Kilonii. 552 pp. 

Fahraeus, O.I. 1839. (Bruchidae) /n Schonherr, C.J. Genera et species curculionidum, cum 
synonymia hujus familiae: species novae aut hactenus minus, cognitae, descriptionibus a 
Dom. Leonardo Gyllenhall, C.H. Boheman, et entomologis aliis. Vol. 5, Part 1: 1-456. 

Gundlach, J. 1891. Contribucion a la Entomologia Cubana. Tome III. Habana. 

Gyllenhal, L. 1833. (Bruchidae), Jn Schoenherr, C.J. Genera et species curculionidum, cum 
synonymia hujus familiae: species novae aut hactenus minus cognitae, descriptionibus a Dom. 
Leonardo Gyllenhal, C.H. Boheman, et entomologis aliis. Vol. 1, Part 1:31-118. 

Jacquelin du Val, P.N.C. 1857. Insectes, Ordre des Coleopteres Linn. /n Sagra, M.R. de la, 
Histoire physique, politique et naturelle de lille Cuba 7:137-328. 

Jekel, H: 1855. Insecta Saundersiana: or characters of undescribed insects in the collection of 
William Wilson Saunders, Esq., F.R.S., F.L.S. Part 1. Van Voorst, London. 

Johannson, B. and C. Linné 1789. Centuria Insectorum in Amoenitates Academicae 6:384- 
415. Palm, J.J. (ed.). Vols. 1-10. Schreiber, J.C.D. Erlangae. 

Kingsolver, J.M. 1970. A synopsis of the subfamily Amblycerinae Bridwell in the West Indies, 
with descriptions of new species. (Coleoptera: Bruchidae). Trans. Amer. Entomol. Soc. 96:469- 
497. 

Kingsolver, J.M. 1972. Synopsis of the genus Stator Bridwell in the West Indies, with descrip- 
tions of new species (Coleoptera: Bruchidae). Proc. Entomol. Soc. Washington 74:219-229. 

Kingsolver, J.M. 1988. Biosystematics of the genus Merobruchus of continental North America 
and the West Indies (Coleoptera: Bruchidae). U.S. Dept. Agric. Tech. Bull. 1744, 63 pp. 

Kingsolver, J.M. and C.D. Johnson. 1978. Systematics of the genus Mimosestes (Coleoptera: 
Bruchidae). U.S. Dept. Agric. Tech. Bull. 1590. 106 p. 

Linnaeus, C. 1758. Systema naturae per regna tria naturae. Edition 10, Vol. 1. 824 p. Holmiae. 


Vol. 108, No. 3, May & June, 1997 221 


Motschulsky, V. 1874. Enumération des nouvelles espéces de coléoptéres rapportés de ses voy- 
ages. Bulletin Société Imperial Naturelle de Moscou 46:203-252. 

Nilsson, J.A. and C.D. Johnson. 1990. A new species of palm bruchid from Cuba and a rede- 
scription of Caryobruchus gleditsiae (L.) (Coleoptera: Bruchidae: Pachymerinae). Coleopt. 
Bull. 44:50-59. 

Say, T. 1831. Descriptions of North American Curculionides and an arrangement of some of our 
known species agreeably to the method of Schoenherr. New Harmony, Indiana. 30 p. 

Schaeffer, C.F.A. 1904. New Bruchidae with notes on known species and list of species known 
to occur at Brownsville, Texas, and in the Huachuca Mountains, Arizona. Sci. Bull. Mus. 
Brooklyn Inst. Arts & Sciences 1:291-306. 

Schaeffer, C.F.A. 1907. New genera and species of Coleoptera. Jour. New York Entomol. Soc. 
12:197-236. 

Sharp, D. 1885. Bruchidae. Biologia Centrali-Americana. Coleoptera 5: 437-504. 

Suffrian, E. 1870. Verzeichniss der von Dr. Gundlach auf der Insel Cuba gesammelten Riisselkafer. 
Archiv fiir Naturgeschichte 36:50-234. 

Teran, A.L. and J.M. Kingsolver. 1977. Revision del genero Megacerus (Coleoptera: Bruchidae). 
Opera Lilloana 25:1-287. 

Udayagiri, S. and S.R. Wadhi. 1989. Catalog of Bruchidae. Memoirs Amer. Entomol. Inst. 
45:1-301. 


BOOKS RECEIVED AND BRIEFLY NOTED 


NAME THAT INSECT. A GUIDE TO THE INSECTS OF SOUTHEASTERN 
AUSTRALIA. 1997. T.R. New. Oxford University Press. 194 pp. Cloth $22.95. 


Also ppbk., price not shown. 
An introductory guide to the Orders and major families of insects in the region, with se- 
lected examples of species to illustrate their diversity and biology. 


THE BIOLOGY OF INSECT OVERWINTERING. 1993. Ppbk. 1995. S.R. 
Leather, K.F.A. Walters, and J. S. Bale. Cambridge University Press. 253 pp. 
Hardbk. $89.95. Ppbk. $29.95. 


Although most insects in temperate climates spend a large proportion of their life in an 
overwintering stage, the study of insect overwintering has been surprisingly neglected. Yet, the 
study of overwintering can offer an insight into the development of insects. This book provides a 
comprehensive account of the various forms of insect overwintering that highlights important 
areas of economic interest. 


MEDICAL ENTOMOLOGY FOR STUDENTS. 1996. M.W. Service. 
Chapman and Hall. 278 pp., Ppbk. 


This book is aimed at students, whether they be physicians, nurses, health officials and 
workers, or students working on a masters’ degree in parasitology or medical entomology. Groups 
considered include mosquitoes, blackflies, sandflies, biting midges, horseflies, tsetse-flies, house 
and stable-flies, fleas, lice, bedbugs, cockroaches, ticks and mites. For each group there is pro- 
vided text on external morphology, life cycle, ecology, behavior and habits, medical importance, 
and control. 


222 ENTOMOLOGICAL NEWS 


CERCOPOIDEA TYPES OF SPECIES DESCRIBED BY 
EDMUND SCHMIDT IN THE U.S. NATIONAL 
MUSEUM OF NATURAL HISTORY, WITH 
LECTOTYPE DESIGNATIONS 
(HOMOPTERA: CERCOPOIDEA)! 


Ai-Ping Liang2 


ABSTRACT: Syntypes and “cotypes” of 35 Cercopoidea species and subspecies described by E. 
Schmidt and recently found in the U.S. National Museum of Natural History are documented. 
Seventeen lectotypes are newly designated. Colsa kriigeri (Schmidt) is emended as C. kruegeri 
and synonymized with C. cavata Walker. The following new combinations are established: 
Liorhina affinis (Schmidt), L. flaviscutellata (Schmidt), L. pulchra (Schmidt), L. similis (Schmidt), 
and L. taeniaticollis (Schmidt); all are transferred from Clovia. 


Between 1906 and 1932, Edmund Schmidt described many new species 
and genera of Cercopoidea. The great majority of Schmidt’s type specimens 
were formerly kept in the Stettiner Museum, but were removed in 1945, after 
the devastation of Stettiner during World War Two. They are now located in the 
Institute of Zoology of the Polish Academy of Sciences (IZPAS), in Warsaw. 

Part of the type series of 32 species and subspecies described by Schmidt 
were recently found and examined in the National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. The depositories of these syntypes 
were never mentioned in Schmidt’s original descriptions. 

The USNM collection contains authentic Schmidt specimens as evidenced 
by Schmidt’s handwritten determination and type labels. Their authenticity as 
syntypes was verified by comparing the label data on the specimens with the 
original descriptions. 

The deposition of the Schmidt material in Washington might be the result of 
the acquisition of the C.F. Baker collection by the USNM. While he was a 
professor at the University of the Philippines, Baker sent specimens he col- 
lected in Singapore, Borneo, and the Philippines to Schmidt for identification. 
Part of the material was later returned to Baker, together with syntypes of spe- 
cies for which Schmidt had duplicate examples. Baker later donated all of his 
personal collection to the National Museum of Natural History. At present, one 
specimen among the series for each species or subspecies bears the species 
name handwritten on a large black-bordered red label. I attribute this labeling 
to C.F. Baker, as evidenced by the label format and handwriting. 

The purpose of the present paper is to document fully the existing type 


| Received July 29, 1996. Accepted September 18, 1996. 


2 Department of Entomology, Institute of Zoology, Academia Sinica, 19 Zhongguancun Lu, Beijing 
100080, PR China. 


ENT. NEWS 108(3): 222-228, May & June, 1997 


Vol. 108, No. 3, May & June, 1997 223 


material. Lectotype and paralectotype designations are made where appropri- 
ate. One incorrect original specific spelling is emended. Also, one new specific 
synonymy and four new species combinations are reported. Most species of 
Cercopoidea can be identified accurately only by using diagnostic characters in 
the males, particularly the structures of the male genitalia. Therefore, in most 
cases the male was designated as the lectotype when syntypes existed. The 
female was designated as the lectotype when no male was present among the 
known syntypes. Lectotype and paralectotype designations were not made for 
the female syntypes of 15 species and subspecies at this time because the male 
syntypes of these species and subspecies exist in the Institute of Zoology, Pol- 
ish Academy of Sciences, Warsaw. 

The list of types is arranged alphabetically by species name. Each name is 
followed by the original generic combination, author, date, page reference, and 
subsequent synonymy or different generic name combination. To document the 
historical status of the syntypes, information from each was recorded exactly as 
given on labels, with (1), (2), (3), and so forth, indicating the sequence of labels 
on the pin from top to bottom. Lastly, my hand printed red lectotype or yellow 
paralectotype label is attached to each specimen so designated. 


affinis, Clovia, Schmidt, 1922a: 9. Here transferred to Liorhina Stal, New 
Combination. 

Lectotype C — (1) ISOLE BATU 1896-97 H. Raa p. (2) (pink label) Typus; (3) E. Schmidt 
(4) Clovia affinis Schmidt GC Edm. Schmidt determ. 1921 (5) (black-bordered red label) 
Clovia affinis Schm. 
apicalis, Simeliria, Schmidt, 1909c: 290. 

i 2 Syntype — (1) Goenoeng Sitoli, Nias, H. Rolle Berlin SW 11 (2) (pink label) Type 
(3) Simeliria apicalis Schmidt Q Edm. Schmidt determ. 1908 (4) (black-bordered red label) 
Simeliria apicalis Schm. 

The male syntype of this species is in the IZPAS. 
borneensis, Leptataspis, Schmidt, 1911: 90. 

Lectotype CO - (1) Nord-Borneo Waterstradt (2) (pink label) Type (3) Leptataspis 
borneensis Schmidt Q Edm. Schmidt determ. 1910 (4) (black-bordered red label) Leptataspis 
borneensis Schm. 
breddini, Trichoscarta, Schmidt, 1910: 85. 

Lectotype 9 - (1) Soekaranda, Januar 1894 Dohrn (2) (pink label) Type (3) Trichoscarta 
Breddini [sic] Schmidt 9 Edm. Schmidt determ. 1910 (4) (black-bordered red label) 
Trichoscarta breddini Schm. 
camerunensis, Clovia, Schmidt, 1922b: 176. 

1 Q Syntype - (1) Kamerun Barombi Conradt (2) (pink label) Typus (3) Clovia 
camerunensis Schmidt Q Edm. Schmidt determ. 1921 (4) (black-bordered red label) Clovia 
camerunensis Schm. 

The male syntypes of this species are in the IZPAS. 
cochleatum, Ectemnonotum, Schmidt, 1909c: 318. 

1 2 Syntype - (1) Sumatra, Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) 
Ectemnonotum cochleatum Schmidt Q Edm. Schmidt determ. 1909 (4) (black-bordered red 
label) Ectemnonotum cochleatum Schm. 

The male syntypes of this species are in the IZPAS. 


224 ENTOMOLOGICAL NEWS 


dohrni, Ectemnonotum, Schmidt, 1909c: 304. 

Lectotype CO - (1) Soekaranda Januar 1894 Dohrn (2) (pink label) Type (3) Ectemnonotum 
dohrni Schmidt CG Edm. Schmidt 1909 (4) (black-bordered red label) Ectemnonotum dohrni 
Schm. 
excellens, Ectemnonotum, Schmidt, 1909c: 311. 

1 Q Syntype - (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) 
Ectemnonotum exellus [sic] Schmidt Q Edm. Schmidt determ. 1909 (4) (black-bordered red 
label) Ectemnonotum excellens Schm. 

The male syntypes of this species are in the IZPAS. 
flavipes, Eoscarta, Schmidt, 1925b: 40. 

Lectotype C - (1) Sandakan Borneo Baker (2) (pink label) Cotypus (3) Eoscarta flavipes 
Schmidt GC Edm. Schmidt determ. 1923 (4) (black-bordered red label) Eoscarta flavipes 
Schm. 

Paralectotype CO - (1) Sandakan Borneo Baker (2) 17884 (3) (pink label) Cotypus (4) 
Eoscarta flavipes Schmidt O Edm. Schmidt determ. 1923. 
flaviscutellata, Clovia, Schmidt, 1922a: 4. Here transferred to Liorhina Stal, 
New Combination. 

Lectotype 9 - (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Typus (3) Clovia 
flaviscutellata Schmidt Q Edm. Schmidt determ. 1921 (4) (black-bordered red label) Clovia 
flaviscutellata Schm. 
formosula, Leptataspis, Schmidt, 1911: 98. 

Lectotype C - (1) Soekaranda Januar 1894 Dohrn (2) (pink label) Type (3) Leptataspis 
formosula Schmidt CO Edm. Schmidt determ. 1910 (4) (black-bordered red label) Leptataspis 
formosula Schm. 
fornax, Leptataspis, Schmidt, 1911: 96. 

Lectotype & — (1) New Guinea (2) (pink label) Type (3) Leptataspis fornax Schmidt C 
Edm. Schmidt determ. 1910 (4) (black-bordered red label) Leptataspis fornax Schm. 
fruhstorferi, Leptataspis, Schmidt, 1927: 10. 

1 Q Syntype — (1) Tonkin, Than-Moi, Juni-Juli, H. Fruhstorfer (2) (pink label) Typus 
(3) Leptataspis fruhstorferi Schmidt Q Edm. Schmidt determ. 1923 (4) (black-bordered red 
label) Leptataspis fruhstorferi Schm. 

The male syntypes of this species are in the IZPAS. 
haglundi, Literna, Schmidt, 1920: 107. 

1 2 Syntype — (1) Kamerun Barombi Conradt (2) (pink label) Type (3) Literna haglundi 
Schmidt Q Edm. Schmidt determ. 1920 (4) (black-bordered red label) Literna haglundi 
Schm. 

The male syntypes of this species are in the IZPAS. 
kriigeri, Sialoscarta, Schmidt, 1906: 279; Colsa kriigeri, Lallemand, 1949: 
91, pl. iv, fig. 8. A junior synonym of Colsa cavata Walker, 1858: 343, New 
Synonymy. 

Lectotype CO — (1) Dohrn Sumatra Soekaranda (2) (pink label) Type (3) Sialoscarta 
kriigeri Schmidt GC Edm. Schmidt determ. 1906 (4) (black-bordered red label) Sialoscarta 
kriigeri Schm. 

Schmidt’s (1906) original spelling kriigeri is here emended to kruegeri (ICZN, Art. 32c), 
Emendation. 

This species was erroneously synonymized with Colsa concinna (Jacobi) by Lallemand 
GL91255121)): 
lutea, Mioscarta, Schmidt, 192S5a: 36. 

Lectotype 9 — (1) Mt Makiling Luzon Baker (2) 21179 (3) (pink label) Cotypus (4) 


Vol. 108, No. 3, May & June, 1997 225 


Mioscarta lutea Schmidt Q Edm. Schmidt determ. 1923 (5) (black-bordered red label) 
Mioscarta lutea Schm. 
marginalis, Plinia, Schmidt, 1919: 381. 

1 Q Syntype — (1) Sumatra, Soekaranda, Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Plinia marginalis Schmidt Q Edm. Schmidt determ. 1919 (4) (black-bordered red label) 
Plinia marginalis Schm. 

1 Q Syntype — (1) Sumatra, Soekaranda, Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Plinia marginalis Schmidt Q Edm. Schmidt determ. 1919. 

The male syntypes of this species are in the IZPAS. 
marquardti, Clovia, Schmidt, 1924: 289. 

1 Q Syntype — (1) Cambodja (Friedrich) (2) (pink label) Typus (3) Clovia marquardti 
Schmidt Q Edm. Schmidt determ. 1924. 

The male syntypes of this species are in the IZPAS. Lallemand (1940: 146) synony- 
mized this species with Lallemandia navigans (Jacobi). 
nox, Clovia, Schmidt, 1922a: 1. 

Lectotype CO — (1) Sumatra Soekaranda Dr. H. Dohrn. S. (2) (pink label) Typus (3) 
Clovia nox Schmidt GC Edm. Schmidt determ. 1921 (4) (black-bordered red label) Clovia 
nox Schm. 
penskyi, Clovia, Schmidt, 1922a: 5. 

1 Q Syntype — (1) Sumatra Soekaranda Dr. H. Dohrn. S. (2) (pink label) Typus (3) 
Clovia penskyi Schmidt 9 Edm. Schmidt determ. 1921 (4) (black-bordered red label) Clovia 
penskyi Schm. 

The male syntypes of this species are in the IZPAS. 
pulchra, Clovia, Schmidt, 1922a: 3. Here transferred to Liorhina Stal, New 
Combination. 

Lectotype GO — (1) SUMATRA SI-RAMBE XII.90-III.91 E. MODIGLIANI (2) (pink 
label) Typus (3) Clovia pulchra Schmidt OC Edm. Schmidt determ. 1921 (4) (black-bordered 
red label) Clovia pulchra Schm. 
punctipennis, Plinia, Schmidt, 1919: 380. 

Lectotype O — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Typus (3) Plinia 
punctipennis O Schmidt Edm. Schmidt determ. 1919 (4) (black-bordered red label) Plinia 
punctipennis Schm. 
pygmaea, Eoscarta, Schmidt, 1909b: 240. 

Lectotype Q — (1) Java (2) (pink label) Type (3) Eoscarta pygmaea Schmidt Q Edm. 
Schmidt determ. 1908 (4) (black-bordered red label) Eoscarta pygmaea Schm. 
roseinervis, Eoscarta, Schmidt, 1925b: 39. 

Lectotype 9 —- (1) Singapore, Coll. Baker (2) (pink label) Cotypus (3) Eoscarta 
roseinervis Schmidt Edm. Schmidt determ. 1923. 

Paralectotype Q — (1) Singapore Coll. Baker (2) 17885 (3) (pink label) Cotypus (4) 
Eoscarta roseinervis Schmidt Q Edm. Schmidt determ. 1923. 
rugosum, Ectemnonotum, Schmidt, 1909c: 313. 

1 Q Syntype — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) 
Ectemnonotum rugosum Schmidt Q Edm. Schmidt determ. 1909 (4) (black-bordered red 
label) Ectemnonotum rugosum Schm. 

The male syntypes of this species are in the IZPAS. 
similis, Clovia, Schmidt, 1922a: 12. Here transferred to Liorhina Stal, New 
Combination. 

1 Q Syntype — (1) Java K. Fruhstorfer S. (2) (pink label) Typus (3) Clovia similis Schmidt 
© Edm. Schmidt determ. 1921 (4) (black-bordered red label) Clovia similis Schm. 

The male syntypes of this species are in the IZPAS. 


226 ENTOMOLOGICAL NEWS 


taeniaticollis, Clovia, Schmidt, 1922a: 13. Here transferred to Liorhina Stal, 


New Combination. 

Lectotype 9 — (1) Balabac (2) E. Schmidt (3) (pink label) Typus (4) Clovia taeniaticollis 
Schmidt 9 Edm. Schmidt determ. 1921 (4) (black-bordered red label) Clovia taeniaticollis 
Schm. 
taeniatifrons, Clovia, Schmidt, 1922b: 178. 

1 Q Syntype —- (1) IS. S. THOME Agua-Ize XII.1900, 400-600 m. L. Fea (2) E. Schmidt 
(3) (pink label) Typus (4) Clovia taeniatifrons Schmidt Q Edm. Schmidt determ. 1921 (5) 
(black-bordered red label) Clovia taeniatifrons Schm. 

The male syntypes of this species are in the IZPAS. 
testaceicollis, Leptataspis, Schmidt, 1911: 88. 

1 Q Syntype — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) 
Leptataspis testaceicollis Schmidt Q Edm. Schmidt determ. 1910 (4) (black-bordered red 
label) Leptataspis testaceicollis Schm. 

The male syntypes of this species are in the IZPAS. 
tricolor borneensis, Suracarta, Schmidt, 1909a: 182. 

1 Q Syntype — (1) Nord-Borneo, Waterstradt (2) (pink label) Type (3) S. Tricolor [sic] 
subsp. borneensis Schmidt Q Edm. Schmidt determ. 1907 (4) (black-bordered red label) 
Suracarta tricolor subsp. borneensis Schm. 

The male syntypes of this species are in the IZPAS. 
tricolor rubroplagiata, Suracarta, Schmidt, 1909a: 177. 

Lectotype CO — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) S. 
tricolor subsp. rubroplagiata Schmidt CO Edm. Schmidt determ. 1907. 

Paralectotype 9 — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) S. 
tricolor subsp. rubroplagiata Schmidt Q Edm. Schmidt determ. 1907 (4) (black-bordered 
red label) Suracarta tricolor rubroplagiata Schm. 

Paralectotype Q — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) S. 
tri. rubroplagiata var. punctata Schmidt Q Edm. Schmidt determ. 1907. 

Paralectotype 9 — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) S. 
tri. rubroplagiata var. 2 punctata Q Edm. Schmidt determ. 1907. 

Paralectotype 9 — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) S. 
tri. rubroplagiata var. 3 punctata Q Edm. Schmidt determ. 1907. 
trimaculata, Considia, Schmidt, 1909b: 239. 

1 Q Syntype — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Type (3) Considia 
trimaculata Schmidt Q Edm. Schmidt determ. 1908 (4) (black-bordered red label) Considia 
trimaculata Schm. 

The male syntypes of this species are in the IZPAS. 


The following six specimens (representing three Schmidt species) labeled 
“cotypus” in the USNM are not designated as part of the type series, because 
the 1923 date on their determination labels suggests that Schmidt did not have 
these specimens when he described these species in 1911. 


costalis, Leptataspis, Schmidt, 1911: 91. 

1 O “Cotype” — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Leptataspis costalis Schmidt OC Edm. Schmidt determ. 1923 (4) (black-bordered red label) 
Leptataspis costalis Schm. 


Vol. 108, No. 3, May & June, 1997 227 


1 Q “Cotype” — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Leptataspis costalis Schmidt Q Edm. Schmidt determ. 1923. 
similis, Leptataspis, Schmidt, 1911: 87. 

1 O “Cotype” — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Leptataspis similis Schmidt C Edm. Schmidt determ. 1923 (4) (black-bordered red label) 
Leptataspis similis Schm. 

1 Q “Cotype” — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Leptataspis similis Schmidt Q Edm. Schmidt determ. 1923. 
sumatrana, Leptataspis, Schmidt, 1911: 85. 

1 GO “Cotype” — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Leptataspis sumatrensis [sic] Schmidt CO Edm. Schmidt determ. 1923 (4) (black-bordered 
red label) Leptataspis sumatrensis [sic] Schm. 

1 9 “Cotype” — (1) Sumatra Soekaranda Dr. H. Dohrn S. (2) (pink label) Cotypus (3) 
Leptataspis sumatrensis [sic] Schmidt Q Edm. Schmidt determ. 1923. 


ACKNOWLEDGMENTS 


I am grateful to Richard C. Froeschner and Thomas J. Henry for their kind assistance during 
my visit to the National Museum of Natural History, Washington, D.C. I thank Lee Herman and 
Randall T. Schuh of the Department of Entomology, American Museum of Natural History, New 
York, for several useful discussions during the preparation of this work. I also thank Howard P. 
Boyd, Lewis L. Deitz, Richard C. Froeschner, Lee Herman and M. W. Nielson for reading the 
manuscript and for their valuable comments in improving the contents of the paper. Support for 
this research was provided by a President Research Fellowship from the Academia Sinica. My 
travel to the USNM was funded by the Theodore Roosevelt Memorial Fund, Postdoctoral Fel- 
lowship Program, American Museum of Natural History, New York. 


LITERATURE CITED 


Butler, A.G. 1877. Description of three homopterous insects in the collection of the British 
Museum. Ann. Mag. Nat. Hist. (4)19: 311-312. 

International Commission on Zoological Nomenclature. 1985. International Code of Zoo- 
logical Nomenclature. 3rd ed. International Trust for Zoological Nomenclature, London, Uni- 
versity of California Press, Berkeley. xx + 338 pp. 

Lallemand, V. 1912. Homoptera, Fam. Cercopidae. Genera Insect., 22(1 43): 1-167 pp., pls. 1-8. 

Lallemand, V. 1940. Troisieme note sur les Cercopides. Bull. Ann. Soc. Entomol. Belg. 80: 135- 
151. 

Lallemand, V. 1949. Revision des Cercopinae (Hemiptera-Homoptera). Premiére partie. Mém. 
Inst. r. Sci. nat. Belg. (2)32: 1-193; pls. 1-4. 

Metcalf, Z.P. 1960. A bibliography of the Cercopoidea (Homoptera: Auchenorrhyncha), 
Raleigh, N.C.: North Carolina State College, 262 pp. 

Metcalf, Z.P. 1961. General catalogue of the Homoptera. Fasc. VII, Cercopoidea. Part 3. 
Aphrophoridae, Raleigh, N.C.: North Carolina State College, 600 pp. 

Metcalf, Z.P. 1962. General catalogue of the Homoptera. Fasc. VII, Cercopoidea. Part 2. 
Cercopidae, Raleigh, N.C.: North Carolina State College, 607 pp. 

Schmidt, E. 1906. Beitrag zur Kenntnis der Cercopiden. Die Gattung Sialoscarta Jacobi. Stettin. 
Entomol. Ztg. 67: 276-280. 

Schmidt, E. 1909a. Neue und bekannte Gattungen und Arten der Subfamilie Cercopinae Stal des 
indoaustralischen Faunengebietes, ein Beitrag zur Kenntnis der Cercopiden. (Hemiptera- 
Homoptera.) Stettin. Entomol. Ztg. 70: 146-187. 


228 ENTOMOLOGICAL NEWS 


Schmidt, E. 1909b. Neue Cercopiden. Stettin. Entomol. Ztg. 70: 239-244. 

Schmidt, E. 1909c. Neue und bekannte Gattungen und Arten der Subfamilie Cercopinae Stal des 
indoaustralischen Faunengebietes, ein Beitrag zur Kenntnis der Cercopiden. (Hemiptera- 
Homoptera, Stettin. Entomol. Ztg. 70: 284-324. 

Schmidt, E. 1910. Neue Gattungen und Arten der Subfamilie Cercopinae Stal, ein Beitrag zur 
Kenntnis der Cercopiden (Hemiptera-Homoptera.) Arch. Naturgesch. 76: 53-112. 

Schmidt, E. 1911. Neue und bekannte Gattungen und Arten der Subfamilie Cercopinae Stal des 
indoaustralischen Faunengebietes, ein Beitrag zur Kenntnis der Cercopiden. (Hemiptera- 
Homoptera.) III. Stettin. Entomol. Ztg. 72: 52-129. 

Schmidt, E. 1919. Beitrage zur Kenntnis aussereuropaischer Zikaden.(Rhynchota Homoptera). 
V. Die Arten des Genus Plinia Stal (Subfam. Aphrophorinae), ein Beitrag zur Kenntnis der 
Cercopiden. Stettin. Entomol. Ztg. 80: 379-382. 

Schmidt, E. 1920. Beitrage zur Kenntnis aussereuropaischer Zikaden. (Rhynchota Homoptera). 
IX. Zwei neue Arten des Cercopidengenus Literna Stal von Kamerun nebst Bemerkungen 
zum Genus Pogonorhinella Schmidt. Arch. Naturgesch. 85: 107-109. 

Schmidt, E. 1922a. Neue Arten des Cercopiden-Genus Clovia Stal (Rhynchota, Homoptera). 
Zool. Meded. 7: 1-26. 

Schmidt, E. 1922b. Beitrage zur Kenntnis aussereuropdischer Zikaden (Rhynchota Homoptera). 
XVI. Drei neue Arten des Cercopiden Genus Clovia Stal von Afrika. Arch. Naturgesch. 88: 
176-179. 

Schmidt, E. 1924. Neue Zikaden-Gattungen und Arten. Entomol. Mitt. 13: 285-297. 

Schmidt, E. 1925a. Fiinf neue Zikadenarten (Forts f.). Societas Entomol. 40: 35-36. 

Schmidt, E. 1925b. Fiinf neue Zikadenarten (Schluss). Societas Entomol. 40: 39-40. 

Schmidt, E. 1927. Zur Kenntnis der Cercopiden des indoaustralischen Faunengebietes (Homop- 
tera). Entomol. Rundschau 44: 9-11. 

Walker, F. 1858. Addenda. In List of the specimens of homopterous insects in the collection of 
the British Museum 1858: 308-369. 


BOOKS RECEIVED AND BRIEFLY NOTED 


DROSOPHILA CELLS IN CULTURE. 1997. G. Echalier. Academic Press. 
702 pp. Hard $135.00. 


This volume summarizes some thirty years of experience in the handling of in vitro cultured 
cells of Drosophila cells. Its main emphasis is on gene transfer methodology, cell responses to 
heat shock, hormonal regulation of genes, and on the expression and mobility of transposable 
elements. 


THE EVOLUTION OF SOCIAL BEHAVIOR IN INSECTS AND 
ARACHNIDS. 1997. J.C. Choe & B.J. Crespi. eds. Cambridge University Press. 
541 pp. Hard $105.00, Ppbk. $47.95. 


The purpose of this book is to explore the causes of the diversity of animal social systems 
from the viewpoint of behavioral ecology as well as the causes of the transitions between coop- 
eration and competition, commensalism, or parasitism. In a series of twenty-four discourses, the 
many contributors seek to explain under what conditions such behavior as parental care, allo- 
parental care, and other forms of altruism among adults have evolved from the individually self- 
ish life histories so common in animals, and how such behavior, once evolved, can be lost. 


Vol. 108, No. 3, May & June, 1997 229 


AMBLYCERUS SCHWARZI (COLEOPTERA: 
BRUCHIDAE) ATTACKING THE SEEDS OF THE 
TROPICAL-ALMOND TERMINALIA 
(COMBRETACEAE) IN CUBA 


Julio Genaro!, John M. Kingsolver2 


ABSTRACT: A bruchid beetle, Amblycerus schwarzi, described in 1970 from the West Indies, is 
recorded for the first time in Terminalia catappa, a new host and host family for the species. 


The tropical-almond terminalia, Terminalia catappa L. (almendro de la 
India) (Combretaceae), is indigenous to the Old World tropics and is culti- 
vated as an ornamental in Cuba (Sauget and Liogier, 1953). The fruit is edible 
and the seed contains a valuable oil, not unlike that of the true almond, Prunus 
dulcis (Miller) Webb. The bruchid, Amblycerus schwarzi Kingsolver, has a 
wide distribution in the West Indies (Kingsolver, 1970). Bruner et al. (1975) 
listed insects that affect the plant in Cuba but do not mention any that feed 
upon the fruits. Martorell (1964) listed only a scolytid beetle infesting seeds of 
tropical-almond in Puerto Rico. Kingsolver (1970) listed for A. schwarzi three 
host plants. These belong to two plant families: Hippomane mancinella L. and 
Ricinis communis L. (Euphorbiaceae) and Tectona grandis L. (Verbenaceae). 

The purpose of this paper is to cite a new host plant record for A. schwarzi 
and to illustrate the damage to fruits of 7: catappa caused by the larvae of the 
beetle. 

Observations were made in June of 1994, June and December of 1995, and 
January of 1996 on 37 trees of tropical-almond terminalia at Brisas del Mar and 
La Veneciana, Guanabo, at the north coast of Havana City, Cuba. Dry fruits 
were gathered from beneath the trees and placed in bags for transport and later 
study. Diameters of beetle emergence holes were measured, and dry fruits with- 
out signs of infestation were kept to await adult emergence. Beetles were iden- 
tified by the junior author (JMK). 

Beetles affected the dry fruits. The larvae feed in the seed without com- 
pletely consuming it, then cut a hole in the seed wall. Larvae pupate inside the 
seed and adults emerge through the hole. Emergence holes of beetles in dry 
fruits (Fig. 1) that remained on the ground provided entry of other invertebrate 
species (i.e. Dermaptera, Coleoptera, Araneae) that used the cavities as refugia 
or as feeding places. From | to 5 beetles (x = 1.8, SD = 1.0, N = 70) emerged 


1 Museo Nacional de Historia Natural, Obispo #61, esquina officios, Plaza de Armas, Habana 
Vieja 10100, Cuba. 


2 Florida State Collection of Arthropods, P.O. Box 147100, Gainesville, FL 32614-7100 USA. 


ENT. NEWS 108(3): 229-230, May & June, 1997 


230 ENTOMOLOGICAL NEWS 


from each fruit. Average diameter of emergence holes was 2.6 mm (SD-0.4, 
range: 2.0-3.5 mm, N = 44). 

A microlepidopteran Blastobasis (probably a new species) (Blastobasidae), 
identified by David Adamski (Smithsonian Institution, Washington, D.C., also 
emerged from the fruits after the larva had partly consumed the seed. Its co- 
coon was spun inside the cavity. Martorell (1964) cited moths of the families 
Olethreutidae and Pyralidae feeding in the fruits of T: catappa in Puerto Rico. 

Voucher specimens are deposited in the Museo Nacional de Historia Natu- 
ral de Cuba, Florida State Collection of Arthropods, Gainesville (beetles), and 
the Smithsonian Institution (moths). 


Fig. 1. Dry fruits of tropical-almond terminalia showing the emergence holes of Amblycerus 
schwarzi. 
ACKNOWLEDGMENTS 


The first author thanks the RARE Center for Tropical Conservation, Philadelphia, Pennsy]- 
vania for generously providing financial support during a visit to several museums in the United 
States. 

LITERATURE CITED 


Bruner, S.C., L.C. Scaramuzza, and A.R. Otero. 1975. Catalogo de los insectos que atacan a 
las plantas ecénomical de Cuba. Segunda edicién. Academia de Ciencias de Cuba, 399 p. 

Kingsolver, J.M. 1970. A synopsis of the subfamily Amblycerinae Bridwell in the West Indies, 
with descriptions of new species. Trans. Amer. Entomol. Soc. 96:469-497. 

Martorell, L.F. 1964. Annotated food plant catalog of the insects of Puerto Rico. Agric. Exper. 
Sta., Univ. Puerto Rico, 303 p. 

Sauget, J.S. and E.E. Llogier. 1953. Flora de Cuba. Vol. III. Contrib. Ocas. Mus. Hist. Nat., 
Colegio La Salle 13:1-502. 


Vol. 108, No. 3, May & June, 1997 231 


SEASONAL DISTRIBUTION OF EMBOLEMIDAE 
(HYMENOPTERA) IN CENTRAL AND 
NORTHERN ARKANSAS! 


C.N. Lewis, J. B. Whitfield 


ABSTRACT: The seasonal distribution of the two eastern U. S. species of embolemid wasps 
(Ampulicomorpha confusa and Embolemus nearcticus) in the Ouachita Mountains and Ozark 
Plateau of central and northern Arkansas is estimated from Malaise trap collections at 12 sites. In 
this limited study A. confusa appeared early in the season, whereas E. nearcticus was active 
during most of the summer season. Discontinuous morphological variation among the sampled 
male individuals of Embolemus was evident, possibly indicating the presence of undiagnosed 
species. 


Wasps of the family Embolemidae are seldom encountered by collectors 
despite their widespread distribution in North America and many other parts 
of the world. About 10 species are known in 2 genera (Finnamore and Broth- 
ers, 1993); in eastern North America two species are currently recorded, 
Ampulicomorpha confusa Ashmead and Embolemus nearcticus (Brues) 
(Krombein, 1979). Although little is known of the biology of embolemids, the 
species in both genera possibly are external parasitoids of homopterans. 
Ampulicomorpha confusa has been reared from fulgoroid nymphs living un- 
der bark in North America (Bridwell, 1958; Krombein, 1979; Wharton, 1989). 
Otherwise, the only available information is that wingless females of other 
species have been reported to have been recovered from ant nests (Donisthorpe, 
1927) and mammal burrows (Heim de Balsac, 1935) in Europe. 

Malaise trap surveys for braconid wasp diversity in the pine-hardwood 
forests of the Ouachita Highlands (Ouachita National Forest) by C. N. Lewis 
(CNL), Malaise collecting in the same areas to test trapping efficiencies by C. 
E. Carlton (CEC; LSU, Baton Rouge, LA), and Malaise trapping to monitor 
parasitoids in the oak-hickory woodlands of the southern Ozark Plateau by J. 
B. Whitfield (JBW) incidentally yielded 81 embolemid wasps spanning a vari- 
ety of localities and dates. Below we summarize observations we made on spe- 
cies identity and seasonal distributions in the two sampled mountain regions. 


MATERIALS AND METHODS 
In 1993, CEC ran 8 Malaise traps in 4 locations in the Ouachita Highlands 
1 Received July 29, 1996, Accepted January 31, 1997. 
2 Department of Entomology, University of Arkansas, Fayetteville, AR 72701. 


ENT. NEWS 108(3): 231-235, May & June, 1997 


232 ENTOMOLOGICAL NEWS 


Figure 1. Two individuals of Embolemus nearcticus from a single trap sample, showing a large 
male and a moderately small one (the smallest were about half the size of the smaller one shown 
here). 


of west central Arkansas as part of a testing program for arthropod sampling 
techniques. During the same year, JBW ran one Malaise trap in a mostly sec- 
ond-growth oak-hickory forest in the Ozark Plateau near the summit of Mt. 
Sequoyah on the east side of Fayetteville, Washington County, Arkansas, as a 
part of a sampling regime for oak herbivores and their parasitoids. The follow- 
ing year, CNL collected insects with Malaise traps in 9 locations (7 of which 
yielded embolemids) in the Ouachita National Forest as part of a braconid 
diversity assessment of mixed pine-hardwood forests that have been managed 
with several different timber harvest strategies by the U. S. Forest Service. 
The embolemids that were incidentally collected in these studies provide us 
with some seasonal and distributional information on the genera. 


Collection sites. ARKANSAS: Montgomery Co., a pooled sample of locations near Mt. 
Ida (CEC collector). Montgomery Co., (1) Womble 1658, NW Mt. Ida; (2) Womble 1651, 9 km 
SW Mt. Ida; (3) Caddo 23, Walsh off FS 471; (4) Oden 1119, Hwy 270 at CR 527; in Scott Co., 
(5) Poteau 1284, S. Parks off CR 19; and in Yell Co., (6) Jessieville 605, 3 km NE Aly off W 
36900 and (7) Cold Springs 284, 6 km NE Blue Ball off FR 3070 (CNL collector). Washington 
Co., Mt. Sequoyah, E. Fayetteville (JBW collector). 


Vol. 108, No. 3, May & June, 1997 233 


Townes style (Townes, 1972) Malaise traps manufactured by Golden Owl 
Publishers, Lexington Park, MD, were used in all 3 studies. JBW and CEC 
harvested their traps at weekly intervals, while CNL harvested her 9 traps for 
one synchronized week each month from late May to late August 1994. All 
samples were collected in 95% ethanol; no attractants were used in the traps. 

Trap samples were brought to the laboratory and identified initially to hy- 
menopteran family. Embolemids were determined by referring to the keys and 
descriptions of Ashmead (1893), Finnamore and Brothers (1993), Reid (1941), 
Richards (1939) and Wharton (1989). 


RESULTS 


A total of 81 male embolemids were sampled in the trapping regimes out- 
lined above. No females were collected; in many species, females are wing- 
less and not often sampled in Malaise traps. One specimen, collected the week 
of 5 May 1993 at Mt. Sequoyah, Washington County (the Ozark site) was 
Ampulicomorpha confusa Ashmead; all others apparently were Embolemus 
nearcticus (Brues) although they showed marked size differences (see below). 
Table 1 summarizes the catches by month through the trapping season. The 
single individual of A. confusa appeared earlier in the season than any sampled 


Size variation in Embolemus 


Number of individuals 


1a kar eho 2.0 2.3 275 2.7 2093.1 3.3 3.5 
Body length in mm 


Figure 2. Bimodal distribution of body size in Embolemus males collected in the traps. The 
larger “morph” also showed discontinuous variation in antennal form and eye size (see test). 


234 ENTOMOLOGICAL NEWS 


specimens of E. nearcticus. In addition, the more southerly Ouachita High- 
lands traps recovered E. nearcticus earlier in the season than the Ozark site. 
The observed abrupt end to the apparent flight season of E. nearcticus at the 
Ouachita sites is likely an artifact of the late-August final sampling period for 
the more numerous CNL traps; however, the earlier appearance of this species 
at those sites is not likely an artifact because all traps at all sites were set up 
early enough to sample the beginning of the flight season. 

Our Embolemus (putatively nearcticus) collections displayed a diversity 
of body sizes, color patterns and head morphologies (see Fig. 1 for an example 
of the size variation). The more easily quantifiable patterns we observed are 
the following. The distribution of body lengths (range 1.8 to 3.4 mm) among 
the wasps appears bimodal, suggesting the existence of two “morphs” (Fig. 2). 
Within the larger “morph,” two forms appear to be present: the first with the 
first flagellomere 1.4 to 1.5x as long as the scape, and with the eye approxi- 
mately 0.45 to 0.5x the (dorsoventral) height of the head, and the second with 
the first flagellomere 0.9 to 1.1x as long as the scape, and the eye 0.55 to 0.6x 
the height of the head. Whether these variants represent the presence of more 
than one species or represent phenotypic effects due to different hosts is not 
possible to say at this point. Clearly more host data would be of considerable 
aid in understanding these wasps. 


Table 1. Embolemid collection dates and locations. Seasonal distribution of Embolemus 
nearcticus (Brues) in Arkansas. The Ouachita Highlands totals are pooled from 11 localities; the 
Ozark totals are from a single locality (see Materials and Methods for localities). All collected 
individuals were males. 


Location Date 
Collector Arkansas 5-11 6-14 7-14 8-17. 9-24 10-2 


JBW (1993) Wash. Co. 1 2 2 | 
CEC (1993) Mont. Co. i4l 


CNL (1994) Mont. Co. 

(1) Wom 1658 2 2 l 
(2) Wom 1651 5 3 15 
(3) Caddo 23 | 
(4) Oden 1119 l 

Scott Co. 
(5) Poteau 1284 l | 

Yell Co. 
(6) Jessieville 605 
(7) Cold Sprs 284 l 4 3 


to 
oo 
~ 


! Collection represents a pooled sample of 4 locations. 


Vol. 108, No. 3, May & June, 1997 235 


DISCUSSION 


No definite host records have been reported for E. nearcticus. Since previ- 
ous records of embolemids are from homopterans living under bark of pines 
and oaks or from ant nests or mammal burrows, it may be that the hosts of E. 
nearcticus are similarly cryptically located. 

Embolemus appeared in our samples later in the season than the Ampulico- 
morpha individual. Since only a single specimen of A. confusa was collected, 
the temporal disparity may not be significant. Wharton (1989), however, stated 
that he has collected A. confusa from April through June in central Texas, 
whereas in the same traps Embolemus appeared from May to October; thus, a 
later but overlapping flight season for Embolemus seems best supported by the 
available evidence. 


ACKNOWLEDGMENTS 


We would like to thank C. E. Carlton, Louisiana State University, Baton Rouge, LA for 
allowing us to use insects he collected while evaluating arthropod sampling techniques. We very 
much appreciate site maps of the Ouachita/Ozark National Forest provided by the USDA Forest 
Service, Southern Forest Experiment Station, Monticello, AR, with special thanks to James B. 
Baker. T. J. Kring and D. C. Steinkraus, as well as two anonymous reviewers, provided useful 
comments on an earlier draft of this manuscript. This study was funded by the Department of 
Entomology of the University of Arkansas at Fayetteville. 


LITERATURE CITED 


Ashmead, W. 1893. A monograph of the North American Proctotrypidae. Bull. U. S. Nat. Mus. 
45: 1-472. 

Bridwell, J. C. 1958. Biological notes on Ampulicomorpha confusa Ashmead and its fulgoroid 
host (Hymenoptera: Dryinidae and Homoptera: Achilidae). Proc. Entomol. Soc. Wash. 60: 
23-26. 

Donisthorpe, H. St. J.K. 1927. The guests of British ants: Their habits and life histories. Routledge 
and Sons, London, England. 268 pp. 

Finnamore, A. T. and D. J. Brothers. 1993. Chapter 7. Superfamily Chrysidoidea, pp. 130-160. 
In: H. Goulet and J. T. Huber. eds., Hymenoptera of the World: An Identification Guide to 
Families. Research Branch, Agriculture Canada Publication 1894/E, 668 pp. 

Heim de Balsac, H. 1935. Ecologie de Pedinomma rufescens Westwood; sa présence dans les 
nids des micromammiféres (Hym. Embolemidae). Revue Francaise d’Entomologie 2: 109- 
125 

Krombein, K. V. 1979. Superfamily Bethyloidea, pp. 1203-1251. Jn: K. V. Krombein, P. D. 
Hurd, Jr., D. R. Smith and B. D. Burks. eds., Catalog of Hymenoptera in America North of 
Mexico, Vol. II. Apocrita (Aculeata). Pp. 1199-2210. Smith. Institut. Press, Washington, D. C. 

Reid, J. A. 1941. The thorax of the wingless and short-winged Hymenoptera. Trans. R. Entomol. 
Soc. Lond. 91: 367-446. 

Richards, O. W. 1939. The British Bethylidae (s. /.) (Hymenoptera). Trans. R. Entomol. Soc. 
Lond. 89: 185-344. 

Townes, H. 1972. A light-weight malaise trap. Entomol. News 83: 239-247. 

Wharton, R. A. 1989. Final instar larva of the embolemid wasp, Ampulicomorpha confusa (Hy- 
menoptera). Proc. Entomol. Soc. Wash. 91: 509-512. 


236 ENTOMOLOGICAL NEWS 


ERGATANDROMORPHISM IN ODONTOMACHUS 
CLARUS (HYMENOPTERA: FORMICIDAE)! 


Juan A. Rodriguez-Garza~ 


ABSTRACT: An ergatandromorph of Odontomachus clarus was collected in Villa de Arriaga, 
San Luis Potosi, Mexico. The specimen is predominantly that of a worker, but the head is notice- 
ably male on the right half, worker on the left half. Abnormal coloration in abdomen is present. 


In ants, as in many other Hymenoptera, all the fertilized eggs become 
females (both queens and workers) while the unfertilized eggs become males. 
Gynandromorphism is a phenomenon of genetic origin. Combinations of male 
and female characteristics may occur laterally & dorso-ventrally, arteropos- 
teriorly or in patches (mosaics). In social insects, the presence of castes pro- 
vides a terminology based on male and female combinations: a queen-male 
combination is called a gynandromorph, a soldier-male combination is a diner- 
gatandromorph and a worker-male combination is an ergatandromorph (Donis- 
thorpe 1929, Wheeler 1937); Berndt and Kremer (1982) postulated heat shock 
as a possible cause of ergatandromorphism. 

Aberrant forms of Odontomachus chelifer (Latreille) and O. bauri Emery 
{under the name O. haematodus) are mentioned by Wheeler (1928) but in both 
cases their abnormalities were caused by parasites. Therefore, the ergatan- 
dromorph briefly described in the next paragraphs is interesting because it is 
the first example of ergatandromorphism in the genus Odontomachus. 

The description of the ergatandromorph is based on Brown’s diagnosis of 
the genus (1976) with special emphasis on cranial relief. The queen and work- 
ers are superficially similar, 7 mm long, but the queen has the mesosoma larger, 
thus is easily differentiated. In both castes, the gaster is black and the rest of the 
body orange reddish without the constriction in the gaster that occurs in most 
other ponerines. 

The sexes differ greatly in head shape. The workers and the queen have a 
pear-shaped head and large mandibles. The male has weak, small mandibles 
and the head is more or less rounded. Also, the eyes and antenna are very differ- 
ent in both sexes, principally in the relative length of the scape and in the num- 
ber of flagellar segments 

The specimen described was collected by me in Villa de Arriaga, San Luis 
Potosi, Mexico, on November 4th, 1984, and stored in 70% ethyl alcohol for 


| Received September 28, 1994. Accepted January 13, 1997. 


2 Division de Ciencias e Ingenieria, Universidad de Quintana Roo, 77000, Chetumal, Q. Roo, 
Mexico. 


ENT. NEWS 108(3): 236-238, May & June, 1997 


Vol. 108, No. 3, May & June, 1997 237 


six years. The mouthparts are damaged. It is deposited in the entomological 
collection of the Colegio de Postgraduados, Montecillos, Mexico. 

The specimen is predominantly a worker, with a morphological abnormal- 
ity in the right half of the head and abnormal coloration in propodeum, petioie 
and gaster (Fig. 1, B). The head is distorted by the diminution of the right side 
(Fig. 1, A). Despite this reduction, the specimen still shows the characteristic 
cranial relief of a worker. The right temporal prominence is well defined, but 
neither the antennal fossa nor the extraocular depression are well defined, there- 
fore the ocular prominence is not defined. The eye is that of a male and in the 
ventral part of the ocular area the coloration is black. The antenna is as in males 
(i.e. it has a short scape, pedicel and an | 1-segmented flagellum). The mandible 
is as in normal males; weak and short, its reduced size allowing us to see the 


Fig. 1. Odontomachus clarus. A — Frontal view of the ergatandromorph head. B — Dorsal view of 
the ergatandromorph, without appendages. 


238 ENTOMOLOGICAL NEWS 


hypopharaux and right maxilla. On the left side, the eye, antenna and mandible 
are fairly normal, and the cranial relief is typical of workers of this species. 

I did not observe any escape behavior at the moment it was collected, only 
erratic and slow movements. 


LITERATURE CITED 


Berndt, K.P and G. Kremer. 1982. Heat shock-induced gynandromorphism in the pharaoh’s 
ant, Monomorium pharaonis (L.). Experientia 38: 798-799. 

Brown, W.L. Jr. 1976. Contribution toward a reclassification of the Formicidae. Part V1: 
Ponerinae, tribe Ponerini: Subtribe Odontomachiti. Section A. Introduction, Subtribal char- 
acters Genus Odontomachus. Studia Entomol. 19: 67-172. 

Donisthorpe, H. 1929. Gynandromorphism in ants. Zool. Anz. {Festband) 52: 92-96. 

Wheeler, W.M. 1928. Mermis parasitism and intercastes among ants. Jour. Exp. Zool 50: 165- 
23i- 

Wheeler, W.M. 1937. Mosaics and other anomalies among Ants. Harvard Univ. Press, Cam- 
bridge, MA. 95 p. 


BOOKS RECEIVED AND BRIEFLY NOTED 


THE BUTTERFLIES OF COSTA RICA AND THEIR NATURAL HISTORY, 
Vol. II: Riodinidae. 1997. P.J. deVries. Princeton University Press. 288 pp., 25 
plates, 59 figs. Cloth $90.00. Ppbk. $29.95. 

This 6 x 9” field guide provides the first detailed treatment of over 250 species of Costa 
Rican butterflies in the family Riodinidae. The taxonomy, distribution, and natural history of 
each taxon is discussed in detail, plus sections on ecology, evolution, behavior, symbioses with 
ants, caterpillar acoustical calls, systematics, collecting and preserving, host-plant relationships, 
and the comparative diversity of riodinid butterfly faunas. 


OLFACTION IN MOSQUITO-HOT INTERACTIONS. 1996. Ciba 
Foundation. John Wiley & Sons. 331 pp. Cloth $84.95. 


A compilation of sixteen papers presented at the Symposium on Olfaction in mosquito-host 
interactions in collaboration with the World Health Organization at the Ciba Foundation, Lon- 
don, 31 Oct. - 2 Nov. 1995. 


THE THERMAL WARRIORS. STRATEGIES OF INSECT SURVIVAL. 1996. 
B. Heinrich. Harvard University Press. 221 pp. Hard $27.00. 


For insects, the struggle to keep their body temperatures in a range suitable for activity is a 
matter of life and death. The ingenious survival strategies that have evolved to control their 
temperatures is the focus of this book. Heinrich tells of bees that shiver to keep their flight 
motors warmed up for the next takeoff, grasshoppers that pant, cicadas that “sweat”, and numer- 
ous other insects with astonishing strategies for survival. Written in the same engaging prose as 
earlier Heinrich offerings. 


Vol. 108, No. 3, May & June, 1997 239 


ABUNDANCE AND SEASONAL ACTIVITY 
OF CANTHARIDAE, LAMPYRIDAE AND LYCIDAE 
(COLEOPTERA) IN A RASPBERRY PLANTATION 
AND ADJACENT SITES IN 
SOUTHERN QUEBEC (CANADA)! 


Claire Levesque, Gilles-Yvon Levesque2 


ABSTRACT: A total of 185 Cantharidae (four taxa), 403 Lampyridae (eight species) and only 18 
Lycidae (four species) adults were collected with pitfall traps and flight intercept traps during 
1987-1989 in a raspberry plantation and adjacent sites in southern Québec. The most common 
species at the soil surface of the raspberry plantation was Cantharis rufa, an adventive species in 
North America. The most abundant species in flight traps in open sites near raspberry plants were 
Cantharis rufa and Pyropyga decipiens. Ellychnia corrusca was the most abundant species cap- 
tured by both methods at the woods-field boundary and in a pine woods. Cantharis rufa activity 
occurred mainly in June during the period of raspberry flowering, and this species probably 
overwintered as larvae. Adults of Ellychnia corrusca were active from May to October. mainly in 
May-June, and adult overwintering was probable. 


Hill (1952) recorded 137 species of insects on cultivated raspberries (Ru- 
bus idaeus L.) in Scotland, including four Cantharidae species, particularly 
Cantharis nigricans Miller and Malthinus flaveolus Paykull. In spite of their 
abundance, they do not appear to play an important part in the ecology of the 
plantation. Recently, Gordon and Woodford (1994) reported the first record of 
damage to raspberry plants by Cantharidae: in May 1991, the stems of expand- 
ing fruiting laterals of red raspberry plantations in Scotland were extensively 
chewed by adults of Cantharis obscura L. Adults of C. obscura can also dam- 
age young tree shoots of Pinus silvestris L. in Poland (Chobotow 1993). Nev- 
ertheless, adults of many Cantharidae species feed on nectar and pollen, and 
some Cantharis L. and Podabrus Westwood species also feed on insect larvae 
(Andow 1982, Wheeler 1977). 

Little is known about the biology of adult Cantharidae, Lampyridae and 
Lycidae within raspberry plantations and other small fruit crops in North 
America. Over a three-year period (1987-1989), we collected these three fami- 
lies in a raspberry plantation and adjacent sites in southern Québec. We now 
present results on the abundance and seasonal activity of adult Cantharidae, 
Lampyridae and Lycidae. 


MATERIALS AND METHODS 


The beetles were collected from early May through late October during a 


1 Received September 3, 1996. Accepted December 3, 1996. 
2 291 rue des Diamants, Fleurimont, Québec, Canada JIG 4A1. 


ENT. NEWS 108(3): 239-244, May & June, 1997 


240 ENTOMOLOGICAL NEWS 


three-year period, in a monocultural raspberry farm at Johnville (45°26'N, 
71°41'W, about 240 m a.s.1.), near Sherbrooke, in southern Québec, Canada. 
We sampled from the Boyne cultivar in this conventionally cultivated planta- 
tion (about 7 ha, on sandy soil). Levesque and Levesque (1992) presented 
detailed information about the study sites, including a sketch-map of the rasp- 
berry farm. 

