Perlesta armitagei n. sp. (Plecoptera: Perlidae): More cryptic diversity in darkly pigmented Perlesta from the eastern Nearctic Grubbs, Scott A. Dewalt, R. Edward Zootaxa 2018 2018-06-29 4442 1 83 100 8N8PQ Grubbs & Dewalt, 2018 Grubbs & Dewalt 2018 [151,370,1825,1851] Insecta Perlidae Perlesta GBIF Animalia Plecoptera 6 89 Arthropoda species armitagei sp. nov.  http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:TaxonName:502858 ( Figs. 6‒11)  Description. Male forewing length 8.0‒10.0 mm (n = 29). Head pale yellow with a dark brown subquadrate ocellar patch and a small, light brown triangular patch anterior to the median ocellus, usually separated by a pale M-line; epicranial suture extends laterally, slightly beyond lateral ocelli ( Figs. 6a, 6c, 6e, 6g). Pronotum brown with a faint yellow medial stripe ( Figs. 6a, 6c, 6e, 6g). Female forewing length 10.0‒ 11.5 mm(n = 25). Color pattern similar to male but typically lighter and with less ocellar and epicranial suture pigmentation ( Figs. 6b, 6d, 6f, 6h). Wing membrane and veins amber except for pale white costal region. Femora brown dorsally extending to articulation with tibiae ( Figs. 6a‒6h), dorsum of tibiae brown.   FIGURE 5.  Perlesta cinctipes. a–b, Arkansas, unknown locality; c–f, Missouri, South Moreau Creek; a, c, male, head + pronotum, dorsal profile; b, d, female, head + pronotum, dorsal profile; e, right paraproct, lateral view, 180X; f, right paraproct, lateral view, 330X.  Male( Figs. 6‒8). Sternum 9 ca. evenly and sparsely clothed by long hairs across most of the segment ( Fig. 7f). Abdominal terga brown, sterna yellow. Cerci pale yellow proximally, brown distally. Tergum 10 mesal sclerites brown, sensilla basiconica restricted to posterolateral margins with interspersed long hairs ( Figs. 7d‒7e). Paraprocts long and slender, bearing a prominent mesoapical tooth in lateral aspect and directed anteriorly, rounded apically ( Figs. 7a, 7c‒7d), not visible in caudal aspect ( Fig. 7b, 7e). Penis tube + sac long, caecum prominent and thumb-like ( Figs. 8a‒8c), sac nearly as long as tube ( Figs. 8a‒8b, 8e), lateral sclerite often lightly sclerotized, located in proximal ¼ of tube ( Fig. 8a), dorsal patch broad basally, narrowing apically throughout length of sac, completely covering caecum ( Figs. 8d‒8f).  Female( Figs. 6, 9). Subgenital plate, ca. 1/2 width of 8th abdominal sternum ( Fig. 9a), lacking pigmentation; lobes subtruncate and rounded both medially and laterally ( Figs. 9b‒9d), with some populations with a slight concavity near medial corners ( Figs. 9b, 9d); lobes bordered posteriorly by a diffuse row of stiff bristle-like hairs; separated by an oval-shaped notch that is expanded medially ( Figs. 9b‒9d).  Egg( Fig. 10). Oval. Collar distinctly stalked, wide, ribbed, and flanged at apex ( Figs. 10a‒10c). Chorion smooth with very fine shallow impressions visible only under high magnification ( Figs. 10a, 10d). Micropyles located ca. ¼ from anterior pole ( Fig. 10d).  Larva. Unknown.    Type material. Holotypemale ( INHS): USA,  Ohio,  Ross Co., Deer Creek,  15 kmNNE Chillicothe, 39.46120, -83.02120,  28 June 2010, S.A. Grubbs.  