S. B. Peck.
The Eyeless Catopocerus Beetles (Leiodidae) of Eastern North America.
Psyche 81:377-397, 1974.

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THE EYELESS CATOPOCERUS BEETLES (LEIODIDAE) OF EASTERN NORTH AMRI'CA*
BY STEWART B. PECK
Department of Biology, Carleton University Ottawa, Ontario, Canada KIS 5B6
The Leiodid beetle genus Catopocerus Motschoulsky 1869 is composed exclusively of small (1.8-4.5 mm body length), eyeless, wingless, partially depigmented inhabitants of moist forest soil and duff, and occasionally of caves. The genus, the only member of the subfamily Catopocerinae, is known to occur only in North America. The distribution of the genus is principally the unglaciated mountain forests of the eastern and western portions of the continent. Hatch (1957) has reviewed the nine described species from western North America. These range from the San Francisco area northward in coastal forests to Sitka, Alaska. Undescribed species occur in this region and also in southern California, Arizona, and Colorado (per- sonal data). The known and new western species will be covered in a future paper which will also include data from an assemblage of over 2000 specimens collected throughout Oregon by Ellen Benedict of Portland State University.
This paper reports on the two described eastern North American species, and describes three new eastern species. The genus possesses a character common to the family; the eighth antenna1 segment is smaller than either the seventh or ninth segments (except in C. pusio Horn of California in which the seventh and eighth segments are equally small). They may be distinguished from other leiodids by a combination of the 'following characters: eyeless- ness, winglessness, oval shape, dorsal-ventral flattening, open procoxal cavities, five segmented abdomen, and separated metacoxae (Peck, 1973:w).
In the field, series of specimens were taken from moist forest litter, soil, and well-rotted logs. The debris was sifted in the field through a one-half inch mesh screen to remove large objects. The sifted debris was carried and stored in large plastic bags until it could be processed in "Berlese" funnels. When processed, three liters of litter were placed in each funnel (45 cm high, 30 cm across the top, with the ^Manuscript received by the editor December 17, 1974. 377




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3 78 Psyche [September-December
screen 10 cm from the top) on a double layer of cheesecloth sup- ported by the screen. A 40 watt bulb was used over the litter. Two thin wood strips held the top of the funnel above the rim of the bottom and provided ventilation and for the escape of excess moisture. Beetles stopped falling into the collecting bottle at the bottom of the funnel in less than 12 hours. During periods of maximum operation, up to 24 funnels were used, each holding three liter samples of sifted litter for twelve hours. Thus, up to 144 liters of sifted litter could be processed each day.
From 1967 to 1974, in search of Catopocerus and other leiodid beetles, I have extracted the fauna in the above method from 4361 kg of sifted forest litter from the eastern United States. Most of the beetles extracted have been deposited with the CNCI and FMNH, and the arthropod residues are in the FMNH. Barber's Fluid (Valentine, 1942) was used as a preservative under the 'funnels. Species determination can be reliably made only by examination of the male genitalia. This examination is facilitated by the Barber's Fluid which does not harden the tissues of the beetles as does alcohol. The few specimens found under rocks were collected into Barber's Fluid. Some specimens were collected in caves at rotted pig liver bait, and in carrion baited pitfall traps (Peck, 1973). Specimens were cleaned of adhering debris by an ultrasonic cleaner. Dry material was relaxed in boiling water. Dissections were made with minuten needles, in alcohol. Genitalia were observed in alcohol, as temporary glycerine mounts, and dry. Specimens were mounted on points using an alcohol soluble glue. Illustrations were prepared from temporary glycerine mounts.
Measurements are given in millimeters, and were made using a calibrated ocular micrometer disc. The following abbreviations are used in the paper: HW, head width; PW, pronotal width; PL, ps-onotal length ; EL, elvtral length ; and EW, elytral width. Lengths were measured along the midline.
Elytral lengths are from the apex
of the scutellum. Widths are maximum widths. Rounded surfaces are measured as chords of arcs.
The following abbreviations are used to indicate the sources of the material examined: AMNH, American Museum of Natural History; ANSP, Academy Natural Sciences Philadelphia (Horn colln., now in MCZ) ; CAS, California Academy of Sciences; CM, Carnegie Museum ; CNCI, Canadian National Collection of Insects ; FMNH, Field Museum of Natural History ; INHS, Illinois Natural History Survey; MCZ, Museum Comparative Zoology, Harvard University; SBP, Stewart B. Peck; SVAM, St. Vincent Archabbey Museum