The ground surface-active beetles were caught with pitfall traps in the fol- 
lowing sites: (1) a raspberry row planted in 1978 (old plants), (2) a raspberry 
row planted in 1985 (young plants), (3) a woods-field boundary (boundary), 
and (4) an adjacent wooded site dominated by eastern white pine, Pinus strobus 
L. (pine woods). Pitfall traps consisted of glass jam jars (450 ml, 6.5 cm diam- 
eter at the top) partially filled with 100 ml of 4% formalin. In the plantation 
(sites 1 and 2), traps were inserted into the soil beneath the canopy as close to 
the cane of raspberry plants as possible. A plywood cover (20 by 20 cm) was 
placed 2.5 cm above the trap to avoid flooding the trap, prevent excessive 
formalin evaporation and capture of flying beetles. In each site, 20 traps were 
set in a row (5 m apart) and were emptied weekly. 

In addition, we studied beetles flying close to the ground with flight inter- 
cept traps at four sites: (A) an open site near the center of the plantation, about 
20 m from old plants; (B) an open site near a pond, about 5 m from young 
raspberry plants; (C) a woods-field boundary; and (D) a pine woods. These 
traps were not located between rows of raspberry plants because of grower’s 
activities and public access during harvest. Flight traps were modified from 
the large-area “window” trap design promoted by Peck and Davies (1980). 
Each consisted of a gray 1.5 mm mesh window screen (1.22 m height, 1.52 m 
width, about 1.85 m2 of surface) fastened to a wooden frame. The frame itself 
was suspended by two lateral triangular wooden supports (1.83 m at the base, 
1.25 m height), 2-4 cm over a set of two galvanized metal pans (25 by 61 cm at 
the top, 7.5 cm deep) which were placed directly on the ground. The insects 
were caught in the pans partially filled with 2% formalin solution into which a 
few drops of detergent were added. We installed one flight trap in each site; the 
pine woods trap (D) was only operated in 1988 and 1989. Samples were col- 
lected twice a week and were pooled weekly. 

In all traps, formalin was used as a killing and preserving agent as well as 
to prevent escape and predation, in spite of its potential selective effect as 
repellent or attractant to some beetle species (Adis, 1979). 


RESULTS AND DISCUSSION 
Abundance of Cantharidae, Lampyridae and Lycidae. We collected a 


total of 185 Cantharidae (four taxa), 403 Lampyridae (eight species) and 18 
Lycidae (four species). Only one captured taxon is known to be an adventive 


Vol. 108, No. 3, May & June, 1997 241 


species in North America, Cantharis rufa L. (McNamara 1991). 

Pitfall trapping resulted in the collection of 63 Cantharidae (two taxa), 140 
Lampyridae (four species) and 6 Lycidae (two species) (Table 1). The most 
common species at the soil surface were C. rufa in the raspberry plantation, 
and Ellychnia corrusca (L.) at the woods-field boundary and in the pine woods 
(Table 1). 


Table 1. Total catches of adults in pitfall traps (1987-1989) at Johnville, Québec. 


Family and taxon Old Young Boun- Pine Total 
plants plants dary woods 

Cantharidae 

Gantharisirufaite eee 13 26 2 0 41 

ROdaDIUsiSPP sesso oie ee 0 0 13 9 22 

Lampyridae 

Ellychnia‘corrusca (L)) 2. 32. 12 2 = 4 l 62 66 133 

Pyropyga decipiens (Harris)......... 0) 3 0 0 3 

Eucidotacatra (Oliv) 2222s oe 0 1 | 0 D 

Photuris pennsylvanica (DeGeer) .... 0 0 2 0 2 

Lycidae 

Geletesrbasalisite@xe a tee Gee ere 0 0 ! 3 4 

Dictyopterus aurora (Herbst) ........ 0) 0) 0) D Z 


Total catches of beetles in flight intercept traps comprised 122 Cantharidae 
(four taxa), 263 Lampyridae (seven species) and 12 Lycidae (four species) 
(Table 2). Adults of C. rufa and Pyropyga decipiens (Harris) flew almost ex- 
clusively in the two open sites A and B near raspberry plants, while E. corrusca 
flew mainly at the boundary and in the pine woods. 

For the most frequent Cantharidae in Finland meadows, a vegetation with 
grasses or a small woods bordering at grass-covered surfaces represent the 
most preferential habitats (Wrede 1963). It is apparently also true for many 
Cantharidae captured at Johnville. 

Ellychnia corrusca is acommon woodland species, usually found in shady 
locations in Ohio and in Ontario (Chénier and Philogéne 1989; Marvin 1965), 
whereas P. decipiens is collected in open grassland fields near small streams or 
ponds in Ohio (Marvin 1965). Our observations on habitat of these two spe- 
cies at Johnville agreed with previous ones. 

Adults of Lycidae from Johnville occurred almost exclusively at the bound- 
ary and in the pine woods (Tables | and 2). Dictyopterus aurora (Herbst) was 
captured only in spring (from May to mid-June). 

Chénier and Philogéne (1989), capturing beetles in Ontario with traps of 
three designs, believed that Lampyridae use tree trunks as rest or display sites. 
In this study, we collected by pitfall traps many adults of two diurnal species, 


242 ENTOMOLOGICAL NEWS 


Table 2. Total catches of adults in flight intercept traps (1987-1989) at Johnville, Québec. 


Family and taxon Open Open Boun- Pine Total 

site site dary woods 

near near 

center pond 

(A) (B) (C) (D) 
Cantharidae 
Cantharisrufaniss arse rregereniesous,- oie 56 40 3 0) 99 
CanthayiS SPD sarc. Or ce ae l 5 0 , 8 
Podabrusssppiee coe ceca e ec 4 0 6 1 11 
Chauliognathus pennsylvanicus 

(DEGEER) Serine ec et eat Ane ae | 3 0 0 4 

Lampyridae 
Elivehnia corrusca (Ib) e..- 1s a or 3 13 131 83 230 
Pyropypa decipiens (Harris)......... 17 7 0 0 24 
Ructaoigatra(Ouv) persed see ee 0 2 1 1 4 
Pyractonema linearis LeC........... 1 0 0 2 
Pyractonema angulata (Say) ........ 0 0 1 0 1 
Pyractonema borealis (Randall)..... . 0 l 0 0 l 
Photinus obscurellus LeC. .......... 0 l 0 0 l 
Lycidae 
Dictyopterus aurora (Herbst) ........ 0) 0 2 4 6 
GeletesioasalisiWie@a mre ee oc 0 0) l 2 3 
Plateros lictor(Newm.) ...........-.- 0 0 1 ] 2 
Calopteron terminale (Say) ......... 0 1 0 0) | 


4 not sampled in 1987. 


C. rufa and E. corrusca; both species probably searched for a protective site 
under the plywood cover placed 2.5 cm above the trap. 


Seasonal Activity of Three Abundant Species. Adults of Cantharis rufa were 
caught by pitfall traps from May until July during the three-year study, but 
mainly in June (Fig. 1). They flew in June-July only and particularly in June. 
The period of raspberry flowering occurred in June and we observed adults on 
raspberry flowers in the plantation sporadically. We suggest that, during their 
mating period, adults of C. rufa could play a minor role in raspberry pollini- 
zation. The sex ratio for trapped adults of this cantharid varied with the method: 
19: 2.2 C in flight traps and 2.4Q: 1C in pitfall traps. We suspect that the 
epigeal activity of females was linked to an oviposition behavior ensuring an 
egg shelter against dessication and predators. This species probably overwin- 
tered as larvae, such as observed in other cantharids (Aitchison 1979). 

We collected adults of Ellychnia corrusca in pitfall and flight traps from 
May through October in the three years (Fig. 1). The first capture peak oc- 


Vol. 108, No. 3, May & June, 1997 243 


Cantharis rufa Ellychnia corrusca 
60 FLIGHT TRAPS FLIGHT TRAPS r 60 
50 L 50 
7 r 
a 40 1987 per 
=| 
HH i988 
ii 30 L 30 
Ww 1989 
om 20 L 20 
ve 
©). .6 L 10 
om 
Lu 
rua) 
= 30 
=) 30 PITFALL TRAPS 
Zz 
20 20 


MAY JUN JUL AUG SEP OCT MAY JUN JUL AUG SEP OCT 


Fig. 1. Seasonal abundance of Cantharis rufa and Ellychnia corrusca adults in pitfall and flight 
traps at Johnville, Québec. 


curred in May-June and is possibly linked to the activity of overwintering beetles 
during their mating period. An autumnal second flight activity peak has been 
observed in September 1989, possibly associated with the new generation dis- 
persal. Females and males were active during the same period and the sex ratio 
of adults collected by both methods was generally close to one. This species is 
most abundant in early spring and fall in Ohio (Marvin 1965), and E. corrusca 
adults occur on trunks of trees from April until November in Québec (Chagnon 
and Robert 1962). Our results agreed with previous observations. 

We collected three adults of Pyropyga decipiens at the soil surface in the 
raspberry plantation during July 1988. Adults from Johnville flew from June 
to August, but mainly in July (18 of 24 catches in flight traps). This lampyrid 
species was collected from 20 June to 1 August in Ohio (Marvin 1965). This 
species probably overwintered as larvae. 


ACKNOWLEDGMENTS 


We appreciate the help of J. McNamara (Centre for Land and Biological Resources Re- 
search, Agriculture Canada, Ottawa, Ontario) for identifications and confirmations of species 
collected in this study. We are grateful to two anonymous reviewers for their comments improv- 
ing the manuscript. We thank Michel Couture and Lucie Labrecque, owners of “La Framboisiére 
de l’Estrie, enr.” at Johnville (Québec). This study was partially supported by the Fonds F. C. A. 
R. (Québec). 


244 ENTOMOLOGICAL NEWS 


LITERATURE CITED 


Adis, J. 1979. Problems of interpreting arthropod sampling with pitfall traps. Zool. Anz. 202: 
177-184. 


Aitchison, C. W. 1979 Winter-active subnivean invertebrates in southern Canada. II Coleoptera. 
Pedobiologia 19: 121-128. 

Andow, D. 1982. Miridae and Coleoptera associated with tulip tree flowers at Ithaca, New York. 
New York Entomol. Soc. 90: 119-124. 

Chagnon, G. and A. Robert. 1962. Principaux coléoptéres de la province de Québec. Seconde 
édition. Les Presses de l’Université de Montréal. Montréal, Canada. 440 pp. 

Chénier, J. V. R. and B. J. R. Philogéne. 1989. Evaluation of three trap designs for the capture 
of conifer-feeding beetles and other forest Coleoptera. Can. Entomol.121: 159-167. 

Chobotow, J. 1993. Cantharidae (Coleoptera) of pine forests in Poland. Fragmenta Faunistica 
36: 147-156. 

Gordon, S. C. and J. A. T. Woodford. 1994. Cantharid beetle feeding damage to Rubus plants in 
eastern Scotland J. Hortic. Sci. 69: 727-730. 

Hill, A. R. 1952. A survey of insects associated with cultivated raspberries in the east of Scot- 
land. Entomol. Mon. Mag. 88: 51-62. 

Levesque, C. and G.-Y. Levesque. 1992. Epigeal and flight activity of Coleoptera in a commer- 
cial raspberry plantation and adjacent sites in southern Québec (Canada): Introduction and 
Nitidulidae. Great Lakes Entomol. 25: 271-285. 

Marvin, D. E., Jr. 1965. A list of fireflies known or likely to occur in Ohio; with special notes on 
species of Ellychnia (Lampyridae: Coleoptera). Ohio J. Sci. 65: 37-42. 

McNamara, J. 1991. Superfamily Cantharoidea, pp. 188-195, Jn Y. Bousquet (ed.), Checklist of 
beetles of Canada and Alaska. Research Branch, Agric. Canada, Publ. 1861/E, Ottawa. 

Peck, S. B. and A. E. Davies. 1980. Collecting small beetles with large-area “window” traps 
Coleopt. Bull. 34: 237-239. 

Wheeler, A. G., Jr. 1977. Studies on the arthropod fauna of alfalfa. V1l. Predaceous insects. Can. 
Entomol.109: 423-427. 

Wrede, H. 1963. Uber das Auftreten der Cantharis- und Rhagonycha-Arten (Coleoptera, 
Cantharidae) auf siidfinnischen Wiesenbéden. Notulae Entomol. 43: 77-91. 


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ew species of Xenos (Strepsiptera:Stylopidae) from 
Arizona, with discussion of other North American 
species J.L. Cook, B. Mathison 


Description of pupa of Cyrnellus fraternus (Trichoptera: 
Polycentropodidae), with notes on variation 
in pupal case construction Z.B. Johnson, J.H. kennedy 


Description of mature larvae of Microdynerus exilis 
and M. timidus (Hymenoptera: Vespidae) 
J. Tormos, J.D. Asis, S.F. Gayubo, F. Torres 


Linnaean species of Conops (Diptera: Conopidae, Muscidae, 
Sciomyzidae, Syrphidae, & Tachinidae), with designa- 
tions of lectotypes i. Christian Thompson 


Studies on some moth flies (Diptera: Psychodidae), with 
the first record of Clogmia albipunctata in central 
Europe Doreen Werner 


Contribution to the systematics of the genus Cheleocloeon 
(Ephemeroptera: Baetidae) C.R. Lugo-Ortiz, W.P. McCafferty 


Notes on some fleas (Siphonaptera) from Amazonas and 
Bahia States, Brazil M.W. Hastriter, N.E. Peterson 


New distributional record for Haideoporus texanus 
(Coleoptera: Dystiscidae), a stygobiontic beetle 
from the Edwards aquifer, Texas D.E. Bowles, R. Stanford 


Synaptonecta issa (Heteroptera: Corixidae): first New 
World record of an Asian water bug in Florida 
J.T. Polhemus, R.P. Rutter 


New state and U.S. records and distributional notes 


for Heteroptera J.T. Polhemus 
New records for scarab beetles (Coleoptera: Scara- 

baeidae) from North Dakota & Minnesota Paul K. Lago 
Annotated checklist of Mecoptera (Scorpionflies) 

of Arkansas H.W. Robison, G.W. Byers, C.A. Carlton 


SCIENTIFIC NOTES: 
Observation of burying beetle, Nicrophorus tomentosus 


(Coleoptera: Silphidae) burying a mole Paul P. Shubeck 
Mating behavior of Dasymutilla occidentalis 

(Hymenoptera: Mutillidae) Jeffery K. Tomberlin 
Name adjustments and a new synonym for North 

American Ephemeroptera species W.P. McCafferty 
New records of notonectids (Heteroptera) for 

Pennsylvania A.M. Yeakel, E. Larsen 


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Vol. 108, No. 4, September & October, 1997 245 


A NEW SPECIES OF XENOS FROM ARIZONA, WITH 
DISCUSSION OF OTHER NORTH AMERICAN SPECIES 
(STREPSIPTERA: STYLOPIDAE)! 


Jerry L. Cook2, Blaine Mathison 


ABSTRACT: Xenos kifunei, a new species of Strepsiptera in the family Stylopidae, is described 
from Arizona. During its developmental stages Xenos kifunei is a parasite of Polistes comanchus 
navajoe. This becomes the sixth recognized Xenos species from North America, all parasites of 
Polistes species. The status of other North American Xenos is discussed. 


The genus Xenos was described by Rossius (1793) and is distinguished 
from other genera of Stylopidae by several key characters. Xenos males have 
four antennal segments, the last two are flattened and much longer than the 
first two, and the first radial wing vein is either continuous, or, if broken, with 
the apical segment in line with the proximal. All North American species have 
maxillary palps that are shorter than the mandibles. Female Xenos have four, 
or rarely five, genital openings, compared to five in other Stylopidae. Although 
not a distinguishing character, Xenos species have thus far been found only in 
hosts from the family Vespidae. Prior to this description, there were 37 recog- 
nized species, most parasitic on species of Polistes. All North American spe- 
cies are parasites of Polistes. Valid North American species include Xenos peckii 
Kirby 1813; Xenos nigrescens Brues 1903; Xenos pallidus Brues 1903; Xenos 
hunteri (Pierce 1909); and Xenos rubiginosi (Pierce 1909). 

The North American species of Xenos were last reviewed by Bohart (1941). 
In this revision, Bohart synonymized ten species with either X. pallidus or X. 
peckii and recognized only these two species as being valid and distinct. Bohart 
did not comment on three other named species (X. nigrescens, X. hunteri, and 
X. rubiginosi), but instead left their status as questionable. Since Bohart’s revi- 
sion, no new North American species of Xenos have been described, nor has 
any taxonomic work been carried out on this group. 

All specimens of the new species of Xenos described below were recovered 
from Polistes comanchus navajoe Cresson, collected in Arizona, USA. We ex- 
amined three females and five males and found them to have characters suffi- 
ciently different from other species of Xenos to warrant this description. Meas- 
urements in this description are presented as a range from these specimens. 


1 Received March 3, 1997. Accepted March 22, 1997. 
2 Department of Entomology, Texas A & M University, College Station, TX 77843-2475, USA. 
3 Department of Entomology, University of Arizona, Tucson, AZ 85721, USA. 


ENT. NEWS 108(4): 245-252, September & October, 1997 


SEP 29 1097 | 
LIBRARIES 


~ 


246 ENTOMOLOGICAL NEWS 


Xenos kifunei, Cook and Mathison, NEW SPECIES 
(Figures 1 A-C, 2, & 3 A-H) 
(all measurements in mm) 


Female Description. Cephalothorax dark brown posteriorly. Anterior areas orange-tan (Fig. 1B). 
First abdominal segment dark brown and tan. Remaining abdominal segments tan with a some- 
what darker dorsal region (Fig. 1A). There is some variation in coloration between specimens, 
some are much darker than others but the pattern of coloration is consistent in all specimens. 
Total body length 6.38 to 7.40. Greatest body width, measured across the abdomen, 1.90 to 2.13. 
Cephalothorax length 1.22 to 1.24; its greatest width (behind the first abdominal spiracles) 1.35 
to 1.43. Width at first abdominal spiracles 1.24 to 1.32. First abdominal spiracle lateral and 
somewhat ventral. Distance between mandibles 0.23 to 0.24. Mandibles shaped as in figure 1C. 


Fig. 1A-C. Xenos kifunei, NEW SPECIES, adult female. A. dorsal habitus. B. dorsal cephalotho- 
rax and part of first abdominal segment. C. Right mandible. 


Vol. 108, No. 4, September & October, 1997 247 


Abdomen with one genital opening on each of segments II-V, and one specimen has a very small 
genital opening on abdominal segment VI. 


Male description. Body coloration brown, ranging from almost black to light tan or yellow. 
Total body length 3.14 to.3.53. Head somewhat dumb bell-shaped, with large, dark eyes (Fig. 2). 
Antennae (Fig. 3A) typical of Xenos; third and fourth segments approximately the same size and 
covered with numerous sensory cups, the latter absent from segments I and II. Antennal segment 
lengths; I = 0.10, II = 0.06, III = 0.87, and IV = 0.83. Vertex of head pointed, triangular. Man- 
dibles and maxillae shaped as in Fig. 3F. Mandibles crossing in front of mouth, yellow, clear, 
length 0.26 to 0.30. Maxillae and palps yellow to light brown, combined length 0.21 to 0.25. 
Legs yellow ventrally, light brown dorsally. Tarsal segment I on prothoracic leg with one large 
sensory cup on its ventral surface. Femur of metatarsal leg somewhat bowed. Shapes of leg 
segments as in Fig. 3B, 3C, and 3D. Prothoracic leg measurements; coxa = 0.33 0.34, 
trochantofemur = 0.40-0.44, tibia = 0.43-0.47, tarsi: I = 0.22-0.23, I] = 0.17-0.19, II] =0.11-0.14, 
IV = 0.15-0.21. Mesothoracic leg measurements; coxa = 0.39. trochantofemur = 0.56-0.58, tibia 
= 0.48-0.52, tarsi: I = 0.18-0.19, Il = 0.12-0.14, IIIf = 0.10, IV = 0.14-0.17. Metathoracic leg 
measurements; trochanter = 0.21-0.25, femur = 0.52-0.55, tibia = 0.41 -0.47, tarsi I = 0.17-0.19, 
II = 0.1-0.16, IN = 0.11-0.14, IV 0.14-0.19. Fore wing club-shaped with one vein (R) to half its 
length, total wing length 0.33-0.55. Hind wing (Fig. 3E) clear, except for shaded area between Sc 


O.5mm 


Fig. 2. Xenos kifunei, NEW SPECIES, male habitus. 


248 ENTOMOLOGICAL NEWS 


and R veins. Sc vein heavy, reaching half-way to wing margin; R, and R, divided by gap half as 
long as R,; R, and R, appear to be in line, R, is proximal, R, is distal; R, three times length of R,; 
R , slightly longer than R,; R, five times as long as R,; MA = CuA, = =CuA,; CuP heavy; no veins 
reach wing margin. Thoracic segments as in Fig. 2. “Abdomen brown, slender. Abdominal seg- 
ment X dark brown, triangular in dorsal view (Fig. 2). Dorsal abdominal segment IX triangular 
(Fig. 2), prescutum shape distinctive, shaped as in Fig. 2. Aedeagus (Fig. 3G) clear-yellow, bowed 
with sharp dorsal projection, no ventral projection or basal hump, thickest in middle of shaft, 
length 0.25-0.28. One male cephalotheca of puparium was observed (Fig. 3H). Cephalotheca 
dark brown, total size = 1.23 X. 0.56, distance between maxillae = 0.12, distance between man- 
dibles = 0.20, distance between antennae = 0.51. 


Triungulin larvae. unknown. 


Holotype. Female, in alcohol; Catalina Mountains, Pima County, Arizona; June 11 1965; col- 
lected by F. Werner, the host is Polistes comanchus navajoe Cresson. Deposited in the United 
States National Museum. 


Allotype. Male, in alcohol, Sycamore Canyon, T 105, R 13, Pima Co., Arizona, June 24, 1961, 
collected for the Arizona Survey, the host is Polistes comanchus navajoe. Deposited in the United 
States National Museum. 


Paratypes. Host of all paratypes is Polistes comanchus navajoe. One female, pinned; Molino 
Canyon in the Catalina Mountains, Pima County, Arizona, October 16, 1994, collected by Blaine 
Mathison, deposited in the University of Arizona Insect Collection (UAIC). One male, in alco- 
hol, Sycamore Canyon, T 105, R 13, Pima Co., Arizona, June 24, 1961, collected for the Arizona 
Survey, deposited in UAIC. One male, in alcohol, Catalina Mountains, Pima Co., Arizona, Oct. 
7, 1984, collected by Olson, in the collection of the senior author (JLC). 


Host voucher. A host, Polistes comanchus navajoe, is deposited as a voucher specimen at UAIC. 


Etymology. This species is named in honor of Teiji Kifune, for his contribution 
to the study of Strepsiptera, including the descriptions of several species of 
Xenos. 


Diagnosis. Xenos kifunei most closely resembles the other five North American 
species. It is distinct from all Xenos in the shape of the prescutum (Fig. 2). It 
differs from all other North American species in having no basal hump on the 
shaft of the aedeagus (Fig. 3 G). 

Xenos kifunei most closely resembles Xenos peckii. Females differ in the 
pigmentation pattern of the cephalothorax. Xenos kifunei has a pigmented area 
that extends slightly anteriorly in the center (Fig. 1). The pigmented area is 
strongly emarginate in the middle in X. peckii. The teeth of the mandible are of 
about equal length in X. kifunei, but the inner tooth of the mandible of X. peckii 
is much longer than the outer tooth. The cephalothorax of X. kifunei is more 
narrowed anteriorly, giving it a somewhat triangular appearance compared to 
a round-shaped cephalothorax in X. peckii. In male X. kifunei, the R5 wing 
vein is much shorter than Rq while X. peckii have Rq4 and R5 veins almost 
equal in length. Male mandibles of X. kifunei are completely clear yellow but 
they are black at the base in X. peckii. 

Xenos kifunei differs from Xenos pallidus in several characters. Male X. 


Vol. 108, No. 4, September & October, 1997 249 


0.5 


Fig. 3A-M. Xenos kifunei, NEW SPECIES, male characters. A. Right antennae. B. Right protho- 
racic leg. C. Right mesothoracic leg. D. Right metathoracic leg. E. Right hind wing with veins 
labeled (following designation by Kinzelbach 1972). F. Mandibles, crossing, and maxillae with 
palps. G. Genital capsule showing aedeagus. H. Cephalotheca of male puparium. 


250 ENTOMOLOGICAL NEWS 


kifunei have a total length of 3.14 to 3.53 mm. compared to 2.25 to 2.75 mm in 
X. pallidus. Lengths of Strepsiptera are quite variable but there is a significant 
difference between these two species. Hind wing veins are heavy and pig- 
mented in X. kifunei but not so in X. pallidus. There is a distinct gap in R1 and 
R2 wing veins in X. kifunei but the two appear almost as one vein in X. pallidus. 
Xenos kifunei has a CuP vein about equal to R1 in length, while CuP is much 
shorter (nearly absent) in X. pallidus. The females of these species have minor 
differences, but no easily recognizable characters to separate them. 

Xenos kifunei males are significantly smaller than is recorded for Xenos 
nigrescens (3.14 - 3.53 vs. 4.5 mm) and have clear yellow mandibles com- 
pared to those of nigrescens, which are black at the base. Wings of X. kifunei 
have heavy, dark wing veins compared to pale veins with posterior veins obso- 
lete in X. nigrescens. Females of X. kifunei have a pigmented area that covers 
less than the posterior half of the cephalothorax and mandibles with both teeth 
nearly equal in length. Xenos nigrescens has a pigmented area covering at 
least 2/3 of the cephalothorax and the inner tooth of the mandible is much 
larger than the outer. 

In males of X. kifunei the palps equal the maxillae in width and the man- 
dibles are nearly straight. In X. hunteri, the palps are about half as wide as the 
maxillae and the mandibles are strongly curved. The cephalothorax of female 
X. kifunei is about as wide as long, with less than the posterior half pigmented. 
The female cephalothorax of X. hunteri is longer than wide, with the posterior 
3/4 pigmented. 

Only the female of Xenos rubiginosi is known. Xenos kifunei has a cepha- 
lothorax about as wide as long, spiracles lateral and somewhat ventral, and 
pigmentation covering less than the posterior half. The cephalothorax of X. 
rubiginosi is much longer than wide, has dorsal first abdominal spiracles, and 
pigmentation covering the posterior 2/3 of the cephalothorax. 


DISCUSSION 


Xenos kifunei becomes the 38th valid species in this genus and the sixth 
recognized species in North America. The number of North American species 
in this genus is likely to change, however. The validity of half the recognized 
species, Xenos nigrescens, Xenos hunteri, and Xenos rubiginosi, is in question 
(Bohart 1941). It is also possible that one or more synonymized species may 
be valid. 

The validity of X. nigrescens is uncertain and no type specimens were des- 
ignated by Brues (1903). Brues gave Polistes rubiginosus Lepeletier as the 
host of X. nigrescens. Polistes rubiginosus has since been synonymized with 
Polistes carolina (Linnaeus), which is the host of Xenos peckii. The character 
that best distinguishes between X. nigrescens and X. peckii is the nature of the 
wing veins. The wing veins of X. peckii are very stout and distinctly darkened, 


Vol. 108, No. 4, September & October, 1997 251 


whereas the veins of X. nigrescens are weak and barely visible. The female of 
X. peckii has a black pigmented posterior cephalothorax with a strong, square 
emargination in the center dorsally, while in X. nigrescens this pigmented bound- 
ary forms a straight line. We have examined three females and one male 
strepsipteran from P. carolina hosts, collected in central Texas, and found them 
all to closely match the description given by Brues of X. nigrescens (type lo- 
cality, Austin Texas). The specimens we examined were from different loca- 
tions within the same geographical region as the type locality and we found 
characters to be consistent among the collections. Our specimens were also 
collected several years apart. The question remains whether these Texas speci- 
mens are a distinct species or a local variation of X. peckii. The total number of 
specimens examined combined with those from Brues’ description amounts to 
five males and four females. Thus, too few are known to make meaningful 
decisions with such minor differences. However, there is almost no variation 
in those we have examined. 

Xenos hunteri is represented by a female type (USNM cat. no. 10115) and 
a male description given by Pierce (1909), but has not been reported since. 
This species was described from an undetermined species of Polistes, reported 
by Pierce to be “near Polistes minor Palisot de Beauvois”. The type locality 
was Victoria, Texas. The actual Polistes host species that Pierce made this 
description from is uncertain, but is likely either Polistes dorsalis dorsalis 
(Fabricius) or Polistes exclamens exclamens Viereck. The reason for this as- 
sumption is that Bequaert (1940) stated that wasps frequently called P. minor 
in the United States are usually P. fuscatus hunteri Bequaert (later synony- 
mized with P. dorsalis dorsalis) or P. exclamens. Both of these species are 
reported from the type locality of X. hunteri. The female type of X. hunteri 
does appear to have characters distinctive from other species of Xenos, but 
with only one specimen available variation is unknown. The male reputedly 
has characters that differ somewhat from other species of Xenos, but reference 
specimens are unavailable for comparison. There seems to be enough charac- 
ter differences with other Xenos to maintain X. hunteri as a separate species at 
this time. Of significant difference are the male mouthparts of X. hunteri, whose 
palp is half as thick as the maxillae, a character found in no other North Ameri- 
can Xenos. 

The status of X. rubiginosi is in some doubt. Pierce (1909) listed the host 
of X. rubiginosi as P. rubiginosus which has since been synonymized with P. 
carolina, the host of X. peckii. However, the description of the shape of the X. 
rubiginosi female cephalothorax appears to be outside the range of X. peckii. 
The uncertainty of species validity occurs because X. rubiginosi is known only 
from the type female (USNM cat. no. 10119). No males have been reported. 
With both species described from P. carolina and only a single specimen of X. 
rubiginosi suggests that this might be an aberrant specimen of either X. peckii 


252 ENTOMOLOGICAL NEWS 


or X. nigrescens. More specimens from P. carolina need to be surveyed to 
determine the status of X. rubiginosi. 

Another aspect of Xenos taxonomy that is in need of review is the syn- 
onyms created by Bohart (1941). In some of the synonymized species, Bohart 
was forced to make decisions using small numbers of specimens. Some of 
these synonymies are clearly correct, but for some of the species to be syn- 
onyms, there would have to be vast character variation within X. peckii and X. 
pallidus. There is, without doubt, some range in the characters of these spe- 
cies. With more specimens collected from reported hosts of the synonymized 
species such as Polistes apachus Saussure and Polistes metricus Say, one or 
more of these synonyms may be reinstated as valid species. 


ACKNOWLEDGMENTS 


We thank David Maddison and Carl Olson, both of the University of Arizona, for allowing 
us access to these specimens. We thank Roger Gold for providing facilities and support for our 
study. We thank Teiji Kifune of Fukuoka University, Japan and Robert Wharton of Texas A&M 
University for their reviews and helpful suggestions in preparing this manuscript. We also thank 
two anonymous reviewers for their aid in bringing this manuscript to its final form. 


LITERATURE CITED 


Bequaert, J. C. 1940. An introductory study of Polistes in the United States and Canada with 
descriptions of some new North and South American forms (Hymenoptera; Vespidae). 
J. N. Y. Entomol. Soc., 48(1): 1-30. 

Bohart, R M. 1941. A revision of the Strepsiptera with special reference to the species of North 
America. Univ. Calif. Pub. Entomol., 7(6): 91-160. 

Brues, C. R. 1903. A contribution to our knowledge of the Stylopidae. Zool. Jb. (Anat., Ontog.), 
18(2): 241-270. 

Kinzelbach, R. K. 1971. Morphologische Befunde an Facherfliiglern und ihre phylogenetische 
Bedeutung (Insecta: Strepsiptera). Zoologica, 41: 1-256. 

Pierce, W. D. 1909. A monographic revision of the twisted winged insects comprising the order 
Strepsiptera Kirby. U. S. Nat. Mus., Bull. 66: 1-232. 

Rossius, P. 1793. Observation de M. Rossi sur un noveau genre d’Insecte, voisin des Ichneu- 
mons. Bull. Soc. Philomatique, |: 49. 


SOCIETY MEETING OF MARCH 2, 1997 


Dr. Owain Edwards 
USDA-ARS Beneficial Insects Introduction Research Lab, Newark, Delaware 


IS ADAPTATION AFTER RELEASE NECESSARY FOR SUCCESSFUL 
CLASSICAL BIOLOGICAL CONTROL? 


Dr. Edwards began by pointing out that, historically, biological control researchers have assumed 
that natural enemies must adapt after introduction into novel environments. In fact, lack of adap- 
tation is often presented as the reason for failed establishment. If parasitoid populations from 
diverse environments differ with respect to fitness traits, this would suggest that adaptation would 
likely be important after release into a novel environment. 


(Continued on page 258) 


Vol. 108, No. 4, September & October, 1997 253 


DESCRIPTION OF THE PUPA OF CYRNELLUS 
FRATERNUS (TRICHOPTERA: 
POLYCENTROPODIDAE), WITH NOTES ON 
VARIATION IN PUPAL CASE CONSTRUCTION! 


Zane B. Johnson, James H. Kennedy2 


ABSTRACT: The pupa of Cyrnellus fraternus is described for the first time, from the Elm Fork 
of the Trinity River, Texas. Study of this species in the Elm Fork has shown two distinct pupal 
case morphologies. These cases are described from field-collected and reared material. 


Banks (1905) originally described Cyrnellus fraternus from a single fe- 
male collected at Plummer’s Island, Maryland. Nyctiophylax marginalis Banks 
and Cyrnellus zernyi Mosely are considered junior synonyms of this species 
(Ross 1944; Flint 1964). Complete bibliographic citations for C. fraternus can 
be found in Flint (1971) and Nimmo (1986). 

Based on associated material, Ross (1959) included the larva of Cyrnellus 
in his generic key, but no published description of the larva was available until 
provided by Flint (1964). The pupa and pupal case of C. fraternus have re- 
mained undescribed. 

Cyrnellus fraternus is widespread in the United States, recorded from Ne- 
braska and Texas, east to Florida and New York, and north to Minnesota. It has 
been recorded from Mexico and the Amazon River Basin in South America 
(Nimmo 1986). Wiggins and Mackay (1978) categorized Cyrnellus as collec- 
tor-filterers of warm, lotic waters of eastern deciduous forest biomes. Larvae 
of C. fraternus, although usually occurring in large rivers, also live in smaller 
streams, lakes and reservoirs (Ross 1944; Flint 1964; Wiggins 1996). Sublette 
(1957) reported collections of the larvae (as C. marginalis) from the gravel 
and sand benthos of Lake Texoma in Oklahoma and Texas. Selwyn Roback 
observed the larvae inhabiting both rivers and lakes, living on wood and rock 
and tolerating current regimes from standing to rapidly flowing (Flint 1964). 
Information on the range of stream orders, stream widths, substratum types 
and flow permanency of the habitats of C. fraternus in the Interior Highlands 
of North America was provided by Moulton and Stewart (1996). 

In this paper we describe the pupa of C. fraternus and the variation in 
structure of the pupal case. Morphological terminology for the pupal descrip- 
tion follows that of Resh (1976). Specimens used for descriptions are from 
field-preserved samples and material reared in 37.85 | aquaria. Material exam- 


! Received January 30, 1997. Accepted March 21, 1997. 
2 Department of Biological Sciences, University of North Texas, Denton, Texas 76203. 


ENT. NEWS 108(4): 253-258, September & October, 1997 


254 ENTOMOLOGICAL NEWS 


ined is deposited in the collection of the University of North Texas, and 
preserved in 70% ethanol. 


Cyrnellus fraternus (Banks) 


Pupal Description. Length 5-6 mm. Labrum with 2 groups of long anterolateral setae, 
each with 3 setae (Fig. 1); 2 groups of posterolateral setae, each with 2 long, | short setae. 
Mandibles thin, sickle-shaped, bases stout with 2 short lateral setae (Fig. 2). Paired 
presegmental plates on III each with 5-6 denticles; IV each with 5-7 denticles; V each with 
7 denticles; VI each with 12-13 denticles; VII each with 5-8 denticles; VIII-each with 4-6 
denticles. Paired postsegmental plates on V each with 10 denticles (Fig. 3). Lateral gills on 
abdominal segments II-VII; single on II, bifid on III-V, single on VI-VII. Anal rods short 
and rounded, each with 10-11 setae (Fig. 4). 

Pupal Case. Length 7.5-9 mm. Cylindrical to ovoid in shape, constructed of sand grains 
and various-sized detrital particles attached to a silk interior (Figs. 5-6). 

Material Examined. U.S.A., Texas, Denton Co., Elm Fork of Trinity River 1 km up- 
stream of bridges at FM 428, ca. 4 mi E Aubrey, 13-vii- 1995, Z. B. Johnson and R. E. Cook, 
1 pharate male, 2 pupae; same but 10-ix-1995, Z. B. Johnson and J. D. Csekitz, 1 pupa; same 
but 08-x-1995, Z. B. Johnson and B. T. Hall, | pupal case; same but 21-xii-1995, Z. B. 
Johnson and J. D. Csekitz, 2 cases from reared specimens; same but 26-i-1996, Z. B. Johnson 
and R. E. Cook, | pharate male, reared; same but 31-i-1996, 1 pharate female, reared. 


DISCUSSION 


Moulton and Stewart (1996) reported C. fraternus immatures inhabiting a 
variety of mineral substrata, ranging from sand/mud to cobbles. In the Elm 
Fork, C. fraternus larvae and pupae were collected from submerged wood, or 
submerged snags, in an 8 km study area bordered by bottomland hardwood 
forest and pasture land. Larvae of C. fraternus were commonly found on snags 
with water flow ranging from standing to 0.2 m/s. Larvae were found in their 
greatest densities on wood covered with dense silt and/or filamentous algae. 
Roback (in Flint 1964) observed C. fraternus larvae on rocks, living under “an 
amorphous silk mat usually silt covered.” Our observations from the Elm Fork 
support Roback’s observations. 

Of 7 pupal cases examined from the Elm Fork, 2 basic types with some 
variation (Figs. 5-7) were constructed by C. fraternus. The most common case 
structure (Fig. 5) consisted of sand and small detrital particles attached to a 
silken interior. Other materials included fine particulate wood debris and scler- 
ites of other aquatic insect larvae. A variant case that included rock fragments 
as well as sand, 1 Hydropsychidae lateral head capsule sclerite, 4 pronotal 
sclerites and additional sclerites from appendages was also observed (Fig. 6). 
A second case type (Fig. 7) is known from a single field-collected pupa and 
from cases produced by terminal instar larvae on Elm Fork submerged wood 


Vol. 108, No. 4, September & October, 1997 295 


Figs. 1-3. Cyrnellus fraternus pupa. 1. Labrum, dorsal view. Scale bar= 300 um. 2. Right man- 
dible, dorsal view. Scale bar = 300 uum. 3. Abdomen, dorsal view with inset showing segmental 
plates with denticles. Scale bar = | mm. 


256 ENTOMOLOGICAL NEWS 


in laboratory rearing tanks. This case type had some sand and many various- 
sized wood particles attached to a silken interior. Wood particles tended to be 
irregularly placed, except for some elongate twigs located laterally. Cases 
collected from the sides of rearing aquaria are considered to have been produced 
under artificial conditions, but the field-collected specimen confirms the 
diversity of cases constructed by C. fraternus under natural conditions. None 
of the cases studied resemble the silken retreats (illustrated in Wiggins 1996) 
constructed by the larvae. It is not known if the larval retreat is used in the 
construction of the pupal case. 

Field collections of Cyrnellus fraternus pupae were relatively rare in the 
EIm Fork, even in samples containing numerous Sth instar larvae. One pharate 
male and 3 pupae were found in preserved submerged wood samples. Surveys 
of Elm Fork habitats have shown that C. fraternus does not inhabit the sand- 
based benthos (Z. B. Johnson, unpublished data). However, we speculate that 
the terminal instar larvae leave the snags to pupate. Sand is a main material in 
pupal case construction, but is not found consistently on submerged wood. 
Observed variation in case morphology is attributed to availability of building 
materials. 


Fig. 4. Cyrnellus fraternus pupa anal rods, dorsal view. Scale bar = 300 um. 


Vol. 108, No. 4, September & October, 1997 257 


Figs. 5-7. Cyrnellus fraternus pupal cases. 5. Typical case comprised of sand grains and detritus 
attached to a silken tube. 6. Case that includes larval sclerites. 7. Variant case predominantly 
comprised of irregularly-arranged wood particles. Scale bar = 2 mm. 


258 ENTOMOLOGICAL NEWS 


ACKNOWLEDGMENTS 


We thank Luciana Nurman for assistance in producing the illustrations used in this paper. 
We thank Stephen R. Moulton, II and Boris C. Kondratieff for reviewing this manuscript. Sug- 
gestions from John C. Abbott and David Houghton improved an early draft of the manuscript. 
The comments of 2 anonymous reviewers were appreciated. 


LITERATURE CITED 


Banks, N. 1905. Descriptions of new Nearctic neuropteroid insects. Trans. Am. Entomol. Soc.32: 
1-20. 

Flint, O. S., Jr. 1964. Notes on some Nearctic Psychomyiidae with special reference to their 
larvae (Trichoptera). Proc. U. S. Natl. Mus. 115(3491):467-481. 

Flint, O. S., Jr. 1971. Studies of Neotropical caddisflies, XII: Rhyacophilidae, Glossosomatidae, 
Philopotamidae, and Psychomyiidae from the Amazon Basin (Trichoptera). Amazoniana 3: 
1-67. 

Moulton, S. R., II, and K. W. Stewart. 1996. Caddisflies (Trichoptera) of the Interior High- 
lands of North America. Mem. Am. Entomol. Inst. 56: 1-313. 

Nimmo, A. P. 1986. The adult Polycentropodidae of Canada and adjacent United States. Quaest. 
Entomol. 22:143-252. 

Resh, V. H. 1976. The biology and immature stages of the caddisfly genus Ceraclea in eastern 
North America (Trichoptera: Leptoceridae). Ann. Entomol. Soc. Am. 69: 1039-1061. 

Ross, H. H. 1944. The caddisflies, or Trichoptera, of Illinois. Illinois Nat. Hist. Surv. Bull. 23: 
1-326. 

Ross, H. H. 1959. Trichoptera. Pp. 1024-1049. In: W. T. Edmondson (ed.). Freshwater biology, 
2nd ed. John Wiley & Sons, Inc., New York, NY. 1248 pp. 

Sublette, J. E. 1957. The ecology of the macroscopic bottom fauna of Lake Texoma (Denison 
Reservoir), Oklahoma and Texas. Am. Midl. Nat. 57:371-402. 

Wiggins, G. B. 1996. Larvae of the North American caddisfly genera, 2nd ed. University of 
Toronto Press, Toronto. 457 pp. 

Wiggins, G. B., and R. J. Mackay. 1978. Some relationships between systematics and trophic 
ecology in Nearctic aquatic insects, with special reference to Trichoptera. Ecology 59:1211- 
1220. 


(Continued from page 252) 


He then described two studies which tested these notions. In the first, populations of the 
Russian wheat aphid parasitoid Aphelinus asychis (Hymenoptera: Aphelinidae) were collected 
from Antibes and Montpellier in France, Spain, Morocco, Kazakstan, China and Greece. Crosses 
of these samples indicated that though there were significant levels of variation in fitness within 
populations, there was not significant variation between populations. This suggests that any of 
these populations could be introduced to another location without need for adaptation. 

The second study concerned the European corn borer parasitoid Macrocentrus grandii (Hy- 
menoptera: Braconidae), which was introduced into North America from both France and Korea. 
Current evidence suggests that the Korean introduction established. An analysis of the environ- 
ments of Korea and Delaware, which was in the center of the original release area, suggests that 
the greatest need for adaptation would have been due to climatic differences. Adult M. grandii in 
Delaware would experience extreme temperatures (32-38°C) significantly more often than those 
in Korea, suggesting that adaptation to high temperatures may have been necessary after release. 
Laboratory experiments have shown that adult M. grandii from Delaware have significantly greater 
survivorship at 36°C and 38°C than do those from Korea. This suggests that adaptation after 


release has occurred. : 
(Continued on page 304) 


Vol. 108, No. 4, September & October, 1997 259 


DESCRIPTION OF MATURE LARVAE OF 
MICRODYNERUS EXILIS AND M. TIMIDUS 
(HYMENOPTERA: VESPIDAE)! 2 


J. Tormos, J.D. Asis, S.F. Gayubo, F. Torres? — 


ABSTRACT: The mature larvae (prepupae) of Microdynerus exilis and M. timidus are described. 
The number and arrangement of the sensilla and setae on the labrum and the sensilla on the 
epipharynx are the characters which best provide differentiation of these species from the other 
previously described from this genus: M. nugdunensis. 


Current knowledge of the preimaginal states of Eumeninae is very frag- 
mentary. Although the subfamily includes more than 2,500 species (Yamane 
1990), the mature larvae of only 39 have been described (important references: 
Enslin 1921; Janvier 1930; Micheli 1930, 1934; Maneval 1939; Reid 1942; 
Grandi 1961; Evans 1977; and Kojima 1991). In this paper we describe the 
prepupae of two species of Microdynerus Thomson, obtained in 1992 and 1993, 
during a study of the fauna of rubicolous species in the northern subplateau of 
the Iberian Peninsula: Microdynerus exilis (Herrich-Schaeffer) and M. timidus 
(Saussure). The mature larva of only one species has been described from this 
genus: M. nugdunensis (Saussure) (Giordani Soika 1934; Grandi 1961). 

The method employed to prepare the specimens was similar to that used by 
Asis et al. (1994). The terminology of larval morphology follows Evans (1987). 

In the measurement of certain structures we use the following abbrevia- 
tions: d= diameter, h = height, 1 = length, w = width. 


RESULTS 


The nest of Microdynerus exilis (1 = 68 mm; d = 2 mm) was located in a 
cane stem at Salduero'(Soria). It had a vestibular cell, initial and final plugs, 
and 4 brood cells (x = 10 mm; range = 9.5-10.5 mm) separated by mud septa, 
all with a cocoon. The prepupa of cell 1 was preserved for study (voucher 
number 93070101). Two females emerged from cells 2 and 4 in March 1994. 
The pupa in cell 3 was dead. 

A nest of Microdynerus timidus (1 = 78 mm; d = 2.5 mm) was obtained 


! Received December 2, 1996. Accepted February 23, 1997. 
2A grant from the DGICYT (PB91-0351-C02-02) supported the study. 
3 Unidad de Zoologia, Facultad de Biologia, Universidad de Salamanca, 3707 1-Salamanca, Spain. 


ENT. NEWS 108(4): 259-264, September & October, 1997 


260 ENTOMOLOGICAL NEWS 


from a cane stem at Boniches (Cuenca). The nest had a vestibular cell, initial 
and final plugs (1 = 4 mm), and 6 brood cells ( x = 12.5; range = 8-18.5 mm) 
separated by mud plugs. All cells had a cocoon. The cocoon did not com- 
pletely enclose the prepupa, and excrement was present in its interior basal 
zone. The prepupa of cell 6 was preserved for study (voucher number 
92030101). Four females emerged from cells 2 to 5 and one male from cell 1, 
in March, 1993. 


Descriptions of prepupae 


Microdynerus exilis (Herrich-Schaeffer) 


General aspect (Fig. 1). Body fusiform (1 = 7.0 mm, w = 2.0 mm), with dorsum of the thoracic 
and first 8 abdominal segments divided into two annulets by a transverse crease. Anus a trans- 
verse slit. Pleural lobes weakly developed. Integument with scanty and scattered minute setae (1 
= 10 um). 

Spiracles (Fig. 2) with walls of atrium smooth; opening into subatrium unarmed; subatrium (d = 
40 tum) narrower than atrium (d = 65 pm). 

Cranium (Fig. 3) (w = 0.96 mm, h = 0.89 mm), with scattered setae (1 = 5 um) and setigerous 
punctures. Coronal suture ill-defined; parietal bands absent. Antennal orbits circular (d = 90 
tim), with 3 small sensilla. Clypeus with scanty punctures (d = 10 um). Labrum (Fig. 3a) (w = 
450 um) bilobed, with 21 short conical sensilla (w = 5 um) on each side. Epipharynx (Fig. 3b) 
with few small spinules medioventrally (w = 5 um) and 3 sensilla on each side. 

Mouthparts. Mandibles (1 = 320 um, w = 240 pm) weakly tridentate, brown pigmented, upper 
tooth truncate apically. Maxillae with few setae (1 = 15 um) on external surface; maxillary palpi 
(1 = 65 um; w = 50 pm) with 4 apical sensilla; galeae (1 = 80 um; w = 30 um), sharply narrowing 
at apex. Labium (1 = 190 um, w = 230 um), with short palpi (1 = 30 um, w = 35 pm), blunt 
apically and with 4 apical sensilla; prementum with setae (1 = 10 xm) ventral to the palpi; spin- 
neret a transverse slit (1 = 140 um) with strongly raised lips. 


Microdynerus timidus (Saussure) 


General aspect (Fig. 4). Body fusiform (1 = 7.0 mm, w = 1.9 mm), similar to M. exilis. 
Spiracles (Fig. 5) with walls of atrium smooth; opening into subatrium unarmed, subatrium 
(d = 30 pm) narrower than atrium (d = 40 um). 

Cranium (Fig. 6) (w = 0.9 mm, h = 0.75 mm), with scattered setae (1 = 10 sm) and punctures. 
Coronal suture not very distinct; parietal bands absent. Antennal orbits circular (d = 50 pm), with 
3 small sensilla. Clypeus with a few setae (1 = 10 pm). Labrum (Fig. 6a) (w = 450 pm) bilobed, 
with 10 short conical sensilla (w = 5 pm) and 5 setae (1 = 10 pm) on each side. Epipharynx (Fig. 
6b) with small spinules medioventrally, and 4 sensilla (w = 5 um) on each side. 

Mouthparts. Mandibles (1 = 270 um, w = 200 um) and maxillae as described for M. exilis, 
maxillary palpi (1 = 50 wm; w = 35 um) with 4 apical sensilla; galeae (1 = 105 ym; w = 25 ym) 
long, pointed at apex. Labium (1 = 180 tm, w = 260 ym) as in the species previously described. 


Vol. 108, No. 4, September & October, 1997 261 


= 


0.1 mm 


FIGURES 1-3.- Mature larva (prepupa) of Microdynerus exilis (Herrich-Schaeffer): 1, Profile; 
2, Anterior thoracic spiracle (atrium, subatrium and tracheal trunk); 3, Cranium in frontal view 
(Labrum [a], Epipharynx[b]). 


262 ENTOMOLOGICAL NEWS 


FIGURES 4-6.- Mature larva (prepupa) of Microdynerus timidus Saussure: 4, General aspect; 5, 
Anterior thoracic spiracle (atrium, subatrium and tracheal trunk); 6, Cranium in frontal view 
(Labrum (a), Epipharynx (b)). 


Vol. 108, No. 4, September & October, 1997 263 


DISCUSSION 


Reid (1942) and Grandi (1961) established several characters to differenti- 
ate eurnenine larvae from those of the rest of the Vespidae: labrum almost as 
wide as the clypeus; distance from the antennae to the bases of the mandibles 
less than that from the center of the anterior edge of the labrum to the center of 
a line joining the bases of mandibles; and deeply bilobed labrum. All of these 
characters are well defined in the two species studied in this paper. Moreover, 
the genus Microdynerus could be characterized among eumenine wasps by the 
presence of very long galeae, narrowing towards the apex. 

Although a certain morphological uniformity is noted when comparing the 
larva of the species described previously (M. nugdunensis) with those described 
in this article, differences do exist (Table I). The most important ones are mor- 
phology, number and arrangement of the sensilla on the labrum; the number of 
sensilla on the epipharynx; and the presence/absence of setae between labial 
palpi. Additional differences are found in: the number of the setae on the ex- 
ternal margin of the maxillae; in the morphology of the mandibles; and in the 
presence/absence of a circle of spines in the opening to the subatrium. 

Each of the three species are differentiated as follows: 

- M. exilis: by having the labrum without setae, only with short conical sensilla 
(Fig. 3a) [setae are present in M. nugdunensis and M. timidus (Fig. 6a) (Table 
I]. 

- M. nugdunensis: a) maxillae with numerous setae on external side [few setae 
are present in M. exilis (Fig. 3) and M. timidus (Fig. 6) (Table I)]; b) opening 
into subatrium with spines [in M. exilis (Fig. 2) and M. timidus (Fig. 5) the 
spiracular atrium has no collar-like processes (Table I)]; and c) setae between 


Table I.- Characters used to differentiate mature larvae of Microdynerus (present: x; absent: -): 
(1) Labrum with setae; (2) Epipharynx with two sensorial areas, each one with 3 sensilla; 
(3) Maxillae with few setae on external surface; (4) Galeae long, sharply narrowing at apex; 
(5) Labium with setae between the labial palpi; (6) Spiracular atrium with collar-like processes. 


Species 1 2 3 4 5 6 
M. exilis ~ Xx X x ms "S 
M. nugdunensis x X is x X x 


M. timidus X - x Xx = = 


264 ENTOMOLOGICAL NEWS 


the labial palpi [no setae in M. exilis (Fig. 3) and M. timidus (Fig. 6) (Table I]. 
- M. timidus: by having 4 sensilla in each of the sensorial areas of the epipharynx 
(Fig. 6a) [M. exilis (Fig. 3) and M. nugdunensis (Fig. 6) have two sensorial 
areas, each one with 3 sensilla (Table I)]. 


ACKNOWLEDGMENTS 


We are indebted to H.E. Evans (Colorado State University, U.S.A.), J.M. Carpenter (Ameri- 
can Museum of Natural History, U.S.A.), J. Kojima (Ibaraki University, Japan) and Sk. Yamane 
(Kagoshima University, Japan) for their comments on the manuscript. 


LITERATURE CITED 


Asis, J.D., Tormos, J. & S.F. Gayubo. 1994. Biological observations on Trypoxylon attenuatum 
and description of its mature larva and its natural enemy Trichrysis cyanea (Hymenoptera: 
Sphecidae: Chrysididae). J. Kans. Entomol. Soc., 67 (2): 199-207. 

Enslin, E. 1921. Beitrage zur Kenntnis der Hymenopteren II. 3. Biologie von Symmorphus sinuatus 
F. 4. Biologie von Ancistrocerus trifasciatus F. Deutsch. Ent. Zeitschr., 279-285. 

Evans, H.E. 1977. Notes on the nesting behavior and immature stages of two species of 
Pterocheilus (Hymenoptera: Eumenidae). J. Kans. Entomol. Soc., 50 (3): 329-334. 

Evans, H.E. 1987. Order Hymenoptera. In: F.W. Stehr (ed.) Jmmature Insects. pp. 597-710. 
Kendall/Hunt Publishing Company. Dubuque. Iowa. 

Giordani-Soika, A. 1934. Etudes sur les larves des Hyménoptéres. Ann. Soc. Ent. France, 103: 
337-344. 

Grandi, G. 1961. Studi di un entomologo sugli imenotteri superior. Bolletino dell ‘Istituto di 
Entomologia dell ‘Universita di Bologna, XXV: 1-659. 

Janvier, H. 1930. Recherches biologiques sur les predateurs du Chili. Ann. Sci. net. Zool., 10: 
235-254. 