Paratypes: USA,  Indiana, Bartholomew Co., East Fork White River, Azalia Bridge,  1.5 kmSW Azalia, 39.0849, -85.8598,  11 June 2000, S.A. Grubbs, 3 males, 3 females( WKUC);   Daviess Co., East Fork White River,  14 kmS Washington, 38.5447, -87.2138,  7 June 2009, S.A. Grubbs, 8 males, 1 female( WKUC);   Greene Co., West Fork White River, Rte. 157, 2 km SE Worthington, 39.1109, -86.9632,  7 June 2009, S.A. Grubbs, 3 males, 16 females( WKUC);   Harrison Co., Blue River,  6 kmNE Leavenworth, Stagestop Canoe Access Site, Harrison-Crawford State Forest, 38.2153, -86.2718,  18 May 2000, S.A. Grubbs, 15 males, 7 females( WKUC); same but  9 June 2000, S.A. Grubbs, 2 males, 1 female( WKUC);  Ohio River, Leavenworth, 38.1966, -86.3504,  28 June 2002, S.A. Grubbs, 3 males( WKUC);   Jackson Co., Little Salt River, Houston, 39.0342, -86.1679,  22 June 2008, R.E. DeWalt, 1 male( INHS);   Martin Co., East Fork White River,  2 kmNNE Shoals, 38.7013, -86.7674,  7 June 2009, S.A. Grubbs, 2 males, 4 females( WKUC);   Washington Co., Blue River, Rte. 150, Fredericksburg, 38.4338, -86.1918,  28 June 2010, S.A. Grubbs, 3 males, 3 females( WKUC).   Kentucky,  Adair Co., Russell Creek, nr. Rte. 768, Milltown, 37.1235, -85.4046,  19 June 2008, S.A. Grubbs, 2 males, 5 females( WKUC);   Green Co., Green River, at mouth of Big Pitman Creek,  7.5 kmSE Summersville, 37.2851, -85.5819,  22 May 2002, S.A. Grubbs, 4 males, 1 female( WKUC);  Big Pitman Creek, Narrows of Pitman, 5 kmWNW Greensburg, 37.2833, -85.5576,  21 May 2002, S.A. Grubbs, 4 males, 5 females( WKUC).   Ohio,  Ross Co., same as holotype but  1 July 2007, S.A. Grubbs, 1 female( WKUC); same but  28 June 2010, S.A. Grubbs, 7 males, 9 females( WKUC).   Pennsylvania,  Crawford Co., French Creek, Meadville Access Site, 3 miSE Meadville, 41.5914, -80.1449,  10 June 1998, S.A. Grubbs, 14 males, 9 females( WKUC); same but  1 July 1998, S.A. Grubbs, 1 male( WKUC).   Additional material examined. Indiana,  Harrison Co., Blue River, White Cloud, 38.2290, -86.2254,  25 May 1949, W.E. Ricker, 1 male( CNCI).   Kentucky,  Fayette Co., Lexington,  7‒9 June 1970, 1 male( DCTC).   Ohio,  Butler Co., Seven Mile Creek, Hwy 127 Collinsville, 39.5050, -84.5946,  28 May 1953, A.R. Gaufin, 1 male, 1 female( BYU);  Seven Mile Creek, New Miami, 39.4316, -84.5441,  28 May 1953, A.R. Gaufin, 1 male( BYU);   Coshocton Co., Beaver Run,  2 kmNW Warsaw at Twp. 348 Bridge, 40.3495¸ -82.0134,  26 June 1999, S.W. Chordas IIIand J. Thompson, 1 male( INHS);  Mohawk Creek,  0.5 kmE Mohawk Village at Co. Rd. 82 Bridge, 40.3205, -82.0741,  26 June 1999, S.W. Chordas IIIand J. Thompson, 2 males( INHS);  Killbuck Creek,  2 kmNNW Randle at Co. Rd. 28 Bridge, 40.3338, -81.9457,  20 July 1999, S.W. Chordas IIIand J. Thompson, 1 male( INHS);   Miami Co., Stillwater River, Hwy 718, 40.0521, -84.3659,  3 June 1953, A.R. Gaufin, 1 male( BYU);   Warren Co., Little Miami River, Morrow, 39.3568, -84.1288,  28 June 1952, A.R. Gaufin, 1 male( BYU).   Pennsylvania,  Crawford Co., French Creek, SR 1002 bridge, Venango, 41.7716, -80.1083,  10 June 1991, S.A. Grubbs, 16 females( WKUC).   Etymology.This species is named in honor of Dr. Brian J. Armitage, former Director of the OhioBiological Survey and now “retired” in Boquete,  Panama. Brian tirelessly performed design, layout, and other production activities for several books through the OhioBiological Survey and Caddis Press. He improved our understanding of the modern day composition and distribution of aquatic insects within Ohioand secured funding for statewide surveys of aquatic insects that trained several young entomologists. The common name “Brian’s Stone” is proposed for this species ( Stark et al. 2012).   