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1 9741 Peck - Catopocerus 379
Collection, Latrobe, Penn.; UK, University of Kansas, Entomology Collection; USNM, United States National Museum. The general collector may occasionally encounter Catopocerus under large rocks and logs in mountain forests. Large series are most usually taken in the spring and fall and only by extraction from soil and litter with the Tullgren modification of a Berlese funnel. Some species have been found in caves in the eastern interior low plateaus. Here they are usually under rocks in organic soil near cave entrances. The use of carrion bait has been useful in attracting the beetles in caves.
Food probably consists of organic debris and associated yeasts and fungi. Several species of Catopocerus have been repeatedly taken on subterranean fungi in Oregon and Washington (Fogel and Peck, in MS) and the eastern species may have similar habits. A laboratory colony of the beetles reproduced and developed on bakers yeast on moist soil. C. hamiltoni was taken congregated around a dead beetle larva, and the many beetles taken on carrion bait in caves were un- doubtedly attracted to the carrion for the purposes of feeding and/or oviposition.
Only one certain case of sympathy is known. Thirty-four C. ulkei and I I C. appalachianus were taken together in 10 kg of litter near Whitmer, W. Va.
Reproduction may occur throughout the year in caves, but in forest litter is probably most active in the spring. Larvae and teneral adults have been found most commonly in early spring in litter. One species has been reared from eggs. At I ~ O C , the egg stage lasts about 17 days, the larval stage about 29 days, and the pupal stage 20 days. Three species have been found in caves in Alabama and Tennessee. However, this association with caves is only 'facultative, for these soil- inhabiting (edaphobitic) beetles probably accidentally enter caves, but once there can survive and reproduce. One of these species, C. appalachianus is found at higher elevations in the Appalachians near the soil surface under rocks and in litter. If the preferred environment of the beetles is a montane type of stable, cool, and moist environment, this will be found at lower elevations deeper in the soil. Collecting of surface soil and litter has never taken the beetles at the lower elevations of the Cumberland Plateau or Highland Rim of Alabama or Tennessee. Thus, in the cave region, the beetles' normal soil habitat must be too deep to be



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Psyche [September-December




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19741 Peck - Catopocerus 381
normally sampled, and this deep soil habitat will bring the beetles into 'frequent contact with rock crevices filled and partly filled with soil, which may open into caves with their cool and moist conditions. Hence, in the southern Appalachians, caves can be a more convenient sampling site for soil inhabitants which are usually hidden under a concealing thickness of soil. Other such montane soil inhabitants that are infrequently found at lower elevations except in caves are Anil- linus carabids (Barr, 1969) and Arianops pselaphids (Barr, 1974). Collecting in prolonged wet weather or during winter months might find all of these beetles closer to the surface in the 'Cumberland Plateau or Highland Rim regions of Alabama and Tennessee. Artificial key to the Catopocerus
........................................ Ia. Body length 4rnm or over C. hamiltoni ........................................................ I b. Body length 3mm or under 2
2a. Hind tibia with setal bearing excavation on dorsal posterior sur- .................................................... ~ face (fig. 2) C. alabamae, n. sp.
2b. Hind tibia with setae only, no excavation on dorsal posterior .......................................................................................... surf ace 3
3a. Third antenna! segment clearly longer than second, five clearly larger than four and six; northeastern Alabama C. jonesi, n. sp. 3b. Third antenna1 segment as long as or only slightly longer than the second, five the same size as four and six; widely distributed ...................................................................................................... 4 4a. Tip of aedea,gus curved downward, forming 80å to lmO angle ....................
with ventral surface of aedeagus (Fig. 4) C. ulkei
4b. Tip of aedeagus only slightly curved downward, angle with ven- tral surface of aedeagus I 35O or greater (Fig. 6) ...................... .................................................................. C. appalachianus, n. sp. Catopocerus hamiltoni (Horn)
Fig. 3 ; Map 2
Pinodytes hamiltoni Horn, 1892:45. Lectotype here designated as female in MCZ, Horn col!n. (no. 3027), seen. Type locality: "vicinity of Allegheny City", Pennsylvania.
Figures 1-11. 1, habitus of Catopocerus appalachianus. 2, metatibia of C. alabamae. 3, aedeagus of C. hamiltoni. 4, side of aedeagus of C. ulkei. 5, ventral tip of aedeagus of C. ulkei. 6, side of aedeagus of C. uppala- chianus. 7, ventral tip of aedeagus of C. appalachianus. 8, side of aedeagus of C. jonesi.
9, ventral view of aedeagus of C. jonesi. 10, side of aedeagus of C. alabamae. 11, ventral view of aedeagus of C. alabamae, tip incom- plete because of damage in dissection. Scale line for figs. 3-11 only.