Kojima, J. 1991. Descriptions of mature larvae of some solitary wasps (Insecta: Hymenoptera: 
Vespidae, Sphecidae). Publ. Itako Hydrobiol. Stn., 5: 5-12. 

Manevyal, H. 1939. Notes sur les hyménoptéres (6éme sér.). Ann. Soc. Entomol. France, 108: 49- 
108. 

Micheli, L. 1930. Note biologiche e morfologiche sugli imenotteri (II). Mem. Soc. Ent. Ital., 9: 
46-66. 

Micheli, L. 1934. Note biologiche e morfologiche sugli imenotteri. Boll. Soc. Ent. Ital., 66: 246- 
252: 

Reid, J,A. 1942. On the classification of the larvae of the Vespidae. Trans. Roy. Ent. Soc. Lond., 
92: 285-331. 

Yamane, Sk. 1990. A revision of the Japanese Eumenidae (Hymenoptera, Vespoidea). Ins. 
Matsum., 43: 1-189. 


Vol. 108, No. 4, September & October, 1997 265 


LINNAEAN SPECIES OF CONOPS (DIPTERA: 
CONOPIDAE, MUSCIDAE, SCIOMYZIDAE, 
SYRPHIDAE, & TACHINIDAE), WITH 
DESIGNATIONS OF LECTOTYPES! 


F. Christian Thompson 


ABSTRACT: Linnaeus described 13 species in the genus Conops, which are now placed in the 
families Conopidae, Muscidae, Sciomyzidae, Syrphidae and Tachinidae. A study is presented of 
the types and other material of these species in his collection. Lectotypes are designated for 10 
names (C. vesicularis Linnaeus 1761, C. macrocephala Linnaeus 1758, C. aculeata Linnaeus 
1761, C. flavipes Linnaeus 1758, C. ferruginea Linnaeus 1761, C. petiolata Linnaeus 1761, C. 
atomaria Linnaeus 1767, C. testacea Linnaeus 1767, C. buccata Linnaeus 1758 & C. 
subcoleoptrata Linnaeus 1758). Three new synonyms (C. macrocephala = Physocephala niara 
De Geer, C. petiolata = Physocephala rufipes Fabricius, C. testacea = Myopa extricata Collin) 
and one new combination (C. atomaria = Euthycera atomaria) are proposed. 


Names are the keys to knowledge as they serve as short tags for complex 
objects. In biology, scientific names are tags for species as well as groups of 
species. These names mean that all organisms that have the same name share at 
least some characteristics in common. The scientific naming system in biology 
began with Linnaeus who perfected his system through a series of books that 
attempted to classify all living things into one natural system. He entitled these 
works Systema Naturae, the system of nature. Subsequent biologists have fol- 
lowed the system established by Linnaeus. So, as our current system of scien- 
tific names is derived from Linnaeus, the need exists to re-examine the initial 
state of the system. This paper does that as that system relates to a small group 
of flies. 

Linnaeus recognized 10 groups (genera) of flies. One of these groups was 
Conops, based on flies that had elongate mouthparts. In his final edition (12th, 
1767) of his system of nature, Linnaeus included 13 species in the genus Conops. 
Common farm pests, such as the stable and horn flies, as well as parasites of 
bees and wasps, scavengers in cow dung, etc., were included in Conops. As 
there is a need to fix the identity of Conops testacea Linnaeus (see Camras 
1994), all the species that Linnaeus included in Conops are reviewed. Fortu- 
nately, the actual specimens on which Linnaeus based his species are preserved 
in London (For details on the Linnaean Collection, its curation and history, see 
Day & Fitton 1978; on Linnaean insect pins, see Mikkola 1983). 

Two Linnaean Conops species are currently considered nomina dubia 
(Chvala & Smith 1988). Their identities are here resolved. Linnaeus defined 
another two Conops species broadly, the definitions of these are here restricted 


! Received August 22, 1996. Accepted March 22, 1997. 


2 Systematic Entomology Laboratory, USDA, NHB-168 Smithsonian Institution, Washington, 
D.C. 20560. 


ENT. NEWS 108(4): 265-272, September & October, 1997 


266 ENTOMOLOGICAL NEWS 


by lectotype designation. I have designated lectotypes for what are today unique 
specimens following the recommendation (73F) of the /nternational Code of 
Zoological Nomenclature. Each species included in Conops is listed, in the 
order that it appears in the 12th edition, along with its current status (family, 
valid name) and an indication of the voucher material available in the Linnaean 
Collection. For Conopidae, I have followed the species concepts used by Smith 
(1969) and Chva4la (1961, 1963 & 1965). 


1) rostrata Linnaeus 1758: 604. Syrphidae, Rhingia rostrata (Linnaeus). A Q 
and 2 OO’. Both O'C are Smith specimens. See Thompson, et alii 1982: 159. 


2) calcitrans Linnaeus 1758: 604. Muscidae, Stomoxys calcitrans (Linnaeus). 
Lectotype C’, paralectotypes Q . See Pont (1981: 168) for designation and full 
information. 


3) irritans Linnaeus 1758: 604. Muscidae, Haematobia irritans (Linnaeus). 
Lectotype 9, paralectotype 9 . See Pont (1981: 169) for designation and full 
information. 


4) vesicularis Linnaeus 1761: 468. Conopidae, Conops vesicularis Linnaeus. 
A single CO’, here designated as lectotype. This specimen corresponds to the 
current concept of the name. 


5) macrocephala Linnaeus 1758: 604. Conopidae, Physocephala nigra (De 
Geer). A single Q , which is here designated lectotype. In the most recent catalog 
(Chvala & Smith 1988: 252), this name is stated to be ‘’probably a senior 
synonym of Conops vesicularis Linnaeus.” Much paper was wasted on whether 
Moses Harris’ misidentification (Harris 1776) of this species was an independent 
proposal, and, hence, a valid name for the species now known as Rhingia 
campestris Meigen 1822 (Collin 1946, 1947, 1948; Goffe 1946, 1947, 1948, 
1949). Given the confusion over this epithet, macrocephala is best left as a 
forgotten name and current usage of nigra be continued. 


6) aculeata Linnaeus 1761: 468. Conopidae, Dalmannia aculeata (Linnaeus). 
A single CO here designated as lectotype. This specimen pyran to the 
current concept of the name. 


7) flavipes Linnaeus 1758: 604. Conopidae, Conops flavipes Linnaeus. A single 
CO with no head is here designated as lectotype. This specimen corresponds to 
the current concept of the name. 


8) ferruginea Linnaeus 1761: 468. Conopidae, Sicus ferrugineus (Linnaeus). 
A single Q here designated as lectotype. This specimen corresponds to the 
current concept of the name. 


9) petiolata Linnaeus 1767: 1005. Conopidae, Physocephala rufipes (Fabricius). 
Three C'C’, one of these is clearly a specimen subsequently added by Smith as 


Vol. 108, No. 4, September & October, 1997 267 


it is labelled as from “Angl.”, another probably also a Smith specimen as it is 
on a different and non-Linnaean pin. The third C’ here designated as lectotype. 

In 1761, Poda (1761: 118) described Empis petiolata, a species that is clearly 
a conopid. Unfortunately, whether this name applies to a Conops or Physo- 
cephala cannot be determined from the original description and, unfortunately, 
the types are lost (Thompson & Pont 1994: 37). For pragmatic reasons, I con- 
sider Poda’s species to belong to the current concept of Physocephala. Hence, 
the Poda name becomes the senior homonym of the Linnaean name. Thus, 
Conops rufipes Fabricius 1781 may remain the valid name for the species first 
described by Linnaeus as petiolata. 

Chvala & Smith (1988: 256) list a Conops petiolata Donovan (1808: pl. 451) 
as an available name and a synonym of rufipes Fabricius. However, Donovan 
clearly and correctly identified the Linnaean species giving the appropriate 
citation to the name in the then most recent edition of the Systema Naturae 
(13th; Gmelin 1790). If subsequent workers had been as careful as Donovan 
then today I would not have been forced to twist my interpretation of the 
literature to suppress the Linnaean name and preserve current usage! 


10) atomaria Linnaeus 1767: 1005. Sciomyzidae, Euthycera atomaria, probably 
a senior synonym for Euthycera chaerophylli (Fabricius). A single 9 remains, 
but it is in poor condition. The specimen was identified as Euthycera by Cogan 
when the collection was recurated in 1974. As a modern revision of Euthycera 
is needed (Rozko$ny 1984), the nomenclatural implication of the identity of 
this Linnaean name is left for future workers. 


11) testacea Linnaeus 1767: 1006. Conopidae, Myopa testacea (Linnaeus). A 
single Q here designated as lectotype. The specimen agrees with the current 
concept of Myopa testacea of Chvala (1965), but seems to run in Collin (1959) 
either to extricata Collin or testacea Linnaeus as the specimen has a mixture 
of the characteristics used by Collin to distinguish those species. The specimen 
is clearly reddish anterior to the scutellum and has sparse, inconspicuous black 
facial pile. The palps are yellow. Unfortunately, David Clements (personal 
communication), who is revising the genus Myopa, has confirmed the specimen 
to be a pale representative of extricata Collin. 

The real question of what testacea Linnaeus is goes beyond what the syntype 
in the Linnaean Collection is. Linnaeus also included in his concept of testacea, 
Sicus ferrugineus Scopoli and the then unnamed species (potential type of) 
Stomoxoides Schaeffer. When the species Sicus ferrugineus Scopoli was 
included in Conops, the epithet was identical to that of Linnaeus’ ferrugineus. 
Hence, Linnaeus was forced to rename the species. Thus, specimens of either 
of the two different species could be designated as the type of the name Conops 
testacea Linnaeus and then the species represented by that type would become 
the first included species in the genus Stomoxoides (and thus becoming the 
type species of Stomoxoides by subsequent monotypy as was recognized by 


268 ENTOMOLOGICAL NEWS 


Coquillett (1910: 609). To maintain current usage, I select the specimen in the 
Linnaean Collection to be lectotype. This action preserves the current 
interpretation of Sicus ferrugineus Scopoli. The genus group name, 
Stomoxoides, which some authors believe is an available (and valid) name, 
would become the senior synonym of Myopa Fabricius (1775) except as noted 
below. 

There is no basis for the assumption that Stomoxoides Schaeffer (Schaeffer 
1766b: pl. 120) is an available name. Schaeffer rejected the binominal system 
of Linnaeus. His work Elementa Entomologia is best considered binary3.as 
that was the nomenclatural system he followed in his other works (for 
example, his cones [Schaeffer 1766a-1779]). Elementa includes no references 
to species, only orders and genera. The use of Stomoxoides in the Icones is 
clearly binary. These species taxa of the Jcones only received available 
binominal names from Panzer (1804), but Panzer used Myopa, not Stomoxoides. 
One could make a tedious and pedantic argument that since the Elementa does 
not deal with the species category, there is no evidence within the Elementa 
itself as to whether Schaeffer’s nomenclature would be consistent with the 
Principle of Binominal Nomenclature or not, hence new genus-group names 
are available from it under Article Ilc(i). But why do so? The historical record 
is clear, Schaeffer used a binary system of nomenclature and regardless of 
whether Stomoxoides is available or not, the name will remain a synonym, 
either an objective junior synonym of Sicus or suppressed subjective senior 
synonym of Myopa (see below). 

Collin (1959) was undoubtedly correct in identifying the species figured 
by Schaeffer as Sicus ferrugineus Scopoli. 


12) buccata Linnaeus 1758: 605. Conopidae, Myopa buccata (Linnaeus). Three 
specimens (1 C2 Q ) are associated with this name, one is apparently a Smith 
addition, the other two are undoubtedly Linnaean specimens. The one male, 
associated with the Linnaean name label, is Myopa fasciata Meigen. The 
females are buccata of current authors. The female without antennae (one of 
the Linnaean specimens) is here designated as lectotype and has been so 
labelled. 


13) subcoleoptrata Linnaeus 1767: 1006. Tachinidae, Phasia subcoleoptrata 
(Linnaeus). A single CO is present and is clearly a syntype because it is on a 
Linnaean pin. The specimen belongs to Phasia and appears to agree with the 
current concept. 


There are 2 additional specimens in box 23 that are labelled as from old boxes 
196 and 197. 


The Linnaean species of Conops were re-evaluated as there is a question of the 
status of the genus-group name Myopa (Camras 1994). Unfortunately, this 


3 Binary nomenclature is the system of using a uninominal name for the genus and a polynominal 
name for the species. 


Vol. 108, No. 4, September & October, 1997 269 


proposal to the International Commission on Zoological Nomenclature was 
riddled with errors, many of which have already been noted (Sabrosky 1994, 
Wheeler 1994). Unfortunately, no one in their rush to resolve an old problem 
bothered to carefully review what earlier workers had done. Linnaeus always 
attempted to synthesize all previous work in his Systema Naturae. His treatment 
(fig. 2) of Sicus, Sicus ferrugineus Scopoli and Stomoxoides Schaeffer was 
reasonable within the context of his times. Had subsequent workers followed 
Linnaeus, this current work as well as the application (Camras 1994) to the 
Commission would have been unnecessary. 


Beyond the errors noted by Sabrosky and by Wheeler, the following should 
also be corrected. 

Camras and others want to interpret history to be convenient: Sicus 
ferrugineus Scopoli is clearly an independent and new proposal, which has 
nothing to do with the previously described species, Conops ferrugineus 
Linnaeus. Like all systematists, Scopoli gave citations to earlier use when he 
cited available names (see for example, under Sicus buccatus; fig. 1). 

Camras (1994) stated that Fabricius included Sicus in the synonymy of 
Myopa when he established Myopa, which is a rather generous interpretation 
of the facts. Fabricius gave no synonymy for genus-group names, but he did 
equate, in his species synonymy, Conops ferruginea Linnaeus with Sicus 
ferrugineus Scopoli. That is, in retrospect, a correct species synonymy, and, 
thereby, the genus-group names [given the subsequent type-species designa- 
tion of Sicus] are synonyms. However, as noted above, Linnaeus had previ- 
ously considered his ferruginea distinct from Scopoli’s. 

The statement (Camras 1994) that the genus-group name Stomoxoides 
Schaeffer “was subsequently only” used by Schaeffer is clearly wrong as 
Linnaeus and Coquillett treated the name as indicated above. 

Unless the plenary powers are invoked, the invalid designation of Coquillett 
(1910: 605) of a non-originally included species (Conops ferruginea Linnaeus 
1761) as the type species of Sicus Scopoli 1763 remains invalid. As there has 
never been a valid type designation for Sicus, I hereby designate the second 
originally included species, Sicus ferrugineus Scopoli 1763, as the type spe- 
cies. This type species is currently recognized by the name Sicus ferrugineus 
(Linnaeus 1761). 

If one considers Stomoxoides Schaeffer to be an available name (I do not 
as it appears only in works that are not binominal), then the question remains 
as to what is its type species. Under the current rules of nomenclature, because 
the genus name was published without any included species in the sense of 
named species, the first subsequently included species become the original 
included species (ICZN, Art. 69a(i)1). By including Stomoxoides in the spe- 
cies synonymy of Conops testacea, Linnaeus was the first worker to define 
Stomoxoides by subsequent monotypy. As the Linnaean species testacea was 


ENTOMOLOGICAL NEWS 


eee 


Os armatum roitro unifeto: vagina rigida, 
porrecta, longa, e medio refracta & inflexa, 
bali palpigera. 


1094. Sicus Ferrugineus. 
— long. lin. 34. 

Diagn. Antenoz feta brevi laterali. Abddo- 
men teres, incurvum, ferrugineum. 

Habicat io pratis, & etiam in fylvis. 

Antenne ferruginez. Frons aurea. Thorax 
fufco-rufus. Ale immaculate. Abdomen (pirdlicer 
incurvum. Roftri vagina apice binda. 


1005. Sicus Buccatus. 
— long. lin. 3. 
Linn. Sy{t. Nat. p. 605. Conops n. 6. 
F.un. Svec. 2. 1905. 

Diagn. Facies velicularis, aiba. Abdomen in- 
curvum pedesque rufL 

In pratis. 

Antenne rife, fetrriz. Oculi fufci. Alz 
pund&o medio fuico. Abdcom21 apice muaculis la- 
terajibus Cigeraicentidus ut:injue (3-4). Pedes 
rufi. Geuca pallidiora, 

Cs TABA 


1006 INSECTA DIPTERA. Afilus. 


teftacea. 11. C. antennis fetariis teftacea, abdomioe fubovato ha- 

mofo, facie veficulari alba, alis hyalinis. 

Scop. carn. 1004. Sicus ferrugineus. 

Scheff. elem. s. 120. Stomoroides. 

Habitat in Europa auftrali. Afcanias. 

Corpus ferrugineum. Abdomen wom cylindricum, in- 
flexum. Ala byalina vemis farrugincis. StasuraC. 
buccate. 


buceata. 1a. C. antennis fetariis, abdomine hamofo grifco, facie 
veficulari alba, alis nebulofis. Fw. /rec. 1905. ® 
Scop. carm. 1cos. Sicus buccatus. 
Habitat im Europa. 


fubcoleo- 13. C. antennis fetariis, abdomine fubferrugineo, alis pre. 
ptrata. mortis externe craffioribus. 
Hobitat Upfalix. Carel. Chriftiernin, p.m. juvenis, 
Refers infellum coleoptratum. Antenna fetaria, uti 
Caput, Thorax, Pedes mufca domefiice. Abdomen 
mazis rotundatum, ferrugineum apice migro. Alz 
ereAa, abdomine vix lomgiores, quai pramorfa, la 
tifime, craffiores quam in altis, quafi uftulate ex 
fufco &F albc; pe peregrina facies. 


Figs. 1-2. Taxonomic descriptions. 1. Page from Scopoli (1763) treating Sicus and its included 


species. 2. Page from Linnaeus (1767) treating Conops testacea Linnaeus. 


Vol. 108, No. 4, September & October, 1997 27h 


clearly a composite of at least two species, which was not resolved until the 
present lectotype designation, the appropriate type species of Stomoxoides re- 
mains unresolved. As the illustration provided by Schaeffer is clearly of 
ferrugineus Linnaeus 1761 and the first included species is here restricted to 
testacea Linnaeus (sensu its lectotype), there is the problem of misidentification 
of the type species. The International Commission on Zoological Nomencla- 
ture needs (under Art. 70b) to rule whether the type is Sicus ferrugineus Linnaeus 
or Conops testacea Linnaeus, of which I would recommend the former. 


ACKNOWLEDGMENTS 


I thank Brian Pitkin, the Natural History Museum (formerly the British Museum [Natural 
History]), London (BMNH) and Michael Fitton, the Linnaean Collection, London (LSL); for 
permission to study material in their care. 

Without the advice of various specialists on conopid flies, this paper would not have been 
possible, so I am indebted to: Sidney Camras, Field Museum of Natural History, Chicago; David 
Clements, Cardiff, Wales; Milan Chvala, Charles University, Prague; and Kenneth G. V. Smith, 
retired from the British Museum (Natural History) [=The Natural History Museum], London. 

I also thank Neal L. Evenhuis, Bishop Museum, Honolulu; James Pakaluk, Richard E. White, 
Curtis W. Sabrosky, and Manya B. Stoetzel of the Systematic Entomology Laboratory, USDA, 
Washington; and Wayne N. Mathis of the Smithsonian Institution (USNM), Washington, for 
their critical review of the manuscript. 


LITERATURE CITED4 


Camras, S. 1994. Case 2881. Sicus Scopoli, 1763 and Myopa Fabricius, 1775 (Insecta, Diptera): 
proposed conservation by the designation of Conops buccata Linnaeus, 1758 as the type 
species of Myopa. Bull. Zool. Nomencl. 51: 31-34. 

Chvyala, M. 1961. Czechoslovak species of the subfamily Conopidae (Diptera: Conopidae). Acta 
Univ. Carol. Biol. 1961: 103-145. 

Chvyala, M. 1963. A review of the conopid flies of the genus Sicus Scop. (Diptera, Conopidae). 
Acta Univ. Carol. Biol. 1963: 275-282. 

Chvala, N. 1965. Czechoslovak species of the subfamilies Myopinae and Dalmanniinae (Diptera, 
Conopidae). Acta Univ. Carol. Biol. 1965: 93-149. 

Chvyala, M. and K. G. V. Smith. 1988. Family Conopidae. Pp. 245-272. In Soos, A. (ed.), Cata- 
logue of Palearctic Diptera. Vol. 8, Syrphidae — Conopidae. 363 pp. Akademiai Kiddo, 
Budapest. [1988.09.20]. 

Collin, J. E. 1946. ‘Rhinaia macrocephala’ not a valid name in Diptera (Syrphidae). Ent. Rec. J. 
Var. 58: 136-137. 

Collin, J. E. 1947. On suggested proposals for altering the names of two species of Syrphidae 
(Diptera), and on alterations in entomological names generally. Ent. mon. Mag. 83: 274-276. 
{1947.11.28] ; 

Collin, J. B. 1948. On the interpretation of Article 31 of the International Rules of Nomencla- 
ture. Ent. mon. Mag. 84: 235 236. [1948.10.11]. 

Collin, J. E. 1959. The British species of Myopa (Dipt., Conopidae). Ent. mon. Mag. 95: 145- 
151. [after 1959.09.10]. 

Coquillett, D. W. 1910. The type-species of the North American genera of Diptera. Proc. U. S. 
natn. Mus. 37: 499-647. [1910.08.04]. 

Day, M. C. and N. G. Fitton. 1978. Re-curation of the Linnaean Hymenoptera (Insecta), with 
a reassessment of the taxonomic importance of the collection. Biol. J. Linnaean Soc. 10: 
181-198. 


4 Titles and other details for older works are abbreviated; full titles and information on these 
works can be found in Thompson & Pont. 


272 ENTOMOLOGICAL NEWS 


Donovan, E. 1808. The natural history of British insects ... vol. 13, 74 + 4 pp., pls. 433-468. 
Rivington, London. 

Gmelin, J. F. 1790. Caroli a Linne, Systema naturae ... 13th ea., vol. 1 (5): 225-3020. G. E. Beer, 
Lipsiae [=Leipzig]. 

Goffe, E. R. 1946. The Syrphidae (Diptera) of Moses Harris, 1776. Ent. mon. Mag. 82: 67-86. 
[1946.03.30]. 

Goffe, E. R. 1947. Some notes on Syrphidae (Diptera). Ent. mon. Mag. 83: 195-197. [1947.08.28]. 

Goffo, E. R. 1948. Musca macrocephala Harris, 1776, and Scaeva arcuata Fallen, 1817 (Dipt., 
Syrphidae). Ent. mon. Mag. 84: 53-55. [1948.04.01]. 

Goffe, E. R. 1949. The position of Musca macrocephala Harris, 1776, and of certain other names 
of Syrphidae (Diptera) under Article 31 of the Rules. Ent. mon. Mag. 85: 35-37. [1949.03.11]. 

Harris, M. 1776. An exposition of English insects ... Decad III, pp. 73-99, pls. 21-30. Robson 
Co., London. 

Linnaeus, C. 1758. Systema naturae ... Ed. 10, Vol. 1. 824 pp. L.Salvii, Holmiae [=Stockholm]. 
[1758.01.01]. 

Linnaeus, C. 1761. Fauna Svecica ... Editio altera, auctior [49] + 578 pp., 2 pls. L. Salvii, 
Stockholmiae [= Stockholm]. (after 1761.07.28, date of preface). 

Linnaeus, C. 1767. Systema naturae ... Ed. 12 (revised.). Vol. 1, Pt. 2, pp. 533-1327. L. Salvii, 
Holmiae [=Stockholm]. [1767.06.14]. 

Mikhola, R. 1983. Diagnostic insect pins: Some problems of the Linnaean insect collection 
solved. Antenna 7: 16-17. 

Panzer, G. W. F. 1804. D. Jacobi Christiani Schaefferi Iconum Insectorum circa Ratisbonam 
indigenorum Enumeratio Systematica. xvi + 260 pp. Palmii, Erlangae [= Erlangen]. 

Poda [von Neuhaus], N. 1761. Insecta mused Graecensis ... 127 + [xii] pp., 2 pls. Widmanstadii, 
Graecii [= Graz] [1761.09.03]. 

Pont, A. C. 1981. The Linnaean species of the families Fannidae, Anthomyiidae and Muscidae 
(Insecta: Diptera). Biol. J. Linnaean Soc. 15: 165-175. [1981.02.23]. 

Rozkoény, R. 1984. The Sciomyzidae (Diptera) of Fennoscandia and Denmark. Fauna Ent. Scand. 
14, 224 pp. 

Sabrosky, C. W. 1994. Comments on the proposed conservation of Sicus Scopoli, 1763 and 
Myopa Fabricius, 1775 by the designation of Conops buccata Linnaeus, 1758 as the type 
species of Myopa (Insecta, Diptera), and on the proposed rejection of Coenomyia Latreille, 
1796. Bull. Zool. Nomencl. 51: 259-260. 

Schaeffer, J. C. 1766a. Icones Insectorum ... 3 vole., xx + 280 + xiv pp., 280 pls. Breitfeld, 
Regensburg. 

Schaeffer, J. C. 1766b. Elementa Entomologica. [7 + 8 + 17 pp.,] + 135 plates with facing 
legends. Weissianis, Regensburg. 

Scopoli, J. A. 1763. Entomologia carniolica ... 421 pp. Trattner, Vindobonae [=Vienna] [before 
1763.06.23]. 

Smith, K. G. V. 1969. Diptera. Conopidae. Handbk ident. Brit. Ins. 10 (3a), 19 pp. [1969.03.04]. 

Thompson, F. C. and A. Pont. 1994. Systematic database of Musca names (Diptera). Theses 
Zool. 20, 221 pp. 

Thompson, F. C., J. R. Vockeroth and M. C. D. Speight. 1982. The Linnaean species of flower 
flies (Diptera: Syrphidae). Mem. ent. Soc. Washington 10: 150-165 [1982.09.30]. 

Wheeler, T. A. 1994. Comments on the proposed conservation of Sicus Scopoli, 1763 and Myopa 
Fabricius, 1775 by the designation of Conops buccata Linnaeus, 1758 as the type species of 
Myopa (Insecta, Diptera), and on the proposed rejection of Coenomyia Latreille, 1796. Bull. 
Zool. Nomencl. 51: 260-261. 


Vol. 108, No. 4, September & October, 1997 273 


STUDIES ON SOME MOTH FLIES (DIPTERA: 
PSYCHODIDAE), WITH THE FIRST RECORD OF 
CLOGMIA ALBIPUNCTATA IN CENTRAL EUROPE! 


Doreen Werner2 


ABSTRACT: In the course of an ecological study of waste disposal facilities, the psychodid 
fauna of terrestrial habitats on refuse or garbage was investigated. Species of this family are 
important components of the edaphic insect fauna of these biotopes. The fauna was sampled at 
five sites by means of emergence traps, and material was reared under natural conditions. Eco- 
logical notes are given. Six species of Psychodidae were found and one of these, Clogmia 
albipunctata, is recorded for the first time from Germany. The most abundant species was Psychoda 
parthenogenetica. with 88.8% of the individuals sampled. No moth flies were reared at two of 
the five sites. 


Because of their anthropogenic characteristics, rubbish tips and dumps have 
featured regularly as a field of interest for zoologists and have also been the 
subject of faunistic and ecological studies. However, they are still relatively 
under-researched biotopes, and many basic questions remain unanswered. It 
has only recently been accepted that these sites consititute a complex ecosys- 
tem, with conditions that vary greatly within very short periods of time and 
which exist only for a short time because of continuous interference and modi- 
fication. The insects that inhabit these biotopes have frequently been the sub- 
ject of ecological examinations but have not been as intensively researched as 
other groups of soil animals. In the following paper the results are given of a 
study of Psychodidae that live on dumps. In general, the larvae of the 
Psychodidae and their ecology are better known than those of other Diptera, 
but they have not been examined in detail. The preimaginal stages of Psycho- 
didae are generally found in terrestrial, aquatic and dendro-limnic habitats. 


STUDY AREA 


This investigation was carried out in the Berlin area at several waste dis- 
posal facilities which employ different methods of waste management. The 
dumps at site 1, Schwanebeck, and site 2, Wernsdorf, manage refuse from 
Berlin and use a method of dense packing. These are so-called “Class II sites”. 
Approximately 70% of the waste is household waste, 20% household-related 


1 Received July 29, 1996. Accepted December 28, 1996. 


2 D. Werner, Museum fiir Naturkunde, Institut fiir Systematische Zoologie, InvalidenstraBe 43, 
10115 Berlin, Germany. 


ENT. NEWS 108(4): 273-282, September & October, 1997 


274 ENTOMOLOGICAL NEWS 


rubbish, and 10% is estimated to be communal waste. Following the windrow 
method, the refuse is immediately crushed and is then stacked as densely as 
possible so as to avoid empty spaces between the layers. After this treatment, 
the surface is covered usually with clay- or sandy soil to prevent rubbish being 
blown away and/or the air being polluted by dust or odors. Samples were col- 
lected from waste that was approximately 4-6 weeks old. 

The tip at site 3, Eggersdorf, is an unmanaged tip onto which not only 
household waste, building rubble, faeces and sewage are dumped but also in- 
dustrial waste. The survey for Diptera was carried out on deposits that were 3 
years old. 

At site 4, Falkenberg facility, biological waste from fruit growers and mar- 
ket gardeners (i.e. horticultural and agricultural waste) is composted. The or- 
ganic substances are treated in a manner that guarantees high biological 
degradability. The refuse is shredded and broken down, and is then stacked in 
windrows. The specific treatment of these windrows, with careful temperature 
and moisture control, ensures a constant process of decay. The traps for the 
Diptera survey were set on a new windrow. 

Since the dumps and tips discussed so far differ both in their components 
and their size, and thus in the overall conditions they provide for decomposi- 
tion, the investigation was extended to include a heap which contained the 
kitchen and garden waste from a private household. This was in an isolated 
position in Waldsieversdorf (site 5) in the Markische Schweiz region, about 
45m above sea level, and was about two years old when the survey was carried 
out. The heap remained undisturbed during the period of the investigation, 
with neither chalk nor other soil constituents added. 


SAMPLING METHODS 


The use of pyramid-shaped traps (photoeclectors) for the compilation of 
species inventories has proved very effective (Funke 1971). They have been 
used for ecological studies with increasing frequency in recent years. This 
method was used to collect the Diptera emerging from the soil. Most of the 
emerging Diptera behaved with positive phototaxis and flew into the transpar- 
ent plastic box filled with ethylene-glycol. Supplementary samples were col- 
lected using Barber traps (pitfall traps), colored dishes, or baited traps. The 
sampling period was from March 1993 to April 1994. The traps were used in a 
three-week cycle in which they were emptied at the end of the first week, 
removed, and then set up again after two weeks in order to avoid any effect 
from large climatic variations (Grimm et al. 1975). Furthermore the photo- 
eclectors were made of a material with a special fiber to reflect light and avoid 
overheating. 


Vol. 108, No. 4, September & October, 1997 DiS 


RESULTS 


Adult Psychodidae are small Diptera of compact or stocky, slightly hunch- 
backed build. They are characterized by their dense vestiture of hairs or scales. 
These features are responsible for their restricted and short flight ability, and 
also explain why they are found mainly in the immediate vicinity of their breed- 
ing sites. Predominantly Psychoda species hold their wings roof-shaped over 
the abdomen, a tactic to hide in small crevices. Climatic factors, for example 
temperature or dampness of the substrate, have a strong influence on the oc- 
currence of Psychodidae. Three of the study sites presented almost ideal con- 
ditions for Psychodidae. These were the dumps in Schwanebeck and Wernsdorf 
and the compost heap in Waldsieversdorf. 

On the dumps, the heterogeneous structure of the various components of 
rubbish, garden waste, and household waste offered protection from unfavor- 
able climatic conditions, for example from wind and rain, as the flies matured. 
Furthermore, there is a special microclimate in the windrows, characterized by 
an atmospheric humidity of 95-100% and a high soil temperature, caused mainly 
by energy released through fermentation and by the effect of the sun. 

When any sort of decaying organic material is added to these already fa- 
vorable conditions, flies can breed in huge numbers. Adults move over their 
feeding or breeding substrates by running or hopping or in short bursts of flight. 
They were only rarely collected in direct sunlight. Moth flies are crepuscular 
creatures, with only a short phase of activity in the morning, whereas they can 
be observed for several hours in the evening when they may even fly to a 
height of almost 1.5 metres. However, the moth flies observed during this project 
always remained close to their breeding medium. 

Psychodid larvae can be arranged into different ecological groups: they 
are found in stagnant or flowing, clean or dirty, fresh or salt water, mud, mosses, 
sap or soil (Vaillant 1971). In addition to larvae with a mydobiontic mode of 
life (particularly the genera Pericoma, Telmatoscous and Clytocerus), the lar- 
vae of many species, and in particular those of the genus Psychoda which has 
a saprobiontic mode of_life, are found in and on rotten plant material in ma- 
nure pits, sewage, cesspits and dumps. In general the larvae need a firm sub- 
strate, damp rotten organic material, and permanent access to atmospheric 
oxygen. 

This investigation produced a total of 3033 moth flies, which belong to a 
relatively small number of species: 6 species in 4 genera, all in the subfamily 
Psychodinae. Breeding populations of all these species were found at the sites 
listed above. 2262 specimens were caught in traps, and 771 using bait. The 
following species were found (nomenclature follows Wagner 1990): 


Satchelliella trivialis (Eaton 1893), Tinearia alternata (Say 1824), Psychoda 
albipennis (Zetterstedt 1850), Psychoda cinerea (Banks 1894), Psychoda 
parthenogenetica (Tonnoir 1940), and Clogmia albipunctata (Williston 1893). 


276 ENTOMOLOGICAL NEWS 


The dominant species was the cosmopolitan Psychoda parthenogenetica. 
Its life-cycle depends mainly on the available food supply, temperature and 
humidity. The time from egg-laying until adult eclosion is in the region of 22 
to 30 days at a temperature of 20-22°C. 

Psychoda parthenogenetica also breeds in baits. It has bred and hatched 
particularly successfully in damp rabbit droppings, rotting potatoes, cabbage, 
manure and carrots. A few specimens also have bred in rotten celeriac, soggy 
bread, ground coffee dregs and agar culture mixed with cow’s blood. When 
several substrates are available, it prefers dung as its basic food and breeding 
medium. This range of food substances is more extensive than has previously 
been mentioned in the literature on this species. Vaillant (1971) refers to the 
remarkably euryoecious mode of life of Psychodinae larvae. 

Although adults of Psychoda parthenogenetica visit liver and excrement, 
no egg-laying on these media took place either on the baits at the dumps or in 
the laboratory. 

As arule, larvae were found on the preferred baits in dark and shady slits 
and cracks, with the tube-like siphon projecting out of the soft development 
medium. 

The pupae are light brown, and are found in large numbers on the surface 
and at the edges of the substrates. Unlike other families of Nematocera, they 
are not connected to the last larval skin. The pupal stage lasts 3 to 5 days at 20- 
225C: 

At all stages this relatively short life-cycle is subject to strong natural in- 
fluences, mainly climatic variations. Delays are therefore possible in each of 
the individual developmental stages. A reduction in humidity and tempera- 
tures below 10°C greatly delayed or even held up the development of the 
preimaginal stages. Despite this, however, Psychoda parthenogenetica is able 
to develop and emerge even at low temperatures. In mild winters the first adults 
emerge as early as January, and only temperatures below 0°C stop their growth 
completely (R. Wagner pers. comm.). The larvae and pupae of Psychoda 
parthenogenetica are illustrated in Fig. 1. 

Adult moth flies were frequently infested by mites, 6 to 8 parasites being 
commonly found on a single moth fly, but they seem to have no influence on 
adult activity. Infestations are rarely found on those individuals which develop 
in or close to streams. 

The seasonal occurrence of Psychoda parthenogenetica at the various study 
Sites is shown in Figs 2a-c. Only the moth flies collected in photoeclector soil 
traps and in the Barber traps placed beneath the eclectors are included in these 
figures. Psychoda parthenogenetica was found constantly throughout the pe- 
riod of the investigation. From the figures it can be seen that moth fly activity 
began in March. Several generations developed in the biotopes under study, 
with the generations overlapping continuously throughout the season. A maxi- 


Vol. 108, No. 4, September & October, 1997 D0 


mum in the intervals between generations was noted in late summer and au- 
tumn. Wagner (1979) also reported that this species has a continuous seasonal 
distribution, reaching a maximum in autumn at the Breitenbach stream. 

The species collected at the various study areas are listed in Table 1. 

Compared with Psychoda parthenogenetica, Psychoda albipennis was rare 
although greater numbers had been expected. This species is usually found 
wherever there is decaying organic matter, for example alongside streams and 
brooks, silage, and mushroom farms. The larvae live in the faeces of various 
vertebrates (Wagner 1977). Whereas Satchell (1947) observed Psychoda 
albipennis on decaying substances and reaching a maximum in October, the 
species was only found in the course of this investigation to develop in the 
spring at the Waldsieversdorf compost heap. It is a bisexual species, and so it 
may not be possible for it to reach the high population densities that Psychoda 
parthenogenetica reaches (R. Wagner pers. comm.). Psychoda albipennis was 


Fig. 1: Larvae and pupae of Psychoda parthenogenetica. 


278 ENTOMOLOGICAL NEWS 


20 


Total (%) 


0 


te) 
DEP MED OO NH HAM OY YS gh gg” 
oP Pe PO hh HH OM ON DY QM OS 
Be ee gre BR a aN OY ons ee aa Oe 


Date 


Fig. 2a: Seasonal distribution of Psychoda parthenogenetica at the Schwanebeck tip 


10 


x 9: a YY SP GM OS 
PE PED _ EO NH H_E \ So 8 
PS Pp SPP Spr” PP errr Pirin’ 


Date 


Fig 2b: Seasonal distribution of Psychoda parthenogenetica at the Wernsdorf tip. 


Vol. 108, No. 4, September & October, 1997 279 


30 


° 
BS 
A 
== 
3 
~~ 20 
ae 
10 
0 
5 9: Lee 
SB? Pe PD EM H_H_ OD O'S o>: ws Sh Oo 
ASSESS x \ ro YM Og 
- Sa Via Foi ag 


Date 


Fig. 2c: Seasonal distribution of Psychoda parthenogenetica at the Waldsieversdorf compost 
heap. 


Total (%) 


10 


os? ok S&S wo oA ow DDD DD AS got VY oh 
SOG NS SY MY _ PS 
SPV VE DT MP Qh ADT BT ABT OT AT YD ne an SOA > ay OF AD? 


Date 


Fig. 3: Seasonal distribution of Tinearia alternata at the Schwanebeck tip. 


280 ENTOMOLOGICAL NEWS 


Table |: Species collected at the various study areas. 


Area found 


Eggersdorf | Falkenberg 


Number of 
individuals total 


Waldsievers- 
dorf 


Species 
Schwanebeck 


Satchelliella 


trivialis 


Tinearia 


alternata 


Psychoda 


albipennis 


Psychoda 


cinerea 


Psychoda 
partheno- 


genetica 


Clogmia 


albipunctata 


reared from potatoes and cabbage. Weber (1993) also recorded this species in 
potatoes, together with Psychoda minuta. It was not possible to rear this spe- 
cies in other types of decaying material. 

Psychoda cinerea is a multivoltine and euryoecious species with larvae 
that live in a wide range of conditions, for example on the bacterial slime in 
domestic household drains. Because of this it is doubtful whether they are able 
to survive extreme temperature fluctuations (R. Wagner pers. comm.). Speci- 
mens of Psychoda cinerea hatched from compost and from the Schwanebeck 
dump, but only in August and September and only in small numbers. Satchell 
(1947) found this species on farms, but never in high numbers. 


Vol. 108, No. 4, September & October, 1997 281 


Tinearia alternata, another cosmopolitan species, also requires decaying 
organic matter for its development. Vaillant (1971) refers to numerous types 
of larval nutrition. These range from, among others, saprobionts, coprobionts, 
urinobionts, psammabionts of various kinds in water from bryomadicoles 
to mycobionts. In the present investigation, specimens were found at the 
Schwanebeck and Wernsdorf dumps as well as on the Waldsieversdorf com- 
post heap. Although the species occurred throughout the entire period of the 
investigation, the maximum development was in May and August although it 
was also able to continue into September. The seasonal distribution of Tinearia 
alternata at Schwanebeck is shown in Fig. 3. Tinearia alternata developed 
and hatched from various baits, such as celeriac, carrots, potatoes, oak leaves, 
rabbit droppings and manure. Skidmore (1991) reported the occurrence of 
Psychoda cinerea and Psychoda parthenogenetica on cow dung. 

Clogmia albipunctata is a ubiquitous species of the tropics and subtropics 
whose larvae use all kinds of rotten organic matter for their basic food and 
breeding medium. This species has been known for many years to be widely 
distributed (Duckhouse 1989). It was previously known from Cuba and from 
various parts of tropical Africa. The present specimens emerged from samples 
of manure from the Schwanebeck dump in August 1993. The pH-value of both 
samples was between 8.2 and 8.3. Species from other families of Diptera were 
also developing in these manure samples, for example Syrphidae and Scatop- 
sidae. Clogmia albipunctata is here recorded for the first time in Germany. It 
may be that special microhabitats enable the eggs and larvae of this species to 
survive winter outside the tropics and subtropics. 

The larvae of the West Palaearctic Satchelliella trivialis commonly de- 
velop alongside brooks, streams and small rivers, in damp marginal soil or in 
mud mixed with plant remains. This species is highly tolerant of decaying 
matter (Vaillant 1979), and under optimum conditions there are two genera- 
tions per year (Wagner 1979). Satchelliella trivialis is adapted to slightly brack- 
ish water and water with a high organic content of vegetable origin. In addition 
to adult flies collected with a suction trap, Satchelliella trivialis hatched from 
manure samples from the Schwanebeck dump, as did Clogmia albipunctata. 
Its occurrence was restricted to August and September. In central Europe there 
are two generations per year, while in Scandinavia the species is univoltine. 
Under optimum conditions in the Mediterranean area there are 3 to 4 genera- 
tions. The specimens were found together with Tinearia alternata. Vaillant 
(1981) also includes Psychoda cinerea in this group. At the Schwanebeck dump, 
Psychoda cinerea was found at the same time but not in the same sample. 

It is surprising that no Psychodidae were found at either the Falkenberg or 
the Eggersdorf dumps. At Eggersdorf it is possible that conditions are too het- 
erogeneous or that the sites are too small or too temporary for moth fly devel- 
opment. At Falkenberg, as a result of the specific treatment of the layers, levels 


282 ENTOMOLOGICAL NEWS 


of temperature and humidity were established which should have been very 
favorable for these flies. There was constant humidity, atmospheric oxygen 
for respiration, and ample organic substances for food and breeding grounds. 
However, microbial breakdown of organic material led to a very high level of 
heat within the layers. As the process of decay continued, the extreme tem- 
peratures fell, but, when compared with its immediate environment and with 
the other dumps, this site has the most favorable temperatures. 

It is possible that Psychodidae avoid these extreme temperatures, which 
are tolerated or even preferred by many other families. It is also possible that 
these features are of too short a duration for the development of moth fly lar- 
vae and pupae. The preimaginal stages need almost constant conditions through- 
out the life-cycle. 

Further investigations will be necessary to examine fully the habitat and 
environmental requirements of this family. 


ACKNOWLEDGMENT 


I would like to thank R. Wagner, Limnologische FluBstation of the Max-Planck-Institut 
of Limnology for his assistance with the determination of difficult material. 


LITERATURE CITED 


Duckhouse, D.A. & D.J. Lewis. 1989. Family Psychodidae. In: Evenhuis, N.L. (eds): Cata- 
logue of the Diptera of the Australasian and Oceanan Region. - Bishop Museum Press. |- 
SS: 

Funke, W. 1971. Food and Energy Turnover of Leaf-eating Insects and their Influence on Pri- 
mary Production. Ecol. Stud. 2: 81-93. 

Grimm, R, Funke, W. & J. Schauermann. 1975. Untersuchungen an Tierpopulationen in 
Waldékosystemen. Verh. Ges. Okol. Erlangen. 1974: 77-87. 

Satchell, G.H. 1947. The ecology of the British species of Psychoda (Diptera, Psychodidae). 
Ann. appl. Biol. 34: 611-621. 

Skidmore, P. 1991. Insects of the British cow-dung community. Field Studies Council. 21: 1- 
166. 

Vaillant, F. 1971. Psychodidae In: Lindner, E.: Die Fliegen der Palaearktischen Region. 9d: 1- 
48, 

Vaillant, F. 1979. Psychodidae. In: Lindner, E.: Die Fliegen der Palaearktischen Region. 9d: 
239-270. 

Vaillant, F. 1981. Psychodidae. In: Lindner, E.: Die Fliegen der Palaearktischen Region. 9d: 
271-310. 

Wagner, R. 1977. Zur Kenntnis der Psychodidenfauna des Allgaus. Nachrichtenblatt Bayer. 
Entomol. 26 (2): 23-28. 

Wagner, R. 1979. Psychodidenstudien im Schlitzerland. Schlitzer Produktionsbiologische Studien. 
Arch. Hydrobiol. Suppl. 57 (1): 38-88. 

Wagner, R. 1990. Family Psychodidae, In: Sods, A. & Papp, L. (eds.): Catalogue of Palaearctic 
Diptera. 2: 11-65. — Akadémiai Kiad6, Budapest. 

Weber, G. 1993, Die Nematocera (Insecta: Diptera) eines klarschlammgediingten und schwer- 
metallbelasteten Ackers: OkGlogie und Larvalbiologie. PhD thesis, Technical University of 
Braunschweig (FRG), 1993. 


Vol. 108, No. 4, September & October, 1997 283 


CONTRIBUTION TO THE SYSTEMATICS OF THE 
GENUS CHELEOCLOEON (EPHEMEROPTERA: 
BAETIDAE)!: 2 


C. R. Lugo-Ortiz, W. P. McCafferty 


ABSTRACT: The Afrotropical genus Cheleocloeon (Ephemeroptera: Baetidae) is shown to con- 
stitute a small, but widespread, monophyletic group of species that are distinguished by an ante- 
riorly convex medial process of the second segment of the labial palps and long, poorly denticu- 
late tarsal claws. The genus is hypothesized to belong to the Bugilliesia complex of genera on the 
basis of the basally bulbous second segment of the male genital forceps and general morphology 
of the larval tarsal claws and gills. Cheleocloeon dimorphicum, n. comb., and C. excisum, n. 
comb., are transferred from the genus Afroptilum, and are distinguishable among species of 
Cheleocloeon by the presence of hindwings in male adults and modifications of the medial pro- 
cess of the second segment of the labial palps. The commonly collected larval stage of C. excisum 
is redescribed, and new figures are provided to illustrate previously overlooked characters of the 
species. 


Wuillot and Gillies (1993) erected the genus Cheleocloeon for two distinc- 
tive species of small minnow mayflies (Ephemeroptera: Baetidae) from Guinea: 
C. carinatum Wuillot and C. yolandae Wuillot. Those authors distinguished 
larvae of Cheleocloeon by the long, distally acute medial process of the sec- 
ond segment of the labial palps (Wuillot and Gillies 1993: Figs. 6, 15), ab- 
sence of hindwingpads, elongate first pair of gills (Wuillot and Gillies 1993: 
Figs. 9, 18), and relatively long, poorly denticulate tarsal claws (Wuillot and 
Gillies 1993: Figs. 8, 17). They stated, however, that adults of Cheleocloeon 
could not be confidently separated from other baetid adults with single mar- 
ginal intercalaries in the forewings. 

Our examination of the baetid fauna of Africa has revealed that Afroptilum 
dimorphicum (Soldan and Thomas), from Algeria, and A. excisum (Barnard), 
from southern Africa, belong in Cheleocloeon. Both species were originally 
described in the genus Centroptilum Eaton but later assigned to the dimorphicum 
group of Afroptilum Gillies by Gillies (1990). The inclusion of these two spe- 
cies in Cheleocloeon expands the concept of the genus because they have cer- 
tain characteristics not present in the type of the genus, C. yolandae. Here, we 
provide a revised diagnosis of Cheleocloeon and discuss the newly observed 
variability in the genus. We also redescribe the larval stage of C. excisum and 
provide new figures showing characteristics overlooked by Barnard (1932). 
The material examined is housed in the Albany Museum (AM), Grahamstown, 
South Africa, and the Purdue Entomological Research Collection (PERC), West 
Lafayette, Indiana. 


! Received December 9, 1996. Accepted January 28, 1997. 
2 Purdue Agricultural Research Program Journal No. 15288. 
3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 


ENT. NEWS 108(4): 283-289, September & October, 1997 


284 ENTOMOLOGICAL NEWS 


Cheleocloeon Wuillot and Gillies 


Cheleocloeon Wuillot and Gillies 1993:213. 


Diagnosis. Larvae of Cheleocloeon are distinguished by the anteriorly con- 
vex medial process of the second segment of the labial palps (Figs. 6, 7; Soldan 
and Thomas 1985: Fig. 5; Wuillot and Gillies 1993: Figs. 6, 15) and the long, 
poorly denticulate tarsal claws (Figs. 8, 9; Soldan and Thomas 1985: Fig. 8; 
Wuillot and Gillies 1993: Figs. 8, 17). Male adults have the second segment of 
the genital forceps basally bulbous (Wuillot and Gillies 1993: Fig. 2), and, when 
hindwings are present, these are long and have two longitudinal veins and a 
hooked costar process located near the basal third (Barnard 1932: Fig. 14e). 
These adult characteristics, however, will not allow them to be distinguished 
from all other baetid genera with single marginal intercalaries in the forewings. 


Type species. Cheleocloeon yolandae Wuillot. 


Included species. 


Cheleocloeon dimorphicum (Soldan and Thomas), n. comb. 
Centroptilum dimorphicum Soldan and Thomas 1985:180 (larva; male subimago; 
female adult). 
Afroptilum dimorphicum (Soldan and Thomas): Gillies 1990:99. 
Cheleocloeon excisum (Barnard), n. comb. 
Centroptilum excisum Barnard 1932:224 (larva; male, female subimagos; male, 
female adults). 
Afroptilum excisum (Barnard): Gillies 1990:99. 
Cheleocloeon carinatum Wuillot 
Cheleocloeon carinatum Wuillot, in Wuillot and Gillies 1993:214 (larva; male 
adult). 
Cheleocloeon yolandae Wuillot 
Cheleocloeon yolandae Wuillot, in Wuillot and Gillies 1993:214 (larva; male adult). 


Distribution. Algeria (C. dimorphicum); Guinea (C. carinatum, C. yolandae); Lesotho 
(C. excisum); South Africa: Eastern Cape, KwaZulu Natal, Mpumalanga, Western Cape (C. 
excisum). 


Discussion. We consider the anteriorly convex medial process of the sec- 
ond segment of the labial palps (Figs. 6, 7; Soldan and Thomas 1985: Fig. 5; 
* Wuillot and Gillies 1993: Figs. 6, 15) and long, poorly denticulate tarsal claws 
(Figs. 8, 9; Soldan and Thomas 1985: Fig. 8; Wuillot and Gillies 1993: Figs. 8, 
17) to be autapomorphies that cladistically define Cheleocloeon. This mono- 
phyletic group is now known to include species with larvae that have either an 
apically blunt or pointed medial process of the second segment of the labial 
palps, and that have male adults with or without hindwings. Cheleocloeon 
dimorphicum and C. excisum differ from C. carinatum and C. yolandae in the 
morphology of the medial process of the second segment of the labial palps. In 


Vol. 108, No. 4, September & October, 1997 285 


C. excisum, that process is apically pointed (Figs. 6, 7), but it is not as long as 
in C. carinatum (Wuillot and Gillies 1993: Fig. 15) or C. yolandae (Wuillot 
and Gillies 1993: Figs. 6). In contrast to those three species, C. dimorphicum 
has a process that is somewhat short and apically blunt (Soldan and Thomas 
1985: Fig. 5). Only the male adults of C. dimorphicum and C. excisum have 
hindwings, but both males and females of C. carinatum and C. yolandae have 
hindwings. The hindwings of the male alate stages of C. excisum are some- 
what long and possess two longitudinal veins and a hooked costal process 
located near the basal third (Barnard 1932: Fig. 14e). Male adults of C. 
dimorphicum are not known; however, the hindwings were described by Soldan 
and Thomas (1985) based on male subimago hindwings dissected from 
mature larvae, and their morphology is similar to that of C. excisum. 

The inclusion of C. dimorphicum and C. excisum in Cheleocloeon consid- 
erably extends the known geographic range of the genus. Not only is the genus 
widespread in Africa (see Distribution, above), but it occurs at low and high 
altitudes and in arid, semiarid, and humid biomes (Soldan and Thomas 1985, 
Palmer et al. 1993, Wuillot and Gillies 1993). 

Wuillot and Gillies (1993) suggested that Cheleocloeon and Demoulinia 
Gillies are sister groups due to the shared absence of hindwings, pointed me- 
dial process of the second segment of the labial palps, and long, poorly den- 
ticulate tarsal claws. We cannot agree based on that assessment alone. Irre- 
spective of the fact that hindwings are now known to be present in some spe- 
cies of Cheleocloeon, the absence of hindwings in Baetidae has been shown to 
be a generally unreliable characteristic for inferring common ancestry and 
generic constraints due to the numerous instances of convergence among un- 
related species throughout the family (McCafferty and Waltz 1990). The mor- 
phology of the medial process of the second segment of the labial palps is 
actually quite different in Cheleocloeon and Demoulinia. In Demoulinia, that 
process is anteriorly straight and distally convex (Crass 1941: Fig. 27d), not 
anteriorly convex as in Cheleocloeon (Figs. 6, 7; Soldan and Thomas 1985: 
Fig. 5; Wuillot and Gillies 1993: Figs. 6, 15), and it is not always pointed in 
Cheleocloeon (Soldan and Thomas 1985: Fig. 5). Furthermore, the presence 
of long, poorly denticulate tarsal claws in Cheleocloeon and Demoulinia may 
or may not be commonly derived because such claws are found in many unre- 
lated taxa in Baetidae (e.g., Apobaetis Day, Paracloeodes Day, Potamocloeon 
Gillies, Pseudocentroptiloides Jacob). 

Cheleocloeon apparently belongs to the Bugilliesia complex of genera de- 
fined by Lugo-Ortiz and McCafferty (1996) because the second segment of 
the male genital forceps is basally bulbous (Wuillot and Gillies 1993: Fig. 2). 
Furthermore, within the hypothesized cladogram of the Bugilliesia complex, 
Cheleocloeon would be in a position intermediate between the two most an- 
cestral genera of the complex, Potamocloeon and Afrobaetodes Demoulin 


286 ENTOMOLOGICAL NEWS 


(Lugo-Ortiz and McCafferty 1996: Fig. 1). Cheleocloeon shares reduction of 
the tarsal claws (Fig. 9) and loss of the dorsal gill flaps (Figs. 11, 12) with 
Afrobaetodes and other more apotypic genera of the Bugilliesia complex; how- 
ever, it does not share the absence of the third segment of the male genital 
forceps with that same group of genera. Thus, we hypothesize a sister relation- 
ship of Cheleocloeon with those genera, and the relatively basal phyletic deri- 
vation mentioned above. Although the larval characteristics and lack of hindwings 
in Demoulinia are inconclusive with respect to deducing its relationships, as 
noted above, there remains the possibility that the genus may also prove to 
belong to the Bugilliesia complex. Unfortunately, the male genital forceps fig- 
ured by Crass (1947) are too schematic to allow a definitive analysis, and we 
have not been able to secure adult male specimens for examination. 


Cheleocloeon excisum (Barnard), n. comb. 


