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Descriptions of the nymphs of Eastern North American species of cultus (plecoptera: perlodidae).

Abstract.--The nymphs of eastern North American stonefly genus Cultus are described. Previous misidentifications and putative nymphal descriptions are reviewed. A key separating the late instar nymphs is also provided.

Key words: Plecoptera, Perlodidae, Cultus, nymphs, eastern North America.

INTRODUCTION

The Nearctic genus Cultus is represented in eastern North America by two species; C. decisus (Walker, 1852) and C. verticalis (Banks, 1920) (Stark et al., 1988; Kondratieff, 2004). Additionally Stark et al. (1988) recognized two disjunct geographic forms of C. decisus, C. d. decisus typically occurs in larger streams and rivers of Pennsylvania, New York, Michigan, Quebec, Ontario, and New Brunswick, whereas C. d. isolatus (Banks, 1920) is known from larger streams and rivers of the southern Appalachians from Georgia, North Carolina, and Virginia (Stark et al., 1988; Kondratieff, 2004). Cultus verticalis has been recorded from small to medium sized streams of the Appalachians from Quebec south to North Carolina.

Claassen (1934) provided the first putative nymphal description of C. verticalis from Neversink, Sullivan County, New York. Yet, after examination of this specimen in the Cornell University Insect Collection (CUIC), it appears that this description pertains to C. d. decisus. The biology of the eastern Cultus remains poorly known (Stewart and Stark, 2002). Minshall and Minshall (1966) examined the supposed life history of C. decisus in Morgan's Creek, Meade County, Kentucky. It is probable that their information pertains to Diploperla robusta Stark and Gaufin, 1974, as no recent published records of Cultus exist for this state (Tarter et al., 1984; Tarter et al., 2006). However, there is a female specimen of C. verticalis in the C. P. Gillette Museum of Arthropod Diversity from Wayne County, in eastern Kentucky.

Nymphs of Cultus can be separated from other North American Perlodinae genera by the combination of the following characters; lacinia bidentate with few marginal setae, pronotal setae restricted to 2-3 at corners, and mesosternal Yarms well developed, reaching posterior corners of furcal pits, transverse suture connecting furcal pits absent (Stewart and Stark, 2002). The purpose of this paper is to provide the first comparative descriptions of the nymphs of C. d. decisus, C. d isolatus, and C. verticalis.

Cultus decisus (Walker) (Figs. 1A, B, 2A, B)

Cultus decisus decisus, Stark et al., 1988: 91.

Cultus decisus isolatus, Stark et al., 1988: 92.

General body color yellow with brown markings. Length of terminal instar 12-14 mm. Antennae yellow, antennomeres with apical circlets of stout setae. Head wider than pronotum, yellow with conspicuous dark transverse band anterior to the epicranial suture, anterior portion of transverse band with indistinct M-shaped pattern, two light spots anterior and posterior to median ocelli; labrum yellow and with a complete fringe of short marginal setae; epicranial suture light brown with light yellow to white colored maculations; occipital row of setae lacking, a few short post-ocular setae present. Lacinia triangular, bidentate, with 2 marginal setae near base of subapical tooth, 10-12 basal submarginal setae, and 2-3 axillary setae; terminal tooth of lacinia 0.45 times the total outer length of lacinia, subapical tooth 0.43 times the length of the terminal tooth (N = 5) (Fig. 1). Left mandible with row of acanthae along base of molars (Fig. 2). Width of pronotum more than two times the length, transversely ovoid, and narrower than the head; border of pronotum with two or three setae at corners, a distinct black grove around the edges and with a light median diamond-shaped maculation (Figs. 2A and B). Wing pads diverge 35 degrees from body midline, and with light bands of pigmentation. Mesothorax and metathorax with light diffuse pigmentation in the center and darker shading on the perimeter. Legs mostly yellow dorsally with light brown maculations on the anterior 1/3 of the femur, and on the posterior 1/3 of the tibia; femur, tibia and tarsus with a conspicuous fringe of long silky setae on the dorsal surface; femora and tibia with stout setae variously arranged on the dorsal surface. Abdominal terga brown with light, median, transverse bands on each segment, a posterior abdominal fringe of stout setae present on all abdominal segments, interrupted medially on sternum 8 in both sexes, abdominal sterna yellow, occasionally with dark mottling. Cerci brown, shorter than abdomen, segments with apical circlets of short setae, and a dorsal fringe of long silky setae.

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

Cultus verticalis (Banks) (Fig. 1C).