Diagnosis.Males of  P. armitageipossess a well-developed dorsal caecum ( Figs. 8a‒8c), which initially takes this species to couplet 2 in Stark (2004, pg. 88). The first choice in couplet 2 reads “epicranial suture arms extend well beyond ocelli as a distinct dark line” ( Stark 2004). Couplet 2, however, can be problematic for specimens that are lightly pigmented either due to (a) bleaching following long-term wet storage, or (b) adults preserved while still teneral. Five midwestern and Interior Highland species exhibit this characteristic:  P. adena,  P. baumanni Stark, 1989,  P. cinctipes,  P. fusca Poulton & Stewart, 1991, and  P. xube. Material of  P. adena( Figs. 1a, 1c, 1e),  P. cinctipes( Figs. 5a, 5c), and  P. xube( Figs. 2a, 2c, 2e) examined in this study exhibit this characteristic.  Perlesta armitagei( Figs. 6a, 6c, 6e, 6g) and  P. browni( Figs. 3a, 3c), however, generally lack this feature. Males of  P. armitageiwill key most closely to  P. decipiens( Walsh, 1862) in Stark (2004, see couplet 12)and Grubbs & DeWalt (2012, a modified couplet 12), with a paraproct spine visible in lateral view and directed forward ( Figs. 7a, 7c‒7d). Although the dorsal aedeagal patch of  P. armitagei( Figs. 8d‒8e) is similar to  P. decipiens( Stark 2004, his Fig. 7.306), the mesoapical paraproct tooth of  P. armitagei( Fig. 7a, 7c) is easily contrasted from the antepical paraproct tooth of  P. decipiens( Stark 1989, his Figs. 35‒36, 38‒41).  The paraprocts of  P. armitagei( Fig. 7c) more closely resemble those of  P. cinctipes( Figs. 5e‒5f) and  P. browni( Figs. 4a‒4c) but are not as tubular or elongate as either species.  Perlesta armitageiis more readily distinguished from  P. cinctipesand  P. browniby aedeagal characteristics. The aedeagi of  P. armitageiand  P. cinctipesare similar in that both species have a long tube + sac and well-developed caecum. The dorsal patch of  P. cinctipesis very narrow ( Stark 2004, his Fig. 300) whereas the dorsal patch of  P. armitageiis markedly wider ( Figs. 8d‒8e). Although the epicranial suture arms of  P. browniextend little laterad of the ocelli ( Figs. 3a, 3c), and similar to  P. armitagei( Figs. 6a, 6c, 6e, 6g), the aedeagi of the two species are distinct. The aedeagal tube + sac of  P. browni( Stark 1989, his Fig. 80; Stark 2004, his Fig. 7.297) is much shorter than that of  P. armitagei( Figs. 8a‒8b).   FIGURE 9.  Perlesta armitagei  sp. nov., female, subgenital plate, ventral profile. a, Ohio, Deer Creek, 95X; b, Ohio, Deer Creek, 170X; c, Indiana, Blue River, 190X; d, Kentucky, Russell Creek, 170X. One additional male characteristic appears worthy of further study and may provide diagnostic information. On  P. brownithe posterolateral corners of the male 9th sternum are clothed by a dense matting of long hairs ( Fig. 4d). These are visible under standard light microscopy and appear brush-like and denser than the remaining portions of the segment. In contrast, the male 9th sternum of  P. armitageiis only sparsely clothed with long hairs ( Fig. 7f). The female of  P. armitageican be confidently identified if mature eggs are present and associated with a male with a fully extruded aedeagus. Females will key to couplet 14 in Stark (2004), a dichotomy with  P. shubuta Stark, 1989and  P. decipiens.  