Holomorphology And Systematics Of The Eastern Nearctic Stonefly Genus Remenus Ricker (Plecoptera: Perlodidae) Author Verdone, Chris J. Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado, 80523, U. S. A. E-mail: verdonec @ gmail. com verdonec@gmail.com Author Kondratieff, Boris C. Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado, 80523, U. S. A. E-mail: verdonec @ gmail. com verdonec@gmail.com text Illiesia 2018 14 5 81 125 journal article http://doi.org/10.5281/zenodo.4757871 0aee75cf-6abf-4f6d-9fa1-ae8d320bc0eb 1854-0392 4757871 AAEA5971-4C18-45C2-AA83-5DD3D622315D Remenus daniellae Verdone & Kondratieff , sp. n. ( Figs. 42–67 ) http://lsid.speciesfile.org/urn:lsid: Plecoptera .speciesfile.org: TaxonName:502852 Material examined: Holotype ♂ : Tennessee , Sevier Co. , tributary to Le Conte Creek , Twin Creeks Uplands Research Lab , GRSM , N 35.68706 , W 83.50096 , 16 May 2017 , C. Verdone , B.C. Kondratieff ( NMNH ) . Paratypes : North Carolina , Haywood Co. , Ball Branch , Old Cataloochee Turnpike , GRSM , N 35.71817 , W 83.09251 , 25 May 2016 , C. Verdone , B.C. Kondratieff , 2♂ , 3♀ ( CSUIC ); Same location, 14 May 2017 , [emerged 18 May 2017 ], C. Verdone , B.C. Kondratieff , 1♂ , 3♀ , 3 exuvia ( CSUIC ); Same location, 20 May 2017 , C. Verdone , D. Fuller , 3♂ , 2♀ ( CSUIC ); Right Fork Cove Creek , Rte 284, [ N 35.62104 , W 83.05193 ], 23 May 1993 , B.C. Kondratieff , R . F. Kirchner , 2♂ , 1♀ ( CSUIC ). Swain Co. , Collins Creek , Collins Creek Picnic Area , Hwy 441, GRSM , N 35.56752 , W 83.09251 , 14 May 2017 , C. Verdone , B.C. Kondratieff , 1♂ ( CSUIC ); Gunna Creek , at confluence with Spence Cabin Branch , GRSM , N 35.55120 , W 83.73220 , 3 June 2003 , B.D. Heinold , C. Favret , 1♂ ( INHS ); Proctor Branch , Twentymile Creek Trail, GRSM , N 35.48558 , W 83.83684 , 5 June 2003 , B.D. Heinold , 1♂ ( INHS ) . Tennessee , Blount Co. , Anthony Creek , E Cades Cove Campground , Anthony Creek Trail at 3 rd footbridge going upstream, GRSM , N 35.58680 , W 83.75160 , 26 May 2001 , R . E. DeWalt , B.D. Heinold , 1♂ , 2♀ ( INHS ). Sevier Co. , Greenbriar Cove , Smoky Mtns. , [ N 35.70704 , W 83.38294 ], 15 June 1939 , A.C. Cole , 1♂ ( INHS ); Le Conte Creek , Gatlinburg , [ N 35.70164 , W 83.51361 ], 14 June 1940 , T .H. Frison , 1♂ , 3♀ ( INHS ); Le Conte Creek , ATBI Plot , Twins Creek , GRSM , [ N 35.68500 , W 83.49888 ], 8 May–25 May 2010 , C. R . Parker , 2♂ ( CSUIC ); Little Laurel Branch , Ramsey Cascade Trail. GRSM , N 35.70270 , W 83.35654 , C. Verdone , B.C. Kondratieff , 4♂ , 4♀ ( CSUIC ); tributary to Le Conte Creek , Twin Creeks Uplands Research Lab , GRSM , N 35.68706 , W 83.50096 , 16 May 2017 , C. Verdone , B.C. Kondratieff , 11♂ , 9♀ ( CSUIC ) . Fig. 41. Remenus daniellae sp. n. distribution of examined material. Additional material not paratypes . North Carolina, Haywood Co., Ball Branch, Old Cataloochee Turnpike, GRSM , N 35.71817 , W 83.09251 , 14 May 2017 , C. Verdone, B.C. Kondratieff, 3L ( CSUIC ); Mt. Sterling Creek, Old Cataloochee Turnpike, GRSM , N 35.70819 , W 83.09658 , 10 July 1983 , B.C. Kondratieff, 2♀ ( CSUIC ); tributary to Hemphill Creek, 5 km WNW Jonathan, pumphouse spring below Purchase Knob House, GRSM , N 35.58220 , W 83.07370 , 31 May 2003 , R .E. DeWalt, 2L ( INHS ). Tennessee , Sevier Co., Le Conte Creek, Gatlinburg, [ N 35.70164 , W 83.51361 ], 14 June 1940 , T .H. Frison, 6L ( INHS ). Distribution. USA – NC, TN ( Fig. 41 ) Etymology. The patronym honors the senior author’s wife, Danielle M. Fuller, for her valued support and patience. The proposed common name is the “Danielle’s Stripetail”. Fig. 42. Adult male habitus of Remenus daniellae sp. n. , tributary to Le Conte Creek, Sevier County, Tennessee. Male . ( Fig. 