Cultus verticalis, Stark et al., 1988: 95.

General color yellow with brown markings. Length of terminal instar 12-15 mm. Antennae yellow to brown, antennomeres with apical circlets of stout setae. Head slightly wider than pronotum, light brown and with dark transverse band anterior to the epicranial suture; anterior portion of transverse band with indistinct M-shaped pattern, two light spots anterior and posterior to median ocelli; labrum yellow and with a complete fringe of short marginal setae; epicranial suture predominantly light brown with light yellow to white colored maculations on a light brown background; occipital row of setae lacking, a few short post-ocular setae present. Lacinia triangular, bidentate with 2 marginal setae near base of subapical tooth, 10-14 basal submarginal setae, and 2-3 axillary setae; terminal tooth of lacinia 0.55 times the total outer length of lacinia, subapical tooth 0.6 times the length of the terminal tooth (N = 4). Left mandible with row of acanthae along base of molars. Pronotum transversely ovoid, and slightly narrower than the head, with a dark line around the perimeter, 2 to 3 setae at the corner, and a light colored diffuse maculation that is square to triangular shape and located near the posterior margin (Fig. 1C). Wing pads diverge 35 degrees from body midline, and with dark bands of pigmentation. Mesothorax and metathorax with light diffuse pigmentation in the center and darker shading on the perimeter. Legs mostly yellow dorsally with light brown maculations on the anterior 1/3 of the femur, and on the posterior 1/3 of the tibia; femur, tibia and tarsus with a conspicuous fringe of long silky setae on the dorsal surface; femora and tibia with stout setae variously arranged on the dorsal surface. Abdominal terga brown with light transverse bands on the posterior half of each segment. With a posterior abdominal fringe of stout setae present on all abdominal segments, interrupted medially on sternum 8 in both males and females; cerci light brown, slightly shorter than the abdomen, segments with apical circlets of short stout setae, and a dorsal fringe of long setae.

DISCUSSION

Stark et al. (1988) was unable to find reliable characters to distinguish the adults of C. d. decisus and C. d. isolatus. Concurrently, no reliable characters were discovered between the nymphs of the northern and southern populations of this species. Because of the geographical disjunction, however, Stark et al. (1988) suggested that subspecific status be maintained. Adults of C. decisus can usually be distinguished from C. verticalis (Stark et al., 1988). However, no morphological characters were noted at this time to separate the nymphs of these two species (except possibly the slight head, pronotal, and mesonotal patterns observed in the specimens examined, Figs. 1C, 2A, B). Close examination of the pronotal maculations combined with habitat and geographical proximity may provide a reliable means of separating nymphs of these three taxa, and have been used in constructing the following tentative key. The nymphs of C. decisus appear to be restricted to larger rivers, whereas C. verticalis usually occurs in the small to medium sized streams of the Appalachians throughout the range of C. d decisus and C. d. isolatus.

KEY TO LATE INSTAR EASTERN NORTH
AMERICAN CULTUS NYMPHS

1. Median pronotal maculation diamond-shaped,
   located in the middle of the pronotum (Fig. 2);
   usually found in larger streams and rivers        decisus complex 2

-- Median pronotal maculation located near posterior
   margin, not diamond shaped, with diffuse
   pattern (Fig. 1C); restricted to smaller streams
   and rivers of the Appalachians                           verticalis

2. Distribution restricted to Georgia, North
   Carolina, and Virginia                             decisus isolatus

-- Distribution restricted to Pennsylvania, New
   York, Michigan, Quebec, Ontario, and New
   Brunswick                                           decisus decisus


MATERIAL EXAMINED

The following material was examined, including adults. These records augment Stark et al. (1988). The codens are as follows: CSUC, C. P. Gillette Museum of Arthropod Diversity, Colorado State University, CUIC, Cornell University Insect Collection, Ithaca, NY, JE, Personal Collection of Jane Earle.