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382 Psyche [September-December
Diagnosis. The large size, the third antenna1 segment which is 1.5 times or more the length of the second segment, and the shape of the aedeagus serve to distinguish this species. Redescription.
Measurement of lectotype: HW, 0.85. PW, 1.63. PL, 1.23. EW, 1.63. EL, 2.74.
Length 4.1-4.7 mm. Width 1.66-1.73 mm. Color uniform dark reddish-brown, shining.
Shape oblong, moderately elongate, not very convex.
Head without trace of eyes; sparsely punctate. Antennae well supplied with long hairs throughout length, last three segments with dense vestiture; segment I11 I .5 X or more the length di I1 ; I11 slightly wider than 11; IV, V, VI as wide as 11, subglobose, slightly longer than wide; VII, IX, X cup shaped; VII longer than but nar- rower than IX and X; VIII shorter than but slightly wider than VI ; XI conical.
Pronotum widest at middle, sides evenly arcuate; narrower at front than back; hind angles rectangular; front angles rounded; front margin evenly emarginate to receive head ; hind margin weakly convex; disc finely but not closely punctate; covered with fine strigae. Elytra fused; at base slightly wider than base of pronotum; widest at middle; sides slightly arcuate; narrowing in apical third; each with seven striae of close punctures, less distinct at sides and apex, intervals flat, sparsely punctate.
Protibia gradually widening to apex; setose on apical half of inner margin. Mesotibia gradually widening to apex; sharply outward flaring in outer margin at apex; spinose on outer concave margin; inner margin sinuous; setose on apical half o'f inner margin. Meta- tibia gradually widening to apex; inner and outer margin slightly sinuous; setae on apical half of inner margin. Penultimate abdominal segment, 'fourth, with transverse depression on either side. Male protarsomeres not dilated, Abdominal segment five truncate. Aedeagus poorly sclerotized; thin and straight in lateral view; wide and straight in dorsal view; tip dorsoventrally flattened, slightly up- turned at edge, smoothly rounded in dorsal view, 14 setae along edge; parameres separated, longest on ventral surface but not reach- ing tip of aedeagus, partially covering anterior ventral surface and dorsal median surface of aedeagus (fig. 3). Female abdominal segment five rounded.
Note on lectotype. The specimen is damaged. It possesses only two prothoracic and one metathoracic legs. One antenna is missing its terminal segments.




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1974J Peck - Catopocerus 383
Distribution. Southwestern Pennsylvania (map 2). The type locality, Allegheny City, lying on the north shore of the confluence of the Allegheny and Monongahela Rivers, was incorporated into the city of ~ittsbur~h near the beginning of this century. Material examined. Pennsylvania. Allegheny, I female, lectotype 3027 (ANSP) ; 2, (ANSP, CM). Beatty, I female (USNM). Charleroi, 2 (MCZ, UK). Jeannette, vi. 1901, 2 (CM). Pitts- burgh, ix, 2 (CM). St. Vincent, 3 (Ulke colln. in CM ; MCZ). Washington Co., I female (Fall colln. in MCZ). State label only,
I paratype no. 3027 (ANSP) ; I (UK) .
Biology.
The available information on the species has been pre- sented by Hamilton ( 1897).
He found the first specimen in Decem-
ber,
1872, "about a foot under ground beneath a large impacted boulder in a wild mountainous place."
Several individuals were seen
but only one was collected because the ale color and remarkable swiftness led Hamilton to believe they were young roaches. This
and a second specimen found dead by Hamilton the following June on a wooded hillside were those which Horn had for description. Other specimens were taken later by other collectors in winter months under bark, under stones, and by sifting decaying leaves. A group of twelve was taken in late November under a log in a wooded ravine around a large dead elaterid beetle larva upon which they were prob- ably feeding. I have not seen specimens taken since the turn of the century.
Catopocerus ulkei Brown
Fig. 4, 5 ; Map I
Catopocer~~s ulkei Brown 1933; 215. Holotype female in CNC, seen. Type locality ; District of Columbia.
Pinodytes cryftophagoides Mannerheim (in part), Horn, 1880 : 249 ; 1892: 46; Hamilton, 1894: 16. Misidentified. Diagnosis. The species is separated from others by its small size, the subequal second and third antenna1 segments, and the flattened, downward curved tip olf the aedeagus.
Redescription. Length I .4-2.2 mm. Width 0.7 1-0.94 mm. Color uniform dark reddish-brown, shining. Shape oblong. Head with trace of eyes as depigmented spot on side of head in some pale specimens; vertex finely punctulate, finely striolate. Anten- nae well supplied with hairs throughout length; segments VII, IX, X supplied with dense vestiture of long and short hairs; I stout; I1 and 111 equally long; I11 narrower at base than 11; IV, V, VI equal, globose; VII as long as broad; VIII smaller than VII, larger than