Larva. Body length: 5.9-6.2 mm; caudal filaments length: 2.5-2.7 mm. Head: Coloration 
yellow-brown, with vermiform markings on frons. Antennae nearly 2.0x length of head capsule. 
Labrum (Fig. |) with submedial setae and bifurcate anterior marginal setae. Hypopharynx as in 
Figure 2. Left mandible incisors (Fig. 3) with 3 + 3 denticles; prostheca apically denticulate; 
small tuft of simple setae present between prostheca and mola; triangular process at base of mola 
somewhat sharp and elongate. Right mandible incisors (Fig. 4) with 3 + 4 denticles; prostheca 
apically setose; large tuft of simple setae present between prostheca and mola; smaii tuft of setae 
present at base of mola. Maxillae (Fig. 5) with four apical denticles on galealaciniae; three to 
four fine, simple setae present laterally near base of denticles; medial hump with stout, simple 
seta; palps two segmented, extending beyond galealaciniae, palp segment 2 slightly longer than 
segment |. Labium (Fig. 6) with many long, simple setae marginally on glossae and paraglossae; 
palps two segmented; palp segment | subequal to segment 2; segment 2 (Fig. 7) with well- 
developed, distally pointed medial process, and abundant long, robust and short, fine, simple 
setae on surface. Thorax: Coloration yellow-brown, with no distinct pattern. Hindwingpads present 
in males, absent in females. Legs (Fig. 8) pale yellow-brown; femora dorsally with four to six 
short, simple setae and ventrally with six to eight short, simple setae; tibiae dorsally with numer- 
ous minute, fine, simple setae and ventrally with six to eight short, simple setae; tarsi dorsally 
with numerous minute, fine, simple setae and ventrally with 15-17 short, simple setae, somewhat 
increasing in length distally; tarsal claws (Fig. 9) long, nearly 0.5x length of tarsi, with three to 
four poorly developed denticles. Abdomen: Coloration medium brown to cream, with variable 
color patterns. Terga 1, 3, 5, 6, and 9 usually medium brown with various cream markings; other 
terga usually cream with various medium brown markings. Tergal surfaces (Fig. 10) with minute 
scale bases, mostly in pairs; posterior margins with sharp triangular spination. Sterna cream to 
pale yellow. Gill 1 (Fig. 11) elongate, paddle-shaped, poorly tracheated, without marginal serra- 
tions; gills 2-7 (Fig. 12) broadest in middle, well tracheated, with minute marginal serrations 
(Fig. 13). Paraprocts (Fig. 14) with six to seven sharp marginal spines. Caudal filaments cream to 
pale brown; terminal filament subequal to cerci. 


Adult. See description by Barnard (1932). 
Material examined. LESOTHO: Linakeng-Sengu, Sani R, pool above Sani bridge, IX-23- 


1988, P. H. Skelton, larvae (AM); SOUTH AFRICA: Eastern Cape Province: Klein Vispruit at 
Groot Valley, ca 20 km SW of Cradock, X-11-1990, W. P. and N. McCafferty, larvae (PERC); 


Vol. 108, No. 4, September & October, 1997 287 


Figs. 1-14. Cheleocloeon excisum, larva. |. Labrum. 2. Hypopharynx. 3. Left mandible. 4. Right 
mandible. 5. Right maxilla. 6. Labium (left-ventral; right dorsal). 7. Segment 2 of labial palps 
(enlarged). 8. Right foreleg. 9. Right foretarsal claw. 10. Detail of tergum 4. 11. Gill 1. 12. Gill 4. 
13. Detail of gill 4. 14. Paraproct. 


288 ENTOMOLOGICAL NEWS 


Fish R, N of Karoo Sulphur Springs, nr Cradock, XI-10-1990, W. P. and N. McCafferty, larvae 
(PERC); Little Fish R, at R 32, nr Sheldon, XI-11-1990, W. P. and N. McCafferty, larvae (PERC); 
Kap R crossing, Bathurst St. For., 10 km S of Rd N 2, nr Grahamstown, IX-14-1990, W. P. and N. 
McCafferty, larvae (PERC); Blaukrans R at Blaukrans Pass, on Rd 67, between Grahamstown 
and Bathurst, XI-14-1990, W. P. and N. McCafferty, larvae (PERC); KwaZulu-Natal Province: 
Howick Falls, Umgeni R, X-2-1971, G. F. and C. H. Edmunds, larvae (PERC); Umzinkulu R, 
between Underberg and Boesmansnek, 15.5°C, X-2-1971, G. F. and C. H. Edmunds (PERC); 
Krantzkloof Nat. Res., Molweni R, nr Kloof, X-4-1971, G. F. and C. H. Edmunds, larvae (PERC); 
Umlaas R, at Durban waterworks filtration plant, nr Pinetown, X-4-1971, G. F. and C. H. Edmunds, 
larvae (PERC); Umgeni R, above Nagel Dam Impoundment, IX-21-1990, W. P. and N. McCafferty, 
larvae (PERC); Wilge R, at R 714, W of Warden, X-16-1990, W. P. and N. McCafferty, larvae 
(PERC); Molweni R, at Krantzkloof Nat. Res., nr Durban, IX-21-1990, W. P. and N. McCafferty 
and B. Fowles, larvae (PERC); Mpumalanga Province: Buffelspruit at Shalom (Aalwan), 4 km 
W of Badplaas paralleling Rt 38, off Avantune Rd, 1167 m, X-17 1990, W. P. and N. McCafferty, 
larvae (PERC); Sabie-Sand Game Res., Sand R at Londolozi, X-20-1990, W. P. and N. McCafferty, 
larvae (PERC); Kruger Ntl. Pk., Sabie R at Molondozi, X-23-1990, W. P. and N. McCafferty, 
larvae (PERC); Kruger Ntl. Pk., Sabie R, SE corner of Old Rhino Camp, X-24-1990, W. P. and 
N. McCafferty, larvae (PERC); Kruger Ntl. Pk., Sabie R at Lisbon Estates, X-27-1990, W. P. and 
N. McCafferty, larvae (PERC); Kruger Nat. Pk., Olifants R, 15 km from Blacktop Rd, at Fig Tree 
Site, X-29-1990, W. P. and N. McCafferty, larvae (PERC), Kruger Nat. Pk., Olifants R at bridge 
on dirt rd, nr Olifants Camp, X-29-1990, W. P. and N. McCafferty, larvae (PERC). 


Discussion. Barnard (1932) described C. excisum based on larvae and reared 
adults from the Western Cape Province in South Africa. However, his descrip- 
tion of the larval stage is brief and his drawings are schematic and inaccurate. 

Larvae of C. excisum are distinguished from those of C. dimorphicum by 
the distally pointed medial process of the second segment of the labial palps 
(Figs. 6, 7). Larvae of C. carinatum and C. yolandae are generally similar to 
those of C. excisum, but the medial process of the second segment of the labial 
palps of C. carinatum and C. yolandae is longer (Wuillot and Gillies 1993: 
Figs. 6, 15), and their males and females lack hindwingpads. 

Larvae of C. excisum are found in riffles and stony backwaters (Crass 1947, 
Palmer et al. 1993). Palmer et al. (1993) found that they mostly feed on ultrafine 
and fine particulate organic matter (0.5-250 tum), but mature individuals are 
capable of handling relatively large diatoms. Adults emerge primarily during 
the south-temperate winter months (June-August) according to Crass (1947). 


ACKNOWLEDGMENTS 


We thank H. Barber-James and F. de Moor (Grahamstown, South Africa) and N. McCafferty 
(West Lafayette, Indiana) for their assistance. Research funds were provided to WPM by the 
South African Foundation for Research Development and the Anglo-American and de Beers 
Chairman’s Fund. The Albany Museum kindly provided office and laboratory facilities to WPM 
during his stay in South Africa. 


Vol. 108, No. 4, September & October, 1997 289 


LITERATURE CITED 


Barnard, K. H. 1932. South African may-flies (Ephemeroptera). Trans. Roy. Soc. S. Afr. 20: 
201-259. 

Crass, R. S. 1947. The may-flies (Ephemeroptera) of Natal and Eastern Cape. Ann. Natal. Mus. 
11: 37-110. 

Gillies, M. T. 1990. A revision of the African species of Centroptilum Eaton (Baetidae, 
Ephemeroptera). Aq. Insects 12: 97-128. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996. The Bugilliesia complex of African Baetidae 
(Ephemeroptera). Trans. Entomol. Soc. Am., 122: 175-197. 

McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) 
of North and Middle America. Trans. Entomol. Soc. Am. 116: 769-799. 

Palmer, C., J. O’Keefe, A. Palmer, T. Dunne, and S. Radloff. 1993. Macroinvertebrate func- 
tional feeding groups in the middle and lower reaches of the Buffalo River, eastern Cape, 
South Africa. I. Dietary variability. Freshwat. Biol. 29: 441-453. 

Soldan, T. and A. G. B. Thomas. 1985. Centroptilum dimorphicum, sp. n., a new species of 
mayfly (Ephemeroptera: Baetidae) from Algeria. Acta Entomol. Bohemoslov. 82: 180-186. 

Wuillot, J. and M. T. Gillies. 1993. Cheleocloeon, a new genus of Baetidae (Ephemeroptera) 
from West Africa. Rev. Hydrobiol. Trop. 26: 213-317. 


SCIENTIFIC NOTE 


OBSERVATION OF THE BURYING BEETLE, 
NICROPHORUS TOMENTOSUS (COLEOPTERA: 
SILPHIDAE), BURYING A MOLE! 


Paul P. Shubeck2; 3 


On September 28, 1996 at 6:15 P.M. I approached a dead mole which had been lying near a 
garbage container, next to my home, about 3 days. It was breezy, and had been for several hours, 
and for the first time in 3 days I detected the characteristic odor of a decomposing animal. I was 
surprised to see what appeared to be a “bumble bee” flying around the dead mole and I bent 
down for a better look. Close observation revealed that the insect was actually a beetle, Nicrophorus 
tomentosus Weber, which then landed next to the mole and immediately crawled beneath it. 
Possible mimicry by this beetle has been commented on by Milne and Milne (1944) and by 
Shubeck (1971). While observing the mole and beetle I realized that only about 10 centimeters 
from the cadaver was another burying beetle. It was perched upon the upper edge of a stone walk 
with its head pointed downward and the tip of its abdomen, above the pavement, pointed straight 
up. In Pukowski’s European study (1933), Nicrophorus males crawl from a small cadaver to a 
stone or plant and extend their abdomens obliquely and thus release odors to attract a mate. 

At 6:45 P.M. (official sunset) I returned to the scene and found the “calling” individual had 
left its perch and apparently crawled under the mole where there was much activity. The carcass 
was wiggling and I could see the sides of at least 2 beetles as they excavated. The mole had sunk 
at least 1/2 cm along its entire length (about 15 cm). 


(Continued on page 299) 


1 Received October 17, 1996, Accepted March 5, 1997. 
2 Professor Emeritus, Montclair State University. 
3 Present Address: 65 Pleasantview Ave., New Providence, NJ 07974. 


290 ENTOMOLOGICAL NEWS 


NOTES ON SOME FLEAS (SIPHONAPTERA) FROM 
AMAZONAS AND BAHIA STATES, BRAZIL! 2 3 


Michael W. Hastriter4, Norman E. Peterson> 


ABSTRACT: Ten species of fleas collected from 19 mammalian host species are reported from 
the Brazilian States of Amazonas and Bahia. Validity of previously used taxonomic characters 
are considered and new ones presented. 


U.S. Army Medical personnel collected small mammals in Brazil, for bio- 
medical research, from August 1980 to February 1986. Collections included 
548 fleas (10 species) from 164 mammals (19 species). Although the majority 
of specimens were collected from Bahia State, some were collected from the 
State of Amazonas where few records have been established. The purpose of 
this paper is to establish new distributional records, to present relevant char- 
acters to distinguish several species of Polygenis females, and to clarify an 
anatomical inconsistency published by Cerqueira and Linardi (1976) dis- 
tinguishing Polygenis tripus (Jordan 1933) from P. rimatus (Jordan 1932). Data 
representing host/parasite associations, collection localities, and remarks on 
individual flea species follows. Mammal classifications follow Wilson and 
Reeder (1993). 


Study Sites: Abbreviated collection localities and ecological data are provided 
as follows: 


STATE OF AMAZONAS - 

Loc. A: Carauari, Gavide - 4°52°S 66°52°W, elev: 100 meters. 

Loc. B: Tefé, Cidade Jurua - 3°23°S 66°01 “W, elev: 100 meters. Collected in the Amazon 
Basin along the Jurua River in or adjacent to humid tropical forest where the commercially 
important trees have been removed. 


STATE OF BAHIA - 
Loc. C: Caatinga de Moura - 10°59’S 40°45 °W, elev: 600 meters. Collected along streams, 
or marsh in the dry Caatinga region of northeastern Brazil. 


1 Received January 9, 1997, Accepted February 24, 1997. 


2 Supported by U.S. Army Medical Research and Development Command Research Contract 
DAMD 17-88-H-8008. 


3 The opinions contained herein are those of the authors and should not be construed as official 
or reflecting the views of the Department of the Army. 


4 Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, P.O. Box 20200, 
Provo, Utah 84602-0200. 


5 U.S. Army Medical Research Unit, Brazil, Unit 3501, APO AA 34030. 


ENT. NEWS 108(4): 290-296, September & October, 1997 


Vol. 108, No. 4, September & October, 1997 291 


Loc. D: Corte de Pedra - 13°20°S 39°28°W, elev: 200-450 meters. 

Loc. E: Gandu - 13°47°S 39°35°W, elev: 300 meters. 

Loc. F: Trés Bracos - 13°32°S 39°45°W, elev: 200-550 meters. Collected in the Atlantic 
Forest Region approximately 150 kilometers southeast of Salvador. This area has steep hills and 
valleys where cacao is grown. The region was originally covered with a humid tropical forest 
which has been cleared for agricultural purposes. Tall forests remain only on the hilltops. Banana 
and cacao plantations, secondary scrub, pasture and varying degrees of secondary forest are 
close to the houses. 


RESULTS AND DISCUSSION 


Mammalian species are arranged alphabetically followed by the species of 
fleas in descending order of abundance on each specific host. The numbers 
following the flea species indicate the number of fleas collected per number of 
hosts that harbored one or more fleas. Metachirus nudicaudatus (21%), 
Proechimys theringi (18%), and Bolomys sp. (18%) comprised 57% of the 
mammals which harbored fleas. 


Mammalia 


Carnivora (Canidae): Cerdocyon thous (Linnaeus, 1766): Adoratopsylla intermedia intermedia 
(1/1), Ctenocephalides felis felis (1/1). 


Didelphimorphia (Didelphidae): Didelphis albiventris Lund, 1840: A. i. intermedia (9/11), 
Adoratopsylla antiquorum antiquorum (2/11), Polygenis bohlsi jordani (2/11), Polygenis pradoi 
(1/11), Polygenis rimatus (1/11), Polygenis roberti roberti (1/11); Didelphis marsupialis Linnaeus, 
1758: A. i. intermedia (3/3), P. r. roberti (1/3); Marmosa murina (Linnaeus, 1758): A. i. intermedia 
(2/3), A. a. antiquorum (1/3; Marmosa sp.: A. a. antiquorum (1/1); Marmosops parvidens Tate, 
1931: A. a. antiquorum (1/1); Micoureus demerarae (Thomas, 1905): A. a. antiquorum (2/3), P. 
r. roberti (1/3); Metachirus nudicaudatus (Desmarest, 1817): A. i. intermedia (35/35), A. a. 
antiquorum (2/35), Monodelphis americana (Muller, 1776): A. a. antiquorum (4/4); Philander 
opossum (Linnaeus, 1758): A. i. intermedia (2/2). 


Rodentia (Cavidae): Cavia aperea Erxleben, 1777: Polygenis tripus (3/4), P. r. jordani (2/4), A. 
i. intermedia (1/4). 


Rodentia (Echimyidae): Proechimys iheringi Thomas, 1911: Hectiella nitidus (27/29), A. i. 
intermedia (2/29), P. pradoi (1/29); Proechimys longicaudatus (Rengger, 1830): Gephyropsylla 
klagesi samuelis (3/3). E 


Rodentia (Muridae): Bolomys lasiurus (Lund, 1841): P. rimatus (2/5) A. i. intermedia (1/5), P. 
pradoi (1/5), P. tripus (1/5); Bolomys sp.: P. pradoi (17/25), P. rimatus (6/25), A. i. intermedia (2/ 
25), A. a. antiquorum (1/25), P. tripus (1/25); Holochilus brasiliensis (Desmarest, 1819): G. k. 
samuelis (1/2), P. pradoi (1/2); Nectomys squamipes (Brants, 1897): P. r. roberti (5/6), A. a. 
antiquorum (1/6), A. i. intermedia (1/6); Oryzomys capito (Olfers, 1818): P. r. roberti (2/2), H. 
nitidus (1/2); Oryzomys subflavus (Wagner, 1842): A. a. antiquorum (2/2); Oryzomys sp.: P. r. 
roberti (3/7), A. i. intermedia (2/7), A. a. antiquorum (1/7), P. b. jordani (1/7); Oxymycterus sp.: 
P. rimatus (14/14), P. r. roberti (2/14), A. a. antiquorum (1/14); Rhipidomys masticalis (Lund, 
1840): A. a. antiquorum (1/1). 


N 
\O 
Nm 


ENTOMOLOGICAL NEWS 


Siphonaptera 


The number of male and female fleas found on any particular host species 
precedes the host for each locality at which they were collected. 


Pulicidae: Crenocephalides felis felis (Bouché, 1835) — Loc. C: 19, C. thous. 


Ctenophthalmidae: Adoratopsylla (Adoratopsylla) antiquorum antiquorum (Rothschild, 1904) 
— Loc. D: 20,39, M. americana, 1Q each, M. demerarae, Marmosa sp., N. squamipes, O. 
subflavus, and Oryzomys sp.; Loc. F: 49, M. demerarae,; 20, 19, M. murina, 20, M. 
nudicaudatus; 2Q , D. albiventris; 10,19, O. subflavus; 1C each, M. parvidens, Marmosa 
sp., Oxymycterus sp., and R. masticalis, 1Q, Bolomys sp. 

REMARKS: The known distribution of A. a. antiquorum extends from 
Venezuela (Tipton and Machado-Allison, 1972) to southeastern Brazil. 
Guimaraes (1972) found this species widely distributed in the Brazilian States 
of Alagoas, Ceara, Bahia, and Pernambuco mainly on marsupial hosts and 
sporadically on various species of Bolomys, Oryzomys, and Proechimys. 


Adoratopsylla (Tritopsylla) intermedia intermedia (Wagner, 1901) — Loc. C: 20, C. aperea; 
30, B. lasiurus; Loc. D: 240°, 49, M. nudicaudatus; 150,189 , D. marsupialis; 19 each, C. 
thous, N. squamipes, and Oryzomys sp.; Loc. E: 670, 469, M. nudicaudatus; 20°, 39, D. 
albiventris; 10,19 , Bolomys sp.; 10,19, M. murina; 1C', Oryzomys sp.; Loc. F: 6107, 479, 
M. nudicaudatus; 140°, 109, D. albiventris; 10,29, P. opossum; 10,19, P. iheringi. 

REMARKS: Adoratopsylla i. intermedia has the broadest distribution of 
all the species of Adoratopsylla, occurring from Venezuela south through Ar- 
gentina on marsupials. It comprised 60 percent (329/548) of all species col- 
lected. Metachirus nudicaudatus and D. albiventris were the predominant hosts 
in Bahia. Ninety-six percent of the combined species collected in the genera 
Metachirus and Didelphis were infested with A. i. intermedia. Tipton and 
Machado-Allison (1972) reported large numbers of A. i. intermedia from 
Didelphis azarae in Venezuela, while Guimaraes (1972) found none on D. 
azarae in Bahia. It should be noted that D. azarae is synononomized with D. 
albiventris in Venezuela and with Didelphis aurita in Bahia according to 
Hershkovitz (1969) and Wilson and Reeder (1993), respectively. Thus, two 
closely related marsupials, D. albiventris and D. aurita, do not appear to har- 
bor the same common flea, A. i. intermedia. 


Rhopalopsyllidae: Gephyropsylla klagesi samuelis (Jordan and Rothschild, 1923) — Loc. A: 
307,19,H. brasiliensis; 20,19, P. longicaudatus; Loc. B: 39, P. longicaudatus. 
REMARKS: Linardi and Guimaraes (1993) erected Gephyropsylla, a sub- 
genus established by Smit (1987) under Polygenis, to full generic status. Al- 
though G. k. samuelis has been reported from Costa Rica, Panama, Venezuela, 
Colombia, Ecuador, Bolivia and Brazil, this is the first record of its presence 


Vol. 108, No. 4, September & October, 1997 293 


south of the Amazon River in the State of Amazonas. Specimens were col- 
lected from H. brasiliensis and P. longicaudatus along the corridor of the Jurua 
River. According to a study of Venezuelan populations of G. k. samuelis and 
the nominate subspecies by Machado-Allison and McLure (1963), both prefer 
species of Proechimys as hosts. 


Hectiella nitidus (Johnson, 1957) — Loc. F: 320, 410, P. iheringi; 19, O. capito. 


REMARKS: Hectiella, once considered a subgenus of Polygenis, was 
erected to full generic status by Linardi and Guimaraes (1993). Hectiella nitidus 
was found on 27 of 29 P. iheringi examined in the area of Trés Bracos, while 
only a single specimen (probable accidental association) was found on O. capito. 
Although the species was originally described from two females by Johnson 
(1957) and collected from D. marsupialis in Bahia, the male was later de- 
scribed by Linardi and Nagem (1980) from Proechimys dimidiatus in Caratinga, 
State of Minas Gerais, Brazil. Botelho, et al. (1981) also collected specimens 
from the latter area and determined species of Proechimys as the preferred 
host. In the State of Bahia, P. iheringi is clearly the preferred host for this 
species and few other flea species were collected from this host. The distribu- 
tion of H. nitidus is restricted to Brazil. 


Polygenis (Polygenis) bohlsi jordani (Costa Lima, 1937) — Loc. c: 10,19, C. aperea; Loc. E: 
207,19, D. albiventris; \C', M. nudicaudatus; 1Q_, Oryzomys sp. 

REMARKS: This subspecies was collected on the southern edge of its 
known range of northeastern Brazil. Polygenis b. jordani displays little host 
specificity, occurring on numerous host species. Guimaraes (1972) found this 
species the most ubiquitous flea from areas collected in Bahia State to north- 
eastern Brazil and indicated a close association with domestic rats. Because of 
its potential for transmitting plague between the urban and sylvatic mamma- 
lian reservoirs, this species should be considered important during plague out- 
breaks in Brazil. 


Polygenis (P.) pradoi (Wagner, 1937) — Loc. D: 10°,10Q, Bolomys sp.; Loc. F: 807,119, 
Bolomys sp.; 20,19, B. lasiurus; 1C each, D. albiventris and P. iheringi. 

REMARKS: Botelho and Linardi (1980) collected P. (P.) pradoi primarily 
from B. lasiurus, and indicated the most northern limit as Caratinga County, 
Minas Gerais, Brazil. Our collections (also found on B. /asiurus) extend the 
species range north to Corte de Pedra and Trés Bragos, Bahia. It is reported to 
range south to Buenos Aires Province, Argentina, and a disjunct population is 
also reported in southwestern Colombia by Mendez (1977). Linardi (1979) 
thinks a complex of P. (P.) pradoi along coastal Brazil needs revision. Popula- 
tions from Colombia and those extending from Brazil to Argentina should also 
be included in such a revision. 


294 ENTOMOLOGICAL NEWS 


Polygenis (P.) rimatus (Jordan, 1932) — Loc. C: 10,19, B. lasiurus; Loc. D: 40, 39, 
Oxymycterus sp., 10,19, Bolomys sp.; Loc. F: 90’, 99 , Oxymycterus sp.; 49, Bolomys sp.; 
10,19, D. albiventris; 19, B. lasiurus. 

REMARKS: These collection records constitute the northern range of this 
species occurring on numerous cricetine rodents. Polygenis rimatus females 
from Bahia State differ from those examined from Argentina. The duct of the 
spermatheca is characteristically enlarged from its proximal origin to approxi- 
mately 2/3 its length (pars dilatata). The pars dilatata has a greater diameter 
in all specimens examined from Argentina than those from Bahia, but no por- 
tion is sclerotized. The Bahia State populations of P. rimatus possess a dis- 
tinctly sclerotized U-shaped region (Fig. 1) not present in Argentina popula- 
tions examined. The sclerotization abruptly begins, and diminishes as it traverses 
towards the spermatheca. Maceration alters the position and orientation of this 
structure, but the evident rigidity of the obvious sclerotization maintains the 
consistently uniform U-shape. The posterior margin of the 7th sternite is also 
gently rounded, without lobes, undulations, or angular features. 

Many specimens of the following species borrowed from the British Mu- 
seum that might be confused with P. rimatus (P. acodontus, P. axius axius, P. 
axius proximus, P. brachinus, P. litargus, P. occidentalis occidentalis, and P. 
tripus) were examined. Although the spermathecal ducts may vary within each 
species, only P. rimatus, P. tripus and P. brachinus have an enlarged pars 
dilatata. Excluding P. rimatus, neither possessed an area of sclerotization ex- 
cept for a population of P. tripus from Salta Province, Argentina. The latter 
specimens were illustrated by Smit (1987) to separate P. tripus from all the 
species above. Of the material examined from the British Museum (Argentina, 
Bolivia and Brazil), only the Salta Province, Argentina population possessed 
this character, negating its usefulness to distinguish P. tripus from others. In 
general, little or no expansion or sclerotization of the duct of the spermatheca 
exists in P. acodontus (Argentina), P. axius axius (Argentina), P. litargus (Peru), 
and P. o. occidentalis (Brazil). These observations might be considered for 
future revisionary work on Polygenis, which is badly needed. 


Polygenis (P.) roberti roberti (Roths., 1905) — Loc. D: 50’, 49 , Oryzomys sp.; 39 , N. squamipes; 
10,19, M. demerarae; 19, D. marsupialis; 1’, Oxymycterus sp.; Loc. F: 20’, N. squamipes; 
29, 0. capito: 1Q each, D. albiventris, H. brasiliensis, and Oxymycterus sp. 

REMARKS: Linardi (1977) indicated this species to be rather host spe- 
cific, although it was sporadically collected on five mammalian genera. Speci- 
mens were most frequently collected on species of Oryzomys. 


Polygenis (P.) tripus (Jordan, 1933) — Loc. C: 40’, C. aperea; Loc. F: 19, Bolomys sp. 


REMARKS: This species was reported by Linardi, et al. (1984) and 
Cerqueira and Linardi (1977, 1981) as the most common Polygenis species in 


Vol. 108, No. 4, September & October, 1997 295 


Fig. 1. Polygenis rimatus, spermatheca and sclerotized pars dilatata. Fig. 2. P. tripus, eighth 
tergite illustrating “scale-like clear spot’ beneath seventh sternite. 


Belo Horizonte, Minas Gerais, Brazil on Bolomys lasiurus. It is also com- 
monly found on species of Oryzomys, but none were collected from either of 
these usual host species in Bahia. Cerqueira and Linardi (1976) reported a 
diagnostic feature for distinguishing P. tripus females from those of P. rimatus. 
The latter paper indicates (both in text and in photo caption) a “‘scale-like clear 
spot” in the integument of the 7th sternite, and references the same morpho- 
logical feature on the 8th sternite in the English summary. This unique charac- 
ter, keenly observed and reported by these authors, is located on neither the 
7th nor 8th sternite, but rather on the 8th tergite, occurring just beneath the 
dorso-caudal angle of the 7th sternite (Fig. 2). This clear area occurs within a 
heavily sclerotized region of the 8th tergite bearing coarse reticulations in the 
vicinity of the clear area. The clarity of this spot in specimens examined from 
Brazil, Argentina, and Bolivia was distinct in some specimens and obscure in 
others. This seemed to be a function of proper specimen preparation (not over, 
or under-clearing). 


296 ENTOMOLOGICAL NEWS 


ACKNOWLEDGMENTS 


Grateful appreciation is extended to Robert E. Lewis, Iowa State University, Nancy Adams, 
Curator of the Siphonapteran collection, U.S. National Museum of Natural History, and Theresa 
Howard, The Natural History Museum, London, England for providing specimens of Polygenis 
for study; to Paulo R. L. Lago, of the Ministry of Health (SUCAM), Brazil, who assisted in 
mammal and flea collections; and to Clive D. Jorgensen, Glenn E. Haas, and Eustorgio Méndez 
for providing helpful suggestions with the manuscript. 


LITERATURE CITED 


Botelho, J.R. and P.M. Linardi. 1980. Alguns ectoparasitos de roedores silvestresdo municipio 
de Caratinga, Minas Gerais, Brasil. I. Relagdes pulga/hospedeiro. Rev. bras. Entomol., 24(2): 
127-30. 

Botelho, J.R., P.M. Linardi, P. Williams, and R.L. Nagem. 1981. Alquns hospedeiros reais de 
ectoparasitos do municipio de Caratinga, Minas Gerais, Brasil. Mem. Inst. Oswaldo Cruz, 
Rio de Janeiro, 76(1): 57-59. 

Cerqueira, E. J. L. and P.M. Linardi. 1976 . Contribuicgao ao estudo dos Rhopalopsyllidae 
Brasileirés. O uso da mancha clara em sistematica. Bolm. Mus. Hist. nat. Univ. Fed. Minas 
Gerais (Zool.), 22: 1-4. ; 

Cergueira, E.J.L. and P.M. Linardi. 1977. Indices Pulicidianos em Belo Horizonte, Minas 
Gerais, Brazil. Cienc. Cult. Sao Paulo, 29(2): 191-93. 

Cergueira, E. J. L. and P.M. Linardi. 1981 . Relagdes hospedeiro/ parasito em Polygenis tripus 
(Siphonaptera: Rhopalopsyllidae). Bolm. Mus. Hist. nat. Univ. Fed. Minas Gerais (Zool.), 
24: 1-11. 

Guimaraes, L.R. 1973. Contribuigao a epidemiologia da peste endemica no nordeste do Brasil e 
estado da Bahia estudo das pulgas encontradas nessa regiao. Rev. Brasil. Malariol. Doengas 
Trop., 24(1/4): 95-164. 

Johnson, P.T. 1957. A classification of the Siphonaptera of South America, with descriptions of 
new species. Memoir No. 5, Entomol. Soc. Wash., 298 pp. 

Hershkovitz, P. 1969. The evolution of mammals on southern continents VI. The recent mam- 
mals of ‘he Neotropical Region. A zoogeographical and ecological review. Quart. Rev. Biol., 
44: 1-70. 

Linardi, P.M. 1977. Relagoes pulgas/roedores observado nos municipios de Salesdpolis e 
Itapetininga, SP. Bolm. Mus. Hist. nat. Univ. Fed. Minas Gerais, 23: 1-25. 

Linardi, P.M. 1979. Sobre algumas espécies de Rhopalopsyllidae (Siphonaptera) sulamericanas 
integrantes do “complexo pradoi”. Rev. bras. Entomol., 23(2): 99-106. 

Linardi, P.M. and R.L. Nagem. 1980. Descrigao do macho de Polygenis nitidus Johnson, 1957 
(Siphonaptera, Rhopalopsyllidae). Rev. Brasil. Biol., 40(1): 155-57. 

Linardi, P.M., J.R. Botelho, D.P. Neves, and H.C. Cunha. 1984. Sobre alguns ectoparasitos de 
roedores silvestres de Belo Horizonte, MG. Rev. Brasil. Biol., 44(2): 215-19. 

Linardi, P.M. and L.R. Guimaraes. 1993. Systematic review of genera and subgenera of 
Rhopalopsyllinae (Siphonaptera: Rhopalopsyllidae) by phenetic and cladistic methods. J. 
Med. Entomol., 30(1): 161-70. 

Machado-Allison, C.E. and M.T. McLure. 1963. Notas sobre Rhopalopsyllidae III. 
Consideraciones sobre las subspecies de Tiamastus (Gephyropsylla) klagesi (Siphonaptera, 
Rhopalopsylloidea). Acta Biol. Venezuelica, 3(27): 421-36. 

Méndez, E. 1977. Mammalian-Siphonapteran associations, the environment, and biogeography 
of mammals of southwestern Colombia. Quaest. Entomol., 13(2): 91-182. 

Smit. F.G.A.M. 1987. An illustrated catalogue of the Rothschild collection of fleas (Siphonaptera) 
in the British Museum. Volume VII: Malacopsylloidea (Malacopsyllidae and Rhopalo- 
psyllidae). Oxford Univ. Press, 380 pp., Plates V. 

Tipton, V.J. and C.E. Machado-Allison. 1972. Fleas of Venezuela. Brigham Young University 
Sci. Bull., Biol. Ser., 17(6): 1-115. 

Wilson, D.E. and D.M. Reeder. 1993. Mammal species of the world, a taxonomic and geo- 
graphic reference. 2nd ed., Smithson. Inst. Press, Wash., 1206 pp. 


Vol. 108, No. 4, September & October, 1997 297 


A NEW DISTRIBUTIONAL RECORD FOR 
HAIDEOPORUS TEXANUS (COLEOPTERA: 
DYTISCIDAE), A STYGOBIONTIC BEETLE FROM 
THE EDWARDS AQUIFER, TEXAS! 


David E. Bowles2, Ruth Stanford 


ABSTRACT: Haideoporus texanus recently was collected from the Edwards Aquifer at Comal 
Springs, New Braunfels, Texas. This is a significant new distributional record for this rare subter- 
ranean beetle formerly known only from an artesian well in San Marcos, Texas, suggesting the 
species may be more widely distributed in the Edwards Aquifer than previously thought. 


The Edwards Aquifer located in south-central Texas has the highest known 
diversity of endemic subterranean aquatic species in the world (Longley 1981). 
Presently, there are 44 endemic, stygobiontic species reported from this aqui- 
fer (Bowles and Arsuffi 1993, Spangler and Barr 1995). Spangler (1996) de- 
fined stygobiontic species as those found almost exclusively in all their devel- 
opmental stages in subterranean aquatic habitats and exhibiting various mor- 
phological adaptations associated with such habitats. Adaptations to subterra- 
nean habitats may include reduction or loss of eyes, loss of hindwings in some 
insects, greatly elongated sensory setae on the body and appendages, and de- 
creased pigmentation and sclerotization of the integument (Young and Longley 
1976). Represented among the primarily crustacean fauna of the Edwards Aqui- 
fer are three beetles (Coleoptera) including Haideoporus texanus Young and 
Longley (Dytiscidae: Hydroporinae), Comaldessus stygius Spangler and Barr 
(Dytiscidae: Bidessini), and Stygoparnus comalensis Barr and Spangler 
(Dryopidae). Stygobiontic beetles occurring in the United States have been 
reviewed by Barr and Spangler (1992), Larson and LaBonte (1994), Longley 
and Spangler (1977), Spangler and Barr (1995), and Young and Longley (1976). 

Haideoporus texanus previously was known only from an artesian well 
located on the campus of Southwest Texas State University in San Marcos, 
Texas (Young and Longley 1976, Longley and Spangler 1977). The well was 
drilled into the San Marcos pool of the Edwards Aquifer in the late 1800’s and 
has been the source for most of the collections of stygobiontic species from 
the aquifer. Recent collections from natural spring orifices at Comal Springs 
in New Braunfels, Texas, also yielded examples of H. texanus. Comal Springs, 


1 Received January 30, 1997. Accepted March 4, 1997. 


2 Texas Parks and Wildlife Department, 300 C. M. Allen Parkway, Building B, San Marcos, TX 
78666. 


3 United States Fish and Wildlife Service, Ecological Services, 10711 Burnet Road, Suite 200, 
Austin, TX 78758. 


ENT. NEWS 108(4): 297-299, September & October, 1997 


298 ENTOMOLOGICAL NEWS 


the largest spring system in Texas, issues from the San Antonio pool of the 
Edwards Aquifer approximately 24 km south of the artesian well in San Marcos. 
This new collection record shows that H. texanus is more widely distributed in 
the Edwards Aquifer than previously thought. However, the complete extent 
of this species’ distribution in the aquifer remains unknown. The Edwards 
Aquifer extends for approximately 282 km from Brackettville in Kinney County 
northward to Georgetown in Williamson County and varies from eight to 48 
km in width (Klemt et al. 1979, Barker et al. 1994). Numerous springs and 
wells occur in this broad area (Brune 1981) and H. texanus eventually may be 
found at other locations fed by the aquifer. 

Several studies have shown that the Edwards Aquifer is being rapidly de- 
pleted due to excessive groundwater pumping (Longley 1992), and water quality 
is threatened by the encroachment of poor quality, saline water (Perez 1986, 
Barker et al. 1994) and contamination from the surface. Because of these threats, 
some of the unique and diverse aquatic fauna inhabiting the aquifer is at risk of 
extinction (Bowles and Arsuffi 1993). Recently, the United States Fish and 
Wildlife Service published proposed rules for listing some of these species as 
endangered though H. texanus was not among them (Stanford and Shull 1995). 

Three adult specimens were collected by placing Wildco® stream drift- 
nets (363 um mesh) at various orifices at Comal Springs for 24 hours on vari- 
ous sampling dates during 1993-1994. Samples were preserved in the field 
with 70% isopropyl alcohol and the beetles were later removed and pinned. 
Specimens are deposited in the National Museum of Natural History, Wash- 
ington, DC (NMNH), the Essig Museum of Entomology, Berkeley, California 
(EME), and the Texas A&M University Entomological Collection, College 
Station, Texas (TAMU). Immature stages were not collected during this study. 


SPECIMENS EXAMINED: United States, Texas, Comal County, New Braunfels, Landa Park, 
Comal Springs, spring-run 2, D. E. Bowles, and R. Stanford, 24-hour drift net, Oct 1993, 1 CO 
(NMNH); same data, but spring run 3, 26 Jan 1994, 1 9 (TAMU); same data, but Apr 1994, 1 O 
(EME). 


ACKNOWLEDGMENTS 


We thank Paul Spangler, Cheryl Barr and Sharon Jasper for reviewing an earlier draft of 
this paper and for confirming our identification of the beetle. 


LITERATURE CITED 


Barker, R.A., P. W. Bush, and E. T. Baker, Jr. 1994. Geologic history and hydrogeologic set- 
ting of the Edwards-Trinity aquifer system, West-central Texas. U.S. Geol. Surv., Water-Res. 
Investig. Rep. 94-4039, Austin, Texas. 51 pp. 

Barr, C. B., and P.J. Spangler. 1992. A new genus and species of stygobiontic dryopid beetle, 
Stygoparnus comalensis (Coleoptera: Dryopidae), from Comal Springs, Texas. Proc. Biol. 
Soc. Wash. 105:40-54. 


Vol. 108, No. 4, September & October, 1997 299 


Bowles, D. E., and T. L. Arsuffi. 1993. Karst aquatic ecosystems of the Edwards Plateau region of 
central Texas, USA: a consideration of their importance, threats to their existence, and efforts 
for their conservation. Aquatic Conservation: Marine and Freshwater Ecosystems 3:317-329. 

Brune, G. 1981. Springs of Texas, vol. 1. Branch-Smith, Inc., Ft. Worth, Texas. 566 pp. 

Klemt, W. B., T. R. Knowles, G. R. Elder, and T. W. Sich. 1979. Ground-water resources and 
model applications for the Edwards (Balcones Fault Zone) Aquifer in the San Antonio re- 
gion, Texas. Texas Dept. Water Res. Rep. 239: 1-88. , 

Larson, D. J., and J. R. LaBonte. 1994. Stygoporus oregonensis, a new genus and species of 
subterranean water beetle (Coleoptera, Dytiscidae, Hydroporinae) from the United States. 
Coleopt. Bull. 48:371-379. 

Longley, G. 1981. The Edwards Aquifer: Earth’s most diverse groundwater ecosystem? Internat. 
J. Speleology 11:123-128. 

Longley, G. 1992. The subterranean aquatic ecosystem of the Balcones Fault Zone Aquifer in 
Texas — threats from overpumping. pp. 291-300. In: J. A. Stanford and J. J. Simons [eds.], 
Proc. First Intern. Conf. on Groundwater, Am. Water Res. Assoc., Bethesda, Maryland. 

Longley, G., and P. J. Spangler. 1977. The larva of anew subterranean water beetle, Haideoporus 
texanus (Coleoptera: Dytiscidae: Hydroporinae). Proc. Biol. Soc.Wash. 90:532-535. 

Perez, R. 1986. Potential for updip movement of saline water in the Edwards Aquifer, San Anto- 
nio, Texas. U.S. Geol. Surv., Water-Res. Investig. Rep. 86-4032, Austin, Texas. 21 pp. 

Spangler, P. J. 1996. Four new stygobiontic beetles (Coleoptera: Dytiscidae; Noteridae; Elmidae). 
Insecta Mundi 10:241-259. 

Spangler, P. J., and C. B. Barr. 1995. A new genus and species of stygobiontic dytiscid beetle, 
Comaldessus stygius (Coleoptera: Dytiscidae: Bidessini) from Comal Springs, Texas. Insecta 
Mundi 9:301 -308. 

Stanford, R., and A. Shull. 1995. Endangered and threatened wildlife and plants; proposal to 
list three aquatic invertebrates in Comal and Hays counties, Texas, as endangered. Federal 
Register 60:29537-29543. 

Young, F. N., and G. Longley. 1976. A new subterranean aquatic beetle from Texas (Coleoptera: 
Dytiscidae-Hydroporinae). Ann. Entomol. Soc. Am. 69:787-792. 


(Continued from page 289) 


At 7:00 A.M. the following morning (official sunrise had occurred at 6:50 A.M.) I returned 
to the scene and found the interment had been completed. There was no mound or any evidence 
of a burial except that the surface of the soil had been perfectly smoothed over and was even with 
the soil surface adjacent to the site of burial. 

In a previous study (Shubeck et. al., 1981), it had been noted that seasonal activity for N. 
tomentosus virtually ended in October and November. It is possible then, this burial provided 
food and protection for overwintering adult and/or immature individuals. Another previous study 
showed also that this species of Nicrophorus is active diurnally (Shubeck 1971). In the observa- 
tions described in this scientific note, I noted the following: (1) the cruising, searching, and 
“calling” activity of N. tomentosus apparently can continue up to the very end of the diurnal 
period, and (2) the interment of the cadaver can be completed nocturnally. 


LITERATURE CITED 


Milne, L. J. and M. J. 1944. Notes on the behavior of burying beetles (Nicrophorus spp.). 
J. N. Y. Entomol. Soc. 52:311-327. 

Pukowski, E. 1933. Okologische untersuchungen an Necrophorus F. Z. Morph. Okol. Tiere. 
27:518-586. 

Shubeck, P. P. 1971. Diel periodicities of certain carrion beetles (Coleoptera: Silphidae). Coleopt. 
Bull. 25:41-46. 

Shubeck, P. P., N. M. Downie, R. L. Wenzel, and S. B. Peck. 1981. Species composition and 
seasonal abundance of carrion beetles in an oak-beech forest in the Great Swamp National 
Wildlife Refuge (N.J.). Entomol. News. 92:7-16. 


300 ENTOMOLOGICAL NEWS 


SYNAPTONECTA ISSA (HETEROPTERA: 
CORIXIDAE), FIRST NEW WORLD RECORD OF AN 
ASIAN WATER BUG IN FLORIDA! 


J. T. Polhemus2, R. P. Rutter3 


ABSTRACT: The Asian corixid Synaptonecta issa was first discovered in south Florida in Octo- 
ber 1993. A single male was taken in a dipnet sample from an herbaceous depressional wetland 
in St. Lucie County, Florida. Its presence in Florida may be the result of commercial trade in 
aquarium plants. Despite state regulation and federal inspection, there is ample opportunity for 
the importation and subsequent release of foreign aquatic insects. 


Synaptonecta issa (Distant) was discovered in Florida when a single male 
was taken in a dipnet sample from an herbaceous depressional wetland in St. 
Lucie County, south Florida, in October 1993. The wetland was one of two, 
measuring 3.6 and 7.8 h, that had been constructed in the spring of 1990. Both 
wetlands were disked with donor mulch from nearby wetlands to help estab- 
lish aquatic vegetation, and some shoreline was made contiguous with natural 
wetlands to promote water and propagule exchange. Both wetlands were quali- 
tatively sampled again in April 1994, but no S. issa were found; the only corixid 
found was Sigara bradleyi (Abbott). Sampling was conducted again in Octo- 
ber 1995, at which time several adults and immatures of S. issa were collected 
from two locations in the wetland where the species was first found. Water 
depth was 51 cm, and predominant vegetation was maidencane (Panicum 
hemitomon), umbrella grass (Fuirena scirpoidea), fragrant water lily (Vymphaea 
odorata), and the filamentous alga Spirogyra sp. No specimens were recorded 
from the other constructed wetland, although it was only 46 m distant and had 
periodically experienced exchange of surface water. 

In September 1994, a single adult female was collected in a dipnet sample 
from the shoreline of oligotrophic Lake Viola in Highlands County, south cen- 
tral Florida. The predominant vegetation in the sample area was torpedograss 
(Panicum repens), spikerush (Eleocharis sp.), and associated filamentous al- 
gae. Values for selected physiochemical parameters measured at the time of 
sampling were: water temperature 30° C, specific conductance 227 umhos/ 
cm, pH 7.8 (8.2 in March 1994), and alkalinity 34 mg/l. No other corixids 
were collected in the sample. 

A multitude of other habitats in south Florida, with varying water quality 


! Received January 9, 1997. Accepted February 23, 1997. 
2 Colorado Entomological Museum, 3115 S. York St., Englewood, CO. 80110. 


3 Florida Department of Environmental Protection, South District, 2295 Victoria Avenue, Fort 
Myers, Florida 33901. 


ENT. NEWS 108(4): 300-304, September & October, 1997 


Vol. 108, No. 4, September & October, 1997 301 


conditions including rivers, streams, canals, lakes, ponds, and wetlands, have 
been sampled for aquatic macroinvertebrates since 1980 as part of the Florida 
Department of Environmental Protection’s water quality monitoring program, 
but no other specimens of S. issa have been collected. 


Synaptonecta issa (Distant) 


Micronecta issa Distant, 1910:350 (Holotype, macropterous male, Travancore, S. India); Lundblad, 
1933:100 (systematics; distribution); Chen, 1960:102 (key to species); Fernando & Leong, 
1963:548 (dispersal); Leong, 1966:84 (systematics; distribution); Wrobleswki, 1968:775 
(checklist); Fernando & Cheng, 1974:37 (distribution). 

Synaptonecta breddini Lundblad, 1933:109 (Holotype, brachypterous male, Buitenzorg, Java); 
Hutchinson, 1940:398 (synonymy). 

Synaptonecta issa,; Hutchinson, 1940:346, 348, 398, pl. XXXI, figs. 411-415 (systematics; dis- 
tribution; brachypterous form from Burma); Wroblewski, 1972:4, 14 (systematics; distribu- 
tion; redescription, macropterous form); Jansson & Meyer-Rochow, 1990:328 (sound pro- 
duction; distribution). 

Micronecta (Synaptonecta) issa; Wrobleswki, 1967:243 (systematics; distribution). 


Figure |. Synaptonecta issa (Distant). Female, dorsal habitus. Scale bar= 1 mm. 


302 ENTOMOLOGICAL NEWS 


Macropterous females are relatively common in southeast Asia, and are 
often taken in light traps (Fernando & Leong 1963), but specimens of the brac- 
hypterous morph, mostly males, are not common in collections. Wroblewski 
(1967:248) states that, with respect to light trap material, the “immense domi- 
nance of females in ... Synaptonecta issa (Dist.) is easy to explain. The males 
are mostly, or almost exclusively, brachypterous, not able to fly.” The brachypter 
differs radically in appearance from the more common macropter. Distant (1910) 
provided a habitus illustration of the macropterous morph, but the brachypter- 
ous morph, with coleopteroid hemelytra, has never been adequately illustrated 
(see Fig. 1). Lundblad (1933) provided a line drawing of the left hemelytron 
and some other structures (of the synonymous S. breddini). Even when the two 
morphs are compared side by side, it is very easy to believe that they represent 
different species, as Lundlbad (1933) did when he described the synonym, 
Synaptonecta breddini. One of us (JTP), when confronted with the first brac- 
hypterous specimens from Florida, thought it might represent an undescribed 
New World genus (which would be truly remarkable, from well collected 
Florida), but recently he found similar brachypterous specimens in a sample 
from Thailand (Phattalung Province; Sites and Nichols leg.). The male geni- 
talic morphology is diagnostic, as is the low elliptical carina on the vertex of 
the head. 


DISTRIBUTION AND ECOLOGY 


Synaptonecta issa is a widespread species, naturally distributed from Sri 
Lanka and India to Java, Malaysia and Thailand, and recently accidentally 
introduced into New Zealand (Jansson & Meyer-Rochow 1990). 

Jansson & Meyer-Rochow (1990) suggest that the most plausible mode of 
introduction into New Zealand was through eggs attached to aquatic aquarium 
plants brought from Singapore, where the species is common. We believe it 
likely that the species was brought to Florida in the same way. Because the 
Singapore suppliers of aquatic plants undoubtedly ship to many parts of the 
world, this species can be expected to turn up in aquariums anywhere, and in 
the wild in warm climates. 

Aquatic plants imported from southeast Asia enter Florida weekly at Mi- 
ami International Airport, and can include the following species (list provided 
by Brad McLane, State of Florida Aquatic Plant Association, August 1995): 


Singapore Cabomba caroliniana, Crinum thaianum, Echinodorus blerhi, Hygrophila 
difformis, Microsorium pteropus. Vallisneria sp. (Corkscrew) 


Thailand Anubias nana, Barclaya longifolia, Cabomba caroliniana, Crinum thaia- 
num, Cryptocoryne balansae, C. retrospiralis. Vesicularia dubyana 


Sri Lanka Aponogeton crispus, Cryptocoryne wendtii (red), Nymphaea stellata 


Madagascar Aponogeton boivinianus, A. capuroni, A. fenestralis, A. longiplumulosus, 
A. ulvaceus 


Vol. 108, No. 4, September & October, 1997 303 


Most imported plants are nursery grown, but not under quarantine condi- 
tions. Growers dam up streams to allow water to flood their crops. Plants col- 
lected from the wild are generally inferior in appearance and are hence less 
marketable; these plants are trimmed and propagated in nurseries before ex- 
portation (Brad McLane, pers. comm.). USDA inspection of incoming plants 
is limited, as only about 2% of each species is visually inspected for the pres- 
ence of potential pests (e.g., Lepidoptera) and parasite vectors of public health 
concern (e.g., Gastropoda). If a potential pest is found, the entire species lot is 
fumigated with methyl bromide (Tad Dobbs, pers. comm.). There is no inten- 
tional control of nonpest/nonvector species. Synaptonecta issa in any life stage 
would go unnoticed because of its small size, but even if found it would go 
untreated because aquatic hemipterans are presumed to be predatory (although 
most corixids are not) and therefore not of quarantine concern (Tad Dobbs, 
pers. comm.). 

Both the Florida Department of Agriculture and Consumer Services, and 
the Florida Department of Environmental Protection (FDEP) have regulatory 
control over the importation, cultivation, and distribution of aquatic plants in 
Florida. Rules in chapter 62C-52 state that “No person, except aquatic plant 
nurseries regulated by the Department of Agriculture and Consumer Services, 
shall engage in any business activity involving the importation, transportation, 
sale or possession of any aquatic plant species without a permit issued by the 
department....” Florida maintains an Aquatic Plant Locator, which as of June 
1993 (Rogers 1993) listed 250 businesses that supply aquatic plants, imported 
and native. Despite state regulation and federal inspection, there is opportu- 
nity for the importation and subsequent release of aquatic insects. 

This is not the first documentation of an Asiatic aquatic insect becoming 
established in south Florida. Paulson (1978) recorded the presence of an Asi- 
atic dragonfly, Crocothemis servilia (Drury), from Dade County, Florida in 
1977, which by 1993 had spread to six additional counties (Daigle and Rutter 
1984), and is now established in Hawaii as well (Polhemus 1995). 


ACKNOWLEDGMENTS 


Our compliments to Ed Emmons for collecting, as far as we know, the first Florida specimen 
of Synaptonecta issa, and also our thanks for providing the monitoring reports for the constructed 
wetlands. Brad McClane, Tad Dobbs, and Rob Kipker were helpful in providing information on 
the importation and regulation of aquatic plants. We thank R. L. Sites and B. Nichols for making 
available specimens from Thailand. R. T. Schuh and D. A. Polhemus provided constructive re- 
views of the manuscript. JTP carried out this research as a faculty affiliate of the Entomology 
Department, Colorado State University, Fort Collins. 


LITERATURE CITED 


Chen, L. C. 1960. A study of the genus Micronecta of India, Japan, Taiwan and adjacent regions 
(Heteroptera - Corixidae). J. Kans. Entomol. Soc. 33:99-118. 


304 ENTOMOLOGICAL NEWS 


Daigle, J. J. & R. P. Rutter. 1984. New county records for Crocothemis servilia (Dru.) from 
Florida, United States (Anisoptera: Libellulidae). Not. Odonatol. 2: 63. 

Distant, W. L. 1910. The fauna of British India including Ceylon and Burma. Rhynchota 
(Heterotpera - Appendix). Taylor & Francis, London, vol. 5, xii + 362 pp 

Fernando, C. H. & L. Cheng. 1974. A preliminary study on the fauna and distribution of aquatic 
Hemiptera in Malaya and Singapore. Fed. Mus. J. 19:21 -44. 

Fernando, C. H. & C. Y. Leong. 1963. Miscellaneous notes on the biology of Malayan Corixidae 
(Hemiptera: Heteroptera) and a study of the life histories of two species, Micronecta 
quadristrigata Bredd. and Agraptocorixa hyalinipennis (F.). Ann. Mag. Nat. Hist. (13) 6:545- 
558. 

Hutchinson, G. E. 1940. A revision of the family Corixidae of India and adjacent regions. Trans. 
Conn. Acad. Arts Sci. 33:339-476, pls. I-XXXVI. 

Jansson, A. & V. B. Meyer-Rochow. 1990. Sound production in Synaptonecta issa (Heteroptera: 
Corixidae, Micronectinae) - an Asian bug that turned up in a New Zealand aquarium. Appl. 
Entomol. Zool. 25:328-331. - 

Leong, C. Y. 1966. A taxonomic study of the Malayan Corixidae (Hemiptera - Heteroptera) with 
the description of Micronecta malayana sp. nov. Bull. Nat. Mus. Singapore 33:83-90. 

Lundblad O. 1933. Zur Kenntnis der aquatilen und semi aquatilen Hemipteren von Sumatra, 
Java und Bali. Arch. Hydrobiol., 1933, Suppl. Bd. 12, Tropische Binnengewasser 4: 1-195, 
263-498. 

Paulson, D. R. 1978. An Asiatic dragonfly Crocothemis servilia (Drury) established in Florida 
(Anisoptera: Libellulidae). Not. Odonatol. 1: 9-10. 

Polhemus, D. A. 1995. New Heteroptera and Odonata (Insecta) records and range extensions in 
the Hawaiian Islands. In: N. L. Evenhuis and S. E. Miller (eds.), Records of the Hawaii 
Biological Survey for 1994. Part 2: Notes. Bishop Mus. Occ. Pap., 42: 42-43. 

Rogers, M. 1993. Florida aquatic plant locator. Florida Department of Agriculture and Con- 
sumer Services, Tallahassee, FL. 