Cultus decisus decisus (Walker)

New Brunswick: Carleton Co., St. Johns River Jct. Becaquimie Stream HWY 105 Hartland, June 16, 1993, B. C. Kondratieff, R. W. Baumann, 8 males, 8 females, 3 exuvia, (CSUC). Gloucester Co., Nepisiquit River, HWY 360, June 17, 1993, B. C. Kondratieff, R. W. Baumann, 12 males, 21 females, 1 exuvia, (CSUC). Victoria Co., St. Johns River Grand Falls at Light Falls June 15, 1993, B. C. Kondratieff, R. W. Baumann, 1 male, 1 female (CSUC). New York: Clinton Co., Saranac River, Bowen Rd., 44[degrees]39.431'N 73[degrees]43.968'W, May 27, 2006, L. Myers, 2 males, 3 females, 1 exuvia, (CSUC); Saranac River, Bowen Rd., 44[degrees]39.431'N 73[degrees]43.968'W, June 5, 2006, L. Myers, 1 male, (CSUC); Saranac River, River Rd. nr. Morrisonville, 44[degrees]41.559'N 73[degrees]34.286'W, May 28, 2006, L. Myers, 2 males, 1 female, 4 exuvia, (CSUC); Saranac River, River Rd. Nr. Morrisonville, 44[degrees]41.559'N 73[degrees]34.286'W, May 30, 2006, L. Myers, 1 female, (CSUC); Saranac River, River Rd. Nr. Morrisonville 44[degrees]41.559'N 73[degrees]34.286'W, June 2, 2006, L. Myers, 2 females, (CSUC); Saranac River, Soper Rd. NYSEG Substation, 44[degrees]38.387'N 73[degrees]44.802'W, June 2, 2006, L. Myers, 1 male, (CSUC); Saranac River, Strackville Rd., 44[degrees]36.315'N 73[degrees]48.152'W, May 26, 2006, L. Myers, 2 males, 4 females, (CSUC); S. B. Saranac River, Silver Lake Rd., 44[degrees]35.478'N 73[degrees]50.365'W, May 19, 2006, L. Myers, 3 males, 1 female, 1 exuvia, (CSUC); S. B. Saranac River, Silver Lake Rd., 44[degrees]35.478'N 73[degrees]50.365'W, May 23, 2006, L. Myers, 5 males, 4 females, (CSUC); S. B. Saranac River, Silver Lake Rd., 44[degrees]35.478'N 73[degrees]50.365'W, May 17, 2007, L. Myers, 1 nymph, (CSUC); S. B. Saranac River, Union Falls NYSEG Substation, 44[degrees]30.569'N 73[degrees]54.849'W, May 27, 2006, L. Myers, 1 male, (CSUC). Columbia Co., Stockport Creek, Rt 9 Nr. Stockport, 42[degrees]18.852'N 073[degrees]45.297'W, June 24, 2007, L. Myers, B. C. Kondratieff, 1 female, (CSUC). Delaware Co., W. B. Delaware River, at Hancock, April 19, 2006, D. S. Bogar, 5 nymphs (JE). Essex Co., W. B. Ausable River Notch on rt 86 near Wilmington, 44[degrees]21.730'N 73[degrees]50.640'W, May 24, 2006, L. Myers, 3 nymphs, (CSUC); W. B. Ausable River Notch on Rt 86 near Wilmington, 44[degrees]21.730'N 73[degrees]50.640'W, March 13, 2006, L. Myers, 1 nymph, (CSUC); W. B. Ausable River Notch on Rt. 86 near Wilmington, 44[degrees]21.730'N 73[degrees]50.640'W, May 20, 2006, L. Myers, 2 males, 1 female, (CSUC); W. B. Ausable River, Rt 86 in Wilmington, 44[degrees]21.730'N 73[degrees]50.640'W, June 26, 2007, L. Myers, B. C. Kondratieff, females, (CSUC). Sullivan Co., Neversink, May 11, 1929, P. R. Needham, 1 nymph, (CUIC). Pennsylvania: Forest Co., Crane Run, May 10, 2005, M. Passmore, 1 nymph, (JE). Perry Co., Sherman Creek, 3 mi WSW Duncannon, March 31, 2006, J. Reppert, 1 nymph, (JE). Pike Co., Delaware River, Dingmans Ferry, May 29, 1996, B. C. Kondratieff, 12 males, 14 females, 5 exuvia, (CSUC). Tioga Co., Babb Creek, Up Lick Creek, May 20, 1992, Jane Earle, 1 nymph, (JE).

