Glenn K. Morris and Darryl T. Gwynne.
Geographical Distribution and Biological Observations of Cyphoderris (Orthoptera: Haglidae) with a Description of a New Species.
Psyche 85:147-167, 1978.

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PSYCHE
VoI. 85 June-September, 1978 No. 2-3
GEOGRAPHICAL DISTRIBUTION AND BIOLOGICAL OBSERVATIONS OF CYPHODERRIS
(ORTHOPTERA: HAGLIDAE)
WITH A DESCRIPTION OF A NEW SPECIES
BY GLENN K. MORRIS' AND DARRYL T. GWYNNE~ With the ex.ception of Prophalangopsis obscura (F. Walker) from India, Cyphoderris are sole survivors of a primitive orthopteran family, the Haglidae, abundant in the Triassic and ancestral to modern Ensifera (Zeuner, 1939; Ander, 1939; Ragge, 1955; Sharov, 1968).
There are presently two recognized species of Cyphoderris: C. monstrosa Uhler and C. buckelli Hebard. Their most dramatic distinguishing feature is the presence in C. monstrosa, and the absence in C. buckelli, of a prominent ventrally-directed sternal process, shaped like the claw of a hammer and located on the IXth sternum (Hebard, 1934). Specimens of both species have been extensively collected from mountainous areas of the North Ameri- can northwest.
When Uhler established Cyphoderris in 1864 he had before him two adult male specimens. He published body measurements for both of these and there is a substantial size difference e.g. body length 22 mm for one specimen and only 16 mm for the other. These specimens are in the Museum. of Comparative Zoology, Harvard University and we have examined them. The larger has a prominent IErindale College and Department of Zoology, University of Toronto, Missis- sauga, Ontario, L5L lC6, Canada.
2Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado, 80523, U.S.A.
Manuscr@t received by the editor November 15, 1978.



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148 Psyche [June-September
sternal process and is C. monstrosa of present usage; the smaller specimen lacks the process and is C. buckelli of Hebard. It is the C. buckelli specimen which bears a red 'Type 792' label. Uhler's description concentrates on the larger specimen. Thus monstrosa is said to have a "pointed keel-like elevation, projected backwards upon the segment, grooved and emarginated at tip", i.e. a sternal process. He does not indicate that such a structure is absent from the other specimen.
Caudell (1904) described a variety of C. monstrosa which he called Cyphoderris monstrosapiperi. His types, which we have seen, are a male and a female from Mt. Rainier, Washington, housed in the U.S. National Museum. The male has a sternal process identical with that of Uhler's larger specimen and the female has Ander's organs (see below).
In 1922 Fulton collected a series of males in Oregon about 30 mi. southwest of Crater Lake (Fulton, 1930). He compared these with specimens furnished him by E. R. Buckell from southern British Columbia and found that Buckell's specimens lacked a genitalic process. Drawings were sent to Nathan Banks and to Caudell who compared them with the types of monstrosa and piperi. It might now have become apparent that Uhler's types differed in their genitalia and that only the larger was of the same species as piperi. Since the published description applied substantially to the larger specimen one would then have expected it to be designated as monstrosa. But for some reason piperi was given specific status by Fulton and applied to the taxon with the sternal process while Uhler's name was conferred upon the smaller of Uhler's two species. Probably it was at this point that the red type label was appended at Harvard.
Hebard (19341, responding to Uhler's published description, recognized piperi as a synonym of monstrosa and gave the name buckelli to the species without the sternal process. It is clear that he did not examine Uhler's types and was unaware that one of these was his new species. Uhler did not designate a holotype and so in accordance with Article 74 of the code and in the interest of taxonomic stability, we here designate as lectotype the larger of the two specimens in his type series, that possessing the sternal process. This ensures that application of the name monstrosa continues in conformity with present custom.