Wroblewski, A. 1967. Further notes on Micronectinae from Viet-Nam (Heteroptera, Corixidae). 
Bull. Entomol. Pologne 37:229-251. 

Wroblewski, A. 1968. Notes on Oriental Micronectinae (Heteroptera, Corixidae). Bull. Entomol. 
Pologne 38:753-779. 

Wroblewski, A. 1972. Further notes on Micronectinae from Ceylon (Heteroptera, Corixidae). 
Bull. Entomol. Pologne 42:3-52, 3 pis. 


(Continued from page 258) 


Dr. Edwards described an elegant developmental model used to determine the distribution 
of temperatures experienced by adult M. grandii in both Delaware and Korea. Though there were 
significant differences in the temperature distributions between populations, the differences in 
distributions between generations within populations were greater. Laboratory experiments showed 
no difference in the effects of temperature on ovipositional performance between M. grandii 
from Delaware and Korea. This result suggests no adaptation with respect to this fitness trait. At 
both locations, however, M. grandii would be expected to have greater fitness (ovipositional 
performance) in the first generation than in the second. Dr. Edwards is continuing to research 
these questions. 

Several items of entomological news were related, including publication in the Philadelphia 
Inquirer of a report that the last remaining habitat for Regal Fritillary (Speyeria idulia) in Penn- 
sylvania is threatened, and a report from Dave Wright on the earliest ever adult Spring Azure 
(Celastrina lucia of authors) in New Jersey, Feb. 27, in Cape May. 

W. J. Cromartie 
Corresponding Secretary 


Vol. 108, No. 4, September & October, 1997 305 


NEW STATE AND U. S. RECORDS AND OTHER 
DISTRIBUTIONAL NOTES FOR 
HETEROPTERA (INSECTA)! 


J. T. Polhemus2 


ABSTRACT: Buenoa pallens is recorded for the first time in the United States, from Val Verde, 
Co., TX. Fifteen new state records are established, and distributional data are given for six addi- 
tional species. 


The following represent range extensions, or records of seldom-collected 
but previously reported species for the United States. Voucher specimens are 
found in the J. T. Polhemus Collection, Colorado Entomological Museum 
(JTPC), unless otherwise noted; a significant amount of material reported here 
is from the Texas A & M University collection. These state or U. S. records 
were not included in the Catalog of Heteroptera of North America (Henry & 
Froeschner 1988). Several of these species are rather common in the states 
listed and some were collected over 50 years ago; but to my knowledge they 
have never been reported in the literature. 

The collection location (CL) numbers following localities refer to codes 
used by JTP to reference ecological data. Unless otherwise stated, the known 
distributions are based on Henry and Froeschner (1988). 


NEW U. S AND STATE RECORDS 


Enicocephalidae 


Systelloderes biceps (Say) New Record for Kansas 
This species is widespread in the eastern and southern U. S. 


Material examined: KANSAS, Riley Co.: 1 male, 29 May 1968, R. L. Bertwell (University 
of Wyoming, Laramie). 


Lygaeidae 
Ischnodemus falicus (Say) New Record for Colorado 


Common over the eastern half of the U. S., this is the westernmost record 
for this species. 


Material examined: COLORADO, Jefferson Co.: many males & females, marsh, Chatfield 
State Park, 14 May 1992, J. T. Polhemus 


1 Received January 9, 1977. Accepted February 23, 1977. 
2 Colorado Entomological Museum, 3115 S. York St., Englewood, Colorado, 80110, USA. 


ENT. NEWS 108(4): 305-310, September & October, 1997 


306 ENTOMOLOGICAL NEWS 


Reduviidae 
Reduvius personatus Linnaeus New Record for Colorado 
This cosmopolitan species is widespread in the U. S. 


Material examined: COLORADO, Arapahoe Co.: males & females, Englewood, in home 
or at light at home, J. T. Polhemus, on the following dates: 13 July 1984; 21 June 1986; 5 June 
1988; 23 June 1989; 16 Dec. 1989 (nymph found in home, raised to adult over 18 months); 9 July 
1990; 11 July 1993; 2 June 1994; 13 June 1994; 16 July 1994; 6 Aug. 1994; 2 July 1996; 14 July 
1996, between timbers. 


Belostomatidae 
Belostoma bakeri Montandon New record for Colorado 
Previously known from Kansas, New Mexico, and other states further west. 


Material examined: COLORADO, Las Animas Co.: many males & females, pond, head of 
Carrizo Creek, S. of Kim, CL 2729, 26 Aug. 1992, J. T. Polhemus. 


Hebridae 
Hebrus comatus Drake & Harris New record for Colorado 


Previously known from Kansas, New Mexico, and Texas. 


Material examined: COLORADO, Yuma Co.: 2 females, Chief Creek, nr. Wray, el. 1098 
m., CL 2690, 6 June 1992, J. T. Polhemus & C. N. McKinnon; 2 males, 3 females, same data as 
for preceding record, but on seeps at head of Chief Creek, west of Wray, el. 1113 m., CL 2691. 


Hydrometridae 
Hydrometra martini Kirkaldy New records for Nevada and Washington 


Common in the eastern U. S. as far west as Arkansas, Oklahoma, and Texas, 
and previously known but scarce in Arizona, California, Colorado, Idaho, and 
Oregon. The occurrence in Oklahoma was recorded by Schaefer & Drew (1967) 
but overlooked in the Catalog of Heteroptera of North America (Smith 1988). 


Material examined: NEVADA, Lincoln Co.: | male, Crystal Spring, S. of Hiko, el. 1234 m, 
CL 2710, 20 July 1992, J. T. & D. A. Polhemus. WASHINGTON, Grant Co.: 1 male, 1 female, 
Soda Lake nr. O’Sullivan Dam, Columbia Nat. Wildlife Ref., 10 June 1973, D. Corredor, Yakima 
Co., 1 male, 1 female, 8 mi. SW of Tieton RS, Snoqualmie NF, Bear Cr., 11-12 June 1973, 
M. Jackson. 


Mesoveliidae 


Mesovelia mulsanti White New records for Arizona, 
Colorado, and Nebraska 


This species is very widespread in the Western Hemisphere. 


Vol. 108, No. 4, September & October, 1997 307 


Material examined: ARIZONA, Mohave Co.: 1 male, Beaver Lodge, 13 May 1961, J. T. 
Polhemus. COLORADO, Jefferson Co.: 1 female, Lakewood, 14 May 1992, J. T. Polhemus; 
Yuma Co.: 2 males, 2 females, Hale Ponds, 6 mi. E. Hale, CL 1920, 10 July 1987, J. T. Polhemus. 
NEBRASKA, Douglas Co.: 1 female, Omaha, 9 July 1952, J. T. Polhemus.. 


Mesoveiia cryptophyla Hungerford New record for Texas 


Previously known from the eastern U. S., as far west as Iowa, Michigan, 
Mississippi, and Oklahoma. 


Material examined: TEXAS, Collin Co.: 2 males, 2 females, McKinney, Heard Pond, 7 
May 1991, T. Vasarhelyi. 


Naucoridae 
Ambrysus thermarum La Rivers New record for Utah 


This species occurs in Arizona, New Mexico, and three widely separated 
regions of Colorado, including West Creek, near Gateway, on the eastern side 
of the La Sal Mountains (Polhemus, unpubl. data). Mill Creek originates on 
the west side of the La Sal Mountains and flows into Moab, Utah. The record 
for Ambrysus woodburyi from Moab (La Rivers 1951) is a misidentification of 
A. thermarum. 


Material examined: UTAH, Grand Co.: many males & females, Moab, Mill Creek, CL 
2907, 5 Aug. 1993, J. T. Polhemus. 


Notonectidae 
Buenoa margaritacea Torre Bueno New record for Iowa 
Common and widespread in the United States and northern Mexico. 


Material examined: IOWA, Boone Co.: 1 female, Ledges State Park, 28 March 1946, J. T. 
Polhemus; 4 males, 6 females, Ledges State Park, 22 Sept. 1960, J. C. Schaffner (TAMU); Clay 
Co.: 1 female, E. of Peterson, 2 Aug. 1975, J. T. Polhemus; Story Co.: 1 female, Ames, 20 Sept. 
1945, J. T. Polhemus; 3 males, 5 females, Lake Comar, 18 June 1945, J. T. Polhemus. 


Buenoa pallens (Champion) New record for Texas and U. S. 


Anisops pallens Champion, 1901:374. 
Buenoa pallens Kirkaldy, 1904:121. 


Common from central Mexico and the West Indies southward to Chile and 
Peru (Truxal 1953). 


Material examined: TEXAS, Val Verde Co.: many males & females, Dolan Falls Ranch, 
The Nature Conservancy of Texas, pools in side canyon to Dolan Creek, 20 April 1996, J. T. 
Polhemus; same data as for preceding record, but from Leon Spring, 21 April 1996, J. T. Polhemus. 


308 ENTOMOLOGICAL NEWS 


Notonecta lunata Hungerford New record for Indiana 


Widespread in eastern Canada and the northeastern U. S., as far west as 
Michigan. 


Material examined: INDIANA, Monroe Co.: many males & females, Lake Beanblossom, 
7-25 March 1961, J. C. Schaffner; Brown Co.: 1 male, 1 female, Yellowwood S. F., 28 March 
1961, J. C. Schaffner. 


Notonecta undulata Say New record for Colorado 


Previously known from states surrounding Colorado. 


Material examined: COLORADO, Arapahoe Co.: | male, | female, spring fed pond along 
Highline Canal Trail, Cherry Hills Village, 27 May 1996, J. T. Polhemus; Saguache Co.: 2 males, 
1 female, Russell Lakes SWA, 7-8 Aug. 1994, R. Durfee (CSUC). 


Notonecta unifasciata unifasciata Guérin New record for Colorado 
Previously known from California, New Mexico, and Mexico. 


Material examined: COLORADO, Saguache Co.: 2 males, 4 females, Russell Lakes SWA, 
7-8 Aug. 1994, R. Durfee (CSUC). 


Saldidae 
Salda obscura Provancher New record for Utah 


This is a boreal species that, in the southernmost part of its range, inhabits 
relatively dry biotopes in the mountains of the midlatitude western U. S. It was 
previously known from Canada, a few northern states of the U. S., the Rocky 
Mountains of Colorado, and the Ruby Mountains of Nevada (Polhemus, unpubl. 
data). 


Material examined: UTAH, Grand Co.: many males & females, La Sal Mtns., Gold Basin, 
off Geyser Pass Rd., 6 August 1993, J. T. Polhemus, S. Kaneno & M. Sat6. 


OTHER DISTRIBUTIONAL RECORDS 
Gelastocoridae 
Gelastocoris rotundatus Champion Additional record for Texas 


Common in Mexico and Arizona, and occurs in California. To my knowI- 
edge, this is only the third record for Texas, and the first for Val Verde Co. 


Material examined: TEXAS, Val Verde Co.: many males & females, Dolan Falls Ranch, 
The Nature Conservancy of Texas, Dolan Creek, 20 April 1996, J. T. Polhemus. 


Vol. 108, No. 4, September & October, 1997 309 


Notonectidae 
Buenoa marki Reichart Additional record for Florida 


Previously known only from the type series taken in Everglades National 
Park. 


Material examined: FLORIDA, Collier Co.: 9 males, 10 females, ponds in cypress ham- 
mock nr. Turner River at Hwy. 41, 15 Nov. 1989, D. A. Polhemus. 


Buenoa platycnemis (Fieber) Additional record for Texas 


Rather common from Mexico and the West Indies southward to Brazil and 
Peru, and was previously known in the U. S. from a few localities in Texas and 
one in the Florida Keys (Truxal 1953). 


Material examined: TEXAS, Cameron Co.: | male, 1 female, Sabal Palm Grove Sanct., 16 
Oct. 1993, at UV light, Blackmon, Quinn & Riley (TAMU). 


Buenoa speciosa Truxal Additional record for Texas 


Previously known from Mexico and the Davis Mountains of Texas (Truxal 
1953). 


Material examined: TEXAS, Presidio Co.: 2 females, Big Bend Ranch S. N. A., La Sauceda 
(Hdatrs.), 27 April 1991, J. C. Schaffner (TAMU). 


Buenoa uhleri Truxal Additional record for Texas 


Widespread in Mexico, and previously known from one locality each in 
California and Texas (Truxal 1953). 


Material examined: TEXAS, Presidio Co.: 1 male, Big Bend Ranch S. N. A., La Sauceda 
(Hdatrs.), 27 April 1991, J. C. Schaffner (TAMU). 


Ochteridae 
Ochterus banksi Barber Additional record for Texas 


Widespread in the eastern and southern United States, including eastern 
Texas. This is the westernmost record for this species. 


Material examined: TEXAS, Val Verde Co.: 2 males, Dolan Falls Ranch, The Nature Con- 
servancy of Texas, springs along Dolan Creek, 20 April 1996, J. T. Polhemus 


ACKNOWLEDGMENTS 


I am grateful to J. A. Slater, Univ. Connecticut, Storrs, for identification of Ischnodemus 
falicus. 1 am also indebted to the following for the gift or loan of specimens: T. Vasarhelyi, 
Budapest; Scott Shaw, University of Wyoming, Laramie, WY, Boris Kondratieff, Colorado State 
University, Fort Collins, CO (CSUC); J. C. Schaffner and E. G. Riley, Texas A. & M. University, 
College Station, TX (TAMU). I thank D. A. Polhemus and R. T. Schuh for their reviews. 


310 ENTOMOLOGICAL NEWS 


LITERATURE CITED 


Champion, G. C. 1897-1901. Insecta: Rhynchota (Hemiptera - Heteroptera) 2. In: F. D. Godman 
& O. Salvin (eds.), Biologia Centrali Americana (Zoologia), London, xvi + 416 pp. 

Henry, T. J. & R. C. Froeschner. 1988. Catalog of the Heteroptera, or True Bugs, of Canada and 
the Continental United States, E. J. Brill, Leiden, xix + 958 pp. 

Kirkaldy, G. W. 1904. Uber Notonectiden (Hemiptera). Wien. entomol. Ztg. 23:93-135. 

La Rivers, I. 1951. A revision of the genus Ambrysus in the United States (Hemiptera: Naucoridae). 
Univ. Calif. Publ. Entomol. 8: 277-338. 

Schaefer, K. F. & W. A. Drew. 1967. Hydrometridae and Mesoveliidae (Hemiptera) of Okla- 
homa. Southw. Nat. 12: 486-487. 

Smith, C. L. 1988. Family Hydrometridae Billberg, 1820, pp. 156-158. In: T. J. Henry & R. C. 
Froeschner (eds.), Catalog of the Heteroptera, or True Bugs, of Canada and the Continental 
United States, E. J. Brill, Leiden, xix + 958 pp. 

Truxal, F. S. 1953. A revision of the genus Buenoa (Hemptera, Notonectidae). Univ. Kansas Sci. 
Bull. 35:1351-1523. 


SCIENTIFIC NOTE 


MATING BEHAVIOR OF DAS YMUTILLA 
OCCIDENTALIS (HYMENOPTERA: MUTILLIDAE)! 


Jeffery K. Tomberlin2 


Because of the short time required for successful mating to occur in velvet ants (Hymenoptera: 
Mutillidae), very few observations have been recorded (Manley 1977). Dasymutilla occidentalis 
(L.) isa common species, whose mating behavior has not been recorded. From | to 10 July 1996, 
I made field observations on nine occasions during three periods of the day: 0700 - 0900 h 
(morning), 1145 - 1400 h (noon), and 1530 - 1630 h (afternoon) (three for each period). Copula- 
tion was recorded three times during afternoon observations. 

Observations were made in Clemson, South Carolina, in a fallow field which had been plowed 
two weeks prior to observed activity. Soil was dry and sandy with no vegetative growth. Sweet 
potatoes, cantaloupe, and watermelon were planted in a field located at the north end of the 
fallow field. Four to eight males were observed during morning and afternoon sessions, whereas 
female numbers were never above two/session. Neither sex was observed during the noon ses- 
sion. Manley and Spangler (1983) observed similar activity patterns for species of the genus 
Dasymutilla. Maximum daily temperatures ranged from 32 to 35 °C. Males patrolled approxi- 
mately 30 cm above the surface of the field, but returned to shaded areas (2.5 °C cooler) to take 
in water from the irrigation system. 

The behavioral pattern was similar for each mating. One male landed within 20 cm of the 
female and moved towards the female. Following contact with the female, the male grasped her 
dorsally from behind. The female remained still until the male extended and attached his genita- 
lia to the female’s posterior end. Following genital contact, the female began stridulating. Spangler 
and Manley (1978) observed similar mating stridulations for Dasymutilla foxi (Cockerell). 
Dasymutilla occidentalis copulation lasted from 2 to 5 seconds. Once the male removed his 
genitalia from the female, stridulation ceased and the pair moved apart. The male resumed low 


(Continued on page 317) 


| Received September 3, 1996. Accepted September 30, 1996. 
2 Department of Entomology, Clemson University, Clemson, SC 29634-0365. 


Vol. 108, No. 4, September & October, 1997 311 


NEW STATE RECORDS FOR SCARAB BEETLES 
(COLEOPTERA: SCARABAEIDAE) 
FROM NORTH DAKOTA AND MINNESOTA! 


Paul K. Lago 


ABSTRACT: State records are presented for Omorgus punctatus and Cremastocheilus wheeleri 
from North Dakota and for Stephanuca pilipennis and Cremastocheilus nitens from Minnesota. 


Since the publication of the latest survey paper on the North Dakota scarab 
fauna (Lago et al. 1979), I have had the opportunity to do some additional 
collecting in the state and have examined a considerable amount of material 
collected by the students and staff of both the Department of Entomology and 
the Department of Geology, North Dakota State University, Fargo, ND. Dur- 
ing this time, two new records for North Dakota and two from neighboring 
Minnesota have been discovered. Since it is doubtful that a major faunistic 
study of scarabs will be conducted in this area anytime in the near future, I 
wish to add these new records to the North Dakota state list to bring it up to 
date. 

Cremastocheilus wheeleri LeConte occurs in the western Great Plains, and 
is often found in association with ants of the genus Formica (Potts 1945; 
Ratcliffe 1991). Although known to occur as far north as southern Alberta, 
Saskatchewan and Manitoba (Alpert 1994; Bousquet 1991) and as far east as 
eastern Nebraska (Ratcliffe 1991) the species has not been reported from the 
Dakotas. On 19 May, 1981, I collected a single female at Walcott Dunes, 
Richland Co., N.D., in a nest of Formica obscuripes Forel. This area of Richland 
County is characterized by its active sand dunes, but the ant nest, which also 
contained adult Euphoria inda (L)., was located in a grassy area on the edge of 
a mixed stand of green ash (Fraxinus pennsylvanica Marsh.) and bur oak 
(Quercus macrocarpa Michx.). 

Omorgus punctatus (Germar) is a fairly common species found throughout 
the Great Plains, the southwestern U.S.A., and into central Mexico. Based on 
the range presented by Vaurie (1955), the presence of this species in North 
Dakota was considered a distinct possibility by Lago et al. (1979). On 2 
August 1993, David Cuthrell, using a mercury vapor light, collected a single 
male at Burning Coal Vein, Slope County, N.D., confirming our suspicions. 

The distribution of Cremastocheilus nitens LeConte appears to be cen- 
tered around the 100th meridian, the species occurring from southeastern 


! Received November 25, 1996. Accepted January 28,1997. 
- Department of Biology, University of Mississippi, University, MS 38677. 


ENT. NEWS 108(4): 311-312, September & October, 1997 


312 ENTOMOLOGICAL NEWS 


sandhills of North Dakota (Lago et al. 1979) through Okiahoma (Alpert 1994) 
and Missouri (Blackwelder and Arnett 1974). One female, collected by T. L. 
McCabe at Bird Prairie (46.51.20-96.28.40), Clay County, MN, 9 May 1972, 
represents the first record for this species from Minnesota. 

Stephanuca pilipennis Kraatz inhabits sandy, upland habitats in the eastern 
Great Plains from Kansas through North Dakota (Lago et al. 1979; Ratcliffe 
1991). I recently examined two specimens of this species collected by David 
Cuthrell, 2 miles south of Fertile, Polk County, MN, 26 April, 1993. This rep- 
resents the first record for this species east of the Red River of the North. This 
area contains sand prairie habitat very similar to the blowouts in southeastern 
North Dakota where I have previously encountered the species. Known as 
Agassiz Dunes, this is the largest dune field in Minnesota associated with Gla- 
cial Lake Agassiz. The sand blowouts contain creeping juniper (Juniper 
horizontalis Moench). Large bur oaks occur in the area. Other common plants 
include silky prairie clover (Dalea villosum Nutt.), skeletonweed (Lygodesmia 
juncea (Pursh)), two species of blazing star (Liatris spp.), gramma grass 
Bouteloua spp.), and, of course, the bluestems Andropogon gerardii Vitman 
and Schizachyrium scoparium (Michx.). 

The specimens of Cremastocheilus wheeleri and C. nitens reported above 
are housed in my personal collection. The other specimens are part of David 
Cuthrell’s collection. 


ACKNOWLEDGMENT 


I wish to thank David Cuthrell and Tim McCabe for the opportunity to examine their speci- 
mens. In addition, David provided a detailed description of the Agassiz Dunes area and made 
valuable comments concerning the contents of this manuscript. Ed Zuccaro and Brett Ratcliffe 
also provided helpful editorial comments, as did two anonymous reviewers. 


LITERATURE CITED 


Alpert, G.D. 1994. A comparative study of the symbiotic relationships between beetles of the 
genus Cremastocheilus (Coleoptera: Scarabaeidae) and their host ants (Hymenoptera: 
Formicidae). Sociobiology 25 (1): 1 -276. 

Blackwelder, R.E. and R.H. Arnett, Jr. 1974. Checklist of the beetles of Canada, United States, 
Mexico, Central America and the West Indies. Vol. 1, Part 3. The scarab beetles, ant-loving 
beetles, clown beetles and related groups (red version). Biol. Res. Inst. Amer., Inc. Lantham, 
NY. 231 pp. 

Bousquet, Y. 1991. Checklist of the beetles of Canada and Alaska. Agriculture Canada Publ. 
1861/E. Biosystematics Res. Cent., Ottawa, ON. 430 pp. 

Lago, P.K., R.L. Post and C.Y. Oseto. 1979. The Phytophagous Scarabaeidae and Troginae 
(Coleoptera) of North Dakota. North Dakota Insects Publ. 12. (Schafer-Post Series). 131 pp. 

Potts, R.W.L. 1945. A key to the species of Cremastocheilini of North America and Mexico 
(Coleoptera, Scarabaeidae). Bull. Brooklyn Entomol. Soc. 40:72-78. 

Ratcliffe, B.C. 1991. The Scarab Beetles of Nebraska. Bull. Univ. Nebraska St. Mus. 12.333 pp. 

Vaurie, P. 1955. A revision of the genus Trox in North America (Coleoptera, Scarabaeidae). Bull. 
Amer. Mus. Natur. Hist. 106. 89 pp. 


Vol. 108, No. 4, September & October, 1997 31 


Ww 


ANNOTATED CHECKLIST OF THE MECOPTERA 
(SCORPIONFLIES) OF ARKANSAS! 


Henry W. Robison2, George W. Byers>, Christopher A. Carlton4 


ABSTRACT: Collection data are presented for 16 species representing 3 families of Mecoptera 
(scorpionflies and their close relatives) from Arkansas as the first checklist for the state. Panorpa 
submacalosa (Panorpidae) and Hylobittacus apicalis (Bittacidae) are reported from the State for 
the first time. 


Little 1s known about the Mecoptera (scorpionflies and their close rela- 
tives) of Arkansas. This checklist brings together all known records of the 
mecopteran fauna from the state, based on data gleaned from previous litera- 
ture (Byers 1973, 1993, 1996; Carpenter 1931; and Webb et al., 1975), mu- 
seum records, and recent collecting by the authors within the state. Collection 
localities, dates of capture, collector (where known), brief data on habitat, and 
known range of each species are included. Sixteen species representing three 
families of scorpionflies (Mecoptera) were documented from Arkansas includ- 
ing one of Meropeidae, ten of Panorpidae, and five of Bittacidae. Panorpa 
submaculosa Carpenter and Hylobittacus apicalis (Hager) are reported from 
Arkansas for the first time. 

Arkansas includes portions of the Ozark and Ouachita Highlands, which 
are of considerable biogeographic significance as areas of endemism and refu- 
gia for disjunct populations of organisms, particularly those with faunal affini- 
ties to the southern Appalachian Mountains (Ross and Ricker 1971; Mayden 
1985; Carlton and Cox 1989; Poulton and Stewart 1991; Mohlenbrock 1993; 
Robison and Allen 1995; Carlton and Nobles 1996; Moulton and Stewart 1996). 
This paper is part of our continuing effort to document the distribution and 
composition of inadequately known insect taxa in Arkansas and to improve 
our understanding of arthropod distribution patterns in North America. 


Meropeidae 


Merope tuber Newman. Montgomery Co.: 5 mi. NE of Mt. Ida, one male, 9 June 1993, flight 
intercept trap, C. E. Carlton; habitat: moist north slope deciduous forest. Newton Co.: 
Buffalo National River, road to Fitton Cave, 36 05' 30" N, 93 14'54" W, one female, flight inter- 


| Reeeived October 1, 1996. Accepted February 14, 1997. 


2 Dept. of Biology, Southern Arkansas University, Magnolia, AR 71753-5000. Corresponding 
author. 


3 Snow Entomological Museum, Univ. of Kansas, Lawrence KS 66045. 
41 ouisiana State University Agricultural Center, Dept. of Entomology, Baton Rouge, LA 70893. 


ENT. NEWS 108(4): 313-317, September & October, 1997 


314 ENTOMOLOGICAL NEWS 


cept trap, 26-28 lune 1994, C. E. Carlton; same except trail to Fitton cave, 2 km. NW of Erbie 
Campground, 3 females, 26-28 June 1994, C. E. Carlton. Washington Co.: Fayetteville, 15 speci- 
mens: 5-12 Sept. 1992, in flight intercept trap, C. E. Carlton; one male, one female, 20-27 June 
1995, in flight intercept trap, C. E. Carlton. 

Range: N GA to ME west to OK, KS, and MN. 


Panorpidae - Scorpionflies 


Panorpa anomala Carpenter. Lee Co.: St. Francis National Forest, Bear Creek Lake, 5 mi. SE 
Marianna, 16 May 1988, D. W. Webb, one male and one female, sweeping herbaceous vegetation 
along outlet creek, /mpatiens sp. and stinging nettle. Washington Co.: 21 May 1926. 

Range: TN and GA to WI, KS, and AR. 


Panorpa braueri Carpenter. Benton Co.: Bentonville (within city limits), 21 September 1977, G. 
W. Byers; habitat: mints and other herbaceous plants in ravine shaded by white oaks, sycamores, 
elms; stream dry on this date. Washington Co.: 18 September 1975, C. Daggett; 1964, no collec- 
tor; Lake Weddington Park, west of Fayetteville, 20 August 1967, J. Scott; Blue Springs camp- 
ground, south arm of Beaver Lake, 29 May 1974, R. L. Heitzman; Lake Weddington Park, 20 
Aug. 1967, R. L. Heitzman; 10 October 1976 (no other data); 9-10 June 1975, W. D. Wylie; 6 
May 1975, W. D. Wylie; 15 Sept. 1975, Walker; 3 mi. NE of Fayetteville, 3-7 June 1995, C. E. 
Carlton. 
Range: AR and MO. 


Panorpa capillata Byers. Montgomery Co.: 14.4 km. N of Pencil Bluff, 21 May 1978, G. W. 
Byers and C. W. Young; Gap Creek, 2.1 km. SE of Joplin, Ouachita National Forest, 22 May 
1978, G. W. Byers and C. W. Young; 5 mi. NE of Mt. Ida, 25 May 1993, C. E. Carlton; same 
locality, 9 June 1993, C. E. Carlton. Polk Co.: Rich Mountain, 6 July-November 1995, pitfall 
trap, H. W. Robison. 

Range: AL to KY, W to MS, and AR. 

Comments: This is a species of field-woodland ecotones. 


Panorpa choctaw Byers. Johnson Co.: Gilliam Bog, | 0 mi. N of Clarkeville, 11 September 1995, 
H. W. Robison. Newton Co.: Dogpatch, 5 September 1971, A. S. Menke. Montgomery Co.: 9 mi. 
NW of Langley, 17 September 1995, H. W. Robison. Pike Co.: 21 September 1974, P. McLeod. 
Polk Co.: U.S. Hwy. 270 near Scott Co. line, altitude 1160 ft., 27 September 1979, G. W. Byers; 
Bard Springs, 11 October 1995, H. W. Robison; Rich Mountain, jct. of St. Hwys. 88 and 272, 6 
July - 10 November 1995, pitfall trap, H. W. Robison, pitfall trap; Rich Mountain at jet. of St. 
Hwys. 272 and 88, 10 October 1996, H. W. Robison, pitfall trap. Searcy Co.: 20 September 1976, 
R. L. Brown. Scott Co.: Mill Creek at Y-City, altitude 740 ft., 27 September 1979, G. W. Byers; 
Ouachita National Forest, TIN, R30W, Sec. 33, altitude 1020 ft., 27 September 1979, G. W. 
Byers; on shaded, low herbaceous vegetation primarily at edges of woods. Washington Co.: 2 
July 1975, E. Sizemore; 13 September 1976, E. Brack; 29 September 1974, P. McLeod; 13 Sep- 
tember 1974, D. Dunn; 22 September 1974, D. Dunn; 23 September 1975, Graves. 
Range: KY S to GA, W to OK and MO. 


Panorpa helena Byers. Garland Co.: Blue Springs (near Hot Springs National Park), 23 May 
1966. Pope Co.: 17 June 1970, R. Flanigan. Washington Co.: October 1961, H. Roberts; July 
1970, G. W. Wallis. 

Range: GA to MA and W to Manitoba and AR; isolated record from UT (Gurney, 1937, as P. 
venosa). 


Panorpa nuptialis Gerstaecker. Arkansas Co.: I | August 1975, Rhinehart; White River Refuge, 
August 1969, R. L. Brown. Calhoun Co.: 14 October 1993, J. Rader. Columbia Co.: Village, 28 
October 1996, K. Arbuckle. Craighead Co.: Jonesboro, 27 September 1966, Hopkins. Crittenden 


Vol. 108, No. 4, September & October, 1997 315 


Co.: West Memphis, 22 October 1966, no collector. Drew Co.: Tillar, 5 October 1939, E. C. 
VanDyke. Garland Co.: Hot Springs, 4 October 1939, E. C. VanDyke. JeffersonCo.: Pine Bluff at 
UAPB Campus, Lot 129, September 1990 (no other data). Lawrence Co.: Imboden (no other 
data). Nevada Co., 13 October 1976, H. Greenbaum. Nevada Co.: 3 October 1964, G. Wallis. 
Ouachita Co.: 10 mi. N Camden, 1-3 November 1960, M. Verley. Pope Co.: 5 July 1970, R. 
Flanigan. Pulaski Co.: Burns Park, North Little Rock, 22 September 1977, G. W. Byers (col- 
lected together with P. rupeculana; see for habitat). Scott Co.: Mill Creek at Y City, 27 Sept. 
1979, G. W. Byers. Van Buren Co., 2 October 1985, N. Brock. Washington Co.: 25 October 1937, 
no collector; 20 September 1938, no collector; October 1938, no collector; 8 August 1956, D. M. 
Powell; Cove Creek Valley, 1958, M. Hite; October 1960, L. O. Warren; September 1961, M. 
Roberts; October 1961, N. Roberts; 14 October 1962, O. Hite; 1 May 1970, no collector; August 
1971, no collector; 20 June 1975, J. Welch; 30 September 1975, Prichett; 21 May 1976, J. Fitz; 
September 1976, McCormick; 7 October 1984, A. Wilburn; Fayetteville, 17 August 1985, S. 
Tedder. 
Range: AL to MO W to KS, OK, TX. 


Panorpa rupeculana Byers. Miller Co.: 0.5 mi E of Doddridge, 26 April 1987, D. W. Webb, one 
female. Ouachita Co.: 8.7 mi. NE of Camden. 24 Oct. 1957, I. J. Cantrall, T. Cohn, D. Eades. 
Pulaski Co.: Burns Park, North Little Rock, 22 September 1977, G. W. Byers. 

Range: AR, northern LA, MS. 

Comments: At the type locality near Little Rock, thc specimens were collected from low 
growth of honeysuckle, greenbriers, and brambles (Rubus sp.) shaded by oaks, hickory, maple, 
red gum, black gum, and pine. 


Panorpa speciosa Carpenter. Washington Co.: 19 June 1964, A. E. Shumate; Sept. 1961, H. 
Roberts; 21 May 1925, no collector; White River, 0.5 mi. N of Brentwood, 21 May 1978, G. W. 
Byers. 

Range: AR, TN, KY, OH to MN and WI. 


Panorpa submaculosa Carpenter. Johnson Co.: Bull Creek Flats Spring, N of Clarkeville, 25 
May 1993, H. W. Robison, one male; New State Record. Footprint Spring, ca. 10 mi. N of 
Clarkeville, 7 June 1996, G. Leeds, four females in pitfall trap. 

Range: GA to MA and W to AR and WI. 


Panorpa vernalis Byers. Bradley Co.: 14 April 1967, I. Brown (paratype); 1 mi. SE El Dorado, 6 
May 1984, D. W. Webb. 

Range: MS, LA, southern AR. 

Comments: The usual habitat is in somewhat open areas by mixed hardwoods and pine; the 
specimens were found on low growth of honeysuckle, poison ivy, greenbrier with some patches 
of “pine straw” about 2 inches deep. 


Bittacidae - Hangingflies 


Hylobittacus apicalis (Hager). Johnson Co.: N of Clarksville (TI3N, R24W, Sec. 4), 2 June 
1995, G. Leeds. Lee Co.: St. Francis National Forest, Bear Creek Lake, 5 mi. SE Marianna, 16 
May 1988, D. W. Webb, one male, one female, sweeping herbaceous vegetation along outlet 
creek, /mpatiens sp. and stinging nettle. New State Record. 

Range: NC to NY and W to OK and AR. 


Bittacus pilicornis Westwood. Benton Co.: 12 May 1941 (no other data); Johnson Co.: Lake 
Dardanelle, Spadra, 27 May 1978, G. W. Byers; habitat: poison-ivy, wild roses, and herbaceous 
plants. Logan Co.: Magazine Mountain, altitude 1350 ft., 17 May 1989, in black light trap, J. 


316 ENTOMOLOGICAL NEWS 


MacGowan and Q. Fang; Magazine Mountain, 25 May 1978, L. D. Newson; in light trap; (Note: 
A large male, front wing 25 mm, was taken at the light trap and has one pair of pheromone- 
dispersing organs everted. In the left fore wing, R4 divides, then branches coalesce before the 
wing margin. Det. G.W. Byers). Montgomery Co.: 9 mi. NW of Pencil Bluff, 21 May 1978, G.W. 
Byers and C. W. Young; 6.5 mi. NW of Bonnerdale, S. of FSR 476 (Sec. 22, T3S, R23W), 22 
May 1996, H. W. Robison, black light. Union Co.: 1 mi. SE El Dorado, 6 May 1984, D. W. Webb. 
Washington Co.: 4 June 1962, no collector; 18 May 1963 (no data); 5 mi. W. Fayetteville, 15 
May 1986, D. E. Bowles; 15 mi. W. Fayetteville, 16 May 1986, D. E. Bowles; 24-25 May, no 
collector; 15 mi. S Prairie Grove (no date or collector); Devil’s Den State Park, altitude approx. 
1400 ft., 24 May 1977, G. W. Byers. 

Range: FL to Canada and W to MN and TX. 

Comments: Habitat of B. pilicornis is typically herbaceous undergrowth 1-3 ft. high in mixed 
deciduous woods. 


Bittacus punctiger Westwood. Johnson Co.: Spadra, U.S. Corps of Engineers Park on Lake 
Dardanelle, 27 May 1978, G. W. Byers; habitat: these bittacids were taken low in the under- 
growth, 8-12 inches above the ground, while other Bittacus sp. were higher in the shaded vege- 
tation. 

Range: FL to PA and W to MO, OK and TX. 


Bittacus stigmaterus Say. Arkansas Co.: 4 mi. E. Ethel, 3 July 1969, R. L. Brown; 19 July 1969, 
R. L. Brown; 31 July 1969, R. L. Brown; 13 August 1969, R. L. Brown. Polk Co.: Rich Moun- 
tain, jct. of St. Hwys. 88 and 172, 6 July - 10 November 1995, pitfall trap, H. W. Robison. 
Washington Co.: 10 July 1965, H. R. Dodge. 

Range: GA to NY and W to MN and TX. 


Bittacus strigosus Hagen. Carroll Co.: 1 mi. NW Berryville, 27 June 1972, D. W. Webb. Newton 
Co.: Buffalo National River, trail to Fitton Cave, 2 km. NW of Erbie Campground, 26-28 June 
1994, C. E. Carlton. Washington Co.: 22 June 1968. J. Kimbrough; 3 mi. NE of Fayetteville, 
moist deciduous woodland, 20-27 June 1995, C. E. Carlton. 

Range: SC to southern Canada W to Manitoba, MT, WY, and TX. 


Undoubtedly, other species of mecopterans await discovery in Arkansas as 
much additional collecting is needed, particularly in remote mountain areas. 
Such species as Panorpa sigmoides Carpenter, Bittacus occidentis Walker, and 
Bittacus texanus Banks inhabit neighbortng states and possibly occur in Ar- 
kansas. Our studies will continue in an attempt to define further the state 
scorpionfly fauna. 


ACKNOWLEDGMENTS 


Travel funds for Henry Robison were graciously provided by the Southem Arkansas Univer- 
sity Research Committee. Field work by Chris Cariton was funded in part by grants from the 
USDA-Forest Service and Department of Interior, Buffalo National River. Special thanks are 
extended to Jan Rader and Nick Covington, Southern Arkansas University students, for their 
assistance in the field and to Gene Leeds, USDA-Forest Service, Pleasant Hill Ranger District, 
who kindly loaned us several specimens of scorpionflies for this study. 


LITERATURE CITED 


Byers, G. W. 1973. Descriptions and distributional records of American Mecoptera. III. J. Kan. 
Entomol. Soc. 46:362-375. 

Byers, G. W. 1993. Autumnal Mecoptera of southeastern United States. Univ. Kan. Sci. Bull. 
55(2): 57-96. 

Byers, G. W. 1996. Descriptions and distributional records of American Mecoptera. IV. Univ. 
Kan. Sci. Bull. 55 (14):519-547. 


Vol. 108, No. 4, September & October, 1997 317 


Carlton, C. E. and R. T. Cox. 1989. A new species of Arianops from central Arkansas and the 
biogeographic implications of the Interior Highlands Arianops species (Coleoptera: 
Pselaphidae). Coleopt. Bull. 44:365-371. 

Carlton, C. E. and L. S. Nobles. 1996. Distribution of Speyeria diana (Lepidoptera: Nymphalidae) 
in the Interior Highlands of Arkansas, Missouri, and Oklahoma, with comments on conserva- 
tion. Entomol. News 107:213-219. 

Carpenter F. M. 1931. Revision of the nearctic Mecoptera. Bull. Mus. Compar. Zoology, Harvard 
Univ. 72(6):205-277, plates 1-8. 

Gurney, A. B. 1937. A new species of Panorpa from Utah, with notes on other nearctic species 
(Mecoptera). Proc. Entomol. Soc. Wash. 39 (8):222-227. 

Mayden, R. L. 1985. Biogeography of Ouachita Highland fishes. Southwest. Natur. 30(2): 195- 
211i: 

Mohlenbrock, R. H. 1993. Ouachita Mountains, Arkansas. Nat. Hist. 9:22-24. 

Moulton, S. R. and K. W. Stewart. 1996. Caddisflies (Trichoptera) of the Interior Highlands of 
North America. Mem. Amer. Entomol. Insitute 56: 1-313. 

Poulton, B. C. and K. W. Stewart. 1991. The stoneflies of the Ozark and Ouachita Mountains 
(Plecoptera). Mem. Amer. Entomol. Soc. 38: 1-116. 

Robison, H. W. and R. T. Allen. 1995. Only in Arkansas: a study of the endemic plants and 
animals of the state. Univ. Arkansas Press, Fayetteville. 121 pp. 

Ross, H. H. and W. E. Ricker. 1971. The classification, evolution, and dispersal of the winter 
stonefly genus Allocapnia. Illinois Biol. Monographs 45: 1-166. 

Webb, D. W., N. D. Penny and J. C. Marlin. 1975. The Mecoptera, or scorpionflies, of Illinois. 
Illinois Nat. Hist. Sur. Bull. 31(7):251-316. 


(Continued from page 310) 


elevation flight over the field, while the female retreated into surrounding vegetation dragging 
her posterior end on the soil. Following mating, no other males attempted to copulate with the 
female. 

Similar mating behaviors have been observed for different species of Mutillidae (Manley 
and Deyrup 1989; Manley and Taber 1978; Fattig 1936). Males present in mass numbers have 
been observed swarming 20 to 30 cm above the ground, apparently searching for females (Manley 
and Deyrup 1989). However, the number of males I observed was not as high as that observed for 
other species (Manley and Taber 1978). In general, mating of mutillids occurred within a few 
seconds and once a female was mated, no other male attempted further copulation. 


ACKNOWLEDGMENT 


I would like to thank Peter H. Adler and Donald G. Manley for their helpful comments on 
this manuscript. This is Technical Contribution No. 4222 of the South Carolina Agricultural 
Experiment Station, Clemson University. 


LITERATURE CITED 


Fattig, P. W. 1936. An unusual mating of velvet ants (Hymen.: Mutillidae). Entomol. News 47: 
51-52. 

Manley, D.G. 1977. Notes on the courtship and mating of Dasymutilla Ashmead (Hymenoptera: 
Mutillidae) in Calif. Southw. Nat. 21: 543-559. 

Manley, D.G. and M.A. Deyrup. 1989. Notes on the biology of Dasymutilla pyrrhus (Fox) 
(Hymenoptera: Mutillida). J. Entomol. Sci. 24: 53-56. 

Manley, D.G. and G. Spangler. 1983. Observations on daily activity patterns of mutillid wasps 
of the genus Dasymutilla. J. Ga. Entomol. Soc. 18: 235-239. 

Manley, D.G. and S. Taber, III. 1978. A mating aggregation of Dasymutilla foxi in southern 
Arizona. Pan-Pac. Entomol. 54: 231-235. 

Spangler, H.G. and D.G. Manley. 1978. Sounds associated with the mating behavior of a mutillid 
wasp. Ann. Entomol. Soc. Amer. 71: 389-392. 


318 ENTOMOLOGICAL NEWS 


SCIENTIFIC NOTE 


NAME ADJUSTMENTS AND A NEW SYNONYM FOR 
NORTH AMERICAN EPHEMEROPTERA SPECIES! 


W. P. McCafferty 


While researching the original orthography for the list of North American mayflies, as it is 
to appear in volume 4 of Nomina Insecta Nearctica, | discovered that three specific epithets had 
not been corrected to gender agreement when recombined by McCafferty and Waltz (1990) or 
listed again by McCafferty (1996); these names are corrected to Baetis futilis (McDunnough), 
1936 [originally Pseudocloeon futile], Baetis rubrolateralis (McDunnough), 1931 [originally 
Pseudocloeon rubrolaterale), and Baetis virilis (McDunnough), 1923 [originally Pseudocloeon 
virile]. One specific epithet had not been correctly formed by Berner (1940) or corrected subse- 
quently either with its original genus or in recombination; this name is corrected to Procloeon 
viridoculare (Berner), 1940 [originally Centroptilum viridocularis and subsequently P. 
viridocularis]. One specific epithet had not been corrected to gender agreement by McDunnough 
(1926) or any North American authors listing or treating the name subsequently; this name is 
corrected to Cinygma lyriforme (McDunnough), 1924 [originally Ecdyonurus lyriformis and sub- 
sequently C. lyriformis]. One additional specific epithet had been incorrectly formed by 
McDunnough (1939); this name is corrected to Baetis persecutor McDunnough, 1939 [origi- 
nally B. persecuta and subsequently B. persecutus). 

With regard to the first three corrections, the specific epithets are transformed to the mascu- 
line gender from the original neuter to agree with the masculine name Baetis. In the case of P. 
viridoculare, both the original Centroptilum Eaton and subsequent Procloeon Bengtsson generic 
combinations required neuter agreement. In the case of Cinygma Eaton, McDunnough (e.g., 
McDunnough 1924) formed all his epithet names that were associated with this genus, and de- 
rived from the nominative singular, in the incorrect feminine gender. It is clear from Eaton (1885) 
that his genus is neuter. The “a” ending of the name, which is most often indicative of the Latinized 
feminine gender was evidently misleading. Notably, Stenonema Traver is another example of a 
mayfly genus group name ending in “ma” (latinization of ending Greek letters “ac’’) that is 
neuter, not feminine and that has confused workers. The emendation to B. persecutor is based on 
the fact that the correct spelling of the Latin noun in apposition is “persecutor.” 

McCafferty (1993) synonymized Drunella conestee (Traver), 1932 with Drunella tuberculata 
(Morgan), 1911. Further examination of North Carolina material has revealed that Drunella wayah 
(Traver), 1932 also falls within the range of variation of D. tuberculata as discussed by McCafferty 
(1993). This variablility involves, for example, frontal shelf, genae, and tuberculation develop- 
ment. Allen and Edmunds (1962) drew the forefemur of wayah with a forefemoral ridge some- 
what less apparent than that drawn for tuberculata or conestee, but the ridge is nonetheless present. 
No adults have ever been associated with the name D. wayah, and its known distribution is 
consistent with that recorded for D. tuberculata and, more precisely, D. conestee. Therefore, I 
place D. wayah as a junior subjective synonym of D. tuberculata: Drunella tuberculata (Mor- 
gan) [= D. wayah (Traver), n. syn.]. 

(Continued on top of page 320) 


! Received February 22, 1997. Accepted March 21, 1997. 
2 Purdue Agricultural Research Program Journal No. 15325. 
3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 


Vol. 108, No. 4, September & October, 1997 319 


SCIENTIFIC NOTE 


NEW RECORDS OF NOTONECTIDS 
(HETEROPTERA) FOR PENNSYLVANIA! 2 


A. M. Yeakel3, E. Larsen? 


Members of the family Notonectidae are predaceous aquatic insects. Two principal genera, 
Notonecta and Buenoa, are found in the lentic habitats of North America. Four species have been 
recorded from Pennsylvania (Truxal 1953, Polhemus and Polhemus 1988). We report here four 
additional species, two of Buenoa and two of Notonecta, from Pennsylvania. This note is a con- 
tribution to ongoing attempts to document the distribution of Heteroptera of North America, 
north of Mexico (see Henry and Froeschner 1988). 

Buenoa confusa occurs generally across the eastern United States and Canada, but has been 
unrecorded from PA until now (Truxal 1953, Polhemus and Polhemus 1988). Collections of B. 
confusa were made from a permanent man-made drainage pond at the corner of Lancaster Av- 
enue (PA state route 30) and King of Prussia Road, Villanova, Delaware County, PA. This pond is 
0.5 to 1 meter deep around the edges and over 2 meters deep in the center. There is little emergent 
or submerged aquatic vegetation and the diameter of the pond is approximately 25 meters. De- 
spite the pollution and litter from the drainage into this pond, invertebrates are abundant, includ- 
ing other notonectids such as, B. margaritacea and Notonecta undulata. Four individuals (3 
males, 1 female) of B. confusa were found in June, 1995. Sampling efforts in May, 1996 pro- 
duced no individuals of this species, while 4 and 3 individuals of B. confusa were collected in 
September and October, 1996, respectively. The previously recorded distribution in northeast 
North America for B. confusa includes CT, DC, ME, NJ, NY and Nova Scotia (Truxal 1953). 
Collection of this species in PA fills a gap in the distribution record. 

Buenoa scimitra is known from Mexico and the southern half of the United States (Truxal 
1953, Polhemus and Polhemus 1988). To date, this species has not been recorded from any local- 
ity in the northeast. Buenoa scimitra was collected from the same pond described above, 25 
males in September, 1995 and 31 males in October, 1996. Six males of B. scimitra, also collected 
in September 1995, were taken from a temporary pool created by construction activities on 
Villanova University’s west campus. The pool was shallow and muddy, and filled in soon after 
the collection. An additional 8 male B. scimitra were collected in September, 1995 from a deco- 
rative pond, approximately 20 x 40 meters in size, located on the west campus of Villanova 
University. Buenoa margaritacea was also present at both of these sites. Because the females of 
these two species are indistinguishable (Truxal 1953), only the number of males are reported 
here. This record of B. scimitra in Pennsylvania extends the range for this species farther north 
from its previously northernmost record of VA and its presence in southeastern PA suggests that 
it may be found in DE, MD, and NJ as well. 

Notonecta insulata is known from northeast North America, including CT, MD, NJ and NY 
(Hungerford 1933, Polhemus and Polhemus 1988). One individual of N. insulata was collected 
from the previously mentioned drainage pond on the corner of Lancaster Avenue and King of 
Prussia Road, during June, 1993 and eight specimens were collected in October of the same year. 


(Continued on bottom of page 320) 


! Received December 9, 1996. Accepted January 18, 1997. 


2 We thank J. Gelhaus, Academy of Natural Sciences of Philadelphia, J. Polhemus, Colorado 
Entomological Museum, and two anonymous reviewers for comments on an early draft of this 
note. 


3 Department of Biology, 800 Lancaster Avenue, Villanova University, Villanova PA 19085. 


320 ENTOMOLOGICAL NEWS 


(Continued from page 318) 


LITERATURE CITED 


Allen R. K. and G. F. Edmunds, Jr. 1962. A revision of the genus Ephemerella (Ephemeroptera: 
Ephemerellidae). V. The subgenus Drunella in North America. Misc. Publ. Entomol. Soc. 
Am. 3: 147-179. 

Berner, L. 1940. Baetine mayflies in Florida. Fla. Entomol. 23: 33-45, 49-62. 

Eaton, A. E. 1885 (1883-88). A revisional monograph of recent Ephemeridae or mayflies. Trans. 
Linn. Soc. Lond. 2nd Ser.-Zool. 3: 1-352. 

McCafferty, W. P. 1993. Commentary on Drunella tuberculata and Procloeon penulatum 
(Ephemeroptera: Ephemerellidae; Baetidae) in North Carolina. Entomol. News 104: 235- 
239. 

McCafferty, W. P. 1996. The Ephemeroptera species of North America and index to their com- 
plete nomenclature. Trans. Am. Entomol. Soc. 122: 1-54. 

McCafferty, W. P. and R. D. Waltz. 1990. Revisionary synopsis of the Baetidae (Ephemeroptera) 
of North and Middle America. Trans. Am. Entomol. Soc. 116: 769-799. 

McDunnough, J. 1924. New Canadian Ephemeridae with notes II. Can. Entomol. 56: 90-98, 
113-122, 128-133. 

McDunnough, J. 1926. New Canadian Ephemeridae with notes IV. Can. Entomol. 58: 296-303. 

McDunnough, J. 1939. New British Columbian Ephemeroptera. Can. Entomol. 71: 49-54. 


(Continued from page 319) 


One female was collected from the same pond in October, 1996. This site provides another state 
record for N. insulata in the Middle Atlantic States. 

Notonecta lunata is known from northeast (Hungerford 1933) and midwest North America 
(Polhemus and Polhemus 1988, J. Polhemus, pers. comm.). Notonecta lunata was collected from 
the Great Marsh in Nantmeal and Wallace Townships, Chester County, PA. The marsh is approxi- 
mately 33 miles northwest of Philadelphia and is part of the Marsh Creek watershed. Part of the 
marsh property is adjacent to Pennsylvania State Route 100 and Interstate Route 76. This site is 
characterized by small areas of open water surrounded by grasses and sedges, along with interior 
floating plants. Thirteen individuals were collected in July, 1994, numerous individuals were 
collected in July, 1995, 5 individuals were found in September, 1995, and 7 individuals in June, 
1996. This site provides an additional state record for N. /unata in the Middle Atlantic States 
(Hungerford 1933, Polhemus and Polhemus 1988). 

Representative voucher specimens of the four species are deposited in the collection of the 
Entomology Department, Academy of Natural Sciences of Philadelphia, Philadelphia, PA. 


LITERATURE CITED 


Henry, T.J. and R.C. Froeschner. 1988. Catalog of the Heteroptera, or True Bugs, of Canada 
and the Continental United States, E.J. Brill, New York. 

Hungerford, H.B. 1933. The Genus Notonecta of the World (Notonectidae-Hemiptera). Unv. 
Kan. Sci. Bull. 21:5-195. 

Polhemus, J.T. and D. A. Polhemus. 1988. Family Notonectidae Latreille, 1802, The 
Backswimmas pp. 533-540. In: T.J. Henry and R.C. Froescher, Eds., Catalog of the Heteroptera, 
or True Bugs, of Canada and the Continental United States, E.J. Brill, New York. 

Truxal, F.S. 1953. A Revision of the Genus Buenoa (Hemiptera Notonectidae). Unv. Kan. Sci. 
Bull. 35: 1351-1523. 


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perla (Plecoptera: Chloroperlidae) of western 
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neview of Nearctic genus Ceramphis (Coleoptera: 
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Adventive Onthophagus (Coleoptera: Scarabaeidae) in 
North America: geographic ranges, diagnoses, 
and new distributional records E.R. Hoebeke, K. Beucke 


Micksiops, a new genus of small minnow mayflies 
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W. P. McCafferty, C.R. Lugo-Ortiz, H.M. Barber-James 


Maliqua: a new genus of Baetidae (Ephemeroptera) for 
a species previously assigned to Afroptilum 
C.R. Lugo-Ortiz, W.P. McCafferty 


New Orchesella species (Collembola: Entomobryidae) 
from North America Richard J. Snider 


A new species of Tenagobia (Heteroptera: Corixidae) 
from Venezuela J.T. Polhemus, N. Nieser 


A new species of Dacnusa (Hymenoptera: Braconidae) 
from Spain I. Docavo, J. Tormos 


Catocala louiseae, C. grisatra, and C. jair (Lepidop- 
tera: Noctuidae) in North Carolina 
W.J. Cromartie, D.F. Schweitzer 


SOCIETY MEETING OF APRIL 24, 1997 
PUBLISHER’S STATEMENT i 
MAILING DATES - VOLUME 108 
INDEX - VOLUME 108 


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Vol. 108, No. 5, November & December, 1997 321 


ALLOPERLA (PLECOPTERA: CHLOROPERLIDAE) OF 
WESTERN NORTH AMERICA ! 


Mary Leigh Lyon, Bill P. Stark2 


ABSTRACT: Seven species of the stonefly genus Alloperla are recognized in western North 
America. Scanning electron micrographs of epiproct structures and eggs are presented for each 
species, and adults are keyed. 


Adult Alloperla are distinguished from most other chloroperlid genera by 
their pale green or yellow body coloration, the lack of dark markings on the 
head and pronotum, and the reduced anal area of the hind wing with less than 
five anal veins. Several western Alloperla species with median dorsal abdomi- 
nal stripes may be confused with Suwallia, or other chloroperlid genera, but 
these are easily distinguished by male genitalic and female subgenital plate 
characters (Harper & Stewart 1996; Surdick 1985). Presently there are seven 
species of Alloperla known in western North America (Stark et al. 1986). While 
each has been discussed elsewhere (e.g., Baumann et al. 1977; Jewett 1959), no 
single publication has treated all seven species. Therefore, we have used scan- 
ning electron microscopy to develop descriptions of epiprocts and eggs, and we 
present the first key to the adults of all the western Nearctic Alloperla. 