Cultus decisus isolatus (Banks)

Georgia: Fannin Co., Tocca River, March 24, 1976, B. C. Kondratieff, 3 nymphs, (CSUC); Ocoee River, March 25, 1976, B. C. Kondratieff, 12 nymphs, (CSUC). North Carolina: Ashe Co., S. F. New River, Hwy 221 W of Scotsville, 36.47473N, 81.33760W, May 2, 2005, B. C. Kondratieff, R. F. Kirchner, R. E. Zuellig, D. R. Lenat, 2 males, 1 nymph (CSUC). Burke Co., Yadkin River, Grandin Rd., April 18, 2007, B. C. Kondratieff, R. Kirchner, D. Lenat, 1 female, 2 nymphs, 1 exuvia, (CSUC). Caldwell Co., Johns River, Old Johns River Rd., CR1356 1.1 mi. NW Colletsville, 35[degrees]93.374'N 81[degrees]69.087'W, April 18, 2007, B. C. Kondratieff, R. Kirchner, D. Lenat, 1 female, 4 nymphs, 1 exuvia (CSUC); John's River, Old John's River Rd. 3.2 mi upstream of Highway 90 NE of Colletsville 35[degrees]93.379'N 81[degrees]69.036'W, May 3, 2005 B. C. Kondratieff, R. F. Kirchner R. E. Zuellig, D. R. Lenat, 2 females, 1 exuvia, (CSUC). Virginia: Carroll Co., New River, Rt 606, April 19, 1980, Reared April 26, 1980, B. C. Kondratieff, 2 males, 1 nymph, 1 exuvia, (CSUC). Grayson Co., New River Route 94 bridge 3 mi W of Galax, May 18, 1994, B. C. Kondratieff, R.F. Kirchner 1 male, 5 females (CSUC). Pulaski Co., New River, June 8, 1973, J. Wright, 1 nymph, (CSUC).

Cultus verticalis (Banks)

Kentucky: Wayne Co., Beech Fork, Beech Fork Lake State Park, May 20, 1990, B. C. Kondratieff, 1 female (CSUC). Maine: Somerset Co., Moose River, Demo Rd. below Long Pond, June 12, 1993, B. C. Kondratieff and R. W. Baumann, 2 males, 4 females (CSUC). New York: Clinton Co., Cold Brook, Standish Rd. High Banks, 44[degrees]37.148'N 73[degrees]54.191'W, May 26, 2006, L. Myers, 1 nymph, (CSUC); True Brook, True Brook Rd., 44[degrees]38.220'N 73[degrees]46.094'W, March 17, 2006, Reared May 1, 2006, L. Myers, 1 female, (CSUC). Franklin Co., Alder Brook off Route 3, 44[degrees]34.327'N 073[degrees]56.574'W, May 19, 2006, L. Myers, 1 male, (CSUC); Alder Brook off Route 3, 44[degrees]34.327'N 073[degrees]56.574'W, May 26, 2006, L. Myers, 1 nymph, (CSUC); N. B. Saranac River, Loon Lake Rd., 44[degrees]32.356'N 74[degrees]03.089'W, May 18, 2006, L. Myers, 1 female, (CSUC); N. B. Saranac River, Loon Lake Rd., 44[degrees]32.356'N 74[degrees]03.089'W, May 18, 2006, L. Myers, 1 nymph, (CSUC). Oneida Co., BLT, Mile Creek, Moose River Rd. nr. Boonville, 43[degrees]35.707'N 75[degrees]19.677'W, June 5, 2008, L. Myers and R. Mowrey, 1 female (CSUC). Warren Co., Glen Creek, Glen Creek Rd., 43[degrees]35.108'N 73[degrees]52.279'W, June 26, 2007, L. Myers, B.C. Kondratieff, 1 female, (CSUC). Mill Creek, Rt. 8 nr. Weavertown, May 21, 2007, L. Myers, P. Novak, 1 nymph, (CSUC). North Carolina: Avery Co., Linville River, Blue Ridge Parkway, May 18, 1994, B. C. Kondratieff, 3 males, 4 females, 4 exuvia, (CSUC). Burke Co., Linville River, NC-126 bridge 6 mi NE Nebo, 35[degrees]79.510'N 81[degrees]89.044'W, April 21, 2007, B. C. Kondratieff, R. Kirchner, D. Lenat, 1 larva (CSUC). Caldwell Co., Wilson Creek, Ralph Winchester Rd., S. Mortimer, April 18, 2007, B. C. Kondratieff, R. Kirchner, D. Lenat, 3 males, 5 females, 4 nymphs, 10 exuvia, (CSUC). Surry Co., Fisher River, Ladonia Church Rd., April 18, 2007, B. C. Kondratieff, R. Kirchner, D. Lenat, 2 males, 2 females, 1 nymph, 4 exuvia, (CSUC). Wilkes Co., Middle Fork Reddings River, Old NC 16 East of Wilbar, 36[degrees]26.189'N 81[degrees]30.073'W, May 2, 2005, B. C. Kondratieff, R. Zuellig, R. Kirchner, D. Lenat, 6 males, 8 females, 1 nymph, 3 exuvia, (CSUC); Basin Creek, CR1730 2 mi. NW Ashers, July 18, 2007, B. C. Kondratieff, R. Kirchner, D. Lenat, 1 nymph, (CSUC); Elk Creek on Elk Creek Rd. NW of Elkville, 36[degrees]06'23"N 081[degrees]26'16'W May 17, 2004, B. C. Kondratieff, R. Zuellig, R. Kirchner, D. Lenat,1 male, 1 female, (CSUC). Pennsylvania: Lycoming Co., Loyalsock Creek at Barbours Slate Rd. 41[degrees]23'36"N 76[degrees]47'57"W, May 9, 1995, E.C. Masteller, 1 male, 1 female (CSUC). Virginia: Giles Co., Little Stony Creek, Rd. 460 Pembrooke, May 7, 1980, B. C. Kondratieff, 3 males, 1 female, 4 exuvia, (CSUC). Patrick Co., Round Meadow Creek, Highway 614, 2 mi SW of dam June 1, 1989, R. I. Hoffman 1 male 1 female (CSUC). Smyth Co., Route 600, St. Clair Bottom S. Fork Holston River, May 18, 1990, B. C. Kondratieff, 1 male, 3 females (CSUC). West Virginia: Randolph Co., Gandy Creek, Rt. 40 Bridge below Sinke, May 15, 1990, Kondratieff, Welch, 2 males, 20 females, 1 nymph, 27 exuvia, (CSUC).