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Range
males
0.44-0.51
0.81-0.91
0.54-0.63
1.08-1.20
0.80-0.92
1.41-1.58
1.28-1.36
1.72-1.86
1.42-1.56
Psyche
Me an
females males
0.44-0.54 0.49
0.80-0.91 0.85
0.54-0.64 0.59
1.10-1.28 1.15
0.84-0.99 0.88
1.41-1.54 1.48
1.29-1.38 1.32
1.70-1.86 1.77
1.45-1.60 1.50
females
0.50
0.88
0.60
1.12
0.90
1.46
1.34
1.77
1.51
[September-December
Standard Deviation
males
0.024
0.034
0.30
0.052
0.034
0.05
0.03
0.05
0.05
females
0.034
0.054
0.047
0.071
0.045
0.05
0.03
0.04
0.06
Table 1. Variation in a sample of a population of Catopocerus ulkei topotypes from the District of Columbia. 10 males and 10 females. Measurements in millimeters.
VI; IX smaller than X, larger than VII; IX and X broader than long; XI conical.
Pronotum widest just behind middle; sides evenly arcuate; nar- rower at front than back; hind angles slightly obtuse; front angles rounded; front margin slightly concave; hind margin straight; disc finely and sparsely punctulate; finely striolate. Elytra fused; at base slightly wider than base of pronotum; widest at middle; sides slightly curved; narrowing in apical third; surface finely punctate, some punctures vaguely indicating striae; finely striolate; apical declivity smooth.
Protibia widening to apex in distal 2/3; setose on apical half of inner margin; mesotibia gradually widening to apex in apical 2/3; inner margin setose in apical third, faintly sinuous; outer margin spinose; outer apical angle truncate. Metatibia gradually widening to apex; outer margin smooth; inner margin smooth to serate; outer apical angle slightly truncate.
Male first four protarsomeres dilated and supplied with long lateral hairs ; first three mesotarsomeres feebly dilated. Aedeagus in lateral view arcuate; ventral surface of anterior end projecting for- ward and curved downward ; tip dorsoventrally flattened and turned downward forming So0-iooO angle with ventral surface of aedeagus; parameres long and thin, exceeding tip of aedeagus, hinged at aedea- gus base to ventrally deflected elongate basal pieces (figs. 4, 5).



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19741 Peck - Catop ocerus 385
Female tarsomeres not expanded; abdominal segments as in male. Variation. Table I
presents meristic variation for the topotypic population. All populations examined which are assigned to this species have aedeagi 04f the males identical to aedeagi of the topotypic population but for two exceptions. The two samples from south- western North Carolina are 220 air miles to the southwest of the next nearest locality from which males have been collected. These southwestern collections differ in the angle of deflection of the aedea- gal tip, A distinct acute angle of about 80å is formed between the ventral surface of the aedeagus and the tip of the aedeagus. All other populations possess aedeagi in which the angle is usually slightly obtuse. Only rarely is it a right angle. The variation of measurements for a sample of four males and eight females from Joanna Bald, Cherokee County, North Carolina, when compared to that in table I, showed that the size of the south- ern populations is larger, but there is a great deal of overlap within the range of the measurements. Variation of the ratios, when com- pared to the topotypic population shows less difference, and that which does exist is contained within an overlap of one standard deviation from each mean. This numerical data demonstrates little or no difference from the topotypic C. ulkei. Since there is a laxge gap in knowledge of the range of the species between western Virginia and southwestern North Carolina, I am unwilling to interpret the taxo- nomic significance o'f the greater curvature of the tip of the aedeagus in the southwestern North Carolina populations. It may be a distinct subspecies if its distribution is geographically separated from typical C. dkei. Or the population may represent one end or segment of a cline of gradual deflection of the aedeagal tip. Further collecting
may tell.
Distiibution. The species is known to occur in southwestern and southeastern Pennsylvania, the District of 'Columbia, Maryland, North Carolina, Virginia, and West Virginia (map I). Future col- lecting might show it to be continuously distributed along the Pied- mont and eastern edge of the Appalachians to the southern known locality in Cherokee County, North Carolina. Material examined and habitat data. District of Columbia: no- other data, 32. Washington, D. C., I ; 1.5, I ; 5.3, I ; Hubbard & Schwarz. Maryland: Garrett Co.; Deer Park, 4.7, I, Hubbard & Schwarz; 2 mi. E. Keysers Ridge, 2500' elev., 18.vi.1968, IS. Peck, 3 in 220 lbs. log-stump litter. North Carolina: Cherokee-Graham County; Andrews, Joanna Bald, 4700' elev., 26.vii.1967, S. Peck & A. Fiske, 13 in 89 lbs rotted chestnut log; Joanna Bald, 5.viii.1960,