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A third species of Cyphoderris, C. strepitans, is described here. Its distribution lies southeast of both monstrosa and buckelli (Figure 5), populations of strepitans being originally considered as southern range extensions of monstrosa (Alexander, 1935; Willey and Willey, 1963). C. strepitans appears to be most similar in rnorphology and calling song to buckelli but is readily distinguished from the latter species by the structure of the male terrninalia. We have made use of the following abbreviations: ROM/ Royal Ontario Museum, Toronto, Canada; UMMZ/ University of Michi- gan, Museurn of Zoology, Ann Arbor; ANSP/ Acaderny of Natural Sciences, Philadelphia; CNC/ Canadian National Collection, Ot- tawa, Canada; MCZ/ Museum of Comparative Zoology, Harvard University; USNM/ National Museum of Natural History, Srnith- sonian, Washington, D.C.
Cyphoderris strepitans new species
Figures 1 and 2
The specific name refers to the calling song: strepitans (Latin) 'rnaking a great noise'.
Cyphoderris monstrosus, Thotnas (not Uhler), 1876. Proc. Davenp. Acad. Nat. Sci. I, p. 263. Wind River, Wyo.
Cyphoderris monstrosa, Hebard (in part not Uhler), 1934. Trans. Am. ent. SOC. 59, p. 374. Pearl, Col.
Cyphoderris monstrosa, G. Alexander (not Uhler) 1935. Ent. News 46, p. 30. Park Range (Pearl) Col.; ibid 1941. Univ. Col. Studies 1, p. 136. Cyphoderris monstrosa, Willey & Willey (not Uhler) 1963. Ent. News 74, p. 200. Los Pinos Pass, Col.
C.yphoderris monstrosa, Evans (not Uhler) 1970. Bull. Mus. Cornp. Zool. Harv. 140, p. 484. Jackson Hole, Wyo.
HOLOTYPE. Adult 0. Park Range 24.7 mi. west of Cowdrey via Pearl, nr Big Creek Lakes, Jackson Co., Col., U.S.A.; 19 June 1976; Coll. G. K. Morris & D. T. Gwynne (Deposited ROM). DESCRIPTION OF MALE TYPE. Body length (fastigiurn to para- procts in dorsal view) 18.6 rnrn; pronoturn rnid-line length 7.2 rnrn; caudal pronoturn width 7.8 rnrn; maximurn exposed tegrninal length in dorsal view 8.2 rnrn; length in lateral view of left rnetathoracic femur 8.3 rnrn.




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19781 Morris & Gwynne - Cyphoderris 151
ALLOTYPE.
Adult 0, Los Pinos Pass, Saguache Co., Col., U.S.A.; 17 June 1977; Coll. D. T. Gwynne (Deposited ROM). DESCRIPTION OF FEMALE ALLOTYPE. Body length (fastigiurn t0 extrernity of epiproct in dorsal view) 21.5 rnrn; pronoturn mid-line length 5.9 rnrn; caudal pronoturn width 5.4 rnrn; length of ovipositor 2.0 rnrn; length in lateral view of left rnetathoracic fernur 9.6 rnrn. DIAGNOSIS. Adult males of strepitans are similar in size and coloration to buckelli, but readily distinguished by the presence of the sternal process (Figures 3 & 4). Males of both strepitans and buckelli are generally smaller than males of monstrosa. In life they usually lack the vivid pink coloration of monstrosa's venter, their venters being instead cream white. The styli of the IXth sternum are strongly depressed in monstrosa; viewed from above each stylus is sublanceolate and broadest at its base; they are inserted on the larnellate dorsal projection of the IXth sternum at a distance slightly greater than the stylus length. By contrast the styli of strepitans are distally dilated and broadly rounded (mitten-like), gently arcuate and tapering slightly to the base; they are inserted close together immediately adjacent to the mid-line with less than one stylus length between their bases.
The sternal process of monstrosa, viewed in lateral outline, follows a broadly concave arc beyond the base of the styli to where it turns abruptly downward; in strepitans this arc is shorter and much shallower. From the end of the arc the process of monstrosa is more strongly reflexed than in strepitans and is often bent sharply forward at its extrernity.
We are unable at present to distinguish between females of buckelli and strepitans but both of these species may be separated from monstrosa by their lack of the 'stridulatory' organs of Ander (1938). In C. monstrosa these structures are located dorsolaterally at +he junction of abdomen and thorax (Ander, 1938; Kevan, 1954; Dumor:ier, 1963). Each organ consists of a row of robust posteri- orly-directed recurved teeth on the slightly swollen posterolateral edge of the rnetanoturn. The teeth contact a patch of transverse ridges on the first abdorninal tergite during telescoping of the abdominal segment. Ander's organs are present in both sexes and are readily seen in later stadia of irnrnatures. While buckelli sometimes possesses weak thoracic teeth, it never exhibits the ridged