ACKNOWLEDGMENTS

We thank Dave Carlson, Windsor, Colorado for the illustrations. Dr. K. W. Stewart University of North Texas, provided a useful review of this manuscript. Jane Earle, Pennsylvania, provided additional nymphal material for study. Dr. Tim Mihuc, Lake Champlain Research Institute, Plattsburgh State University of New York, provided logistical support for field sampling in New York State. Additionally Dr. Janet Mihuc, Paul Smith's College, Bob Bode, New York State Stream Biomonitoring Unit, and Paul Novak, New York State DEC, provided assistance in the field. This research was made possible by a grant funded by the New York State Biodiversity Research Institute, Albany, New York.

Received 17 December 2008; accepted 6 April 2009

LITERATURE CITED

Banks, N. 1920. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology 64: 299-362.

Claassen, P. 1934. Plecoptera nymphs of North America. Thomas Say Foundation 3: 1-199.

Kondratieff, B. C. 2004. Perlodidae--Perlodinae (The Springflies), pp. 149-180. In: B. P. Stark and B. J. Armitage (eds.). 2004. Stoneflies (Plecoptera) of Eastern North America Volume II Chloroperlidae, Perlidae, Perlodidae (Perlodinae). Ohio Biological Survey Columbus, Ohio. 192 pp.

Minshall, G. W. and J. N. Minshall. 1966. Notes on the life history and ecology of Isoperla clio (Newman) and Isogenus decisus Walker (Plecoptera: Perlodidae). American Midland Naturalist 76: 340-350.

Stark, B. P., S. W. Szczytko and B. C. Kondratieff. 1988. The Cultus decisus complex of Eastern North America (Plecoptera: Perlodidae). Proceedings of the Entomological Society of Washington 90: 9146.

Stewart, K. W. and B. P. Stark. 2002. Nymphs of North American Stonefly Genera (Plecoptera). The Caddis Press, Columbus, Ohio. 510 pp.

Tarter, D. C., D. A. Atkins and C. V. Covell, Jr. 1984. A checklist of the stoneflies (Plecoptera) of Kentucky. Entomological News 95: 113-116.

Tarter, D. C., D. L. Chaffee and S. A. Grubbs. 2006. Revised checklist of the stoneflies (Plecoptera) of Kentucky. Entomological News 117: 1-10.

Walker, F. 1852. Catalogue of the specimens of neuropterous insects of the collection of the British Museum. Part I. Phryganides-Perlides. British Museum, London. 192 pp.

LUKE W. MYERS (1,2) AND BORIS C. KONDRATIEFF (1)

(1) Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado 80523

(2) E-mail address for correspondence: myersl@lamar.colostate.edu
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Author:Myers, Luke W.; Kondratieff, Boris C.
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Date:Apr 1, 2009
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