MATERIALS AND METHODS 


Epiproct samples were taken from museum specimens stored in alcohol. 
The terminal abdominal segments were severed, placed in acetone and agitated 
in an ultrasonic cleaner for one minute. Samples were air dried, placed on double- 
stick copper tape on SEM stubs, coated with gold-palladium, and scanned us- 
ing an AMRAY model 1810 SEM. 

Egg samples were collected by opening the abdomen of the female speci- 
mens and gently dislodging the eggs into alcohol. Eggs were pipetted into ac- 
etone, cleaned in an ultrasonic cleaner for 10-20 seconds, air dried on forceps, 
and placed on the taped specimen stubs. Eggs were coated and scanned follow- 
ing the same procedure uSed for the epiprocts. Female subgenital plates from 
these specimens were examined with light microscopy after clearing in hot 10% 
KOH. 

Epiproct samples were examined from two or more populations for each 
species except A. chandleri; egg samples were taken from specimens repre- 


! Received March 15, 1997. Accepted April 19, 1997. 
2 Biology Department, Mississippi College, Clinton, MS 39058. 


ENT. NEWS 108(5) 321-334, November & December, 1997 


B22) ENTOMOLOGICAL NEWS 


senting a single population. Terminology follows Stark and Szczytko (1988), 
Baumann et al. (1977) and Jewett (1959). Specimens are deposited in the Monte 
L. Bean Museum, Brigham Young University (BYU), the C. P. Gillette Mu- 
seum of Arthropod Diversity, Colorado State University (CSU) and in the Stark 
Collection, Mississippi College (BPS). 


KEYS TO ALLOPERLA OF WESTERN NORTH AMERICA 


Males 
i-Epiproct apex smionthly rounded (Figs: 1-6). 72a ce cied o's Se eueeies ce one eee 2 
Epipnic! apex senate or crenulate (Pigs: 7-16). 60 oon oeee Sale obese eee alae 4 
2. Epiproct widest at mid-length (Figs. 5, 6); dorsal abdominal stripe present; 
known from higher elevations of the Colorado Rocky Mts.................+-: A. pilosa 
Epiproct widest at base or near apex (Figs. 1-4); dorsal abdominal. stripe 
present or absent; known from the Coast, Cascades and Sierra Nevada Mts............ 3 
3. Epiproct dumbbell-shaped (Fig. 3); dorsal abdominal stripe usually present.... A. fraterna 
Epiproct parallel sided (Fig. 1); dorsal abdominal stripe usually absent ...... A. chandleri 
4. Anterolateral homis' present on epiproct\(Figs: 7,10) «..c00 a-iscessmiee ye ees eet ee 5 
Anterolateral horns absent from epiproct (Figs. 15-16) ............... eee ee eee eee 6 
5. Epiproct apex triangular, projecting (Figs. 10-12); dorsal abdominal 
Simi pemusualby:aDSente nyc es ceca etre oc ae eect oer A. medveda 
Epiproct apex truncate (Figs. 7-9); dorsal abdominal stripe usually present .... A. delicata 
6. Epiproct at least twice as long as wide, apex crenulate, basal half pilose (Fig. 13). A. severa 
Epiproct about as wide as long, apex serrate, basal margin pilose (Fig. 15) ..... A. serrata 


Females with eggs 


lee es; collan present, (Figt 25) ee 1 seston fetes me isan oe nee Eee ete 2 
Ess collarabsent (Fig Wie ayoaurs seas toa eS eae teens oe rere reer oe ieee ee eee 5 
2. Forewing length greater than 12 mm; dorsal abdominal stripe present .......... A. pilosa 
Forewing length less than 10 mm; dorsal abdominal stripe absent.................--- 3 
3. Subgenital plate base broad, constricted to a rounded, tab-like apex (Fig. 41) .. A. medveda 
Subgenital plate constricted or unconstricted, apex angular (Figs. 37-40).............. a 


4. Subgenital plate reaches at least center of segment nine with abdomen extended 
(Fig. 43), plate reaches posterior margin of segment nine in uncleared specimens . A. severa 
Subgenital plate barely reaches anterior margin of segment nine with abdomen ex- 
tended (Fig. 39), plate reaches midpoint of segment nine in uncleared specimens A. serrata 

5. Subgenital plate apex angular (Fig. 38); egg chorion coarsely pitted (Fig. 22) .. A. delicata 
Subgenital plate apex rounded (Fig. 37); egg chorion essentially smooth (Fig. 17) ...... 6 

6. Dorsal abdomunal stripe usually present . os. 25,.3'20 as cchaiew.4 aero eed oan eiate A. fraterna 
Dorsal abdominal. stripe usnially absent a i5..5 ce n5c)ma.0) mpaiainr sina souade aie eusiniacr tors A. chandleri 


Vol. 108, No. 5, November & December, 1997 323 


Alloperla chandleri Jewett 


Alloperla chandleri Jewett, 1954b. Holotype male, Miami Ranger Station, Mariposa Co., 
California. 


Epiproct — Length 0.171 mm, basal width 0.067 mm, apical width 0.068 mm, constriction 
width 0.065 mm. Dorsal aspect almost parallel-sided, but slightly constricted mesally and rounded 
apically; apex slightly wider than base (Fig. 1). Epiproct covered with dense mat of thick setae 
except for bare apex. Apical bare area begins at point of greatest width and extends 0.02 mm. 
Setal mat less dense on posterior margin. Lateral aspect rather uniformly wide and subequal to 
dorsal width; margins clothed with setae (Fig. 2). Epiproct stem clothed with fine setae (Figs. 1, 
2). 

Egg — Length 0.33 mm, width 0.18 mm. Outline elongate oval, anterior pole more blunt than 
posterior pole. Micropylar row subequatorial; micropyle orifices sessile. Chorionic surface with 
indistinct hexagonal follicle cell impressions; eclosion line absent. Collar absent (Figs. 19-21). 

Subgenital plate — Posterior margin broadly rounded, extending slightly over anterior mar- 
gin of sternum nine (Fig. 37). 

Material - CALIFORNIA: Sierra Co., Big Springs, Hwy 49 N Sierra City, 24 June 1980, R. 
W. Baumann, J. A. Stanger, 5 0, 6 9 (BYU, BPS). 

Known distribution — CA (Stark et al. 1986). 


Alloperla delicata Frison 


Alloperla delicata Frison, 1935. Holotype male, Oak Creek, Benton Co., Corvallis, Oregon. 
Alloperla delicata: Jewett, 1954a. 


Epiproct — Length 0.123 mm, basal width 0.040 mm, mid-dorsal width 0.074 mm. Dorsal 
aspect ellipsoidal with two anterolateral horns extending from truncate apex (Fig. 7). Dorsal 
surface bare except for medial clusters of thick setae near base. Apex blunt with two large (0.015 
mm long), ventrally directed, anterolateral horns; ca. six small serrations present between horns 
(Fig. 9). Lateral margins fringed with long setae (Fig. 8); stem clothed with dense mat of fine 
setae. 

Egg — Length 0.26 mm, width 0.18 mm. Outline elongate oval, poles similar, collar absent 
(Fig. 22). Micropylar row subequatorial; turretted orifices arise from circular pits (Fig. 23). Chori- 
onic surface prominently pitted with widely spaced, variably sized aeropyles on anterior and 
posterior third; equatorial third smooth; eclosion line absent (Figs. 22, 24). 

Subgenital plate — Posterior margin rounded; plate somewhat triangular, extending slightly 
over anterior margin of sternum 9 (Fig. 38). 

Material - CALIFORNIA: Humboldt Co., East Fork Campground, 4 mi W Willow Creek, 
23 June 1985, B. P. Stark, 1 C, 1 9 (BPS). Grassy Creek, Fieldbrook, 22 May 1982, B. P. Stark, 
D. Ziegler, 1 C (BPS). Marin Co., Redwood Creek, Mt. Tamalpais State Park, 22 April 4987, R. 
W. Baumann, B. P. Stark, C. R. Nelson, S. A. Wells, 4 OC (BYU ). Redwood Creek, Muir Woods, 
21 June 1985, B. P. Stark, 4 O',7 9 (BPS). Nevada Co., Sagehen Creek, University of California 
Biological Station, 19 June 1985, B. P. Stark, 2 0’, 4 9 (BPS). 

Known distribution — AB, BC, CA, ID, MT, OR (Stark et al. 1986). 


324 ENTOMOLOGICAL NEWS 


Figs. 1-6. Scanning electron micrographs of Alloperla epiprocts. 1. A. chandleri, dorsal aspect. 
2. A. chandleri, lateral aspect. 3. A. fraterna, dorsal aspect. 4. A. fraterna, lateral aspect. 5. A. 
pilosa, dorsal aspect. 6. A. pilosa nearly lateral aspect. 


Vol. 108, No. 5, November & December, 1997 325 


Alloperla fraterna Frison 


Alloperla fraterna Frison, 1935. Holotype male, Oak Creek, Benton Co., Corvallis, Oregon. 
Sweltsa fraterna: Illies, 1966. 
Alloperla fraterna: Surdick, 1985. 


Epiproct — Length 0.124 mm, basal width 0.045 mm, apical width 0.03 mm, constriction 
width 0.027 mm. Dorsal aspect dumbbell shaped, constricted area nearer apex than base; apex 
smoothly rounded and bare (Fig. 3); length of bare apical area ca. 0.013 mm. Dorsal surface 
covered with dense mat of long setae (Figs. 3-4). Lateral aspect somewhat flattened at base and 
somewhat enlarged at apex (Fig. 4). Ventral surface and lip area around base without setae; stem 
clothed with dense mat of fine setae. 

Egg — Length 0.28 mm, width 0.17 mm. Outline elongate oval. Micropylar row subequato- 
rial, orifices sessile. Chorionic surface smooth, eclosion line absent. Collar absent (Figs. 17-18). 

Subgenital plate — Posterior margin smoothly rounded, sides slightly convergent. Posterior 
margin extending slightly over anterior margin of sternum nine (Fig. 42). 

Material - CALIFORNIA: Humboldt Co., Hwy 299, 10 mi W Willow Creek, 23 June 1985, 
B. P. Stark, 7 0, 6 9 (BPS). Santa Clara Co., Swanson Creek, Uras Canyon, 4 June 1969, D. G. 
Denning, 4 C (BYU). Siskiyou Co., Castle Creek below Castle Lake, 7 July 1979, B. P. Stark, K. 
W. Stewart, 2 0’, 4 9 (BPS). WASHINGTON: Jefferson Co., Taft Creek, 1 mi W Hoh Visitor 
Center, Olympic National Park, 16 July 1979, B. P. Stark, 1 CO (BPS). 

Known distribution. - AK, BC, CA, OR, WA (Stark et al. 1986). 


Alloperla medveda Ricker 


Alloperla medveda Ricker, 1952. Holotype male, Bear Tooth Creek, Bear Tooth Mountains, 
Carbon Co., Montana. 


Epiproct — Length 0.113 mm, basal width 0.062 mm, subapical width 0.04 mm; the 0.073 
mm base of the triangular apex is formed by two large anterolateral horns (Figs. 10, 12). Dorsal 
aspect somewhat elliptical, surface bare except for medial clusters of thick setae near base (Fig. 
10). Apical margins forward of anterolateral horns, deeply serrate with apices laterally directed 
(Fig. 12). Lateral margins fringed with long setae (Fig. 11); stem clothed with dense mat of fine 
setae. 

Egg — Length 0.27 mm, width 0.19 mm. Outline elongate oval. Micropylar row subequato- 
rial, orifices with circular raised lip (Fig. 26). Anterior and posterior third of chorionic surface 
covered with densely packed minute aeropyles; aeropyles of equatorial third more widely spaced 
(Figs. 25-26). Eclosion line absent. Collar stalked, rim irregularly incised, sides with several 
longitudinal carinae which terminate at shoulder (Fig. 27). 

Subgenital plate — Posterior margin rounded, plate narrow and tab-like, extending over 
anterior margin of sternum nine (Fig. 41). 

Material - MONTANA: Gallatin Co., Hyalite Creek, Squaw Creek Trailhead, 24 July 1979, 
B. P. Stark, K. W. Stewart, R. W. Baumann, 9 OC’, 4 9 (BPS). Granite Co., Butte Canyon Creek, 
25 June 1966, M. L. Miner, 2 CO (BYU). 

Known distribution —- AB, BC, ID, MT, WY, YK (Stark et al. 1986). 


Alloperla pilosa Needham & Claassen 


Alloperla pilosa Needham & Claassen, 1925. Holotype male, Boulder, Boulder Co., 
Colorado. 

Sweltsa pilosa: lies, 1966. 

Alloperla pilosa: Surdick, 1985. 


326 ENTOMOLOGICAL NEWS 


Figs. 7-12. Scanning electron micrographs of Alloperla epiprocts. 7. A. delicata, dorsal aspect. 8. 
A. delicata, nearly lateral aspect. 9. A. delicata, apex (ep = epiproct, If = lateral fringe, se = 
serrations, ho = anterolateral horns). 10. A. medveda, dorsal aspect. 11. A. medveda, nearly lat- 
eral aspect. 12. A. medveda, apex. 


Vol. 108, No. 5, November & December, 1997 327 


Epiproct — Length 0.218 mm, basal width 0.054 mm, mesal width 0.082 mm, apical width 
0.038 mm; length bare apical area 0.013 mm Dorsal aspect somewhat barrel shaped, surface 
covered with dense mat of setae except for basolateral fold (Figs. 5, 6). Stem clothed with dense 
mat of fine setae. 

Egg - Length 0.35 mm, width 0.25 mm. Outline elongate oval. Micropylar row subequato- 
rial, orifices with raised circular lip (Fig. 29). Chorionic surface with obscure hexagonal follicle 
cell impressions, anterior and posterior regions covered with densely packed aeropyles, equato- 
rial third with aeropyles more widely spaced (Figs. 28-29). Collar stalked, rim irregularly in- 
cised, sides with several longitudinal carinae (Fig. 30). 

Subgenital plate — Posterior margin acute, sides convergent. Posterior margin extending 
well beyond anterior margin of sternum nine (Fig. 40). 

Material - COLORADO: Clear Creek Co., Berthoud Pass, 5 August 1973, B. P. Stark, R. 
W. Baumann, 4 GC’, 2 9 (BPS). Larimer Co., Icy Brook, Loch Vale below Timberline Falls, 
Rocky Mountain National Park, 5 July 1988, B. C. Kondratieff, 2 OC’, 2 9 (CSU). same location, 
2 August 1988, B. C. Kondratieff, 4 0, 7 9 (CSU). 

Known Distribution — CO (Stark et al. 1986). 


Alloperla serrata Needham & Claassen 


Alloperla serrata Needham & Claassen, 1925. Holotype male, Moraine Lake, Alberta, 
Canada. 


Epiproct — Length 0.13 mm, width 0.15 mm. Dorsal aspect somewhat triangular (Fig. 15); 
surface bare except for basal fringe of posteriorly directed, long setae; fringe ca. 0.022 mm wide. 
Apex rounded, but with median gap and ca. 12 deep laterally directed serrations (Fig. 15). Stem 
clothed with dense mat of fine setae. 

Egg — Length 0.30 mm, width 0.22 mm. Outline elongate oval. Micropylar row subequato- 
rial, orifices with oval lip (Fig. 32). Chorionic surface densely and uniformly punctate with deep 
aeropyles (Figs. 31-32). Eclosion line absent. Collar stalked, rim irregularly incised, sides with 
several longitudinal carinae (Fig. 33). 

Subgenital plate — Posterior margin acute, outline triangular, margin extending to anterior 
margin of sternum nine (Fig. 39). 

Material — ALBERTA: Moraine Creek, Banff National Park, 27. July 1972, A. R. Gaufin, 
40 (BYU). MONTANA: Gallatin Co., Hyalite Creek, Squaw Creek Trailhead, 24 July 1979, B. 
P. Stark, K. W. Stewart, R. W. Baumannn, 4 C,5 Q (BPS). 

Known distribution — AB, AK, BC, ID, MT, WA, WY, YK (Stark et al. 1986). 


Alloperla severa (Hagen) 


Perla severa Hagen, 1861. Holotype (abdomen missing), Unga Island, Alaska. 

Alloperla elevata Frison, 1935. Holotype male, Floras Creek, Curry Co., Oregon. Syn. Ricker, 
1954. 

Alloperla thalia Ricker, 1952. Holotype male, Gallatin Co., Montana Syn. Ricker, 1954. 


Epiproct — Length 0.11 mm, basal width 0.033 mm, subapical width 0.054 mm. Dorsal 
aspect elongate, slightly inflated at the base, apex rounded but with shallow crenulations (Fig. 
13). Surface bare in apical half, densely pilose in basal half. Stem clothed with dense mat of fine 
setae. 

Egg - Length 0.31 mm, width 0.21 mm. Outline elongate oval. Micropylar row subequato- 
rial, obscure, orifices slanted, lip oval (Fig. 35). Chorion densely and finely punctate throughout 
(Figs. 34-35). Eclosion line absent. Collar stalked, rim irregularly incised, sides with several 
longitudinal carinae (Fig. 36). 


328 ENTOMOLOGICAL NEWS 


- 
.* ‘ 
at SS 
= 


Bi 
; = 4 ) 
4M) bh 70 ri 


Figs. 13-18. Scanning electron micrographs of Alloperla epiprocts and eggs. 13. A. severa, dor- 
sal aspect. 14. A. severa, lateral aspect. 15. A. serrata, dorsal aspect. 16. A. serrata, nearly lateral 
aspect. 17. A. fraterna, entire egg. 18. A. fraterna, chorionic detail showing micropyle. 


Vol. 108, No. 5, November & December, 1997 329 


Figs. 19-24. Scanning electron micrographs of Alloperla eggs. 19. A. chandleri, entire egg. 20. 
A. chandleri, chorionic detail showing micropyle. 21. A. chandleri, follicle cell impressions on 
anterior pole. 22. A. delicata, entire egg. 23. A. delicata, chorionic detail showing micropyle. 24. 
A. delicata, aeropyles on anterior pole. 


330 ENTOMOLOGICAL NEWS 


Figs. 25-30. Scanning electron micrographs of Alloperla eggs. 25. A. medveda, entire egg. 26. A. 
medveda, chorionic detail showing micropyle. 27. A. medveda, collar. 28. A. pilosa, entire egg. 
29. A. pilosa, chorionic detail showing micropyle. 30. A. pilosa, collar. 


Vol. 108, No. 5, November & December, 1997 331 


Figs. 31-36. Scanning electron micrographs of Alloperla eggs. 31. A. serrata, entire egg. 32. A. 
serrata, chorionic detail showing micropyle. 33, A. serrata, collar. 34. A. severa, entire egg. 35. 
A. severa, chorionic detail showing micropyle. 36. A. severa, collar. 


Ww 
Ww 
No 


ENTOMOLOGICAL NEWS 


Figs. 37-43. Female Alloperla subgenital plates. 37. A. chandleri. 38. A. delicata. 39. A. serrata. 
40. A. pilosa. 41. A. medveda. 42. A. fraterna. 43. A. severa. 


Vol. 108, No. 5, November & December, 1997 333 


Subgenital plate Posterior margin acute, outline triangular, margin reaching center of ster- 
num nine (Fig. 43). 

Material - MONTANA: Glacier Co., St. Mary’s River, 2 mi S Babb, 28 July 1966, A. R. 
Gaufin, 18 CO’, 43 O (BPS). Lincoln Co., Fischer River, Hwy. 2, 30 June 1973, A. R. Gaufin, 2 
(BYU). NEVADA: Elko Co., Secret Creek near Secret Pass, 15 June 1974, B. P. Stark 3 C,49 
(BPS). UTAH: Summit Co., Yellow Pine Creek, 8 mi E Kamas, 22 July, 1974, B. P. Stark, R. W. 
Clubb, 1 C (BPS). 

Known distribution - AB, AK, BC, CA, CO, ID, MT, NT, NV, OR, UT, WA, WY, YK 
(Stark et al. 1986). 


DISCUSSION 


The Alloperla of western North America include three sets of similar ap- 
pearing species pairs (A. chandleri-A. fraterna, A pilosa-A. severa and A. 
delicata-A. medveda) and one fairly distinctive species, A. serrata. The eggs 
and epiprocts of each pair are difficult to differentiate using light microscopy. 

Alloperla chandleri and A. fraterna have similar epiproct morphology and 
the presence or absence of a dorsal abdominal stripe may be variable. In our 
material, however, all A. fraterna specimens possessed the stripe, and all A. 
chandleri lacked it. Compared to A. fraterna, the epiproct of A. chandleri is 
longer and broader, the apical bare area is wider and more nearly truncate, and 
the dorsal constriction is much less conspicuous (Figs. 1, 3). In lateral aspect, 
the epiproct of A. chandleri is uniform in width, but in A. fraterna it is con- 
spicuously narrower at the base than at the apex (Figs. 2, 4). The eggs of both 
species are oval, collarless, and without distinctive chorionic sculpturing (Figs. 
17, 19). 

Although A. severa and A. pilosa are usually identified on the basis of size 
and abdominal striping, their epiprocts are difficult to distinguish with light 
microscopy. The dorsal shapes of the epiprocts are very similar, and it may be 
difficult to detect the shallow apical crenulations of A. severa (Figs. 5, 13). 
The epiproct of A. severa is shorter and narrower than in A. pilosa, but the 
most conspicuous difference is in the setation. A. pilosa has only a tiny bare, 
button-like apex to the epiproct (Fig. 5), whereas A. severa has at least the 
apical third bare (Figs. 5, 13). The eggs of these species have similar collars, 
and the chorionic surfaces of both are pitted throughout with aeropyles (Figs. 
28, 34). In A. pilosa the aeropyles are larger and more widely spaced in the 
equatorial area than in the polar areas, but in A. severa the aeropyles are rather 
uniform in size and spacing. 

The dorsal surface of the epiprocts of A. medveda and A. delicata are nearly 
bare, but scattered clusters of thick setae are located along the midline (Figs. 7, 
10). The epiprocts of both species have lateral setal fringes (Figs. 8, 11) and 
conspicuous anterolateral horns (Figs. 9, 12). In dorsal aspect, the epiproct of 
A. delicata is nearly oval, and the area between the anterolateral horns is some- 
what truncate (Fig. 9). In A. medveda, the dorsal aspect diverges to mid-length, 
then constricts to the apex, and a small triangular area projects from between 


334 ENTOMOLOGICAL NEWS 


the anterolateral horns (Fig. 12). A. delicata has a mid-dorsal abdominal stripe 
which is lacking in A. medveda. Eggs of A. delicata are collarless, but the eggs 
of A. medveda have a stalked collar (Figs. 22, 25). 

Alloperla serrata is easily distinguished from all other western Alloperla 
by the button-like epiproct, subequal in length and width, with deep marginal 
serrations (Fig. 15). The basal edge is covered with thick posteriorly-directed 
setae, and the anterior portion is bare. Eggs of A. serrata are oval with a stalked 
collar and large aeropyles (Figs. 31-33). 


ACKNOWLEDGMENTS 


We thank B. C. Kondratieff (Colorado State University) and R. W. Baumann (Brigham Young 
University) for their generous loan of specimens. We also thank R. F. Surdick and B. C. Kondratieff 
for their helpful reviews of an earlier draft of this manuscript. This study was supported in part 
by the Howard Hughes Medical Institute, Undergraduate Biological Sciences Education Pro- 
gram Grant # 71195-538901. 


LITERATURE CITED 


Baumann, R. W., A. R. Gaufin, and R. F. Surdick. 1977. The stoneflies (Plecoptera) of the 
Rocky Mountains. Mem. Amer. Entomol. Soc. 31:1-208. 

Frison, T. H. 1935. New North American species of the genus Alloperla (Plecoptera: 
Chloroperlidae). Trans. Amer. Entomol. Soc. 61:331-344. 

Hagen, H. A. 1861. Perlina, pp. 14-38. In Synopsis of the Neuroptera of North America: with a 
list of South American species. Smithsonian Misc. Coll. 4. 

Harper, P. P. and K. W. Stewart. 1996. Plecoptera, pp. 217-266. In R. W. Merritt & K. W. 
Cummins (Eds.), An introduction to the aquatic insects of North America, 3rd ed. Kendall 
Hunt, Dubuque, Iowa. 

Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich, 82. Walter de Gruyter, Berlin. 

Jewett, S. G. 1954a. New stoneflies (Plecoptera) from western North America. J. Fish. Res. 
Board Can. 11:543-549. 

Jewett, S. G. 1954b. New stoneflies from California and Oregon (Plecoptera). Pan-Pac. Entomol. 
30: 167-180. 

Jewett, S. G. 1959. The stoneflies (Plecoptera) of the Pacific Northwest. Oregon State Mono., 
Studies Entomol. 3:1-95. 

Needham, J. G. and P. W. Claassen. 1925. A monograph of the Plecoptera or stoneflies of 
America north of Mexico. Entomol. Soc. Amer., Thomas Say Found. 2: 1-397. 

Ricker, W. E. 1952. Systematic studies in Plecoptera. Indiana Univ. Pub., Sci. Ser. 18:1-200. 

Ricker, W. E. 1954. Nomenclatural notes on Plecoptera. Proc. Entomol. Soc. British Columbia 
51:37-39. 

Stark, B. P. and S. W. Szczytko. 1988. Egg morphology and phylogeny in Arcynopterygini 
(Plecoptera: Perlodidae). J. Kansas Entomol. Soc. 61:143-160. 

Stark, B. P., S. W. Szezytko, and R. W. Baumann. 1986. North American stoneflies (Plecoptera): 
systematics, distribution and taxonomic references. Great Basin Nat. 46:383-397. 

Surdick, R. F. 1985. Nearctic genera of Chloroperlinae (Plecoptera: Chloroperlidae). Illinois 
Biol. Mono. 54:1-146. 


Vol. 108, No. 5, November & December, 1997 335 


REVIEW OF THE NEARCTIC GENUS CERAMPHIS 
(COLEOPTERA: SCYDMAENIDAE)! 


Sean T. O’Keefe2 


ABSTRACT: The Nearctic genus Ceramphis (Coleoptera: Scydmaenidae) is reviewed and in- 
cludes a single species, C. deformata, from southern California and western Arizona. Habitus 
and male genitalic illustrations are included, as well as illustrations of the female genitalia and 
mouthparts. The phylogenetic placement of Ceramphis within Scydmaenini and as sister group 
of Scydmaenus is discussed. 


Taxonomy of the beetle family Scydmaenidae generally has been neglected 
for North America since the monograph by Casey (1897). Nearly all of the 
genera are in need of review or revision. Some, such as Lophioderus Casey 
(O’Keefe 1996), Euconnus Thomson (s.str) (A. Davies unpublished), and 
Papusus Casey (O’Keefe unpublished), include numerous undescribed spe- 
cies. Others, such as Chevrolatia Jacquelin du Val (O’ Keefe 1997a), Drastophus 
Casey, Noctophus Casey, Xestophus Casey, and Pycnophus Casey (O’ Keefe 
unpublished) include few or no new species. Ceramphis Casey falls into the 
latter category. This review is a continuation of my work on a revision of the 
Nearctic scydmaenid fauna (O’ Keefe 1996, 1997a, 1997b). 

Casey (1897) erected the genus Ceramphis to include a single interesting 
species of scydmaenid, Scydmaenus deformatus (Fig. 1), described by George 
Horn (1885) from southern California. Casey placed Ceramphis in the Eumicrini 
(= Scydmaenini) along with Eumicrus LaPorte (= Scydmaenus Latreille) and 
Acholerops Casey (= Scydmaenus Latreille) based on the presence of elongate 
metatrochanters, arcuate abdominal sutures, a large, vertical pygidium (Fig. 2), 
and short and rounded maxillary palpomere IV (Fig. 4). He justified placing S. 
deformatus in anew genus based on the presence of a metasternum that extends 
from side to side without a visible metepisternal suture, the conical pronotum, 
and the extent to which the prosternum is developed anterior to the procoxae. 
Franz (1985), upon reexamining the few specimens of C. deformata in the Horn 
collection (MCZC), confirmed Casey’s placement of this genus near Scydmaenus 
based primarily on the structure of the male genitalia. 

Casey described but did not illustrate the mouthparts and Franz described 
and inadequately illustrated the male genitalia. I am taking this opportunity to 
redescribe the species, improving upon Casey’s description of the mouthparts 
and Franz’s description of the male genitalia, and describing the female genita- 
lia for the first time. 


1 Received March 3, 1997. Accepted March 22, 1997. 


2 Dept. of Environmental Science, Policy and Management Division of Insect Biology, Univer- 
sity of California, Berkeley, CA 94720. 


ENT. NEWS 108(5) 335-344, November & December, 1997 


336 ENTOMOLOGICAL NEWS 


Ceramphis Casey 


Casey 1897: 538; Franz 1985: 177. 
Type species: Scydmaenus deformatus Horn (by monotypy) 


DIAGNOSIS. Members of Ceramphis can be identified by having an elon- 
gate metatrochanter, a conical pronotum (Fig. 1), a large, vertical, and exposed 
pygidium (Fig. 2), an antennal club composed of two antennomeres, both 
densely covered with short setae, and a posteriorly extended antennomere V 
(Fig. 3). 

Ceramphis deformata (Horn) 
(Figs. 1-9) 


Scydmaenus deformatus Horn 1885, pl. 5, fig. 11. 
Ceramphis deformata (Horn) Casey 1897:539; Csiki 1919:86; Leng 1920:91; Franz 1985:178, 
fig. 11. 


Lectotype female (number 3029) in Horn Collection, MCZC; NEW DESIGNATION. Type 
locality: Los Angeles, California. Label information: “Cal” typed white label; “Lectotype 3029” 
typed red label; “\S. deformatus Horn” handwritten; “MCZ type 34844” handwritten on red label; 
“Ceramphis det. W. Suter 77” handwritten; “Lectotype Q Ceramphis deformata Horn S. O’ Keefe 
des. °97” handwritten on red label. 


DESCRIPTION. Length 1.63-1.83 mm; reddish-brown throughout; vestiture sparse, pale in 
color, erect. 

Head. Rounded, as high as long, slightly longer than wide, 0.41-0.43 mm wide, widest at 
eyes; eyes median in position, convex, ovoid, more than 70 facets; vertex slightly raised with 
sparsely distributed long setae; temples with dense row of short setae behind row of sparsely 
spaced long setae; antennal bases nearly contiguous, separated by a thin septum, covered dor- 
sally by vertex; antennae (Fig. 3) slightly under two-thirds body length; antennomere | 
subcylindrical, slightly compacted dorsoventrally, gradually widened distally, twice as long as 
wide, distal end notched dorsally, interior and ventral surfaces densely covered with short setae; 
antennomere II conical, widest at distal end, 1.5 times longer than wide at distal end; antennomeres 
II-IV subconical, III 1.5 times longer than wide at distal end, IV as long as wide; antennomere V 
modified, posterior margin greatly expanded posteriorly and cup-shaped, twice as wide as IV; 
antennomeres VI-VII subquadrate, subequal, somewhat flattened, slightly wider than long; 
antennomere VIII subquadrate, as long as wide, slightly narrower than either VII or IX; 
antennomere IX subcylindrical, 1.5 times longer than wide, 1.5 times longer than VIII; 
antennomere X tear-drop shaped, widest at distal end, nearly twice as long as wide at distal end, 
nearly twice as long as IX; antennomere XI ovoid, widest near middle, nearly twice as long as 
wide, as wide as X; antennomeres X-XI with denser and shorter setation than preceding 
antennomeres. Maxillae (Fig. 4) with distal margins of galea and lacinia fringed with elongate 
setae; galea subquadrate; lacinia elongate, about three times longer than wide; maxillary palpomere 
II elongate, subcylindrical, four times longer than wide, slightly wider at distal end; maxillary 
palpomere III slightly longer than II, gradually and distinctly expanded distally, distal end twice 
as wide as basal end; maxillary palpomere IV small, rounded, as long as wide at base. Labrum 
(Fig. 5) short, transverse, lateral margins rounded; basal margin broadly emarginate; distal mar- 
gin slightly concave; with four elongate, three medium, and two short setae. Mandible (Fig. 6) 
with apex elongate, curved, with single tooth at base of incisor; base broad; prostheca present, 


Vol. 108, No. 5, November & December, 1997 


Figure 1. Ceramphis deformata (Hom). Habitus, dorsal view. Scale bar = 0.50 mm. 


337 


338 ENTOMOLOGICAL NEWS 


extended entire length from base of mandible to base of incisor, setae elongate, dense. Labium 
(Fig. 7) with mentum subquadrate, with two elongate setae; palpomere II cylindrical, slightly 
longer than broad; palpomere III elongate, tapered at apex, five times longer than wide at base; 
single long and two short setae at apex of paplomere II. 

Pronotum. Distinctly conical, widest at base, sides evenly narrowed to anterior end; 0.48- 
0.53 mm wide at base, 0.48-0.54 mm long; no basal impression or foveae on dorsal posterior 
margin; pronotal vestiture short, sparse, with dense row of short setae along anterior lateral mar- 
gin from dorsum to venter. 

Elytra. Scutellum visible; elytra entire, 0.95-1.10 mm long, 0.85-1.03 mm wide, strongly 
convex, broadly rounded at apex, setation long, erect, moderately dense; without basal fovea; 
humeri evident, but weak. 

Legs. Moderately long; distal half of femora clavate, with short, dense setation on inner 
margins. Protrochanters with anterior margin sharp, anteriorly rounded. Hind femora and coxae 
widely separated by trochanters. 

Venter. Prosternum flat, glabrous, narrowed anteriorly, 1.5 times wider than long, with patch 
of long setae at posterior lateral margin near coxal bases; procoxal margin expanded medially to 
cover bases of profemora; mesepimeron fused to body; mesocoxae separated by carinate 
mesosternum; metasternum large, convex, with moderately dense short setation; hindcoxae sepa- 


MME EM jay 
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CNG ty, 


fr“ 4 


« a rane i" ; , 3 ee z << es eee = = = 
Breau a cANneRS NN Be ene 
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3 Rh 


Figures 2-3. Ceramphis deformata (Hom). Fig. 2. Habitus, left lateral view. Scale bar = 0.50 
mm. Fig. 3. Right antenna, dorsal view. Scale bar = 0.20 mm. 


Vol. 108, No. 5, November & December, 1997 339 


rated by at least their width, subtransverse, not extended to lateral margin; abdominal segment I 
as long as II and III combined; segments II-IV equal in length; segment V half as long as IV; 
segment VI 1.5 times longer than II; sutures rounded apically; pygidium large, convex, vertical 
(Fig. 2). 


Figures 4-7. Ceramphis deformata (Horn). Fig. 4. Left maxilla, posterior view. Fig. 5. Labrum, 
anterior view. Fig. 6. Left mandible, posterior view. Fig. 7. Labium, ventral view. Scale bar = 
0.10 mm. 


340 ENTOMOLOGICAL NEWS 


Male Genitalia. (Fig. 8). Parameres present, fused together along distal half, difficult to 
distinguish from median lobe, extended well beyond median lobe. Median lobe subcylindrical, 
slightly expanded basally, truncate distally, lightly sclerotized ventrally. Endophallus cylindri- 
cal, extended beyond median lobe half length of median lobe, terminated in elongate, slightly 
curved, acuminate process. Ventral-basal portion of median lobe surrounded by sclerotized plate. 

Female Genitalia. (Fig. 9). Ovipositor reduced, faintly sclerotized. Proctiger subtrapezoidal, 
flat, slightly narrowed at distal end. Single sclerotized structure laterally, either paraproct, or 
valvifer, or fused paraproct + valvifer; visible dorsally only at lateral sides, each extended to 
cover one-third of ventral region; coxites absent. Spermatheca spherical; spermathecal duct very 
long, 4 times diameter of spermatheca. 


8a 


8C 


Figure 8. Ceramphis deformata (Horn). Male genitalia; a, dorsal view; b, ventral view; c, right 
lateral view. 


Vol. 108, No. 5, November & December, 1997 341 


Figure 9. Ceramphis deformata (Horn). Female genitalia; a, dorsal view; b, ventral view; c, right 
lateral view. Scale bars = 0.10 mm. 


SPECIMENS EXAMINED. 33. ARIZONA. Graham Co.; 18 specimens, Aravaipa Canyon, 
3,050 ft, 9-II-1977, D.S. Chandler, under rock, (WRSC, FMNH). State record only: | specimen, 
Bolter coll. (INHS); 8 specimens, C.V. Riley coll., (USNM); 1 specimen, (USNM). CALIFOR- 
NIA. Los Angeles Co.; 1 specimen, Los Angeles, Hubbard and Schwartz coll., (USNM). State 
records only: 4 specimens (including lectotype), Horn collection (MCZC). 


DISTRIBUTION. Known from southern California (Los Angeles) and Ari- 
zona. I have been able to locate only about three dozen specimens of C. 
deformata in museum collections. I could find no differences between speci- 
mens from southern California and those from Arizona. 


342 ENTOMOLOGICAL NEWS 


HABITAT AND COLLECTION METHODS. Habitat information is asso- 
ciated only with specimens collected by Don Chandler. These were collected 
underneath rocks in association with Solenopsis sp. (Hymenoptera: Formicidae). 
Specimens of the latter are mounted on points under Chandler’s specimens of 
Ceramphis. 


COMMENTS. Horn (1885) neglected to designate a holotype specimen 
for Ceramphis deformata, nor was one designated by Casey (1897) or Franz 
(1985); therefore, of the four specimens in the type series in the MCZC, I 
chose as the lectotype the specimen with both the handwritten label “S. 
deformatus Horn” and the typed label “Lectotype 3029”. This specimen was 
not in the best condition (head + prothorax separated from the rest of the body 
and right antennomeres VII-XI are missing) but has the distinct characteristics 
of antennomere V, body setation, etc of the others in the series. 

Three statements of Casey (1897) need minor clarification. First, he stated 
that the antennal club is “3-jointed and finely pubescent as in Eumicrus” (Casey 
1897:539). In Ceramphis only antennomeres X and XI] are distinctly enlarged 
and densely covered with short setae. Antennomere IX, though slightly larger 
than VIII and having short setae, is not like those of Scydmaenus, in which 
antennomere IX is distinctly larger than VIII and is as densely covered with 
short setae as either antennomeres X or XI. Second, Casey stated that the 
mesosternum is “apparently not carinate” (Casey 1897:539). In all of the speci- 
mens I have examined, the mesosternum is distinctly carinate, though not so 
much as in other Scydmaenidae. Third, Casey stated the antennae were three- 
fourths as long as the body. In actuality, the antennae are slightly under two- 
thirds the body length. Except for the above listed discrepancies, Casey’s de- 
scription is fairly accurate. 


DISCUSSION 


Ceramphis clearly belongs in the Scydmaenini based on the putative 
synapomorphies of |) metatrochanters elongate, 2) pygidium vertical, 3) 
antennomeres forming antennal club compact and densely covered with short 
setae, 4) antennomeres I notched at distal margin, 5) abdominal sutures curved, 
and 6) maxillary palpomeres IV short and rounded. The exact position of 
Ceramphis within the Scydmaenini is uncertain. 

The tribe Scydmaenini currently includes seven genera: Adrastia Broun 
(1881), Ceramphis Casey, Clavigeroscydmaenus Franz (1986), Eudesis Reitter 
(1881), Palaeoscydmaenus Franz (1975), Pseudoeudesis Binaghi (1948), and 
Scydmaenus Latreille (1802). Scydmaenus is further divided into 31 recognized 
subgenera (Newton, A.F., Jr. and H. Franz, unpublished, World Catalog of the 
Genera of Scydmaenidae (Coleoptera), 23 Oct., 1996 draft). 


Vol. 108, No. 5, November & December, 1997 343 


Palaeoscydmaenus, Eudesis, and Pseudoeudesis are most distantly related 
to Ceramphis, based on the form of the male genitalia. In these three genera, the 
genitalia of males are bulbous and have free parameres (Binaghi 1948, Franz 
1975). In Adrastia, Ceramphis, and most Scydmaenus, the male genitalia are 
elongate and have fused parameres. Males are unknown for Clavigeroscyd- 
maenus (Franz 1986). 

Members of Ceramphis have the autapomorphies of 1) pronotum distinctly 
conical and 2) antennomere V with a posteriorly directed extension. 

The sister group of Ceramphis is to be found within the Scydmaenini; most 
likely it will be either Scydmaenus or a clade included within Scydmaenus. To 
date, there has been no phylogenetic treatment of the 31 recognized subgenera 
of Scydmaenus. Figured male genitalia of members of the Australian subgenera 
Scottiscydmaenus Franz and Heterognathus King (Franz 1975) are quite simi- 
lar in shape to those of Ceramphis, but this similarity may represent a sym- 
plesiomorphy. Unfortunately, there are approximately 450 species currently 
placed as Scydmaenus incertae sedis; therefore, the probable sister group taxon 
of Ceramphis cannot be recognized until further study of Scydmaenus is under- 
taken. For now, I suggest considering Scydmaenus as the sister taxon to 
Ceramphis. 


ACKNOWLEDGMENTS 


I thank David Furth, Phil Perkins, and Michael Kelley (Museum of Comparative Zoology, 
Cambridge, Massachussets, MCZC), Gloria House (United States National Museum, Washing- 
ton, D.C., USNM), Kathleen Methven (Illinois Natural History Museum, Champaign, Illinois, 
INHS), Alfred Newton (Field Museum of Natural History, Chicago, Illinois, FMNH), and Walt 
Suter (Carthage College, Kenosha, Wisconsin, WRSC) for arranging loans of specimens, and 
Michael Kelley for allowing me to examine the lectotype from the LeConte and Horn Collection 
(MCZC). I thank John Doyen, Dave Kavanaugh, Felix Sperling and two anonymous reviewers 
for helpful comments on this manuscript. I also thank Kevin Wiseman for illustrating figure 1. 
The work was funded in part by California Agriculture Experiment Station Grant to Felix Sperling, 
Sigma Xi, Vice Chancellors Grants in Aid, and the Margaret C. Walker Fund. 


LITERATURE CITED 


Binaghi, G. 1948. Revisione delle specie del genere Eudesis e descrizione di un nuovo genere 
(Col. Scydmaenidae). Bollettino della Societa Entomologica Italiana 78: 34-40. 

Broun, T. 1881. Manual of the New Zealand Coleoptera. Part I]. Colonial Museum and Geologi- 
cal Survey Department, Wellington. pp. 653-744. 

Casey, T. 1897. Coleopterological notices, VII. Annals New York Acad. Science 9:285-684. 

Csiki, E. 1919. Scydmaenidae, Pars 70. in: Coleopterorum Catalogus. ed. S. Schenkling. W. 
Junk, Berlin, 106pp. 

Franz, H. 1975. Revision der Scydmaeniden von Australien, Neuseeland und den benachbarten 
Inseln. Denkschriften Osterreichische Akademie der Wissenschaften, Mathematisch- 
Naturwissenschaftliche Klasse 118: 1-312. 


344 ENTOMOLOGICAL NEWS 


Franz, H. 1985. Revision CASEYscher Scydmaenidentypen. Sitzungsberichte der Osterreichische 
Akademie der Wissenschaften, Mathematisch- Naturwissenschaften, Abteilung I, 194(6- 
10):149-186. 

Franz, H. 1986. Monographie der Scydmaeniden (Coleoptera) von Madagaskar (mit Ausschluss 
der Cephenniini). Denkschriften Osterreichische Akademie der Wissenschaften, Mathe- 
matisch Naturwissenschaftliche Klasse 125: 1-393. 

Horn, G. 1885. Contributions to the Coleopterology of the United States, No. 4. Trans. Amer. 
Entomol. Soc. 12:128-172. 

Latreille, P.A. 1802. Histoire Naturelle, Générale et Particuliére des Crustacés et des Insectes. 
Familles Naturelles des Genres. Vol. 3, F. Dufart, Paris, pp. 13-467. 

Leng, C. 1920. Catalogue of the Coleoptera of America, north of Mexico. John D. Sherman, 
Mount Vernon, NY. 470pp. 

O’Keefe, S.T. 1996. Revision of the Nearctic genus Lophioderus Casey (Coleoptera: 
Scydmaenidae). Thomas Say Publications in Entomology: Monographs; Entomol. Soc. Amer. 
Lanham, Maryland, USA. 97 pp. 

O’Keefe, S.T. 1997a Revision of the genus Chevrolatia Jacquelin du Val (Coleoptera: 
Scydmaenidae) for North America. Trans. Amer. Entomol. Soc. In press. 

O’ Keefe, S.T. 1997b. Euconnus longiceps Fall, an odd ant-like stone beetle from the Pacific 
Northwest (Coleoptera: Scydmaenidae). Coleopt. Bull. In press. 

Reitter, E. 1881. Bestimmungs-Tabellen der europdischen Coleopteren. V. Enthaltend die 
Familien: Paussidae, Clavigeridae, Pselaphidae und Scydmaenidae. Verhandlungen der 
Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien 31: 443-592, pl. 19. 


SOCIETY MEETING OF APRIL 24, 1997 


Tom Wood 
University of Delaware, Newark, DE 


Tom Wood began with an illustrated introduction to the stunning and beautiful diversity of 
membracids and a recording of their recently discovered “sounds” or, more correctly, substrate 
vibrations, which, recorded and amplified, could be mistaken for elephant love calls or Bronx 
cheers. He went on to discuss his twenty-five years’ research on speciation by host plant shifts in 
Enchenopa binotata. As far back as 1964, he found that this species feeds on perhaps a dozen 
different trees or shrubs in five different families. The nymphs from different hosts are noticably 
different, and closer examination revealed at least eight sets of distinguishing features among 
hostplant races, including aggregating habits and chemical composition of the egg froth. 

When he began his experiments, he found females clearly prefer to lay eggs on the hostplant 
they grew up on and that mating is similarly mostly confined to individuals from the same host. 
Furthermore, electrophoresis revealed that populations are genetically differentiated. Tom won- 


dered how the isolation was maintained. 
(continued on page 371) 


Vol. 108, No. 5, November & December, 1997 345 


ADVENTIVE ONTHOPHAGUS (COLEOPTERA: 
SCARABAEIDAE) IN NORTH AMERICA: 
GEOGRAPHIC RANGES, DIAGNOSES, AND NEW 
DISTRIBUTIONAL RECORDS! 


E. Richard Hoebeke2, Kyle Beucke> 


ABSTRACT: Five Old World species of the dung beetle genus Onthophagus (nuchicornis, 
depressus, bonasus, gazella, and taurus) are presently recorded in the United States, Canada, and 
Mexico. The current North American range of each, based on numerous literature records, speci- 
mens from private collections, from the insect collections of Cornell University and the Carnegie 
Museum of Natural History, and our own collecting, is mapped. A brief diagnosis, habitus pho- 
tographs, and illustrations of other diagnostic features are provided to aid in accurate identifica- 
tion of these adventive species. We also report the presence of O. gazella in 2 new States in the 
U.S. (Kansas, Tennessee), and of O. taurus in 6 new states in the U.S. (Missouri, New York, 
Pennsylvania, Ohio, Tennessee, West Virginia). 


Species of dung-burying beetles (Scarabaeidae: Scarabaeinae) have been 
intentionally introduced into North America to assist the native dung beetle 
fauna in their efficaceous and rapid removal of livestock feces from pastureland 
(Fincher 1981, Hunter and Fincher 1985). Some of the adventive species of 
the genus Onthophagus Latreille are quickly becoming the most dominant 
members of the dung beetle fauna in pastures in the United States, and the 
geographic range of these species is changing with their dispersal into new 
habitats. 

Prior to 1970, 37 species of Onthophagus were recorded in the United 
States and Canada (Howden and Cartwright 1963), including 2 species3 addi- 
tional species (taurus, gazella, and bonasus) have been either intentionally 
released or discovered in the United States. 

Between 1985-87, 3 other exotic species of Onthophagus [sagittarius (F.), 
binodis Thunberg, and nigriventris d’Orbigny] have been released in Texas to 
aid in the control of the horn fly, but these have not become established (refer 
to Fincher 1990) and they are not discussed further in this paper. 

Of the 5 species of non-indigenous Onthophagus now recorded in North 
America, 4 (nuchicornis, depressus, taurus, and gazella) are securely estab- 
lished and the status of 1 (bonasus) is equivocal. Here, we review and map the 


1 Received February 22, 1997. Accepted April 23, 1997. 
2 Department of Entomology, Cornell University, Ithaca, New York 14853-0901. 
35 Ivy League Lane, Stony Brook, New York 11790. 


ENT. NEWS 108(5) 345-362, November & December, 1997 


346 ENTOMOLOGICAL NEWS 


known geographic distribution of these exotic Onthophagus in the United States, 
Canada, and Mexico based on available records in the literature, and on new 
data gleaned from specimens examined in the private collections of Kyle 
Beucke, Kipling W. Will, Robert A. Androw, and Darren A. Pollock, and from 
the collections of Cornell University (Ithaca, NY) and the Carnegie Museum 
of Natural History (Pittsburgh, PA). We also provide a brief diagnosis and 
habitus illustration of each of these species to aid in their identification. 

The following acronyms are used to indicate deposition of specimens: pri- 
vate collections of Kyle Beucke (KBC), Kipling W. Will (KWC), Robert A. 
Androw (RAAC), and Darren A. Pollock (DAPC); Cornell University Insect 
Collection (CUIC), and Carnegie Museum of Natural History (CMNHC). 


SPECIES ACCOUNTS 
Onthophagus nuchicornis (L.) 


Geographic Range: Onthophagus nuchicornis, acommon and widespread 
dung beetle occuring throughout much of Europe, the British Isles, and west- 
ern Asia, is also widely distributed in North America. All previously recorded 
North American distributional data (taken from Howden and Cartwright 1963; 
Hatch 1953, 1971; Macqueen and Beirne 1975; Morris 1983; Cervenka and 
Moon 1991; and McNamara 1991) and additional locality records taken from 
voucher specimens in the Cornell University Insect Collection and from the 
private collections of D. A. Pollock and R. A. Androw are mapped in Figure 1. 

This Old World dung beetle is firmly established in western Canada (Alberta 
and British Columbia), in the northwestern United States (Idaho, Montana, 
and Washington), in eastern Canada (Ontario, Quebec, Newfoundland, New 
Brunswick, and Nova Scotia), and in a vast area of the northeastem United 
States (Connecticut, Maine, Maryland, Massachusetts, Michigan, Minnesota, 
New Hampshire, New Jersey, New York, Pennsylvania, Rhode Island, Ver- 
mont, and Wisconsin). McNamara (1991) also recorded O. nuchicornis from 
the prairie province of Manitoba. 

This Palearctic species is believed to have been accidentally introduced 
into northeastern North America prior to the 1840s. Melsheimer (1844) de- 
scribed the species Onthophagus rhinoceros from Pennsylvania which was 
later found to be a junior synonym of O. nuchicornis, the Old World species. 
Henshaw (1887) reported this species being taken from cattle droppings in the 
Magdalen Islands in the Gulf of the St. Lawrence in 1881. Later, specimens of 
O. nuchicornis, “doubtless introduced with ballast refuse,” were captured near 
Camden, New Jersey (Bland 1889). By the late 1890s and early 1900s, it was 
reported in the Canadian Maritimes (New Brunswick, Nova Scotia) and Que- 
bec (Evans 1899, Roy 1899, Fletcher 1904). Earliest records of O. nuchicornis 


347 


Vol. 108, No. 5, November & December, 1997 


‘POLIOWY UNON Ul syusoo1yonu sn8vydoyiuC JO UOINQUISIP UMOUY “| “SI4 


C1 si 


Ret 


34 ENTOMOLOGICAL NEWS 


m Ge Pact Northwest molade collections made m southeastern Bntish Co- 
5 2 72 —— a Se "ll seesaw oat OT — Ss ee = a 
lmonies me 1945 (Gatch 1993). The ocommence of O. muckicornis m the Pacific 


cast sEesests mualaple mirodactons. Brown (1940) 


Sy Of Ge cally Sical Gi GES MAMNSTAM Specs Mm cast- 


38.1 mm (males and females). Coloration. Head 


DvEaa ] suriaces. and appendasss black 


Remarks: Orthovheras muctocorms (Fiz. 2) can be Gesimeurshed from other 
North Amencan species by ms moderaiely lee size (6-8 mm) and is black 
head and promote. and tan elytra motied wah black (Fic. 2 

Mimor gngies can be recopmized by the smele. very Gtsimctive. cylmdnica 
mcchan hore amsme irom the wertex of the bead behmd the eyes (Fig. 3). but 


fs mechan hom ss barely madscated m mamor males. In the female. the pronota 


soutien coe Mineman gemma Giicues cnet ceestliee ameell am meena anette a = Sue = 
DELS tS 83 COVES EAD 9D Mais 2a 2 Sota COMA! Prolperance propecis 


bm © Staveiy, 3 VES D. Poliok 1 AB. Car ©) (DAPC) BRITISH COLDMBI4 
ttre £13-1557_G. Stace Sufh, + £ A Deris CLIC) ONTARIO. Guvil. 1¢- VL 
O50. 11-Will-1982 LL Pechuman (©) (0050) ONITED SIATES: INDIANA: Padasiz Co 
jaspeilasis St P_. 2 VE 1988 FP A Antrow 2A WM Bramam(1)(RAAC) MICHIGAN Alleoan 
Co. Saveawck 22-9972 ER Bosak (CUI). Gopebir Co, Wateramest, 4 VIL-1975,1. 
K Letter O) (CUIC). Kalamezep Co. Guill Lake Bal. Ste, VIL-1979 (3) (CUIC). MINNE- 


301+ Buppard Co. Suitpart, 3 41-1979 S-1-1979_17-¥- 1980 (4) BAAC) NEW HAMP- 


rir 


iateral aspect. bode ine fo: Fig. 2 = 


r 


Oninupnigus nucnivornis. 2. Gorsal Gaius, mayor male. 3. mead, mayor male. ironic- 
5.0 mm 


Vol. 108, No. 5, November & December, 1997 349 


SHIRE: Strafford Co., W. Milton, no date (1) (CUIC). MARYLAND: Worcester Co., Assateague 
Isl., 11-V-1957, R. G. Beard (3) (CUIC). NEW YORK: Cayuga Co., Sterling, 18-V-1967, D. G. 
Nielson (1) (CUIC). Essex Co., Mt. MacIntyre top, 19-VI-1941, 27-VI-1941, H. Dietrich (3) 
(CUIC). Suffolk Co., Montauk Pt., no date (1) (CUIC); Shelter Isl., 14-VI-1944, R. Latham (1) 
(CUIC); Stony Brook, Scholar Field, 23-V-1980, J. S. Miller (2) (CUIC). VERMONT: Addison 
Co., no locality, 19-VII-1963, R. A. Morse (1) (CUIC). WASHINGTON: Thurston Co., Rocky 
Prairie, 4 mi. S. Tenino, 10-IV-1976, R. Hagen (1) (CUIC). 


Onthophagus depressus Harold 


Geographic Range: Onthophagus depressus, a species of dung beetle 
native to South Africa, was first collected in the United States in 1937 near 
Vidalia (Toombs Co.), Georgia (Cartwright 1938). Its pathway into North 
America is unknown. A decade after its initial detection, it was discovered near 
Lake Placid (Highlands Co.), Florida (Robinson 1948). By 1995, O. depressus 
was collected in cattle and horse dung, pitfall traps with swine feces as bait, and 
blacklight traps in 27 counties of Georgia and in 5 counties of South Carolina 
(Hunter and Fincher 1996). Harpootlian (1995) also listed O. depressus from 
Aiken and Jasper counties, South Carolina. A collection of O. depressus in 
Torreya St. Park (Liberty Co.), Florida, in June 1982 (reported below), suggests 
that the species is perhaps more widely distributed between the Georgia/South 
Carolina sites in the north and the Highlands Co. locality in peninsular Florida. 
All available distributional records in the literature, and records listed below, 
are mapped in Figure 4. 