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19781 Morris & Gwynne - Cyphoderris
Figure 4.
Lateral view of IXth sternum, A) C. strepitans, B) C. monstrosa, and C) C. buckelli.
patch. No stridulatory function has been established for these structures.
Paratypes. 9 adult m, same data as holotype (ROM, MCZ, & UMMZ); 1 adult Q, same data as holotype (ROM); 4 adult m, Cowdrey, 8.8 mi. west on road to Pearl, 4 June 1978 (G. K. Morris & D. T. Gwynne) (ROM & Colorado State Univ., Ft. Collins); 1 adult 8, Park Range nr Pearl, Col., 17 Aug. 1932 (G. Alexander) (ANSP); 1 adult Q, Pearl, Col., 17 Aug. 1932 (G. Alexander) (Museum of University of Colorado, Boulder, Col.); 7 adult $$, Los Pinos Pass, Col., 17 June 1977 (D. T. Gwynne) (USNM & CNC); 1 irnrn. Q, Los Pinos Pass, Col., 17 June 1977 (D. T. Gwynne) (ROM); 1 adult Q, Dunraven Pass, Yellowstone Nat. Pk, Wyo., 25 June 1930 (E. C. Van Dyke) (UMMZ); 1 adult Q, Jackson Hole Res. Stn, Grand Teton Nat. Pk, Wyo., Aug. 1967 (H. E. & M. A. Evans) (USNM); 1 irnrn. 8, Jackson Hole Res. Stn, Grand Teton Nat. Pk, Wyo. (H. E. & M. A. Evans) (USNM); 1 adult Q, Stratton Exp. Watershed, nr Saratoga, Wyo. (J. M. Schrnid) (USNM).



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Key to Cyphoderris Species
(Males only)
Subgenital plate (IXth sternum) with prominent ventrally- ........................
directed process (Figures 4a, b) .2
Sternal process absent (Figure 4c) ..... C. buckelli Hebard IXth sternum strongly produced posteriorly; sternal process with an angularly forward-bent tip, often appearing terminally cleft and toothed; styli of IXth sternum depressed, sublanceo- late (Figure 3b); Ander's organ ridge-patch present and thoracic teeth robust; fastigiurn often weakly rugose ............... ................................... C. monstrosa Uhler Posterior of sternal process not angular but rounded, never terminally cleft and toothed (Figure 3a); Ander's organ absent or if present, only as weak thoracic teeth; styli of IXth sternum mitten-like; fastigiurn smooth ... C. strepitans new species C. strepitans, as presently known, is confined to the mountains of Colorado and Wyoming (Figure 5). Its distribution is disjunct from that of buckelli and monstrosa. The broad valley of the Snake River isolates it on the northwest from the most southerly populations of Idaho monstrosa; if overlap occurs it must be north of Yellowstone in southern Montana.
C. monstrosa is found from the Canadian Rockies in the south- west corner of Alberta, west through southern British Columbia. It reaches much farther north than buckelli, to Quesnel and to Srnithers B.C. (This latter record exceeds the northern extent of our map and could not be plotted; Srnithers is about 700 miles north of the Canada1U.S. border.) A western arm of monstrosa extends down the Cascades, reaching almost to northern California. A less documented eastern arm crosses western Montana to a cluster of localities in the Salmon River Mts of central Idaho. C. buckelli has a more restricted range. It lies between these arms, overlapping broadly with monstrosa in southern B.C. and extending south through northern Idaho. There are interesting isolated rec- ords from Columbia Falls, Montana and from near Seneca in east central Oregon. Though their distributions overlap substantially we have not found monstrosa and buckelli together at the dernic level.