42 ). Macropterous; forewing length 9.3– 10.4 mm (n = 10) ( Fig. 43 ). Body length, 8.0– 9.8 mm (n = 10). General body color yellow-gold with light brown markings. Dorsum of head typical of genus ( Fig. 44 ). Pronotum light brown, covered in regularly spaced setae, with pale, glabrous rugosities mediolaterally ( Fig. 44 ); medial pale area widest medially ( Fig. 44 ). Abdominal terga with anterior margin darkened ( Fig. 45 ); rarely darkened laterally. Hemitergal lobes short, not separated from 10 th tergum, with long trichoid sensilla and 0–4 minute sensilla basiconica on each lobe ( Figs. 45– 48 ). Epiproct length ~ 170–270 μm; width ~ 82–89 μm (n = 3); epiproct clavate (club- shaped) and lightly sclerotized ( Figs. 45–46 ); covered in dense, thick palmate hair-like spinulae; when produced forward rarely exceeding the anterior margin of the basal anchor; clavate in dorsal and lateral aspects, widest near the apical ¼ and typically bearing a short translucent tube at the apex ( Figs. 49–52 ). Base of epiproct with sparse palmate hair-like spinulae with between 4–10 seta arising from a common base ( Fig. 53 ). Paragenital plates short, rounded, or triangular ( Figs. 45 , 49–50 ). Basal cowl covered in dense lightly pigmented spinulae ( Figs. 45–46 ). Female. Macropterous; forewing length 11.2–11.7 mm (n = 10). Body length, 9.2–10.2 mm (n = 10). General color and morphology similar to the male. Abdominal terga pale, without darkened anterior pigmentation. Subgenital plate broadly rounded ( Fig. 54–56 ), occasionally with a posteromedial emargination ( Fig. 56 ); lightly sclerotized, with regularly spaced setae, extending ½–⅘ over sternum 9; posterolateral margins convex; basolateral margins convergent posteriorly; basolateral crease typically curved, concave posteriorly, extending ~ ¼ length anteriorly into sternum 8 ( Figs. 54–56 ). Ovum. Shape typical of genus ( Figs. 57–60 ). Length 434–435 μm; width 324–355 μm (n =3). Larva. ( Fig. 61 ). Body length 8.3–10.8 mm , (n = 3). Head ( Fig. 62 ), lacinia ( Fig. 63 ), mandibles ( Fig. 64 ), and pronotum ( Fig. 62 ) typical of genus. Mature male larva with a short ovoid process on abdominal tergum 10 outlining the developing epiproct ( Fig. 65 ). Basal cercal segments with whorls of both short and long setae ( Fig. 66 ). Diagnosis . Males of R. daniellae are morphologically similar R. kirchneri , both of which lack a medial dorsal sclerite. However, R. daniellae can be distinguished by details of the epiproct and terminalia. Males of R. daniellae possess an epiproct that is clavate in dorsal and lateral aspects, which is widest towards the apical ¼ and bears a short translucent tube at the apex. Whereas the epiproct of R. kirchneri is dorsoventrally flattened, widest medially or near the basal ¼ and lacks a translucent tube at the apex. Additionally, the new species lacks sensilla basiconica on the 9 th tergum and has 4 or fewer sensilla basiconica on each hemitergal lobe. Remenus kirchneri occasionally lacks sensilla basiconica on the 9 th tergum, but consistently has 12–20 on each hemitergal lobe. Figs. 43–46. Remenus daniellae sp. n. , adult male, tributary to Le Conte Creek, Sevier County, Tennessee. 43. Right forewing. 44. Head and pronotum. 45. Terminalia, dorsal. 46. Terminalia, lateral. Figs. 47–50. Remenus daniellae sp. n. adult male. 47. Hemitergal lobe and sensilla basiconica, Ball Branch, Haywood County, North Carolina. 47–50. Tributary to Le Conte Creek, Sevier County, Tennessee. 48. Hemitergal lobe. 49–50. Epiproct variation, dorsal. Figs. 51–53. Remenus daniellae sp. n. , adult male, tributary to Le Conte Creek, Sevier County, Tennessee. 51. Epiproct, dorsal. 52. Epiproct, lateral. 