Diagnosis: Length. 6.0-7.7 mm (males and females). Coloration. Brownish 
black to black; antennal clubs brown, legs dark reddish brown. 

Remarks: Onthophagus depressus (Fig. 5) can be separated from all other 
North American Onthophagus by the sharply emarginate, bidentate, transversely 
tuberculate clypeus (Fig. 5) and by the rough and granular appearance of the 
pronotum created by the surface with closely-spaced, crescent-shaped punc- 
tures with an elongate flattened tubercle projecting toward their center from the 
anterior margin (Fig. 6). Additional distinguishing characters include the short, 
flattened, stubby yellow setae projecting from each puncture, and the heavily 
punctate tuberculate elytra. 

Males and females exhibit no obvious secondary sexual characteristics. In 
the female, the ventral abdominal segment before the pygidium is much longer 
along the midline as compared to that of the male. The species is keyed in 
Howden and Cartwright (1963). 


Additional United States Records: UNITED STATES: FLORIDA: Highlands Co., Archbold 
Biol. Sta., 18-IV-1969, L. L. Pechuman (1) (CUIC); Archbold Biol. Sta., 8 mi. S. of Lake Placid, 
30-III- 1979, G. Mullen (4) (CUIC); Archbold Biol. Sta., 24-27-III-1984, J. S. Miller (5) (CUIC); 
9-VI-1990, 13,14-VI-1990, R. A. Androw, G. Keeney, and M. Archangelsky (4) (RAAC). Lib- 
erty Co., Torreya St. Pk., 2-VI-1982, M. Brattain (2) (RAAC). 


ENTOMOLOGICAL NEWS 


350 


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Fig. 4. Known distribution of Onthophagus depressus in North America. 


Vol. 108, No. 5, November & December, 1997 351 


Figs. 5-6. Onthophagus depressus. 5, dorsal habitus, male. 6, close-up of pronotal punctation. 
Scale line for Fig. 5 = 5.0 mm. 


Onthophagus bonasus (F.) 


Releases: Onthophagus bonasus, an \ndo-Asian dung beetle native to 
Pakistan, southern Afghanistan, India, Cambodia, Ceylon, and Burma (Balthasar 
1963), was colonized in the laboratory in 1979 and first released at a site in 
east-central Texas (an owner-cooperator cattle ranch in Burleson Co.) during 
the winter of 1980-81 (Fincher 1986, Fincher and Hunter 1989). Additional 
releases of O. bonasus were made in other east-central Texas localities in the 
early 1980s: at one site in Brazos County, July to September, 1982 (Coulson 
1992), and at one site in Grimes County, April to September, 1983 (Coulson 
1994). Releases at unspecified sites in Georgia took place in 1984 (Fincher 
1986). Fincher and Hunter (1989) concluded that the establishment of O. bonasus 
in the United States is equivocal, and that it “has not been recovered more than 
one year after release in any area.” 

Diagnosis: Length. 12-17 mm (males and females). Coloration. Testaceus 
yellow, with the head, pronotum (except a narrow pale margin at the sides and 
base), the elytral suture, the median portion of the metasternum, the upper 
surface of the femora and tibiae, and a large patch on the lower side of each 
femur, greenish-black. The entire surface is suffused with a slight metallic 
luster. 

Remarks: Adults of O. bonasus (Fig. 7) are very similar in shape and 
coloration to those of O. gazella (discussed below), but in general are larger in 
size (12-17 mm vs. 8-13 mm). Onthophagus bonasus can be further distin- 


352 ENTOMOLOGICAL NEWS 


guished by the vertex of the head of major males and females bearing a pair of 
horns (Figs. 7 and 8), forming a backwardly directed crescent, the base slightly 
flattened and granulate, and each horn with a slight basal tooth at the inner 
edge; by a short, acute, erect horn in the middle (Fig. 7) of the frons (forehead) 
in both sexes (median horn absent in male O. gazella); and by the pronotum 
with a slight median groove, and a minute tubercle on each side. Female O. 
gazella bear no horns on the vertex of the head, only a strongly elevated trans- 
verse carina between the eyes. 


Figs. 7-8. Onthophagus bonasus. 7, dorsal habitus, male. 8, head, male, fronto-lateral aspect. 
Scale line for Fig. 7 = 5.0 mm. 


Onthophagus gazella (F.) 


Releases, Establishment, and Geographic Range. Onthophagus gazella, 
an Afro-Asian dung beetle native to much of the hotter, drier parts of Africa 
south of the Sahara, and ranging into Madagascar, Asia Minor, India and Ceylon 
(Balthasar 1963, Tyndale-Biscoe 1990), was first introduced into North America 
in 1972 with releases in Victoria and Kleberg Counties, Texas (Blume and Aga 
1978). This exotic dung beetle was also imported, mass-produced, and released 
at various sites in central and southern California (Anderson and Loomis 1978, 
Legner 1986, Kohlmann 1994). Since its initial introduction into Texas, O. 
gazella has successfully radiated from the release sites and now occurs through- 
out a vast portion of the southern two-thirds of Texas and the southern tier of 


353 


Vol. 108, No. 5, November & December, 1997 


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SS; a ph 


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ey 


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s 


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Fig. 9. Known distribution of Onthophagus gazella in the United States and Mexico. Stars de- 


note release sites. 


354 ENTOMOLOGICAL NEWS 


states from California to Georgia (Fincher et al. 1983, Fincher 1990) (see Fig. 
9). Additional releases were also made in Arkansas, Georgia, and Mississippi, 
and by 1981, this dung beetle was securely established in California, Georgia, 
Louisiana, and Texas (for specific release and recapture localities see Fincher 
et al. 1983). By the end of 1983, O. gazella had been either collected from 
cattle dung, in pitfall traps baited with swine feces, or in a UV light trap from 
new additional counties in Oklahoma, Arkansas, Mississippi, Alabama, and 
Florida (Downie 1984, Hunter and Fincher 1985). Beginning in 1987, O. gazella 
was released by the New Jersey Department of Agriculture at various sites in 6 
counties in northern and southern New Jersey, but was never recovered (R. 
Chianese, pers. commun.). 

By late 1981, O. gazella had expanded its range very rapidly and started 
dispersing into the northeastern Mexican states of Coahuila, Nuevo Leon, and 
Tamaulipas (Fincher et al. 1983, Lago et al. 1984, Barbero and Lopez-Guerrero 
1992). At an astounding pace, it has since invaded a vast area along the Atlan- 
tic and Pacific coasts of Mexico (Kohlmann 1994) and by 1988 it had spread 
into the states of Veracruz, Durango, Guerrero, and Jalisco (Moron et al. 1988, 
Zunino and Halffter 1988, Rivera-Cervantes and Garcia-Real 1991, and Barbero 
and Lopez-Guerrero 1992). Thomas (1993) reported O. gazella from sites in 
lowland rain forest and tropical deciduous forest in the southernmost Mexican 
state of Chiapas. By 1987, O. gazella had even penetrated into Guatemala 
(Kohlmann 1994). 

Diagnosis: Length. 8-13 mm (males and females). Coloration. Very simi- 
lar to O. bonasus; testaceus yellow, with the head, pronotum (except a narrow 
pale margin at the sides and base), the median part of the metasternum, the 
front tibiae, the greater part of the middle and hind tibiae, and a large oval spot 
on the lower surface of the four posterior femora, greenish- or coppery-black. 
The extreme edges of the pronotum, elytra, and all segments of the body and 
legs are also dark. 

Remarks: In shape and coloration, O. gazella (Fig. 10) is extremely simi- 
lar to O. bonasus. but can be separated by being slightly smaller in body size 
(8-13 mm vs. 12-17 mm) and by the lack of a short, acute, erect horn in the 
middle of the forehead of the male (present in male O. bonasus). In the male of 
O. gazella, the vertex bears a pair of slender horns (Figs. 10 and 11), curving 
outward and inclined a little backward, slightly flattened at the base and not 
united; the front of the pronotum is very smooth, glossy, and the declivity is 
crowned by two minute prominences separated by a slight groove. In the fe- 
male, the frons bears a strongly elevated, transverse carina; the front of the 
pronotum is vertical in the middle and produced on each side forming a pair of 
strong, slightly divergent, blunt processes (Fig. 12). 

Additional United States and Mexican Records: All previously reported 
locality records for O. gazella (releases and recaptures) in the literature and 


Vol. 108, No. 5, November & December, 1997 355 


Figs. 10-12. Onthophagus gazella. 10, dorsal habitus, male. 11, head, male, fronto-lateral aspect. 
12, head and pronotum, female, fronto-lateral aspect. Scale line for Fig. 10 = 5.0 mm. 


new and additional records given below are mapped in Figure 9. Specimens 
provided by K. W. Will (Cornell University), R. A. Androw (Carnegie Mu- 
seum) and those sorted from unidentified scarabs in the CUIC provide the 
basis for 2 new state records for the United States [marked with an asterisk 
(*)]. These new distributional data, and other locality records, are as follows: 


356 ENTOMOLOGICAL NEWS 


UNITED STATES: ARIZONA: Pima Co., Continental, 23-30-VII-1992, J. & M. Huether 
(1) (CUIC); 31 km. NW Tucson, 600m., 29-VIII-1988, J. E. Rawlins & S. Thompson (1) 
(CMNHC). Santa Cruz Co., Rock Corral Canyon, 2 mi. SW - 3 Tumacacori, 22-VII-1992, G. C. 
Eickwort (3) (CUIC); 3 km. N. Nogales, 6-1X 1992, L. Wismann (11) (CMNHC). FLORIDA: 
Highlands Co., Archbold Biological Station, 25-III/S-IV-1984 (1) (CUIC); Lake Placid, 22-III- 
1988, M. Ishli (1) (CUIC). Levy Co., Manatee Springs St. Pk., 15-VI-1990, R. A. Androw, G. 
Keeney, M. Archangelsky (1) (RAAC). Seminole Co., Longwood, 5-VII-1987 (3) (RAAC). *KAN- 
SAS: Meade Co., 3 mi. W. Meade, 23-IX-1990, Androw and Brattain (1) (KWC). LOUISIANA: 
Allen Parish, West Bay Game Area, 4-VI-1985, D. Sundberg (2) (RAAC). St. Tammany Parish, 
Slidell, 4-VI-1983, D. Sundberg (5) (RAAC). MISSISSIPPI: Grenada Co., Grenada, 26-VI-1986, 
R. A. Androw (1) (RAAC). Panola Co., no locality, 12-13-VII-1987, K. E. M. Galley, at black 
light (2) (CUIC). Stone Co., 8.4 mi. E. of Wiggins, on Rte. 26, 24-V-1995, K. Will & R. Androw 
(2) (CUIC). NEW MEXICO: Eddy Co., 10-15 mi. S. Carlsbad, 16-VII-1991, Androw and Brattain 
(1 )(KWC). OKLAHOMA: Latimer Co., no locality, 5-VII-1987, K. E. M. Galley, at black light 
(2) (CUIC). *TENNESSEE: Cumberland Co., Grassy Cove, 26-III-1990, R. A. Androw & G. 
Keeney (1) (RAAC). TEXAS: Bastrop Co., 5 km. N. Smithville, 1-V-1983, 16-V-1983, J. E. 
Rawlins (2) (CMNHC). Cameron Co., 10 km. SSE Brownsville, Southpointe Nursery, 11-X 1994, 
R. A. Androw & M. Brattain (1) (CMNHC). Gonzales Co., Palmetto St. Pk., 13-X-1985, R. 
Davidson & J. E. Rawlins (1) (CMNHC). Hudspeth Co., McNary, 8-VII-1986, R. A. Androw & 
D. Heffern (1) (RAAC). Jefferson Davis Co., Davis Mtn. State Park, 28-VI/2-VII-1986, 22-VII- 
1992, R. A. Androw (2) (KWC, RAAC); 7-X-1993, R. A. Androw & S. M. Clark (2) (RAAC); 
Davis Mtns. Resort, 21-VII-1991, R. A. Androw (3) (KWC); vic. Ft. Davis St. Pk., 24-VII-1996, 
Wappes & Huether (2) (CUIC). Medina Co., 8 km. S. Hondo, 26-XI-1978, J. E. Rawlins (1) 
(CMNHC). Palo Pinto Co., Gordon, Rt. 193 at I-20, 12-VIII-1981, R. Davidson (27) (CMNHC). 
San Patricio Co., 12.4 km. NE Sinton, Welder Refuge, 20-VII-1979, D. C. Darling (5) (CUIC). 
Starr Co., 14 mi. E. El Sauz, 10-X-1993, R. A. Androw & S. M. Clark (2) (RAAC). Sutton Co., 
near Sonora, 27-VI-1986, R. A. Androw, D. Heffern & J. Huether (3) (RAAC). Travis Co., Aus- 
tin, Brackenridge Field Lab., 140 in., 13-X-1984, J. E. Rawlins (2) (CMNHC); 1 km. SE Bee 
Cave, 17-X-1985, R. Davidson & J. E. Rawlins (6) (CMNHC); 7-X-1984, J. E. Rawlins (1) 
(CMNHC). Val Verde Co., I-90 at Pecos River, 8-X-1993, R. A. Androw & S. M. Clark (4) 
(CMNHC). Zapata Co., Falcon St. Pk., 15-X-1985, R. Davidson & J. E. Rawlins (6) (CMNHC) 
CHIAPAS: Palenque, approx. 20 km. S. of ruins, to UV & Mercury Vapor, 19-20-VIII-1990, M. 
H. Evans (1) (CUIC). NUEVO LEON: above Cola de Caballo, 2550 ft. alt., to UV, 5-6-VIII- 
1986, M. H. Evans (15) (CUIC). OAXACA: Betanias, 20-VII-1990, 200 m. el., UV light, J. K. 
Liebherr, CAS-CU-UCB Field Exp. (1) (CUIC). VERACRUZ: Est. Biol. “Los Tuxtla”, 24-25- 
VII-1990, 250 m. el., J. K. Liebherr, CAS-CU-UCB Field Exp. (4) (CUIC). 


Onthophagus taurus (Schreber) 


Releases, Establishment and Geographic Range: Onthophagus taurus, 
a common Old World dung beetle native to central and southern Europe, Asia 
Minor, and ranging from Spain to Morocco, Turkey, Iran and Afghanistan 
(Balthasar 1963, Tyndale-Biscoe 1990), was recorded for the first time from 
the United States by Fincher and Woodruff (1975), based on a specimen taken 
in August 1971 in Santa Rosa County, Florida. Fincher and Woodruff (1975) 
suggested that it most likely had been accidentally introduced into the Florida 
panhandle near Pensacola; they proposed the theory of a cattle farmer or rancher 
easily bringing adult beetles back from an overseas trip, or that the beetles 


Vol. 108, No. 5, November & December, 1997 357 


ane 


a OD 
elias 7 


| ad 


O 
ty Ree ae 


tr i 
RYE carenrersetece 


Gren S seen 
Loree 


Onthophagus taurus (Schreber) 


Fig. 13. Known distribution of Onthophagus taurus in North America. Stars denote release sites; 
star/circle denotes a number of unspecified release sites in southern California. 


358 ENTOMOLOGICAL NEWS 


arrived aboard a military vehicle returned from Europe since there are several 
military bases in the Florida panhandle. 

Since its initial detection in the Florida panhandle in 1971, O. taurus has 
spread with remarkable speed westward and northward (Fig. 13). It has been 
recorded (in chronological order) from southwestern Georgia and southeast- 
ern Alabama (Fincher and Woodruff 1975), Stone County, Mississippi (Lago 
1979), Cumberland Island, Georgia (Fincher and Woodruff 1979), additional 
counties in Florida and Georgia, and new coastal localities in North and South 
Carolina (Steiner 1980, Bernhardt 1981), sites in southern Louisiana (Fincher 
et al. 1983), a locality in southern Maryland adjacent to the Potomac River 
(Glaser 1986), and at the Archbold Biological Station in southern peninsular 
Florida (Highlands Co.) (Vulinec and Eady 1993). 

Field releases of O. taurus also have taken place at a number of unspeci- 
fied sites in California and recapture studies have shown that it has success- 
fully overwintered in different parts of the state (Anderson and Loomis 1978). 
Coulson (1994) noted releases of O. taurus at a single site in Grimes County, 
Texas, in April 1983. Moreover, this species was released at several sites in 2 
counties of New Jersey beginning in 1987 by the New Jersey Department of 
Agriculture (Biological Control Group) and was again recovered from these 
counties during 1990-91 (R. Chianese, pers. commun.). 

Diagnosis: Length. 6.0-11.5 mm (males and females). Coloration. Vari- 
able in coloration, but mostly unicolorous dull black; sometimes the head and 
pronotum with a slight metallic reflection. Antennae reddish, with black club, 
and legs blackish or dark reddish-brown. Elytra usually dull black or blackish- 
brown, occasionally reddish-brown. The entire pygidium, or portions of the 
pygidium, and the sides of the abdominal sternites sometimes reddish-brown. 

Remarks: Onthophagus taurus (Fig. 14) can be easily distinguished from 
all American species by two long curved, divergent cephalic horns on major 
males (Fig. 15). However, minor males have a pair of very short, straight horns 
(Fig. 16), or else they are absent. In females, the clypeus and the frons (be- 
tween the eyes) each has a transverse, elevated carina or ridge (Fig. 17). 

New and Additional United States Records: All previously reported 
locality records for O. taurus in the literature and new and additional records 
given below are mapped in Figure 13. Specimens collected by one of us (KB), 
and those provided by K. W. Will (Cornell University) and R. A. Androw 
(Carnegie Museum), provide the basis for 6 new state records [marked with an 
asterisk (*)]. These new and additional distributional data are as follows: 


UNITED STATES: CALIFORNIA: Orange Co., Peter’s Canyon Reservoir, 12-VII-1993, in 
dog dung, K. Beucke (2) (KBC). FLORIDA: Leon Co., Lk. lamonia Landing, 28-III-1986, ex 
black light, E. R. Hoebeke (1) (CUIC). Union Co., 2 mi. S. Baker Co. line, 14-VIII-1976, R. 
Davidson (8) (CMNHC). Volusia Co., Ormond Beach, 25-III-1982, R. A. Androw (3) (RAAC). 
GEORGIA: Crisp Co., Cordele, 31-III-1982, R. A. Androw (7) (RAAC). MARYLAND: Calvert 


Vol. 108, No. 5, November & December, 1997 359 


Figs. 14-17. Onthophagus taurus. 14, dorsal habitus, major male. 15, head, major male, fronto- 
lateral aspect. 16, head, minor male, fronto-lateral aspect. 17, head, female, fronto-lateral aspect. 
Scale line for Fig. 14 = 5.0 mm. 


Co., Chesapeake Beach, 2-VII-1986, J. Green (4) (RAAC). MISSISSIPPI: Oktibbeha Co., Osborn 
Prairie, 16-20-III-1997, C. J. Marshall & K. W. Will, ex baited pitfall traps (3) (CUIC). *MIS- 
SOURI: Reynolds Co., Ellington, 15-VI-1985, M. Brattain (5) (RAAC). NEW JERSEY: Burlington 
Co., Mount Holly, 2-VII-1994, in cow dung, K. Beucke (2) (KBC). *NEW YORK: Suffolk Co., 
Stony Brook, 29-VIII-1993, K. Beucke (1) (KBC); Rocky Point, (burned pine barrens, site #2), 
21-IX-1996, ex chicken pitfall trap, K. Beucke (1) (KBC). *OHIO: Fairfield Co., Barneby Ctn., 


360 ENTOMOLOGICAL NEWS 


12-19-X-1991, ex pitfall/dung, K. Will & R. A. Androw (6) (KWC, CUIC). Pike Co., Jackson 
Twp., Jackson Lk., 5-12-IX-1992, Will & Androw (1) (CUIC). Richland Co., Shelby, Will’s 
Farm, 29-IV-1991, ex dung, K. & I. Will (1) (KWC). Ross Co., Rt. 50 at Scioto River, 4-VII- 
1989, R. A. Androw (1) (RAAC). Scioto Co., Shawnee St. Pk., 26-V-1990, R. A. Androw & G. 
Keeney (2) (RAAC); 20-VII 1990, R. A. Androw (2) (RAAC). *PENNSYLVANIA: Adams Co., 
Gettysburg, 10-VI-1995, in cow dung, K. Beucke (1) (KBC). Berks Co., Wernersville, Stricker 
farm, V-1994, R. A. Byers & G. M. Barker, ex pitfall trap in grazed dairy pasture (1) (CUIC). 
Forest Co., 8.7 km. N. Kellettville, 500m., 8-14-VIII-1995, J. R. Deeds (1) (CMNHC). Hunting- 
ton Co., Tyrone, Hawthorne farm, V-1994, R. A. Byers & G. M. Barker, ex pitfall trap in grazed 
dairy pasture (1) (CUIC). Lawrence Co., New Castle, Harmon farm, VI-1994, R. A. Byers & G. 
M. Barker, ex soil samples in grazed dairy pasture (1) (CUIC); New Castle, Gebhart farm, VI- 
1994, R.A. Byers & G. M. Barker, ex pitfall trap in grazed dairy pasture (1) (CUIC). Miflin Co., 
Belleville, Rodgers farm, V-1994, R. A. Byers & G. M. Barker, ex pitfall trap in grazed dairy 
pasture (1) (CUIC). Warren Co., 8.6 km. S. Irvine, Hedgehog Run, 530 m., 19-27-V-1995, 28-V/ 
2-VI-1995, 8-14-VIII-1995, J. R. Deeds (6) (CMNHC); 7.3 km. SSW Cherry Grove, 520m., 28- 
V/2-VI-1995 (1) (CMNHC); 3-10-VI-1995, C. Bier, J. R. Deeds & T. Schumann (1) (CMNHC); 
6km. E. Cobham, 565 m., 26-VII-1-VIII-1995, J. R. Deeds (1) (CMNHC). *TENNESSEE: Cocke 
Co., Cosby, 13-VI-1987, R. A. Androw, M. Brattain & J. Huether (1) (RAAC). Cumberland Co., 
Grassy Cove, 27-III-1989, R. A. Androw (13) (KWC, CUIC). *WEST VIRGINIA: Hampshire 
Co., 0.5 km. E. North River Mills, Ice Mountain Preserve, 6-19-VII-1995, D. Koenig, W. Zanol 
& C. Young (1) (CMNHC); 1-VIII-20-IX-1995, C. Young & J. E. Rawlins (1) (CMNHC). 


ACKNOWLEDGMENTS 


We thank Kip Will (Cornell University) and Bob Androw (Carnegie Museum of Natural 
History) for reviewing the paper and making available to us new United States records of 
Onthophagus gazella and O. taurus; Kent Loeffler (Cornell University) for providing the habitus 
photographs of the species treated in this paper; R. Chianese (New Jersey Department of Agri- 
culture) for sharing information on releases of exotic Onthophagus in New Jersey; and B. D. Gill 
(Agriculture Canada, Ottawa) and G. T. Fincher (USDA, ARS, FAPRL, College Station, TX) for 
reviewing and providing critical comments on a draft of this paper. 


LITERATURE CITED 


Anderson, J. R. and E. C. Loomis. 1978. Exotic dung beetles in pastures and range land eco- 
systems. Calif. Agric. 32:31-32. 

Balthasar, V. 1963. Monographie der Scarabaeidae und Aphodiidae der palaearktischen und 
orientalischen Region (Coleoptera: Lamellicornia). Band 2: Coprinae (Onitini, Oniticellini, 
Onthophagini). Verlag Tschechoslowakischen Akad. Wiss., Prag. 627 pp. 

Barbero, E. and Y. Lopez-Guerrero. 1992. Some considerations on the dispersal power of 
Digitonthophagus gazella (Fabricius 1787) in the New World (Coleoptera Scarabaeidae 
Scarabaeinae). Trop. Zool. 5:115-120. 

Bernhardt, J. L. 1981. Additional distribution records for Onthophagus taurus Schreber (Co- 
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Bland, J. H. B. 1889. Proceedings of the Monthly Meetings of the Entomological Section of the 
Academy of Natural Sciences of Philadelphia (May 23, 1889). Trans. Am. Entomol. Soc. 16: 
Dex. 

Blume, R. R. and A. Aga. 1978. Onthophagus gazella: Progress of experimental releases in 
south Texas. Folia Entomol. Mex. 29-40:190-191. 


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Brown, W. J. 1940. Notes on the American distribution of some species of Coleoptera common 
to the European and North American continents. Can. Entomol. 72:65-78. 

Cartwright, O. L. 1938. A South African Onthophagus found in the United States. Entomol. 
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Cervenka, V. J. and R. D. Moon. 1991. Arthropods associated with fresh cattle dung pats in 
Minnesota. J. Kansas Entomol. Soc. 64:131-145. 

Coulson, J. R. 1992. Releases of beneficial organisms in the United States and territories - 1982. 
USDA, ARS, Misc. Publ. No.1505. 529 pp. 

Coulson, J. R. 1994. Releases of beneficial organisms in the United States and territories- 1983. 
USDA, ARS. ARS-131. 113 pp. 

Downie, N. M. 1984. Onthophagus gazella Fabricius, new to Florida (Coleoptera: Scarabaeidae). 
Coleopt. Bull. 38:304. 

Evans, J. D. 1899. List of Coleoptera from Halifax, N.S. Can. Entomol. 31:320-321. 

Fincher, G. T. 1981. The potential value of dung beetles in pasture ecosystems. J. Georgia Entomol. 
Soc. 16 (Suppl.): 316-333. 

Fincher, G. T. 1986. Importation, colonization, and release of dung-burying scarabs, pp. 69-76. 
In R. S. Patterson and D. A. Rutz (eds.), Biological Control of Muscoid Flies. Misc. Publ., 
Entomol. Soc. Am. No. 61. 

Fincher, G. T. 1990. Biological control of dung-breeding flies: pests of pastured cattle in the 
United States, pp.137-151. In D. A. Rutz and R. S. Patterson (eds.), Biocontrol of arthropods 
affecting livestock and poultry. Westview Studies in Insect Biology, Westview Press, Inc., 
Boulder, CO. 

Fincher, G. T. and R. E. Woodruff. 1975. A European dung beetle, Onthophagus taurus, new to 
the United States. Coleopt. Bull. 29:349-350. 

Fincher, G. T. and R. E. Woodruff. 1979. Dung beetles of Cumberland Island, Georgia (Co- 
leoptera: Scarabaeidae). Coleopt. Bull. 33:69-70. 

Fincher, G. T. and J. S. Hunter, III. 1989. Overwintering studies on Onthophagus gazella (F.) 
and Onthophagus bonasus (F.) in two different textural classes of soil in east-central Texas. 
Southwestern Entomol. 14:133-138. 

Fincher, G. T., T. B. Stewart, and J. S. Hunter, III. 1983. The 1981 distribution of Onthophagus 
gazella Fabricius from releases in Texas and Onthophagus taurus Schreber from an unknown 
release in Florida (Coleoptera: Scarabaeidae). Coleopt. Bull. 37:159-163. 

Fletcher, J. 1904. Entomological Record, 1903. 34th Ann. Rept. Entomol. Soc. Ont., 1903: 
85-99. 

Glaser, J. D. 1986. Notes on some Maryland Scarabaeidae and Cerambycidae (Coleoptera). 
Coleopt. Bull. 40:355-356. 

Harpootlian, P. J. 1995. Notes and records of Scarabaeidae from the southeastern United States. 
Coleopt. Bull. 49:280. 

Hatch, M. H. 1953. The Beetles of the Pacific Northwest. Univ. Wash. Publ. Biol. 16:1-340. 

Hatch, M. H. 1971. The Beetles of the Pacific Northwest. Part V: Rhipiceroidea, Sternoxi, 
Phytophaga, Rhynchophora, and Lamellicornia. Univ. Wash. Press, Seattle. 661 pp. 

Henshaw, S. 1887. Some European beetles in America. Can. Entomol. 19:159-160. 

Howden, H. F. and O. L. Cartwright. 1963. Scarab beetles of the genus Onthophagus Latreille 
North of Mexico (Coleoptera: Scarabaeidae). Proc. U.S. Natl. Mus. 114:1-135. 

Hunter, J. S., {111 and G. T. Fincher. 1985. Five new state records for the Afro-Asian dung 
beetle Onthophagus gazella (Coleoptera: Scarabeidae). J. Entomol. Sci. 20:24-25. 

Hunter, J. S., If] and G. T. Fincher. 1996. Distribution of Onthophagus depressus: an intro- 
duced species of dung beetle. J. Agric. Entomol.13:319-322. 

Kohlmann, B. 1994. A preliminary study of the invasion and dispersal of Digitonthophagus 
gazella (Fabricius, 1787) in Mexico (Coleoptera: Scarabaeidae: Scarabaeinae). Acta Zool. 
Mex. (n.s.) 61:35-42. 


362 ENTOMOLOGICAL NEWS 


Lago, P. K. 1979. A Mississippi record for an introduced dung beetle, Onthophagus taurus 
Schreber (Coleoptera: Scarabaeidae). Entomol. News 90:152. 

Lago, P. K., J. R. MacDonald and W. H. Cross: 1984. Mexican records for Onthophagus 
gazella Fabricius (Coleoptera: Scarabaeidae). Coleopt. Bull. 38:214. 

Legner, E. F. 1986. The requirements for reassessment of interactions among dung beetles, 
symbovine flies, and natural enemies, pp. 120-131. In R. S. Patterson and D. A. Rutz (eds.), 
Biological Control of Muscoid Flies. Misc. Publ., Entomol. Soc. Am. No. 61. 

Macqueen, A. and B. P. Beirne. 1975. Dung burial activity and fly control potential of 
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107:1215-1220. 

Melsheimer, F. E. 1844. Descriptions of new species of Coleoptera of the United States. Proc. 
Acad. Nat. Sci. Phila., 2 (1846):26-43, 98-118, 134-160, 213-223, 302-318. 

McNamara, J. 1991. Family Scarabaeidae (scarab beetles), pp. 145-158. In Y. Bousquet (ed.), 
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Canada. Publication 1861/E. 

Moron, M.A., C. Deloya, and L. Delgado-Castillo. 1988. Fauna de coledpteros Melolonthidae, 
Scarabaeidae y Trogidae de la region de Chamela, Jalisco, México. Folia Entomol. Mex. 
77:313-378. 

Morris, R. F. 1983. Introduced terrestrial insects, pp. 551-591. In G. R. South (ed.), Biogeogra- 
phy and Ecology of the Island of Newfoundland. W. Junk Publishers, The Hague-Boston- 
London. 

Rivera-Cervantes, L. E. and E. Garcia-Real. 1991. New locality records for Onthophagus 
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Robinson, M. 1948. Remarks on a few Scarabaeidae (Coleoptera). Entomol. News 59:175-177. 

Roy, E. 1899. Notes entomologiques. Nat. Can. 26:177-178. 

Steiner, W. E., Jr. 1980. Additional distribution records for Onthophagus taurus (Coleoptera: 
Scarabaeidae). Coleopt. Bull. 34:332. 

Thomas, D. B. 1993. Scarabaeidae (Coleoptera) of the Chiapanecan forests: a faunal survey and 
chorographic analysis. Coleopt. Bull. 47:363-408. 

Tyndale-Biscoe, M. 1990. Common dung beetles in pastures of south-eastern Australia. CSIRO, 
Australia, Division of Entomology. 71 pp. 

Vulinec, K. and S. P. Eady. 1993. A southern range extension for the introduced dung beetle 
Onthophagus taurus Schreber (Coleoptera: Scarabaeidae). Coleopt. Bull. 47: 129-130. 

Zunino, M. and G. Halffter. 1988. Analisis taxonomico, ecoldgico y biogeografico de un grupo 
americano de Onthophagus (Coleoptera: Scarabaeidae). Monografie IX, Mus. Reg. Sci. Nat. 
Torino. 211 pp. 


BINDING ERROR - Sept. - Oct. ’97 Issue (108:4) 


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Vol. 108, No. 5, November & December, 1997 363 


MICKSIOPS, A NEW GENUS OF SMALL MINNOW 
MAYFLIES (EPHEMEROPTERA: BAETIDAE) 
FROM AFRICA!2 


W. P. McCafferty3, C.R. Lugo-Ortiz>, H. M. Barber-James4 


ABSTRACT: The new genus Micksiops is established for the African species M. bicaudatus 
(Gillies), n. comb. This species is known from Guinea, and undescribed species of Micksiops 
have been seen from Kenya and South Africa. In the larval stage, characters of the labium, the 
single row of denticles, lack of subtending setae on the claws, and the lack of armature on the 
dorsal thorax and abdomen distinguish the new genus from Acanthiops. The condition of the 
claws and caudal filaments, along with several other characteristics, will distinguish the larvae 
of Micksiops from Afroptilum s.s. and its cognates. 


This work is part of the revision of those Afrotropical species of the may- 
fly family Baetidae that historically have been known as Afroptilum Gillies, 
and previous to that as Centroptilum Eaton. Other revisionary works on this 
subject have been contributed by Wuillot and Gillies (1994), McCafferty and 
de Moor (1995), Lugo-Ortiz and McCafferty (1996a,b,c, 1997a,b), and Bar- 
ber-James and McCafferty (1997). 

McCafferty and de Moor (1995) synonymized what had been known as the 
subgenus Afroptiloides Gillies of the genus Afroptilum with the genus 
Acanthiops Waltz and McCafferty. Barber-James and McCafferty (1997) indi- 
cated that the type of Afroptiloides and all species previously assigned to it, 
except one, were assignable to Acanthiops. That one exception is Afroptilum 
bicaudatum Gillies, originally described from adult and larval material from 
Guinea (Gillies 1990). Afroptilum bicaudatum does not share defining 
apomorphies with Acanthiops, as was pointed out by Barber-James and 
McCafferty (1997). This species also cannot be placed with Afroptilum s. s. or 
any other known genera of Baetidae, including those African taxa that now 
contain former species of Afroptilum, i.e., Bugilliesia Lugo-Ortiz and 
McCafferty, Cheleocloeon Wuillot and Gillies, Crassabwa Lugo Ortiz and 
McCafferty, Dabulamanzia Lugo-Ortiz and McCafferty, and Dicentroptilum 
Wuillot and Gillies (see also Lugo-Ortiz and McCafferty 1997b). We therefore 
describe a new genus herein for A. bicaudatum, and other closely related but 
undescribed species that have been seen from Kenya and South Africa. 

The larval type material of A. bicaudatum could not be located at the Brit- 


1 Received April 11, 1997. Accepted May 2, 1997. 

2 Purdue Agricultural Research Program Journal No. 15388. 

3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 
4 Albany Museum, Somerset Street, Grahamstown 6140, South Africa. 


ENT. NEWS 108(5) 363-366, November & December, 1997 


364 ENTOMOLOGICAL NEWS 


ish Museum, where it was to have been deposited. Because this taxon is so 
distinct but unplaceable in the current nomenclature and because there are 
additional closely related species from Africa to be described by other authors, 
we feel it is important to erect a genus at this time. Such a description is also 
one of the final necessary revisions to the species formerly known as Afroptilum. 
We are pleased to name this genus after M. T. (Mick) Gillies, who has contrib- 
uted significantly to our knowledge of the African Ephemeroptera fauna. 


Micksiops McCafferty, Lugo-Ortiz, and Barber-James, NEW GENUS 


Larva. Head: Antennae relatively long, about four times length of head. Labrum (Fig. 1) 
broadly rounded in anterior half with broad emargination in medial third of distal margin. Left 
mandible (Fig. 2) with incisors fused; prostheca robust; tuft of setae absent. Right mandible (Fig. 
3) with two sets of incisors; prostheca somewhat slender; tuft of setae present between incisors 
and mola. Maxillae (Fig. 4) relatively robust; palps two segmented, extending to about apex of 
galealaciniae. Labium (Fig. 5) with glossae shorter than paraglossae; palp segment | subequal to 
segments 2 and 3 combined; palp segment 3 globular and rounded apically, slightly more devel- 
oped medially than laterally. Thorax: Legs (Fig. 6) with femora without villopore, with row of 
stout setae along dorsal margin; dorsal setae well developed on tibiae and tarsi of mid- and 
hindlegs. Tarsal claws (Fig. 7) angular at tip, with single row of distinctive subapical denticles, 
and without pair of long, subtending setae. Abdomen: Lamellate gills (Fig. 8) present on abdomi- 
nal segments 1-7, asymmetrical subobovate, with pinnately branched tracheation. Median cau- 
dal filament not developed; cerci without row of swimming setae along medial margins. 

Male Adult. Head: Eyes well separated and divergent apically. Thorax: Forewing with single 
marginal intercalaries. Hindwings (Fig. 9) narrow; costal process relatively large and compound, 
with straight and more laterally extended basal node and somewhat curved, slightly larger and 
distally oriented distal node. Genital forceps with short basal segment with somewhat developed 
medioapical process (Fig. 10); segment 3 narrow as in Figure 11. 

Type species. Micksiops bicaudatus (Gillies), new combination. 

Included species. 

Micksiops bicaudatus (Gillies) 
Afroptilum (Afroptiloides) bicaudatum Gillies, 1990:121. 

Distribution. Guinea (M. bicaudatus), Kenya (undescribed sp.), and South Africa 
(undescribed: sp.). 


DISCUSSION 


Larvae of Micksiops can be easily told from larvae of Acanthiops, which is 
another African baetid genus having two-tailed larvae and adults with single 
marginal intercalaries in the forewings. Diagnositc characteristics of Micksiops, 
include the well-developed and much broader terminal labial palp segment, 
the unequal glossae and paraglossae, the single rather than double row of claw 
denticles, the lack of subtending claw setae, and the lack of dorsal armature on 
the thorax and abdomen. These characters can also be used to differentiate the 
larvae of Micksiops from larvae of Thraulobaetodes Elouard and Hideux, which 
we believe are very closely related to Acanthiops. The larvae of Micksiops 
may superficially resemble the two-tailed larvae of the African genera 


Vol. 108, No. 5, November & December, 1997 365 


Figs. 1-11. Micksiops bicaudatus [modified from Gillies (1990)]. 1-8. Larva. 1. Labrum (dorsal). 
2. Left mandible. 3. Right mandible. 4. Right maxilla. 5. Labium (ventral). 6. Hindleg (details of 
apex of femur not shown). 7. Hindclaw. 8. Abdominal gill 5. 9-11. Male adult. 9. Hindwing. 10. 
Left basal segment of forceps, ventral. 11. Left segment 3 of forceps, ventral. 


366 ENTOMOLOGICAL NEWS 


Demoreptus Lugo-Ortiz and McCafferty and Tanzaniella Gillies. However, 
those latter genera are quite unrelated to Micksiops, having adults with double 
marginal intercalaries in the forewings and larvae with a femoral villopore 
(see Gillies 1991, Lugo-Ortiz and McCafferty 1997c). 

Adults of other genera are not well enough described at this time to draw 
conclusions with respect to adult diagnosis of Micksiops. The compound cos- 
tal process of the Micksiops hindwings will distinguish it from some of the 
baetids with single marginal intercalaries in their forewings, but not all. 

Barber-James and McCafferty (1997) suggested that M. bicaudatus was 
possibly a species that shared some common ancestry with the genus Acanthiops 
because of the possession of a tibial row of setae in the larvae of M. bicaudatus 
and ancestral species of Acanthiops. However, such setae are variable among 
and within many genera of Baetidae (e.g., Baetis Leach), and Lugo-Ortiz and 
McCafferty (unpublished) have more recently identified an entire grouping of 
genera from Africa and Madagascar that share a double row of claw denticles 
and other characteristics, which contains, among others, Acanthiops and 
Afroptilum sensu stricto. Micksiops would not be a member of this grouping 
unless it could be proven to be an entirely aberrant form. A more definite 
phylogenetic placement of Micksiops can not be made at this time. 


LITERATURE CITED 


Barber-James, H. M. and W. P. McCafferty. 1997. Review and a new species of the African 
genus Acanthiops (Ephemeroptera: Baetidae). Ann. Limnol. 33: 85-92. 

Gillies, M. T. 1990. A revision of the African species of Centroptilum Eaton (Baetidae, Ephe- 
meroptera). Aquat. Insects 12: 97-128. 

Gillies, M. T. 1991. A diphyletic origin for the two-tailed baetid nymphs occurring in East 
Africa stony streams with a description of the new genus and species Janzaniella spinosa 
gen. nov. sp. nov. Pp. 175-187 In: J. Alba-Tercedor and A. Sanchez-Ortega (eds.), Overview 
and strategies of Ephemeroptera and Plecoptera. Sandhill Crane Press, Gainesville, Florida. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996a. Crassabwa: a new genus of small minnow 
mayflies (Ephemeroptera: Baetidae) from Africa. Ann. Limnol. 32: 235-240. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996b. The composition of Dabulamanzia, a new 
genus of Afrotropical Baetidae (Ephemeroptera), with descriptions of two new species. Bull. 
Soc. Hist. Nat. Toulouse 132: 7-13. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996c. The Bugilliesia complex of African Baetidae 
(Ephemeroptera). Trans. Am. Entomol. Soc. 122: 175-197. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1997a. Contribution to the systematics of the genus 
Cheleocloeon (Ephemeroptera: Baetiddae). Entomol. News. 108: 283-289. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1997b. Maliqua: a new genus of Baetidae (Ephe- 
meroptera) for a species previously assigned to Afroptilum. Entomol. News. 108: 367-371. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1997c. A new genus and redescriptions for African 
species previously placed in Acentrella (Ephemeroptera: Baetidae). Proc. Entomol. Soc. Wash. 
99: 429-439. 

McCafferty, W. P. and F. C. de Moor. 1995. South African Ephemeroptera: problems and pri- 
orities. Pp. 463-476. In: L. Corkum and J. Ciborowski (eds.), Current directions in research 
on Ephemeroptera. Canadian Scholars’ Press, Toronto. 

Wuillot, J. and M. T. Gillies. 1994. Dicentroptilum, a new genus of mayflies (Baetidae, 
Ephemeroptera) from Africa. Aquat. Insects 16: 133-140. 


Vol. 108, No. 5, November & December, 1997 367 


MALIQUA: A NEW GENUS OF BAETIDAE 
(EPHEMEROPTERA) FOR A SPECIES PREVIOUSLY 
ASSIGNED TO AFROPTILUM!:2 


C. R. Lugo-Ortiz, W. P. McCafferty 


ABSTRACT: Maliqua (Ephemeroptera: Baetidae), n. gen., is erected for M. plumosa, n. comb., 
a West African species previously assigned to the dimorphicum group of Afroptilum. Larvae of 
Maliqua are distinguished by possessing minute denticles on the tarsal claws. Adults are distin- 
guished by having segment 2 of the male genital forceps medially expanded at base and setose. 
Maliqua is hypothesized to be closely related to Cloeodes on the basis of mouthpart and male 
genitalic similarities. 


Gillies (1990) erected the genus Afroptilum (Ephemeroptera: Baetidae) to 
accommodate all African species previously assigned to Centroptilum Eaton. 
As such, the genus included 30 species in two subgenera, Afroptilum s.s. and 
Afroptiloides Gillies. Gillies (1990) further subdivided Afroptilum s.s. into in- 
formal groupings he called the dimorphicum, sudafricanum, sudanense, and 
tarsale species groups, but he did not provide any phylogenetic rationales for 
those subdivisions. Wuillot and Gillies (1993) later described four additional 
species of Afroptilum that they variously assigned to the tarsale and dimorphi- 
cum groups. 

More recently, species ascribed to Afroptilum have been the subject of con- 
siderable revisionary work. Wuillot and Gillies (1994) erected the genus 
Dicentroptilum for two distinct species previously considered in the 
sudafricanum group. McCafferty and de Moor (1995) synonymized the sub- 
genus Afroptiloides with Acanthiops Waltz and McCafferty, and Barber-James 
and McCafferty (1997) elaborated the rationale for that revision. Lugo-Ortiz 
and McCafferty (1996a,b,c) erected the genera Bugilliesia, Crassabwa, and 
Dabulamanzia for most species in the tarsale and sudanense groups, and Lugo- 
Ortiz and McCafferty (1997a) demonstrated that two species assigned to the 
dimorphicum group are referable to Cheleocloeon Wuillot and Gillies. 
McCafferty et al. (1997) erected the genus Micksiops for a species previously 
assigned to the subgenus Afroptiloides. 

In this paper we erect a new genus for A. plumosum Wuillot, a West Afri- 
can species originally assigned to the dimorphicum group of Afroptilum (Wuillot 
and Gillies 1993). We describe the new genus, hypothesize that it is closely 
related to Cloeodes Traver, and provide guidelines to differentiate the two gen- 


! Received April 11, 1997. Accepted May 2, 1997. 


2 Purdue Agricultural Research Program Journal No. 15378. 
3 Department of Entomology, Purdue University, West Lafayette, IN 47907. 


ENT. NEWS 108(5) 367-371, November & December, 1997 


368 ENTOMOLOGICAL NEWS 


era. Our attempts to secure original material from Paris for examination were 
not successful; however, because the new genus is so distinctive and involves 
one of only a few unresolved species that were previously placed in Afroptilum, 
it is judicious to describe it at this time. 


Maliqua Lugo-Ortiz and McCafferty, NEW GENUS 


Larva. Head: Labrum (Fig. 1) anteriorly broadly rounded, with deep anteromedial notch. 
Left mandible (Fig. 2) with incisors fused; prostheca relatively robust, apically denticulate; small 
tuft of setae between prostheca and mola. Right mandible (Fig. 3) with two broadly based sets of 
inciSors; prostheca somewhat slender, apically with sharp denticles. Maxillae (Fig. 4) relatively 
*obust; palps two segmented, slender, not reaching galealaciniae. Labium (Fig. 5) with glossae 
and paraglossae subequal in length and width; palps three segmented; palp segment | slightly 
longer than segments 2 and 3 combined; palp segments 2 and 3 subequal in length and width; 
palp segment 3 apically rounded. Thorax: Hindwingpads absent. Legs (Fig. 6) with femora with- 
out villopore and with subparallel dorsal and ventral margins; tarsal claws (Fig. 7) with one row 
of minute subapical denticles. Abdomen: Gills (Figs. 8, 9) on abdominal segments 1-7, platelike, 
asymmetrical, anteriorly serrate. Median caudal filament well developed, with abundant short, 
fine, simple setae on both margins; cerci with abundant long, fine, simple setae medially and 
short, stout, simple setae laterally every fourth segment. 

Adult. Thorax: Forewings (Fig. 10) with single marginal intercalaries. Hindwings absent. 
Abdomen: Male genital forceps (Fig. 11) three segmented; segment | short, subcylindrical; seg- 
ment 2 medially expanded at base, with short, fine simple setae medially; segment 3 short, ovoid. 

Type species. Maliqua plumosa (Wuillot), new combination. 

Included species — 

Maliqua plumosa (Wuillot), new combination. 
Afroptilum plumosum Wuillot, in Wuillot and Gillies 1993:276. (larva, male adult). 

Distribution. Guinea, Mali. 


Etymology. The generic name is an arbitrary combination of letters that 
incorporates the name of the country where the type locality of the type spe- 
cies is located. The gender is feminine. 


DISCUSSION 


Larvae of Maliqua are generally similar to those of the Pantropical genus 
Cloeodes, but are distinguished from that genus by the presence of tufts of 
setae between the prosthecae and molae of both mandibles (Figs. 2, 3) and 
minute denticles on the tarsal claws (Fig. 7). Adults of Maliqua are also simi- 
lar to those of Cloeodes, but segment 2 of the male genital forceps of Maliqua 
(Fig. 11) is considerably more expanded than in Cloeodes. Other differences 
that aid in distinguishing larvae of Maliqua from those of Cloeodes are the 
presence in larvae of the latter genus of subproximal arcs of long, fine, simple 
setae on the tibiae [Waltz and McCafferty (1987b): Figs. 1, 7], edentate tarsal 
claws [Waltz and McCafferty (1987b): Fig. 8], and tufts of long, fine, simple 
setae on abdominal sterna 2-6 [Waltz and McCafferty (1987a): Fig. 5; Waltz 
and McCafferty (1987b): Figs. 9, 44]. The tendency of some female adults of 


Vol. 108, No. 5, November & December, 1997 369 


Figs. 1-11. Maliqua plumosa (Wuillot), larva and male adult [modified from Wuillot and Gillies 
(1993)]. 1. Labrum (dorsal). 2. Left mandible. 3. Right mandible. 4. Right maxilla. 5. Labium 
(ventral). 6. Right foreleg. 7. Tarsal claw (pointer to row of denticles). 8. Gill 5. 9. Detail of gill 
5. 10. Forewing. 11. Male genitalia. 


370 ENTOMOLOGICAL NEWS 


Cloeodes to have single marginal intercalary veins in the forewings [Waltz 
and McCafferty (1987a): Fig. 1; Waltz and McCafferty (1987b): Figs. 16, 18, 
19, 22, 33] is also noteworthy. 

Maliqua plumosa cannot be considered to belong to Afroptilum because it 
differs considerably from A. sudafricanum (Lestage), the type species of 
Afroptilum. Moreover, the shape of the claws and presence of a single row of 
denticles on the larval tarsal claws of Maliqua excludes it from consideration 
in a complex of African and Malagasy genera to which Afroptilum belongs 
and that also includes Acanthiops, Centroptiloides Lestage, Dicentroptilum, 
Edmulmeatus Lugo Ortiz and McCafferty, Nesoptiloides Demoulin, and 
Thraulobaetodes Elouard and Hideux (Lugo-Ortiz and McCafferty 1997b). 
That complex is characterized, in part, by the distinctive shape of the claws 
and presence of two subparallel rows of denticles on the larval tarsal claws 
{[Lugo-Ortiz and McCafferty (1997b): Fig. 12]. 

The larval mouthparts are remarkably similar in Maliqua and Cloeodes, 
particularly with respect to the relatively robust maxillae and their somewhat 
short and slender palps [Fig. 4; Waltz and McCafferty 1987b: Figs. 5, 28] and 
the labial palps having segments 2 and 3 subequal in length and width [Fig. 5; 
Waltz and McCafferty 1987b: Figs. 6, 29]. Those larval similarities, plus the 
fundamentally similar medially expanded and setose segment 2 of the male 
genital forceps [Fig. 11; Waltz and McCafferty 1987b: Fig. 34], support the 
hypothesis that the two genera are closely related. It is possible that other 
apomorphies tie Maliqua and Cloeodes, but because the original description 
of M. plumosa by Wuillot and Gillies (1993) is rather brief, they cannot be 
ascertained at this moment. 


LITERATURE CITED 


Barber-James, H. M. and W. P. McCafferty. 1997. Review and a new species of the African 
genus Acanthiops (Ephemeroptera: Baetidae). Ann. Limnol. 33: 85-92. 

Gillies, M. T. 1990. A revision of the African species of Centroptilum Eaton (Baetidae, 
Ephemeroptera). Aquat. Insects 12: 97-128. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996a. Crassabwa: a new genus of small minnow 
mayflies (Ephemeroptera: Baetidae) from Africa. Ann. Limnol. 32: 235-240. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996b. The composition of Dabulamanzia, a new 
genus of Afrotropical Baetidae (Ephemeroptera), with descriptions of two new species. Bul]. 
Soc. Hist. Nat. Toulouse 132: 7-13. 

Lugo-Ortiz, C. R. and W. P. McCafferty. 1996c. The Bugilliesia complex of African Baetidae 
(Ephemeroptera). Trans. Am. Entomol. Soc. 122: 175-197. 

Lugo-Ortiz, C R. and W. P. McCafferty. 1997a. Contribution to the systematics of the genus 
Cheleocloeon (Ephemeroptera: Baetidae). Entomol. News. 108: 283-289. 

Lugo-Ortiz, C R and W. P. McCafferty. 1997b. Edmulmeatus grandis: an extraordinary new 
genus and species of Baetidae (Insecta: Ephemeroptera) from Madagascar. Ann. Limnol., in 
press. 


Vol. 108, No. 5, November & December, 1997 371 


McCafferty, W. P. and F. C. de Moor. 1995. South African Ephemeroptera: problems and pri- 
orities. Pp. 463-476. In: L. Corkum and J. Ciborowski (eds.), Current directions in research 
on Ephemeroptera. Canadian Scholars’ Press, Toronto. 

McCafferty, W. P., C. R. Lugo-Ortiz, and H. M. Barber-James. 1997. Micksiops, anew genus 
of small minnow mayflies (Ephemeroptera: Baetidae) from Africa. Entomol. News. 108: 
363-366. 

Waltz, R. D. and W. P. McCafferty. 1987a. Generic revision of Cloeodes and description of two 
new genera (Ephemeroptera: Baetidae). Proc. Entomol. Soc. Wash. 89: 177-184. 

Waltz, R D. and W. P. McCafferty. 1987b. Revision of the genus Cloeodes Traver (Ephe- 
meropiera: Baetidae). Ann. Entomol. Soc. Am. 80: 191-207. 

Wuillot, J. and M. T. Gillies. 1993. New species of Afroptilum (Baetidae, Ephemeroptera) from 
West Africa. Rev. Hydrobiol. Trop. 26: 269-277. 

Wuillot, J. and M. T. Gillies. 1994. Dicentroptilum, a new genus of mayflies (Baetidae, 
Ephemeroptera) from Africa. Aquat. Insects 16: 133-140. 


(continued from page 344) 


Further research pointed to strong asynchrony in egg hatching among populations, but 
synchrony with the flowering of the hostplants. The brief mating periods, in which females mate 
only once with one of the short-lived males are sufficiently asynchronous to preclude mixed 
matings. Furthermore, it is now known that males have distinctive courtship “songs,” to which 
only females from the same host respond. Other experiments showed that eggs laid on hosts 
other than the females’ own have nearly nil survival and reproduction. They also showed a most 
interesting fact: eggs on the “wrong” host hatched at the time eggs of populations adapted to that 
host hatched, not at the time they would have on the “correct” host. It turned out that egg hatch is 
controlled by the plant’s spring sapflow, which rehydrates the eggs after winter dissication. 

Tom now asked, could a shift to a new hostplant, coupled with asynchronous egg hatch and 
the short mating period, lead to host race formation and eventual speciation? Preliminary trials 
suggested it could, so he has now established a large outdoor experiment, involving twenty- 
eight, 28' x 8' x 8' cages. In these, Enchenopa binotata (he is getting ready to name the host races 
as new species) are confined on mixed and unmixed pairs of four species of Viburnum. He and his 
students, after tens of thousands of individual transplantings of nymphs, have documented non- 
random mating and oviposition among the populations. This success has finally persuaded NSF 
to support the effort. The possibility of sympatric speciation by host shift remains controversial, 
despite the extensive studies of cases such as Rhagoletis pomonella, and his research may do a 
lot to clear matters up. Tom says he looks forward to thirty more years of study of this fascinating 
beast. 

W.J. Cromartie, 
Corresponding Secretary 


372 ENTOMOLOGICAL NEWS 


NEW ORCHESELLA SPECIES 
(COLLEMBOLA: ENTOMOBRYIDAE) 
FROM NORTH AMERICA! 


Richard J. Snider 


ABSTRACT: Three new species of Orchesellinae are described from North America: Orchesella 
gloriosa n. sp. from lichens on granite in Great Smoky Mountains National Park, North Caro- 
lina; Orchesella imitari n. sp. from mosses growing in Parc de la Gaspésie, Quebec; and Orchesella 
texensis n. sp. from cave litter in Williamson and Travis Counties, Texas. The new species are 
separated from those previously described by body chaetotaxy and color patterns. 