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Morris & Gwynne - Cyphoderris
Cyphoderris Cyphoderris
buckelli D rnonstrosa ft
Cyphoderris
strepitans A
Figure 5.
Geographical distribution of Cyphoderris.



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156 Psyche [June-September
However Buckell (1924) states that both species have been seen at Nicola, B.C. "in large numbers during late May feeding together upon flowers of Amelanchier . . .".
We have examined about 300 specimens of monstrosa and 200 of buckelli. In addition to the type, allotype and paratypes of strepi- tans, we have seen 20 or so alcoholic specimens supplied by Dr. R. Willey of the Univ. of Chicago. Dr. Willey's material is from Los Pinos Pass, Col., a locality which he discovered in 1962 (Willey & Willey, 1963).
All plotted localities are based upon actual examination of specimens excepting Wind River, Wyo. This record is taken from Thomas (1876) on the strength of his illustration of an adult male. Though the drawing is small the shape of the sternal process is apparent and marks the specimen as strepitans. A listing of localities is given below for each species.
C. strepitans: WYOMING: Dunraven Pass, Yellowstone N. Pk; Jackson Hole, Grand Teton N. Pk.; Wind River, Fremont Co.; Stratton Exp. Watershed, nr Saratoga, Carbon Co. COLORADO: Park Range, nr Big Creek Lks, Jackson Co.; Cowdrey, 8.8 mi. west, Jackson Co.; Los Pinos Pass, Saguache Co. C. monstrosa: BRITISH COLUMBIA: Srnithers; Quesnel; Chilcotin nr Williams Lk; Lac La Hache; Clinton; Lillooet; Whistler Mt., Garibaldi Prov. Pk; Salmon Arm; Field, Yoho N. Pk; Glacier N. Pk; Monck Prov. Pk; Merritt; Lurnby; Peachland; Fish Lk, nr Summerland; Manning Prov. Pk; Hedley. ALBERTA: Jasper N. Pk; Mt, Eisenhower Cpgrd, Banff N. Pk; Sulphur Mt., Banff N. Pk; Bragg Creek, w. of Calgary; Barrier Lk, Kananaskis Valley; Ka- nanaskis Lks, Kananaskis Valley. MONTANA: Belton, Flathead Co.; 2 mi. s. Elliston, Powell Co. IDAHO: McCall, Valley Co.; Challis, Custer Co.; Beach Cr., nr Bull Trout Lk, Custer Co.; Red Fish Lk, Custer Co.; Centerville, Boise Co.; 22 mi. ne. Idaho City, n. fork Boise R., Boise Co.; Carnas Co.; Arco, Butte Co. WASHINGTON: Lk. Wenatchee St. Pk, Chelan Co.; Entiat R. Trail, Chelan Co.; 2 mi. se. Easton, Kittitas Co.; Stampede, King Co.; Paradise Valley, Mt. Rainier N. Pk.; Berkeley Park, Mt. Rainier N. Pk; Gooseprairie, Yakirna Co.; Trout Lk Cpgrd, Klickitat Co. OREGON: Mt. Hood, Hood R. Co.; Hat Point, Wallowa Co.; Middle Sister, Lane Co. & Deschutes Co.; McKenzie Pass, Lane Co. & Deschutes Co.; Salt Creek Falls, Lane Co.; Waldo Lk, Lane Co.; Lost Lk, Willarnette Nat. For., Linn Co.; North Santiarn R., Linn Co.; Pinehurst,