Perlesta ephelidaGrubbs & DeWalt, 2012also keys to couplet 14, but the subgenital plate of these three species are easily differentiated using standard light microscopy. The subgenital plate notch of  P. armitageiis ovoid and expanded medially ( Figs. 9b‒9d). In contrast, the subgenital plate notches of  P. ephelida(Grubbs & DeWalt 2012, their Fig. 7),  P. shubuta( Stark 1989, his Fig. 91; Stark 2004, his Fig. 7.391), and  P. decipiens( Stark 1989, his Figs. 44, 46, 48; Stark 2004, his Fig. 7.379) are variable but typically v-shaped. The egg is only partially diagnostic for  P. armitagei. The egg of  P. armitageipossesses a wide, well developed, and distally flanged collar ( Figs. 10a‒10c), resembling  P. decipiens( Stark 1989, his Figs. 12‒13; Stark 2004, his Figs. 7.397‒7.398),  P. ephelida(Grubbs & DeWalt 2012, their Figs. 14‒21), and  P. mihucorum Kondratieff & Myers, 2011(their Figs. 9‒14), but easily contrasted from the coarsely and near-completely punctate egg chorion (except for the eclosion line) of  P. cinctipes( Stark 1989, his Fig. 17; Stark 2004, his Figs. 7.394‒7.396).   FIGURE 10.  Perlesta armitagei  sp. nov., egg. a, Kentucky, Russell Creek, entire egg, 270X; b, Kentucky, Russell Creek, anterior pole and collar, 1600X; c, Ohio, Deer Creek, details of collar, 2300X; d, Kentucky, Russell Creek, posterior pole and micropyles, 500X.  Remarks.The typelocality is in southern Ohioat the southern edge of EPA Level IV Ecoregion 55b (Loamy, High Lime Till Plains). All streams where  P. armitageihas been collected are mid-order systems with partially- to fully-open canopies.  Allocapnia granulata( Claassen, 1924),  A. vivipara( Claassen, 1924), and  Taeniopteryx burksi Ricker & Ross, 1968were the only other stonefly species collected at the typelocality.  The plotted distribution of  P. armitagei( Fig. 11b) is conditional since it only includes the specimens examined in this study. This distribution, however, suggests an OhioRiver basin species. We have not found this species during our extensive fieldwork and examination of museum material from Illinois(DeWalt & Grubbs 2011), Michigan(Grubbs et al. 2012), and Ontario(unpublished data). The plotted records from Indianamay represent the western edge the range.  The species reported as  Perlestasp. I-4from Ohioin DeWalt et al.(2012, 2016) is now regarded as  P. armitagei. The specimen determined as  P. cinctipesfrom Kentucky(Fayette Co., Tarter et al.2006), and subsequently carried forward in the updated state checklist ( Tarter et al.2015), was reexamined and determined as  P. armitagei. The specimens determined as  P. cinctipesfrom West Virginia(Monongalia Co., Tarter & Nelson 2006) were not available for study but the location was still plotted ( Fig. 11b) and the inclusion in this map should be considered conditional. 1852083210 2010-06-28 INHS S. A. Grubbs United States of America Ross Co. 39.4612 Deer Creek 1 -83.0212 8 91 1 Ohio holotype 1852083309 2000-06-11 WKUC S. A. Grubbs United States of America Bartholomew Co. 39.0849 Azalia Bridge 6 -85.8598 East Fork White River 8 91 6 3 3 Indiana paratype 1852083211 2009-06-07 WKUC S. A. Grubbs United States of America Daviess Co. 38.5447 East Fork White River 6 -87.2138 8 91 9 1 8 Washington paratype 1852083209 2009-06-07 WKUC S. A. Grubbs United States of America Greene Co. 39.1109 West Fork White River 6 -86.9632 West Fork White River 8 91 19 16 3 Washington paratype 3071119307 2000-05-18 2000-06-09 2000-05-18 WKUC S. A. Grubbs United States of America Harrison Co. 38.2153 Blue River 6 -86.2718 Blue River 8 91 22 7 15 Washington paratype 3071119312 [650,1369,910,935] 2000-06-09 2000-06-09 2000-05-18 WKUC S. A. Grubbs United States of America Harrison Co. Stagestop Canoe Access Site Blue River 8 91 3 1 2 Washington paratype 1852083306 2002-06-28 WKUC S. A. Grubbs United States of America 38.1966 Ohio River 6 -86.3504 8 91 3 3 Ohio paratype 1852083303 2008-06-22 INHS R. E. DeWalt United States of America Jackson Co. 39.0342 Little Salt River 6 -86.1679 Little Salt River 8 91 1 1 Ohio paratype 1852083310 2009-06-07 WKUC S. A. Grubbs United States of America Martin Co. 38.7013 East Fork White River 6 -86.7674 East Fork White River 8 91 6 4 2 Ohio paratype 1852083301 2010-06-28 WKUC S. A. Grubbs United States of America Washington Co. 38.4338 Blue River 6 -86.1918 Rte. 8 91 6 3 3 Washington paratype 1852083207 2008-06-19 WKUC S. A. Grubbs United States of America Adair Co. 37.1235 Russell Creek 6 -85.4046 Russell Creek 8 91 7 5 2 Kentucky paratype 1852083206 2002-05-22 WKUC S. A. Grubbs United States of America Green Co. 37.2851 Green River, at mouth of Big Pitman Creek 6 -85.5819 Green River 8 91 5 1 4 Kentucky paratype 1852083307 2002-05-21 WKUC S. A. Grubbs United States of America Big Pitman Creek 37.2833 Big Pitman Creek 6 -85.5576 Narrows of Pitman 8 91 9 5 4 Kentucky paratype 3071119313 [270,1231,1234,1259] 2007-07-01 2010-06-28 2007-07-01 WKUC S. A. Grubbs United States of America Ross Co. Deer Creek 15 km NNE Chillicothe 8 91 1 1 7 Ohio holotype 3071119302 2010-06-28 2010-06-28 2007-07-01 WKUC S. A. Grubbs United States of America Ross Co. Deer Creek 15 km NNE Chillicothe 8 91 16 9 7 Ohio holotype 3071119310 1998-06-10 1998-07-01 1998-06-10 WKUC S. A. Grubbs United States of America Crawford Co. 41.5914 French Creek 6 -80.1449 French Creek 8 91 23 9 14 Pennsylvania holotype 3071119308 1998-07-01 1998-07-01 1998-06-10 WKUC S. A. Grubbs United States of America Crawford Co. Meadville Access Site French Creek 8 91 1 9 1 Pennsylvania holotype 3071119304 1949-05-25 1970-06-09 1970-06-07 CNCI W. E. Ricker Harrison Co. 38.229 Blue River 6 -86.2254 Co. Rd. 8 91 1 16 1 Indiana 3071119309 [635,1437,1414,1439] 1970-06-07 1970-06-09 1970-06-07 DCTC White Cloud & W. E. Ricker Fayette Co. Lexington Co. Rd. 8 91 1 16 1 Kentucky 3071119305 1953-05-28 1970-06-09 1970-06-07 BYU A. R. Gaufin Butler Co. 39.505 Seven Mile Creek 6 -84.5946 Co. Rd. 8 91 2 1 1 Ohio 3071119314 1953-05-28 1970-06-09 1970-06-07 BYU A. R. Gaufin Harrison Co. 39.4316 Seven Mile Creek 6 -84.5441 Co. Rd. 8 91 1 16 1 Indiana 3071119301 1999-06-26 1970-06-09 1970-06-07 INHS S. W. Chordas III & J. Thompson Coshocton Co. Beaver Run Co. Rd. 8 91 1 16 1 Indiana 3071119311 1999-06-26 1970-06-09 1970-06-07 INHS S. W. Chordas III & J. Thompson Harrison Co. 40.3205 Mohawk Creek 6 -82.0741 Co. Rd. 8 91 2 16 2 Indiana 3071119303 1999-07-20 1970-06-09 1970-06-07 INHS S. W. Chordas III & J. Thompson Harrison Co. 40.3338 Killbuck Creek 6 -81.9457 Co. Rd. 8 91 1 16 1 Indiana 3071119306 1953-06-03 1970-06-09 1970-06-07 BYU A. R. Gaufin Miami Co. 40.0521 Stillwater River 6 -84.3659 Co. Rd. 8 91 1 16 1 Indiana 3071119315 [244,1437,1702,1727] 1952-06-28 1970-06-09 1970-06-07 BYU A. R. Gaufin Warren Co. 39.3568 Little Miami River, Morrow 6 -84.1288 Co. Rd. 8 91 1 16 1 Indiana 1852083208 1991-06-10 1970-06-09 1970-06-07 WKUC S. A. Grubbs Crawford Co. 41.7716 French Creek 6 -80.1083 Co. Rd. 8 91 16 16 1 Pennsylvania