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386 Psyche [September-December
Map 1. East central United States and Catopocerus distributions. Closed dots, C. ulkei.
Open dot, C. alabamae. Closed triangles, C. jonesi. T. C. Barr, I male. Clay-Macon County; Tusquitee Bald, 4.viii. 1960, T. C. Barr & M. C. Bowling, I male. Pennsylvania: x, I. Allegheny County; Pittsburgh, x, 3. St. Vincent, 9. Cambria County; Nicktown, M. Wirtner, 3. Philadelphia County; 28.vi. 191 I, G. M. Greene, I. Westmoreland County; Chestnut Ridge, E. of Youngstown, 27.vi.1961, W. Suter, I in litter under Rhodo- dendron; I I .vii.1961, W. Suter, J. Wagner, D. Reichle, I in forest litter. Jeannette, vii, 2; ix, 3; x, I. Virginia. Carol1 'County; Groundhog Mt., 2900' elev., Blueridge Parkway, 23.vii. 1967, S. Peck & A. Fiske, I female in 46 Ibs. rotted stump and log litter in Rhododendron thicket. Giles County ; Mountain Lake, I-5.vii. I 970, W. B. Muchmore, 8. Madison County; Shenandoah National Park. Dark Hollow Falls, 2.iv.1967, S. Peck, I under rocks. Lewis Mt., 2000' elev., 1o.vi.1967, S. Peck, 104 in rotted logs. Page County; Shenandoah National Park. Hawksbill Mt., 17.x.1954, D. G. Kis- singer, I. Stony Man Overlook, 2.iv.1967, S. Peck, 3 under rocks and 6 in litter. West Virginia. Greenbrier County; outside Ar- buckle Cave, 14.vii.1970, W. B. Muchmore, I in litter. Mercer County; 5.5 mi. N. Princeton, Brush Creek Falls, 13.v.1971, W. Shear, I in cliffside litter. Pocahontas County; W. side 'Cranberry Glades, base of mountain, streamside, under rock, 28.vi.1967, T. C.



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19741 Peck - Catopocerus 387
Barr, I 'female. Hills Creek Falls, 6.v.1970, W. Shear, I ; 1g.vi. 1971, Shear and Platnick, 2. Randolf County; Bickle Knob, 3800' elev., E. of Elkins, ig.vi.1968, S. Peck, 34 in I 79 lbs. of log litter. 5 mi. S. Whitmer, 18.vii.1971, S. Peck, 3000' elev., 34 in litter. (AMNH, CAS, CM, CNC, FMNH, INHS, SBP, SVAM, UK, USNM).
Catopocerus appalachianus New Species
Fig. 6, 7 ; Map 2
Holotype male (MCZ no. 32228) from Balsam Gap, 5500' elev., Mt. Mitchell, Yancey Co., North Carolina, 9.vi.1967, S. Peck. Paratype= all material-liste<L-in section on material examined. Diagnosis.
Most similar to C. ulkei but differing distinctly from it and all other species in the shape of the aedeagus. Usually differ- ing from C. dkei in its larger size and greater EW/PL ratio. Description. Length 2.14-2.86 mm. Width 0.83-1.10 mm. Color uniform dark reddish-brown, shining. Shape oblong.