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Jackson Co.; Union Creek, Jackson Co.; Crater Lk N. Pk, Klarnath Co.; Douglas Co.; Olallie Lk, Marion Co.; Mt. Jefferson, Jefferson Co.
C. buckelli: BRITISH COLUMBIA: Chilcotin; Paul Lk Prov. Pk; Squilax; Salmon Arm; Nicola Lk; 2 mi. s. Merritt; Aspen Grove; Vernon; Lurnby; Kelowna, nr airport; 3 mi. se. Rutland; Okanagan Falls; Rock Creek; Yahk; Ainsworth; Rosebery Prov. Pk; Cran- brook; Boswell, Kootenay Lk. IDAHO: Reeder Bay, Priest Lk, Bonner Co.; Sandpoint, Bonner Co.; 10 mi. s. Coeur d'Alene, Kootenai Co.; Moscow Mt., Latah Co.; Moscow, Latah Co.; Karniah, Lewis Co. WASHINGTON: Newport, Pend Oreille Co.; Palouse, Whitrnan Co.; Pullman, Whitrnan Co. MONTANA: Colum- bia Falls, Flathead Co. OREGON: 3 mi. e. Seneca, Grant Co. HABITAT AND FEEDING BEHAVIOR
The distribution of Cyphoderris corresponds roughly with the Cordilleran forest province (Gleason & Cronquist, 1964). In south- ern British Columbia C. buckelli occurs in the Dry Forest biotic area (Cowan & Guiguet, 1965) characterized by yellow pine (Pinus ponderosa) and at higher elevations, by interior Douglas fir (Pseu- dotsuga menziesii). Amelanchier (serviceberry), Balsamorhiza (ar- row-leaf balsam-root) and Berberis (tall Oregon grape) are common understory plants in this association. In spring the nymphs and adult females of C. buckelli feed upon the flowers of these plants; night collecting at blooms is a good way to obtain specimens. C. buckelli is also found in the Columbia Forest biotic area, the so-called interior wet belt of British Columbia. In 1977 we located large populations adjacent to Kootenay Lake near Boswell and at Rosebery on Slocan Lake.
C. monstrosa does occur in Dry Forest e.g. at Monck Prov. Park in southern British Columbia, but it is typically encountered in Sub- alpine Forest. Lodgepole pine (Pinus contorts) and Englernann spruce (Picea engelmannii) are characteristic of the Sub-alpine biotic area. In the Kananaskis Valley of southern Alberta we observed nymphs and adults to feed upon staminate cones of lodgepole pine (this before the cones reach a 'loose pollen' stage). Consumption was established by identifying cone bracts in the feces of field-caught specimens. Also caged insects were given cones and in most cases overnight they ate large portions.



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158 Psyche [June-September
It is presumably to feed upon staminate cones that C. rnonstrosa nymphs and adult females are observed at dusk climbing high into the trees. A useful method of collection is to search tree trunks with a flashlight just after sunset. The insects are always discovered oriented head upward. More often than not they occur in groups of 2 to 4 on the same trunk, which suggests that they may aggregate during their daytime stay in the leaf litter of the forest floor. David Lightfoot of. Oregon State University has studied C. rnonstrosa at Three Sisters in the central Oregon Cascades. He found this species abundant there in drier, more open stands of lodgepole pine and mountain hemlock (Tsuga rnertensiana) above 5000' elevation. In contrast to our observations of ascent as the evening progresses Lightfoot notes that the singers begin high in the trees (about 6 rn) and gradually occupy lower and lower perches until singing at ground level.
C. strepitans is found in both subalpine forest and high altitude sagebrush prarie. The holotypic site near Big Creek Lakes is an open forest of subalpine fir (Abies concolor) and lodgepole pine at an altitude of 8800'. At Los Pinos Pass, Colorado, strepitans is found in aspen woods adjacent to open areas of prairie. The mature aspen (Populus trernuloides) has an understory of subalpine fir and englernann spruce (altitude 10,200'). Two predominant ground cover plants at both sites are kinnikinik (Arctostaphylos spp.) and a shrubby juniper (Juniperus cornrnunis). In the high altitude (8400') sagebrush (Artemesia tridentata) prairie of North Park, Colorado, the density of singing males appeared to be much greater than in the nearby pine forest of the holotypic site to the east. C. strepitans is also very numerous in the sagebrush areas (altitude 6700') of Grand Teton National Park, Wyoming. In late June, 1978, aggregations of singing males were easily heard while we drove along park roads at night. Thus, C. strepitans may be considered a predominantly sagebrush species although occurring in open forest habitats in the vicinity of sagebrush prairie.
Males of all three species produce a succession of short musical trills, beginning in late evening and continuing well past midnight if weather permits. C. buckelli invariably sing near the ground from low shrubs (knee-height), the bases of tree trunks or on the forest