Orchesella gloriosa, NEW SPECIES 
COLOR DESCRIPTION 


Background white to cream-yellow with blue pigmented color pattern (Figs. | and 2) Head, 
in dorsal aspect, with dark blue pigment forming a band between antennal bases; dorsum with 
interocular maculae in broken pattern extending posteriorly; gena with blue forming an irregular 
band extending to oral region, with “C”’-shape maculae best seen laterally; ANT I with dark blue 
basally and distally, ANT II blue distally with longitudinal blue streaking, ANT III with blue 
basally and distally, ANT IV light blue, darkest apically. Trunk with broken color patterns com- 
posed of dark blue pigment: TH II to ABD II with irregular medial stripe and paramedial stripes; 
ABD III almost entirely blue dorsally; ABD IV and V with medial line continued to mid-ABD V; 
ABD VI dark blue; laterally with blue forming irregular stripes defined by white, (taken to- 
gether, overall pattern could be viewed as forming 5 irregular longitudinal stripes); all micro and 
macro setee with white surrounding sockets giving a stippled-streaking. 


MORPHOLOGICAL DESCRIPTION 


HEAD: eyes 8 + 8 on black patches, ocelli A,B,C,D slightly larger than E and F, G and H subequal, 
G/F ratio ~.S (Fig. 3). ANT IV lacking apical bulb, with Type 4 pin seta (Fig. 4); antennal seg- 
mentation ratio 3.5 :.5 : 6 : 6 with segment II subsegmented (Fig. 5). Labral papillae clearly 
unisetaceous (Fig. 6); labial triangle not observed. LEGS: hind foot complex (Fig. 7) with tenent 
hair shorter than unguis; pretarsus with 2 diagonal, strong bosses; unguis with 2 outer lateral 
teeth, 4 inner teeth, basal teeth more pronounced; unguiculus .52 - .80 times as long as unguis, 
with an outer tooth inserted over 1/2 distance from base. FURCULA: dens 1.08 - 1.22 times as 
long as manubrium; mucronal teeth subequal (Fig. 8). SETAE: Type 5 body setae narrowly fusi- 
form and ciliate for apical .75 of length. Macrochaetae of ABD III with 3 + 3 inner a, 0 + 0 outer 
a, and 2 + 2 m setae (Fig. 9). LENGTH: 2.4 - 2.6 mm. 


TYPES: Holotype (female) preserved in ethanol, 145 paratypes in 95% ethanol, and 26 paratype 
dissection slides prepared with modified polyvinyl alcohol. Holotype and paratypes deposited in 
the Center For Arthropod Diversity Studies, Department of Entomology, Michigan State Univer- 
sity, East Lansing, Michigan. Holotype and paratypes collected from North Carolina, Swain 
County, Great Smoky Mountains National Park, December 15, 1987, from lichens growing on a 
granite boulder, R. J. Snider, collector. 


! Received and accepted August 2, 1997. 
- Department of Zoology, Michigan State University, East Lansing, Michigan 48824. 


ENT. NEWS 108(5) 372-378, November & December, 1997 


Vol. 108, No. 5, November & December, 1997 373 


Figs. 1 & 2 Orchesella gloriosa habitus illustrations. 1. Dorsal view showing color pattern. 2. 
Lateral view showing color pattern. 


DIAGNOSIS 


Orchesella gloriosa n. sp. keys out in Christiansen & Bellinger (1980-81) 
to Orchesella zebra Guthrie. Color pattern will easily separate the two species. 
In addition, the chaetotaxy pattern of ABD III for O. zebra has 4-6 + 4-6 inner 
a, 1(0) + 1(0) outer a, and 3 + 3 m setae. O. gloriosa has 3 + 3 inner a, 0 + 0 
outer a, and 2 + 2 m setae. The claw of O. gloriosa is more robust, ungual and 
unguicular teeth are sharply defined. Chaetotaxy of abdominal segments II 
and IV are also very different. O. gloriosa keys out poorly in Stach (1960) 
between O. zebra Guthrie and O. xerothermica Stach. Color pattern alone will 
separate it from those species. 


374 ENTOMOLOGICAL NEWS 


Figs. 3-9. Orchesella gloriosa morphology. 3. Left eyepatch. 4. Pin seta of antennal segment IV. 
5. Antenna. 6. Labral papillae. 7. Hind foot complex. 8. Mucro, lateral view. 9. Abdominal seg- 
ment III, dorsal chaetotaxy. Figs. 10-16, Orchesella imitari morphology. 10. Left eyepatch. 11. 
Pin seta of antennal segment IV. 12. Antenna. 13. Labial triangle, setal pattern. 14. Hind foot 
complex, 15. Mucro, lateral view. 16. Abdominal segment III, dorsal chaetotaxy. 


Orchesena imitari, NEW SPECIES 
COLOR DESCRIPTION 


Background cream-white to light orange with Tuscan red (deep purple brown) pigmented 
color patterns (Figs. 17 and 18). Head, in dorsal aspect, with dark pigment forming a band be- 
tween antennal bases with circumantennal extensions; genal area with light dusting of purple; 


Vol. 108, No. 5, November & December, 1997 375 


antennal segments distally ringed with Tuscan red and light pigmental dusting becoming pro- 
nounced from base to apex. Trunk dorsally with Tuscan red rings or bands that do not quite 
terminate at pleural region. TH II with light anterior band leaving most of segment background 
color; TH III with wide anterior band extending over 3/4 of segment; ABD I with band 1/2 length 
of segment; ABD II band extends 3/4 length of segment; ABD III with wide band extending 
more than 3/4 length of segment; ABD IV with band covering most of segment; ABD V with 
wide band; ABD VI mostly cream-white to light orange with occasional pigment spots. In dark 
specimens a “T”-shaped pattern is formed connecting segments V and V1. Legs with broken 
pigment spots on precoxae and coxae, rest of legs dusted with light purple becoming darker 
distally, especially tibiae. Furcula and collophore without pigmentation. Many specimens showed 
pigment reduction to the point of only a scattering of light orange with an overlay of common 
body setae defining where the color pattern should be. 


Figs. 17 & 18. Orchesella imitari habitus illustrations. 17. Dorsal view showing color pattern. 
18. Lateral view showing color pattern. 


376 ENTOMOLOGICAL NEWS 


MORPHOLOGICAL DESCRIPTION 


HEAD): eyes 8 + 8 on black patches, ocelli A and B larger than D, E, F, G while H and C are 
smaller and subequal, ratio of G/F ~ .85 - .90 (Fig.10). ANT IV lacking apical bulb, with Type 1 
pin seta (Fig. 11); antennal segmentation ratio 3.5 : 4.5 : 5 : 6 with segment II subsegmented (Fig. 
12). Labral papillae strongly unisetaceous as in gloriosa (Fig. 6); labial triangle setae as illus- 
trated (Fig. 13). LEGS: hind foot complex (Fig. 14) with tenent hair subequal to unguis; unguis 
with 2 outer lateral teeth, 4 inner teeth; unguiculus .48 - .60 times as long as unguis, with an outer 
tooth inserted approximately two thirds distad from base. FURCULA: dens 81 - 1.11 times as 
long as manubrium; mucronal teeth subequal (Fig. 15). SETAE: Type 5 body setae narrowly 
fusiform and ciliate for apical .80 of length. Macrochaetae of ABD III with 3 + 3 inner a, 0 + 0 
outer a, and 3 + 3 m setae (Fig. 16). LENGTH: 1.9 - 2.7 mm. 


TYPES: Holotype (female) preserved in ethanol, 128 paratypes in 95% ethanol, and 13 paratype 
dissection slides prepared with modified polyvinyl alcohol. Holotype and 122 paratypes depos- 
ited in the Center For Arthropod Diversity Studies, Department of Entomology, Michigan State 
University, East Lansing, Michigan. 6 paratypes deposited in the Université de Montreal, Que- 
bec. Holotype and paratypes collected from Canada, Prov. Quebec, Parc de la Gaspésie, Mines 
Madeleine, July 3, 1996, from moss growing in an exposed location at edge of mixed mesophytic 
forest. Additional paratypes taken from the following localities: Rte 299 at Chute Ste. Anne, July 
2, 1996, from mosses growing along footpath and wet spots; Les Lac Castors, July 3, 1996, from 
moss overhanging trail margin; Lac Cascapédia, July 4, 1996, from Sphagnum moss; Lac aux 
Americains, trail to Gite du Mont-Albert, nr. parking lot, July 4, 1996, from Sphagnum moss, (R. 
J. & R. M. Snider, collectors. 


DIAGNOSIS 


Orchesella imitari n. sp. keys out in Christiansen and Bellinger (1980-81) 
to Orchesella hexfasciata Harvey and superficially resembles that species. How- 
ever it can be easily separated from that species by color pattern, and type 1 
pin seta. O. imitari keys out in Stach (1960) between Orchesella leucocephala 
Stach and Ombesella montana Stach. While O. leucocephala is very similar in 
pattern to O. imitari, it has ABD I mostly black banded, imitari is cream- 
yellow with a very narrow band anteriorly; ABD IV of O. leucocephala has 
light areas dorsally, O. imitari is solid dark purple. 


Orcheselia texensis NEW SPECIES 
COLOR DESCRIPTION 


Background white with blue pigmented patterns (Figs. 19 & 20). Head with interantennal 
spot spreading to circumantennal extensions; blue stripe originating at posterior edge of eyepatch 
and ending at occiput; antennal segments with apical rings, light blue dusting becoming darker 
apically on ANT IV. Trunk with 5 longitudinal, broken lines: medial line begins at posterior edge 
of TH II and terminates at posterior edge of ABD IV; paramedial lines begin 1/4 distance from 
anterior of TH II and end at ABD V,; lateral lines originate at anterior of TH II and end at ABD V. 
Legs without pigmentation except trochanter have broken blue patterns. 


Vol. 108, No. 5, November & December, 1997 377 


Figs. 19 & 20. Orchesella texensis diagramatic illustrations from slide mounted specimens. 19. 
Dorsal view showing color pattern. 20. Dorsal view showing reduced color pattern. 


MORPHOLOGICAL DESCRIPTION 


HEAD: eyes 8 + 8 on black patches, ocelli G and half diameter of A-F, G/F ratio .65 - .75 (Fig. 
21). ANT IV lacking apical bulb, with Type 2 pin seta (Fig. 22); antennal segment ratio | : 2: 2.7 
: 2.7, segment II subsegmented (Fig. 23). Mouthparts not observed. LEGS: hind foot complex 
(Fig. 24) with tenent hair subequal to ungual length; unguis with 2 outer lateral teeth and 4 inner 
teeth, unguiculus .93 - .96 times as long as unguis, with outer tooth inserted 3/4 distance from 
base. FURCULA: dens 1.25 - 1.57 times as long as manubrium; mucronal teeth subequal, condyle 
pronounced (Fig. 25). SETAE: Type 5 body setae, narrowly fusiform and ciliate for apical .77- 
.85 of length. Macrochaetae of ABD III with 3 + 3 inner a, 0-1 + 0-1 outer a, and 2(3) + 2(3) m 
setae (Fig. 26). CAPS: Length 2 mm. 


TYPES: Holotype (female) on slide prepared with Marc Andre’s fluid, Texas, Williamson County, 
Jug,cave, May 18, 1989, J. Reddel & M. Reyes, colr. Paratypes taken from the following locali- 
ties: Williamson County, 2 on slide, Raccoon cave, March 16, 1990, litter, J. Reddell & M. 
Reyes, colr.; Travis County, | on slide, Goat cave, January 23, 1991, litter, J. Reddell & M. 
Reyes, colr.; 2 on slide, | in ethanol, Travis County, Hideout cave, January 31, 1991, litter, J. 
Reddell & M. Reyes, colr. Holotype and paratypes deposited in the Museum of Comparative 
Zoology at Harvard University, Cambridge, Massachusetts. 


378 ENTOMOLOGICAL NEWS 


Figs. 21-26. Orchesella texensis morphology. 21. Left eyepatch. 22. Pin seta of antennal segment 
IV. 23. Antenna. 24. Hind foot complex. 25. Mucro, lateral view. 26. Abdominal segment III, 
dorsal chaetotaxy. 


DIAGNOSIS 


Orchesella texensis, sp. keys out in Christiansen & Bellinger (1980-81) to 
O. celsa Christiansen & Tucker. It differs from that species by color pattern, 
O. celsa has the middorsal line not well defined and lacking on ABD IV and V. 
In addition, O. texensis has type 2 pin seta while O. celsa has type 4. In Stach 
(1960) O. texensis keys out to Orchesella xerothermica Stach. This species 
from Poland and Ukraine has the mid-dorsal line originating on the posterior 
of TH II like O. texensis, but not reaching beyond ABD I and it is a large 
species, 4 - 6.2 mm. 


ACKNOWLEDGMENTS 


Special thanks are offered to Kenneth A. Christiansen of Grinnell College, Grinnell, lowa 
for his loan of the O. texensis specimens and critical review of this report. And thanks to Robert 
D. Waltz, Indiana Department of Natural Resources, Indianapolis, Indiana for his review. Thanks 
to Laurent LeSage, Agriculture Canada, Biosystematic Research Center, Ottawa who initiated 
me to the Parc de la Gaspésie Collembola fauna. Very special thanks are extended to Francois 
Landry, Park Biologist, Parc de la Gaspésie for guidance to collection sites known to be produc- 
tive for Collembola; to Renate M. Snider for helping to collect specimens in the field, and to 
Peter H. Carrington for his excellent renderings of O. gloriosa and O. imitari. 


LITERATURE CITED 


Christiansen, K. A. and P. F. Bellinger. 1980-81. The Collembola of North America North of 
the Rio Grande. Grinnell College, Grinnell, IA. 1322 p. 

Stach, J. 1960. The apterygotan fauna of Poland in relation to the world-fauna of this group of 
insects. Tribe: Orchesellini. Acta Mon. Mus. Hist. Nat., Polska Akad. Umiejetnosci, 151 pp. 
+ 15 pl. 


Vol. 108, No. 5, November & December, 1997 379 


NEW SPECIES OF TENAGOBIA 
(HETEROPTERA: CORIXIDAE) 
FROM VENEZUELA! 


J. T. Polhemus2, N. Nieser3 


ABSTRACT: Tenagobia (Baliagobia) castanea sp. n. is described from the Territorio Amazonas, 
Venezuela. 


The genus Tenagobia Bergroth was last revised by Nieser (1977), who 
reviewed previous works and recognized 26 valid species. Since then, Bach- 
mann (1979) described one subspecies. 

The CL number given below refers to ecological data held in collection 
data books with the Polhemus Collection (JTPC). The Nieser Collection is 
noted as (NCTN), the U. S National Museum as (USNM). All measurements 
are in millimeters. 


Tenagobia (Baliagobia) castanea, NEW SPECIES 


Length, male 2.45 - 2.52 mm., female 2.39 - 2.68 mm. 

General color castaneous, head between eyes yellowish, variably infuscated. Hyaline V- 
shaped mark on clavus indistinct, often lacking on posterior part; clavus and corium with hyaline 
guttae, costal margin of hemelytra dark, obscuring darker blotch on middle of corium; hyaline 
stripe on right membrane curved backward, nearly reaching costal margin; left membrane with 
outer half smoky, inner half hyaline, membranal suture distinct. Thoracic venter light to medium 
brown, abdomina! venter medium to dark brown. Legs yellowish, joints and claws darker. 

Head with ocular index of male 1.12-1.20, female 1.10-1.22; sutures of vertex convergent 
behind eyes; laterocaudal angles of vertex perpendicular to slightly obtuse; inner postocular space 
about four times as long as an eye facet. Apicai antennal segment with 2-3 distinctly longer and 
thicker hairs; relative length of antennal segments I:II:III about 2.00: 1 .00:3.25; ratio of width:length 
of apical segment about 3. 

Pronotum short for the genus, lateral margin almost straight, lateral parts of concave poste- 
rior margin somewhat truncate; width, male 0.94-0.95, female 0.96-0.97; proepimera weakly 
expanded, without mustache-like bristles but ventrally with 6 bladelike bristles, a few scattered 
hairs dorsad of proepimeral expansion. Ratio width of pronotum: width of scutellum 1.6-1.8. 

Hemelytra with many minute spines and a few small apical hairs; costal margin without 
spines; ratio, length of pruinose area of embolium: length of hemelytron about 0.2-0.3; only mac- 
ropterous specimens known. 

Fore femur with 2 spines in basal row and | small spinule in apical part, a spine at apical 
1/4 and two apical spines on dorsal face. Pala with 23-24 bristles in lower row, 14-18 in interme- 
diate row and 6 in upper row. Intermediate leg, ratio length of femur:tibia:tarsus:claws about 
8.5:3.0:4.0:2.0. Hind leg, metatrochanter knobby (somewhat less obvious than in subgenus 
Romanagobia), ratio length of femur:tibia:tarsal I:tarsal II:claws about 8:6:6:3:3. Claw appar- 


! Received February 22, 1997. Accepted March 22, 1997. 
2 Colorado Entomological Museum, 3115 S. York St., Englewood, CO. 80110. 
3 Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands. 


ENT. NEWS 108(5) 379-381, November & December, 1997 


380 ENTOMOLOGICAL NEWS 


\ 
= { 
/ ‘\ 
IC \ 


v 
a 


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( 
/ 
1 


Figures | - 4. Tenagobia (Baliagobia) castanea sp. n. 
1. Male abdominal terminalia, dorsal view (scale bar = 0.5 mm). 
2. 3. Male parameres. 2 right, 3 left (scale bar = 0.1 mm) 


4. Female, receptaculum seminis (scale bar = 0.1 mm). 


Vol. 108, No. 5, November & December, 1997 381 


ently with two prongs, the shorter one about half the length of the longer one (measured from 
incision to tip). 

Number of spines and bristles along the lateral margins of abdominal segments: V, 3 spines 
+ | bristle; VI, 2 + 1; VII, 2 (3) + 2 (1); VIII, 3 + 4 (of which 2 are very long). 

Male: paler claw elongate like a very stout bristle; apicodorsal bristle of anterior tibia ab- 
sent. Prostrigil absent; dorsal lobe of left part of tergite VIII broad, bilobed (Fig. 1), with 8 
hooked bristles on apical margin. Parameres characteristic (Figs. 2, 3). 

Female: Length of fore femur subequal to tibia + pala, dorsal tibiotarsal bristle absent. 
Receptaculum seminis simple, ovate (Fig. 4). 

Type material. Holotype, male, Venezuela, Territorio Federal Amazonas, Tobogan area, 
seep pool on rock, CL 2381, 23 Jan. 1989, J. T. Polhemus (USNM). Paratypes, 29 males, 28 
females, 2 nymphs (not paratypes), same data as holotype (JTPC, NCTN, USNM). 


Etymology. The epithet castanea refers to the yellowish-brown color. 


Habitat. The corixid described above was among the many undescribed 
taxa discovered during an expedition in 1989 to the Tobogan Region of the 
Territorio Federal Amazonas, Venezuela, south of Puerto Ayacucho. The 
Tobogan area is named for the smoothly flowing cascade created by the flow 
of the Caio Coromoto over the sloping edge of a large, slick rock shield in the 
jungle. Tenagobia (B.) castanea was collected only from a small, shallow pool 
in rock fed by seeps, in a shallow depression quite removed from the main 
river. 

Comparative notes. In the keys given by Nieser (1977), this species runs 
to the subgenus Baliagobia, previously monotypic. Tenagobia (B.) balia, the 
only other species of the subgenus, is known only from 2 females from the 
upper Rio Negro, Brazil. Tenagobia (B.) balia differs by a lighter color, nar- 
rower receptaculum seminis, lateral margins of pronotum longer and ratio of 
width of pronotum: width of scutellum slightly less. Other Tenagobia species 
have very differently shaped parameres. 


ACKNOWLEDGMENTS 


We are indebted to R. W. Sites and D. A. Polhemus for their constructive reviews. JTP is 
indebted to P. J. Spangler, Smithsonian Institution, Washington, D. C. (USNM), for assisting 
with logisitics in Venezuela. JTP carried out this research as a faculty affiliate of the Department 
of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins. 


LITERATURE CITED 


Bachmann, A. 1979. Notas para una monografia de las Corixidae Argentina (Insecta, Heteroptera). 
Acta Zool. Lilloana 35: 305-350. 

Nieser, N. 1977. A revision of the genus Tenagobia Bergroth (Heteroptera: Corixidae). Stud. 
Neotr. Fauna Envir. 12: 1-56. 


382 ENTOMOLOGICAL NEWS 


A NEW SPECIES OF DACNUSA (HYMENOPTERA: 
BRACONIDAE) FROM SPAIN! 


I. Docavo2, J. Tormos? 


ABSTRACT: Dacnusa rodriguezi, a new species from Spain, is described and compared with 
allied species of the genus. The new species is a parasitoid of Chromatomyia horticola on 
lettuce. 


The subfamily Alysiinae is subdivided traditionally into two tribes, Alysiini 
and Dacnusini (Shenefelt, 1974; Wharton, 1994), whose members are 
endoparasitoids of cyclorrhaphous Diptera. Most of the Dacnusini attack 
agromyzid hosts, and their classification and biology have been studied by 
Griffiths (1964, 1966, 1968, 1984) and Tobias (1986, summary of the Palearc- 
tic taxa with keys to genera and species, translated into English 1995). 

The genus Dacnusa Haliday belongs to the latter tribe, and contains ap- 
proximately 87 Holarctic species. We discovered a new species, described be- 
low, in Jativa (province of Valencia), Spain, reared from Chromatomyia 
horticola (Goureau), a species of agromyzid very common in the Comunitat 
Valenciana (Spain) on cultivated plants (Docavo et al., 1987). 

Terms for body morphology and wing venation follow Griffiths (1964) 
and Wharton (1977, 1986). 


Dacnusa rodriguezi, NEW SPECIES 


Female: Head (Figs. 1, 2, 3) - 1.78-2.0 (x= 1.85) times wider than long, 1.34-1.57 (x=1.45) 
times higher than long; eyes in lateral view 0.7-0.9 (x = 0.8) times as long as temples, slightly 
more closely approximated underneath; head width 1.79-1.95 (x= 1.87) times distance between 
eyes; face fairly smooth, with fine pubescence towards sides and at centre of its foremost part; 
clypeus width 0.62-0.72 (x= 0.69) times distance between eyes; antennae with 20-22 antennomeres; 
mandibles 3-toothed, weakly expanded, 0.28-0.39 (x = 0.33) times length of head, with middle 
tooth blunt; maxillary palpi moderately long. 

Mesosoma (Figs. 2, 3) - 1.21-1.32 (x = 1.26) times longer than high, 1.63-1.90 (x = 1.76) 
times longer than wide; pronotum with a median pit; mesoscutum with dorsal pit, extensively 
smooth, shiny, with pubescence, although longer in its posterior 2/3, covering all its surface; 
notauli weak; prescutellar furrow simple; precoxal suture short, weak, slightly crenulated; 
metapleuron with extended pubescence, towards the posterior coxa; wrinkled propodeum cov- 


1 Received February 12, 1997. Accepted April 10, 1997. 


2 Departamento de Biologia Animal, Biologia Celular y Parasitologia. Facultad de Biologia. 
Universidad de Valencia. C/Dr. Moliner, 50. Burjasot (Valencia). Spain. 


3 Unidad de Zoologia. Facultad de Biologia. Universidad de Salamanca. 3707 1-Salamanca. Spain. 


ENT. NEWS 108(5) 382-388, November & December, 1997 


Vol. 108, No. 5, November & December, 1997 


WwW 
oo 
Ww 


0.5 mm 


2 


Figures | and 2. Dacnusa rodriguezi sp. nov. 1. Head in lateral view, female; 2, Body (except legs 
and wings) in side view, female. 


ENTOMOLOGICAL NEWS 


384 


0.5 mm 


female. 


zi sp. nov. Body in dorsal view, 


Figure 3. Dacnusa rodrigue 


Vol. 108, No. 5, November & December, 1997 


Ww 
oo 
Nn 


C77 oat oc : : oo 
“eG aor ere UD TTTOS eee 


Figure 4. Dacnusa rodriguezi sp. nov. 4a. Anterior right wing, female. 4b. Anterior right wing, 
male. 


386 ENTOMOLOGICAL NEWS 


Figure 5. Dacnusa rodriguezi sp. nov. First tergite in dorsal view, female. 


ered with fine extended pubescence; hind tarsi shorter than hind tibia. 

Wings (Figs. 4a) Pterostigma moderately wide and dark, 1.8-2.0 (x =1.9) times longer than 
metacarp; Im-cu distinctly antefurcal (Fig. 4); Rs sinuate. 

Metasoma (Figs. 2, 3) - First tergite (Figs. 2, 3, 5) 1.3 times longer than wide apically; fairly 
glabrous, with a few fine setae laterally; tergite 3 smooth, without setae on its base; ovipositor 
sheath not extending beyond apical tergite in retracted position. 

Color and size - Head, mesosoma and first tergite black; face black shiny; clypeus dark 
brown; labrum and palpi yellow; antennae dark brown, with yellowish brown scape, base of 
pedicel and annellus; centre of mandibles orange yellow; legs pale yellow, with slightly dark- 
ened tarsi; wings hyaline, with dark pterostigma; second and following tergites yellowish brown, 
becoming darker apically. Body length: 1.49-1.52 mm (x= 1.50 mm). 

Male: Similar to female, but pterostigma wider and dark (Fig. 4b). 

Material examined (deposited in the Fundacion Entomolégica “Juan de Torres Sala” (Docavo 
Collection) (Valencia, Spain)): SPAIN: Valencia: Jativa, 10-VII-1988 (date host capture)/26-29- 
XI-1988 (emergence date of the parasitoids): Holotype, female, from puparium of C. horticola 
(host)/on Lactuca sativa L. (hosts food-plant). Paratypes: 4 females, 3 males, from puparia of C. 
horticola /on L. sativa. 


Vol. 108, No. 5, November & December, 1997 387 


Etymology: This species is dedicated to José Antonio Rodriguez Docavo 
as a token of appreciation for his help in many entomological excursions. 


Notes: This new species is most similar to Dacnusa austriaca (Fischer). D. 
rodriguezi sp. nov. is distinguished mainly by: a) mandibles weakly expanded, 
with middle tooth blunt; b) precoxal suture present; weakly crenulated; c) 
pterostigma much longer than metacarp, and d) first metasomal tergite black. 


This species can be identified by using the keys of Tobias (1995: 226) with 
the following modifications: 


Males 


159 (144) Antennae 22-23 segmented. Mandibles 3-toothed, not expanded. First abdominal 
tergite dark brownish red, 1.7 times longer than wide apically. Stigma narrower than in D. 
melicerta (Fig. 140: 8). Sternauli absent. Body 1.3 mm. Parasite of Liriomyza dracunculi 
Hering, L. artemisicola de Meijere. Center, Central Ural; East Germany; Austria... .... 

AF fey eee sale IRA ie URAC Tere Uslonareel avoids tao nik seer auger aust aeeloke auc D. austriaca Fischer 


159' (144) Antennae 20-22 segmented. Mandibles weakly expanded, with middle tooth blunt. 
First abdominal tergite black, 1.3 times longer than wide apically. Stigma (Fig. 4b). Sternauli 
present. Body 1.5 mm. Parasite of Chromatomyia horticola (Goureau). Spain ......... 

ek. pansy tevege vee Ch sya chas eRe Rat oN edo is Joy sv aVolane cae tee mates afte) na icual te Takemoneeveee D. rodriguezi nov. sp. 


Females 


214 (215) Antennae 21-24 segmented. Mandibles 3-toothed, not expanded. First abdominal 
tergite reddish dark brown, slightly pubescent, 1.7 times longer than wide apically. Stigma 
yellowish dark brown, parallel-sided, few longer than metacarp. Sternauli absent. Body 1.3 
TM) 2ocloc eis clo feeble a avatscatesarenisins Gilat temisrs}s. els oleddlets s\sieleke D. austriaca Fischer 


214'(215) Antennae 20-22 segmented. Mandibles weakly expanded, with middle tooth blunt. 
First abdominal tergite black, fairly glabrous, 1.3 times longer than wide apically. Stigma 
dark brown, much bon ger than the metacarp (Fig. 4a). Sternauli present. Body 1.5 mm .. 

dy Sh its as te tara SGT Set EO SCORN CVG AANG Gocked BuGad cc Suen stators taijetolayor teareLe Pele oiavets D. rodriguezi nov. sp. 


The remaining species of Dacnusini that have been corroborated as parasi- 
toids of C. horticola (Docavo et al., 1987, 1988, 1992; Griffiths, 1984; Spen- 
cer, 1973; Tormos et al., 1989) are: Chorebus canariensis Griffiths; Ch. misellus 
(Marshall); Ch. nana (Nixon); Ch. sativi (Nixon); Dacnusa areolaris (Nees 
von Esenbeck); D. laevipectus Thomson; D. pubescens (Curtis); D. nipponica 
Takada and D. sibirica Telenga. They can be separated from the new species 
described through the keys of Tobias (1995) and Fischer (1994). 

Detailed information on the economic importance and the biology of C. 
horticola has been given by Spencer (1973, 1990). 


388 ENTOMOLOGICAL NEWS 


We wish to thank C. van Achterberg (National Museum of Natural History, Leiden), G. 
Griffiths (University of Alberta), M. Fischer (Naturhistorisches Museum Wien), R. Wharton (Texas 
A&M University) and J. Papp (Hungarian Natural History Museum), for their observations and 
critical reading of the manuscript. We are also very grateful to Ana Sanchis for the drawings, and 
the Fundacion Entomologica “Torres Sala” (Valencia, Spain) for their support in this study. 


LITERATURE CITED 


Docavo, I., Jiménez, R., Tormos, J. & M.J. Verdu. 1987. Braconidae y Chalcidoidea (Hym., 
Apocrita, Terebrantia) parasitos de Agromyzidae (Dipt., Cyclorrapha) en la Comunidad 
Valenciana. Inv. Agrar.: Prod. Prot. veg., 2 (2): 195-209. 

Docavo, I. and J. Tormos. 1988. Aportaciones al conocimiento de los Dacnusini de Espafia (II). 
(Hymenoptera, Braconidae). Boln. Asoc. esp. Ent., 12: 161-163. 

Docavo, I., Tormos, J., Asis, J.D. & S.F. Gayubo. 1994. Dacnusini (Hymenoptera, Braconidae, 
Alysiinae) en la provincia de Valencia (Espafia). Misc. Zool., 16: 105-111. 

Fischer, M. 1994. Untersuchungen tiber Dacnusini der Alten Welt (Hymenoptera, Braconidae, 
Alysiinae). Linzer biol. Beitr., 26/1: 249-28. 

Griffiths, G.C.D. 1964. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae (Diptera). 
I. General questions of taxonomy, biology and evolution. Beitr. Entomol., 14: 823-914. 
Griffiths, G.C.D. 1966. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae (Diptera). 

II. The parasites of Agromyza Fallén. Beitr. Entomol., 16: 551-605. 

Griffiths, G.C.D. 1968. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae (Diptera). 
VII. The parasites of Cerodontha Rondani s.1. Beitr. Entomol., 18: 63-152. 

Griffiths, G.C.D. 1984. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae (Diptera). 
VII. Supplement. Beitr. Entomol., 34: 343-362. 

Spencer, K.A. 1973. Agromyzidae (Diptera) of economic importance. Dr. W. Junk B.V. The 
Hague. pp. 1-418. 

Spencer, K.A. 1990. Host specialization in the world Agromyzidae (Diptera). Kluwer Academic 
Publishers. London. pp. 1-444. 

Tobias, W. I. 1986. Identification key for the insects of the European part of the URSS. Vol. III. 
Part V. Hvmenoptera. Braconidae. pp. 100-105 (key for genera of Alysiinae), 163-231 
(Dacnusini). Akademia Nauk: Leningrad (in Russian, transl. 1995 in English). 

Tormos, J., Gayubo, S.F., Asis, J.D. & M.A.G. Vacas. 1989. Primera contribucion la 
conocimiento de los Braconidae (Hym., Apocrita, Terebrantia) parasitos de Agromyzidae 
(Dipt., Cyclorrapha) en la provincia de Salamanca. Anales de Biologia. Murcia, 15 (4): 
83-86. 

Wharton, R.A. 1977. New World Aphaereta species (Hymenoptera: Braconidae) with a discus- 
sion of terminology used in the tribe Alysiini. Ann. ent. Soc. Am., 70: 782-803. 

Wharton, R.A. 1977. The braconid genus Alysia (Hymenoptera): a description of the subgenera 
and a revision of the subgenus Alysia. Syst. Ent., 11: 453-504. 


Vol. 108, No. 5, November & December, 1997 389 


CATOCALA LOUISEAE, C. GRISATRA AND C. JAIR 
(LEPIDOPTERA: NOCTUIDAE) 
IN NORTH CAROLINA! 


W. J. Cromartie,2 D. F. Schweitzer 


ABSTRACT: Three species of Catocala are newly recorded from North Carolina, all from the 
coastal plain in Bladen County. All are species previously known mainly from localities farther 
south. In addition, the status and distribution of Catocala jair are clarified. 


The lepidopteran fauna of the outer coastal plain of North Carolina is rela- 
tively poorly documented. Recent efforts to obtain a more complete knowledge 
of this area, supported by The Nature Conservancy, the U.S. Forest Service and 
the North Carolina Natural Heritage Program, have found many species not 
previously collected in the state, including numerous species that are found 
mainly in the coastal plain farther south. Surveys in Bladen County, N.C., in 
1992-1994, founa three species of Catocala which fit this latter pattern and 
underscore the unusual diversity of this area. This report is based on occasional 
collecting by the first author. All surveys used blacklight, two bulbs (Sylvania 
350 F15T8/350BL 15 watt) per trap or sheet, and most were run from dark until 
dawn. Sugar bait was used in 1992, but not in 1993 or 1994. Most Catocala 
specimens were taken at light. 

A Single somewhat worn but normal-looking male of Catocala louiseae 
Bauer was collected at a private nature area, 1 km south of Whitehall Road, 
Lake Creek Township, Bladen Co. on 2-3 July 1992 at blacklight. The other 
three closely related Vaccinium (Ericaceae)-feeding species of Catocala, five 
C. gracilis W.H. Edwards (including two form “lemmeri”), one C. andromedae 
(Guenée) and C. sordida Grote, were also taken at this site at blacklight. The 
first two were taken the same night as C. /ouiseae; the third had been taken on 
28-29 May 1992. This is the first known instance of all four of these blueberry 
feeders occurring sympatrically. This record extends the documented range of 
C. louiseae at least 500 km to the northeast. 

Two male Catocala grisatra Brower were collected at Susies Hill, a sand 
ridge north of Johnson Mill Bay in Bladen Lakes State Forest on 25-26 June 
1993. The site was dry forest dominated by Pinus palustris Miller (Pinaceae), 
Quercus laevis Walter (Fagaceae) and Q. incana Bartram, with an understory 
that included Crataegus sp. (Rosaceae) along with lichens and poison sumac. 
This species is generally considered rare (Sargent 1976) and we know of no 
modern records outside of Florida, although the type locality is Athens, Geor- 
gia. C. grisatra feeds on Crataegus. 


! Received January 27, 1997. Accepted March 4, 1997. 
2 Faculty of Natural Sciences and Mathematics, Richard Stockton College, Pomona NJ 08240. 
3 The Nature Conservancy, 1761 Main St., Port Norris NJ 08349. 


ENT. NEWS 108(5) 389-390, November & December, 1997 


390 ENTOMOLOGICAL NEWS 


In 1993 and 1994, Catocala jair Strecker was collected at blacklight on the 
north side of Singletary Lake, just inside the state park boundary: 28-29 June 
1993, four males; 25-26 June 1994, one male; 27-28 June 1994, five males. The 
site was a recently-burned, very xeric, white sand ridge, dominated by medium 
size P. palustris and small (1-3 m) Q. laevis, with a very sparse understory of 
shrubs, grasses and forbs. J. B. Sullivan has also collected three similarly dark 
C. jair in xeric oak scrub in New Hanover County, N.C. on 31 May 1994 and 3 
June 1995 (J.B. Sullivan, pers. comm., determinations by Schweitzer). All North 
Carolina specimens are very dark, similar to the New Jersey specimen illus- 
trated by Sargent (1976, plate 8, fig. 27). Two have the postmedian line very 
straight, like many Florida specimens (Sargent 1976, plate 8, fig. 26). Sargent 
was uncertain whether such dark specimens were conspecific with C. jair. Similar 
dark specimens are found in older collections, mostly from Texas. Smith (1910) 
reported C. jair from New Jersey, but most subsequent authors have discounted 
this record. 

We believe all these specimens are indeed C. jair. The genitalia, which are 
fairly distinctive, are apparently identical in Florida and New Jersey specimens, 
particularly the females (Schweitzer, pers. obs.). Several broods of larvae from 
Citrus County, Florida, and Burlington County, New Jersey, have been reared 
by Dale Schweitzer. Larvae from the two states were virtually identical and 
easily distinguished from those of related species. The habitat in New Jersey is 
also xeric, sandy oak scrub, and larvae occur on shrubby oaks, Q. marilandica 
Muenchhausen and Q. ilicifolia Wangenheim (Schweitzer, pers. obs.). Quercus 
laevis seems likely to be a foodplant in North Carolina. 

We believe that these three species have probably not expanded their ranges 
recently. The most likely explanation of our findings is that they have simply 
been overlooked due to limited collecting, specialized habitats, or, in the case 
of C. grisatra, general rarity. For C. jair, specimens from New Jersey date back 
to at least the 1930’s, so range expansion is highly improbable. 

Most Catocala specimens, including the C. louiseae and one C. grisatra, 
will be deposited in the North Carolina State Museum collection, along with 
other Lepidoptera from these surveys. 


ACKNOWLEDGMENTS 


We thank the late Aubrey Shaw and the staff of Bladen Lakes State Forest and Singletary 
Lake State Park for permission to collect. Dr. Cromartie received funding from the Stockton 
Research and Professional Development Committee in 1992. 


LITERATURE CITED 


Sargent, T.D. 1976. Legion of night. The underwing moths. Univ. Mass. Press, Amherst, 
MA. xiii + 222 pp. 

Smith, J.B. 1910. Insects of New Jersey. Report N.J. State Mus. for 1909. Trenton, NJ. 
pp 15-888. 


Vol. 108, No. 5, November & December, 1997 391 


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MAILING DATES 
VOLUME 104. 1993 
% 


No. Date of issue Pages Mailing Date 
1 Jan. & Feb. 1-76 March 19, 1997 
2 Mar.& Apr. 77-160 May 6, 1997 
| May & June 161-244 July 31, 1997 
4 Sept. & Oct. 245-320 Sept. 16, 1997 
5 Nov. & Dec. 321-396 Nov. 3, 1997 


392 ENTOMOLOGICAL NEWS 


INDEX: Volume 108 


Acari 60 
Acroceridae 190 
Adler, P.H., S.R. Reitz, C.N. Watson 190 
Distrib. & abundance of Eulonchus 
marialiciae 
Aguiar, A.P. 17 
Aldrete, A.N.G., E.L. Mockford 102 
Position of Hemicaecilius & descrip. 
of a n.sp. 
Aleiodes (= Rogas) indiscretus, an 148 
introduced parasite of gypsy moth 
in No. Amer. 
Alloperla of western No. Amer. 321 
Amblycerus schwarzi attacking seeds 229 
of tropical-almond terminalia in 
Cuba 
Amphinemura mockfordi, descrip. of 107 
female, nymph, egg, & redescrip. 
of male 
Announcement 101 
Anthicus unicolor, new predator of 208 
Diuraphis noxia in Turkey 
Anthopotamus verticis, first record 155 
of, & new records of A. distinctus 
in New England 
Aphididae 174, 208 
Appel, A.G. 159 
Aquatic Coleoptera & Hemiptera of 34 
Organ Pipe Cactus Nat’l. Mon., AZ 
Arthropod extraction from soil 203 
cores 
Asis, J.D. 259 
Atopsyche, an.sp. from Costa Rica 161 
Bae, Y.J. 52 
Baetidae 97, 113, 134, 283, 363, 367 
Barber-James, H.M. 363 
Bark beetle outbreak in pine forests 67 
of Sierra de las Minas Reserve, 
Guatemala 
Bauer, S.B. l 
Berytidae NG) 
Beuche, K. 345 
Blahnik, R.J., R.M. Gottschalk 161 


A n.sp. of Atopsyche from Costa Rica 


Blatellidae 
Book Review 
Books rec’d. & briefly noted 


159 


S1s9 
221, 228, 238 


Bowles, D.E., R. Stanford 2977 
New distrib. record for Haideoporus 
texanus, a stygobiontic beetle from 
Edwards aquifer, TX 

Braconidae 66, 148, 382 

Brown, H. P. 179 

Bruchidae 215, 229 

Brunson, M. H. (obituary ) 178 

Burian, S. K. 155 
First record of Anthopotamus 
verticis, & new records of 
A. distinctus in New England 

Byers, G.W. 313 

Caenidae, sister relationship with 52 
Neophemeridae 

Calosoma frigidum, activity & repro- 127 
duction of, in no. Mich. forests 

Cantharidae 239 

Carabidae 127 

Carlton, C.A. 313 

Cassola, F., M.G. Kippenhan 15S 
N. sp. of Oxygonia from Ecuador 

Catocala, three sp. in No. Carolina 389 

Cephidae 24 

Ceramphis, rev. of Nearctic genus 335 

Cercopoidea types of species des- 222 
cribed by E. Schmidt in USNM, 
with lectotype designations 

Chamaemyiidae 174 

Chauliodes rastricornis, a So. 57 
Dakota record for 

Cheleocloeon, contrib. to 283 
systematics of genus 

Chermetidae 19 

Cicindelidae 15 

Clogmia albipunctata, first record 273 
of in cent. Europe 

Cluzobra, new No. Amer. sp. of 151 

Coenagrionidae 77 

Coenonympha tullia inornata, so. 201 


range extension 


ENT. NEWS 108(5): 392-396, November & December 1997 


Vol. 108, No. 5, November & December, 1997 393 


Coher, C. I. 151 
New No. Amer. sp. of genus Cluzobra 


Coleoptera 15929534, G1, 127, 196208; 

213,215, 229, 239, 289; 297, 311, 

335, 345 

Collembola 372 

Conops, Linnean sp. of, with desig- 265 
nations of lectotype 

Cook, J.L., B. Mathison 245 


A n.sp. of Xenos from Ariz., with 
disc. of other No. Amer. sp. 


Corixidae 300, 379 
Corydalidae 57 
Cromartie, W.J., D.F. Schweitzer 389 
Three sp. of Catocala in No. Car. 
Cyrnellus fraternus, descrip: of 253 


pupa, & notes on variation in 
pupal case construction 


Dacnusa, n. sp. of, from Spain 382 

Dasymutilla occidentalis, mating 310 
behavior of 

Davis, L. 183 

Deyrup, M., C. Johnson, L. Davis 183 


Notes on ant Eurhopalothrix flori- 
dana, with descrip. of male 


Dictyoptera 159 
Diplopoda ! 
Diptera 151, 174, 190, 265, 273 
Docavo, I., J. Tormos 382 


N. sp. of Dacnusa from Spain 


Echinobaetis, first adult descrip. 113 
of unusual baetid mayfly genus 
Edmunds, G.F., Jr. 141 
Elmali, M. 
Chamaemyiids as predators of 174 
Diuraphis noxia in Turkey 
Anthicus unicolor, new predator 208 
of Diuraphis noxia in Turkey 
Elmidae 179 
Embolemidae, seasonal distrib. of 231 
in no. & cent. Ark. 
Enallagma, status of some sp. of 77 
Encyrtidae 63 
Entomobryidae 372 
Ephemeroptera 52, 97, 113, 134, 141, 155, 


193, 283, 318, 363, 367 


Ephemeroptera of Spring Creek, Okla. 193 
with remarks on notable records 


Eulonchus marialiciae, distrib. & 190 
abundance of 


Eurhopalothrix floridana, notes on 183 
& descrip. of male 


Fleas from Amazonas & Bahia States, 290 
Brazil 


Formicidae 183, 236 
French, F.E. 209 
Gayubo, S.F. 259 
Genaro, J., J.M. Kingsolver 229 


Amblycerus schwarzi attacking seeds 

of tropical-almond terminalia in Cuba 
George, S.G., J.J. Hoover, H.P. Brown = 179 

Paddlefish, Polyodon spathula, as 

samplers of riffle beetles 


Gottschalk, R.M. 161 


Haack, R.A., G. R-Schwartz 67 
Bark beetle outbreak in pine forests 
of Sierra de las Minas Res., Guatemala 


Haideoporus texanus, a stygobiontic 297 
beetle from Edwards aquifer, TX.., 
with new distrib. record 

Hastriter, M.W. 43 
Report of a morphologically 
hermaphroditic flea & other flea 
anomalies from Morocco 

Hastriter, M.W., N.E. Peterson 290 
Notes on some fleas from Amazonas 
& Bahia states, Brazil 


Hemicaecilius, position of, & des- 102 
crip. of an.sp. frorn Venezuela 
Hemiptera 34 
Heteroptera l75; 300) 305; 3195379 
Heth, R. K. 193 
Hoebeke, E:.R., K. Beucke 345 
Adventive Onthophagus in No. Amer. 
Hofstetter, R.W., K.F. Raffa 63 
New host record for Ovencyrtus 
kuvanae 
Homoptera 174, 208, 222 
Hoover, J. J. 179 
Hydrobiosidae 161 
Hymenoptera 24, 63, 66, 117, 148, 183, 


231, 236, 259, 310, 382 


394 ENTOMOLOGICAL NEWS 


i 


Iftner, D.C. 201 
So. range extension of common ring- 
let, Coenonympha tullia inornata 


Inthavong, C. 60 

Ixodidae 60 

Janus, n.sp. of, from Indonesia 24 

Johnson, C 183 

Johnson, P. J., K.D. Roush, X. Lin Sy 
A So. Dakota record for Chauliodes 
rastricornis 

Johnson, Z.B., J.H. Kennedy 253 


Descrip. of pupa of Cyrnellus fra- 
ternus, with notes on variation in 
pupal case construction 


Karesh, W.B. 60 
Kennedy, J.H. Posy} 
Kingsolver, J.M. 2A A229 
Kippenhan, M. G. 15 
Kondratieff, B.C. 97 
Labiobaetis apache, descrip. of 97 


adults, with add’s. & corrections to 

inventory of Colorado mayflies 
Lachesillidae 102 
Lago, P.K. 311 

New state records for scarab 

beetles from No. Dak. & Minn. 


Lampyridae 239 

Larissimus, first host record for 66 
genus 

Larsen, E. 319 

Larsen, E., C. Olson 34 


Aquatic Coleoptera & Hemiptera of 
Organ Pipe Cactus Nat’!. Mon, AZ 
201, 389 
Levesque, C., G.-Y. Levesque 239 
Abundance & seasonal activity of 
Cantharidae, Lampyridae, & Lycidae 
in a raspberry plantation in Quebec 
Lewis, C.N., J.B. Whitfield 231 
Seasonal distribution of Embolemidae 
in cent. & northern Ark. 
Liang, Ai-Ping 222 
Cercopoidea types of species 
described by E. Schmidt in USNM, 
with lectotype designations 
Lin, X. Si 
Lugo-Ortiz, C.R. 363 


Lepidoptera 


Lugo-Ortiz, C.R., W.P. McCafferty 113 
First adult descrip. of unusual 
baetid mayfly genus Echinobaetis 
Contribution to systematics of 283 
genus Cheleocloeon 
Maliqua, new genus of Baetidae, 367 
for a species previously assigned 
to Afroptilum 
Lycidae 239 
Lyon, M. L., B. P. Stark 321 
Alloperla of western No. Amer. 


Mailing dates - Vol. 108 391 

Maliqua, a new genus of Baetidae, 367 
previously assigned to Afroptilum 

Marin, D.A., J.M. Kingsolver 215 
Prelim. list of Bruchidae of Cuba 

Mathison, B. 245 

May, M. L. 77 


Studies of some species of Enallagma 
McCafferty, W.P. 52, 113, 134, 283, 367 
McCafferty, W.P. 318 

Name adjustments & new synonym for 

No. Amer. sp. of Ephemeroptera 
McCafferty, W.P., G.F. Edmunds, Jr. 141 

Critical commentary on genus 

Siphlonisca 
McCafferty, W.P.,R.K. Heth, R.D. Waltz 193 

Ephemeroptera of Spring Creek, Okla., 

with remarks on notable records 
McCafferty, W.P., C.R. Lugo-Ortiz, 363 

H.M. Barber-James 

Micksiops, a new genus of small, min- 

now mayflies from Africa 
Mecoptera 313 
Megaloptera 57 
Micksiops, a new genus of small, minnow 

mayflies from Africa 


Microdynerus exilis & M. timidus, 259 
descrip’s. of mature larvae 
Miller, R. S. 213 


Female Phengodes feeding, & an 
associated risk 


Mockford, E.L. 102 
Morphologically hermaphroditic flea 43 
& other flea anomalies from Morocco 
Muchmore, W.B. 19 


An unusual new Pachychernes from 
Panama & Mexico 


Vol. 108, No. 5, November & December, 1997 395 


Muscidae 265 
Mutellidae 310 
Mycetophilidae 151 
Nelson, C. H. 107 


Descrip. of female, nymph, egg, & 
redescrip. of male Amphinemura 


mockfordi 

Nemouridae 107 

Neoephemeridae, sister relationship 52 
with Caenidae 

Neoneides muticus: host plants & 175 
seasonality in mid-Appalachian shale 
barrens 

Nicrophorus tomentosus, obs. on bury- 289 
ing a mole 

Nieser, N. 379 

Noctuidae 389 

Notonectidae 319 

Odonata 11, 122 

Odontomachus clarus, ergatandro- 236 
morphism in 

O’Keefe, S.T. 335 
Rev. Nearctic genus Ceramphis 

Olson, C. 34 

Onthophagus, adventive sp. in No. 345 
Amer. 

Ooencyrtus kuvanae, new host record 63 
for 

Orchesella, n.sp. from No. Amer. 372 

Ownership statement 391 

Oxygonia, an.sp. from Ecuador LS 

Pachychernes, an unusual n. sp. from 1g 
Panama & Mexico 

Paddlefish (Polyodon spathula) as 179 
samplers of riffle beetles 

Paeromopodidae ! 

Paiz-Schwartz, G. 67 

Pannota 52 

Pelecinus polyturator, mating 117 
behavior of 

Penteado-Dias, A.M. 66 
First host record for genus Larissimus 

Perlesta xube, new stonefly sp. from 92 
Nebraska 

Peterson, N.E. 290 

Phengodes female feeding, & an 213 


associated risk 


Plecoptera 92 MO S21 


Polhemus, J.T . 305 
New state & US records & other 
distrib. notes for Heteroptera 


Polhemus, J.T., N. Nieser 379 
A n.sp. of Tenagobia from Venezuela 
Polhemus, J.T., R.P. Rutter 300 


Synoptonecta issa: first New World 
record of an Asian water bug intro’d. 


into Fla. 
Potamanthidae 155 
Polycentropodidae 253 
Pseudomops septentrionalis, first 159 
Alabama record of 
Pseudoscorpionida 19 
Psocoptera 102 
Psychodidae 273 
Raffa, K. 63 
Reitz, S.R. 190 
Rhipiphorus luteipennis, notes on 29 
mating behavior of 
Rhodes, H. A . 92 
Robbins, G., et al. 60 


Two noteworthy collections of ticks 
from endangered carnivores in Lao 
People’s Democratic Republic 


Robison, H.W., G.W. Byers, C.A. sii} 
Carlton 
Annotated checklist of scorpionflies 
of Ark. 


Rodriquez-Garza, J.A. 236 
Ergatandromorphism in Odontomachus 
clarus 

Rosenberg, S. 60 

Roulston, T.H., A.G. Appel 159 


First Alabama record of pale-bordered 
cockroach, Pseudomops septentrionalis 


Roush, K.D. 2)// 
Rutter, R.P. 300 
Satyridae 201 
Scarabaeidae Sniles45 
Schonwalter, N. 160 
Schroder, R.F.W., A.M. Sidor 148 


Establish. of Aleiodes (= Rogas) 
indiscretus, an intro’d. parasite of 
gypsy moth in No. Amer. 


Schweitzer, D.F. 389 


396 ENTOMOLOGICAL NEWS 


Sciomyzidae 265 
Scolytidae 67 
Scydmaenidae 335 
Shelley, R.M., S.B. Bauer ] 


New records & species, & taxonomic 
alterations in milliped family 
Paeromopodidae 

Shubeck, P.P. 289 
Obs. on burying beetle, Nicrophorus 
tomensosus, burying a mole 


Sidor, A.M. 148 

Silphidae 289 

Siphonaptera 43, 290 

Siphlonisca, critical commentary 141 
on genus 

Smith, D.R. 24 
A n.sp. of Janus from Indonesia 

Snider, R. J. 372 


New Orchesella from No. Amer. 
Snider, R.M., R. J. Snider 
Activity & reproduc. of Calosoma — 127 
frigidum in no. Mich. forests 
Efficiency of arthropod extraction 203 
from soil cores 
Society meeting reports 76, 182, 252, 344 
Spiroplasma, a new record for Georgia, 209 


& attempted horizontal transmission 
via predation 


Stanford, R. 297 
Stark, B. P. 321 
Stark, B.P., H.A. Rhodes 92 


Perlesta xube, new stonefly sp. 
from Nebraska 


Strepsiptera 245 
Stylopidae 245 
Synaptonecta issa, first New World 300 


record of asian water bug intro’d. 
into Fla. 


Syrphidae 265 

Tachinidae 265 

Tenagobia, a n.sp. of, from 379 
Venezuela 

Tennessen, K. J. 122 
Rate of species descrip’s. in Odonata 

Thompson, F.C. 265 


Linnean sp. of Conops, with 
designations of lectotypes 


Tick collections from endangered carni- 60 
vores in Lao Peoples Democratic Rep. 


Tomberlin, J.K. 310 
Mating behavior of Dasymutilla 
occidentalis 

Tormos, J. 382 

Tormos, J., et al 259 


Descrip. of mature larvae of Micro- 
dynus exilis & M. timidus 


Torres, F. 259 
Trichoptera 161, 253 
Vespidae 259 
Waltz, R.D. 193 
Waltz, R.D., W.P. McCafferty 134 


New generic synonymies in Baetidae 
Wang, T.Q., W.P. McCafferty, Y.J. Bae 52 
Sister relationship of Neoephemeridae 

& Caenidae 


Watson, C.N. 190 


Wedincamp, Jr, J., F.E. French, 209 
R. F. Whitcomb 

A new Spiroplasma record for Georgia, 
& attempted horizontal transmission 
via predation 

Werner, D. 273 
Studies on some moth flies, with 
first record of Clogmia albipunctata 
in cent. Europe 


Wheeler, A.G. Jr. 29 
Notes on mating behavior of 
Rhipiphorus luteipennis 
Neoneides muticus: host plants & NES) 
seasonality in mid-Appalachian 
shale barrens 


Whitcomb, R.F. 209 
Whitfield, J.B. 231 
Xenos, an.sp. from Ariz, with dis- 245 


cuss. of other No. Amer. sp. 


Yeakel, A.M., E. Larsen 319 
New records of notonectids for Penn. 


Zygoptera 177 


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OBRA TAXONONOMICA DEL DR. FRANCESC ESPANOL by A. Viiiolas, O. Escola & J. Vives. 
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