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floor itself. The same is true of C. strepitans. Only C. monstrosa climb high into the trees as the night's signalling progresses. At Monck Park singing heights in excess of 5 rn were common and an hour after sunset collection without climbing trees becomes irn- possible.
The calling songs are generated by tegrninal stridulation. As in Gryllidae the tegmina are morphological mirror-images, both left and right bearing a functional file and scraper. Unlike gryllids however, which maintain a characteristic 'right above' forewing overlap, the overlap of a Cyphoderris male may change during his lifetime and both files take part in his stridulation. Certain Tettigoniidae also have mirror-image tegrnina and two functional files: Megatympanon speculaturn Piza (Listroscelinae) (Riek, 1976), Neduba macneilli Rentz & Birchirn, Neduba sierranus Rehn & Hebard (Decticinae) (Morris et al., 1975). Most tettigoniids have structurally distinct left and right forewings and overlap them 'left above'. In the Neduba species some individuals show left above, some right above. Unlike Cyphoderris they appear to maintain their particular overlap as individuals through life. Both overlaps were represented by Riek's two (pinned) specimens of M. speculaturn. Spooner (1973) analysed the calling song of C. rnonstrosa and describes it as a trill of grylloid (sinusoidal) pulses at a carrier frequency of 13 kHz. He noted substantial variation in the intensity and frequency of pulses and suggested that these changes "reflect irregular switching of tegrnina from top to bottom position". He refers to this habit as "switch-wing singing" and regards it as occurring several to many times in the course of a single trill. Overlap at rest (i.e. between singing bouts) is very infrequently changed in C. buckelli. The overlap of 16 individually-caged males was monitored by examining them once a day during almost 2 weeks. Of 141 checks, only 4 reversals from the immediately previous overlap were observed; the incidence of resting overlap reversal was less than 3%. Thirteen of these males never showed an overlap reversal.
Four C. monstrosa males checked over 5 days, gave similar results: two were never found with reversed overlap (checked respectively 5 and 6 times), one was reversed once in 5 checks and one twice in 6 checks. If Cyphoderris alter overlap several times within a single trill, it is strange that individuals end up so consistently at the same overlap with which they began.



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160 Psyche [June-September
We recorded the calling song of a C. monstrosa specimen (Figure 6, 75-6) before and after damaging with a scalpel, several teeth in the central region of his right tegrnen file. In oscillograrns of post- mutilation recorded song, his use of the damaged file (i.e. right above overlap) was apparent as a drastic mid-pulse drop in arnpli- tude. In one oscillograrn, a portion of which makes up Figure 6 (second trace from bottom), 20 pulses in succession were 'right above'.
Switch-wing singing as suggested by distinctive pulse envelopes within the same trill was only evident in our records on one occasion. A male of C. monstrosa had been released in the immediate vicinity (i.e. within antenna1 range) of a mature female on the observer's hand. He began to sing while walking about on the hand and directing his attention toward the female. His song was recorded and on analysis found to be a trill in which every other pulse was identical in envelope and distinctly different from the intervening pulse i.e. there were two pulse types occurring in alternation without break in the sequence of the trill (Figure 6, bottom trace). This was apparently a courtship song. It is clear that pulse envelopes are highly variable in the genus, though usually quite consistent for a particular recording session of a particular individual. Switch-wing stridulation is probably not an everyday feature of C. monstrosa calling song but it may occur under special circumstances such as courtship. Oscillograrns of normal calling songs are given in Figure 6. The pulses of C. strepitans and C. buckelli are apparently indistinguish- able. They are usually wedge-shaped: each begins with a steep rise to maximum amplitude, then falls steadily to the pulse's end. The pulses of C. monstrosa also have a steep onset but are usually of longer duration. They are drawn out in an uneven envelope near their maximum amplitude before dropping away to silence. Carrier frequency spectra of all three species are highly similar. Specimens were analysed 'live' (i.e. without tape-recording) by