53. Epiproct, palmate hair-like setae. Figs. 54–56. Remenus daniellae sp. n. , adult female subgenital plate variation. 54. Ball Branch, Haywood County, North Carolina. 55. Little Laurel Branch, Sevier County, Tennessee. 56. Tributary to Le Conte Creek, Sevier County, Tennessee. Figs. 57–60. Remenus daniellae sp. n. , ovum, tributary to Le Conte Creek, Sevier County, Tennessee. 57. Ovum, dorsal. 58. Collar, dorsal. 59. Collar, ventral. 60. Ovum, ventral. Females of R. daniellae are most similar to R. kirchneri . Separation of these species may require associated males. Generally, the subgenital plate of R. kirchneri is parallel sided basally, whereas in R. daniellae , the subgenital plate is typically convergent. Based on presently available records, the ranges of these two species do not overlap. Remenus daniellae occurs west of the French Broad River, whereas R. kirchneri inhabits the region to the east ( Fig. 115 ). As such, morphology paired with geographic location should help inform identification of these two similar species. Fig. 61. Larval habitus Remenus daniellae sp. n. , Ball Branch, Haywood County, North Carolina. Figs. 62–65. Remenus daniellae sp. n. , larva. Figs. 62–64. Ball Branch, Haywood County, North Carolina. 62. Head and pronotum. 63. Left lacinia, dorsal. 64. Left mandible, dorsal. 65. Male terminal process (developing epiproct), ventral, tributary to Hemphill Creek, Haywood County, North Carolina. Fig. 66. Remenus daniellae sp. n. , larval cercus, dorsal, Ball Branch, Haywood County, North Carolina. Fig. 67. Remenus daniellae sp. n. , male epiproct variation, dorsal, Ball Branch, Haywood County, North Carolina. Fig. 68. Remenus daniellae sp. n. , type locality, tributary to Le Conte Creek, Great Smoky Mountains National Park, Sevier County, Tennessee. Mature larvae of R. daniellae can be separated from R. kirchneri and R. duffieldi by the presence of long setae on the basal cercal segments. However, this character is shared by R. bilobatus which is sympatric in some locations. Fig. 69. Remenus duffieldi , distribution of examined material and published records. Biological notes. Occasionally, the epiproct of R. daniellae is moderately to highly reduced, or appears flaccid and deflated, possibly due to insufficient hemolymphatic pressure ( Fig. 67 ), an apparent aberrancy not observed in any other species of Remenus . Kondratieff and Nelson (1995) reported R. bilobatus from Haywood County, North Carolina based on two females collected in 1985 from Mt. Sterling Creek in GRSM . This location is 1.1 km from the type locality of R. daniellae . These specimens have been re-examined and have been determined as R. daniellae based on the subgenital plate morphology, habitat similarity and proximity to the paratype locality at Ball Branch. No life history or biological studies have been conducted on this species. Remenus daniellae is known only from 12 locations in GRSM in North Carolina and Tennessee . This species has been documented from 1 st –3 rd order streams with drainage areas ranging from 0.47–11.07 km 2 . The average elevation of occurrence localities is 875.1 m (SD ± 285.7 m ). Based on the material examined, emergence occurs from mid-May to mid-July. Existing records are limited, but this species probably inhabits many other streams within GRSM , which possesses more than 3400 km of high quality stream habitats. The type locality ( Fig. 68 ) of this new species is located in what was the heart of the Chimney Tops 2 fire that burned more than 46 km 2 in December 2016 . The effects of this event on the aquatic macroinvertebrate community are unknown; however, stonefly species richness at the type locality six months later was relatively high. Other stoneflies collected with the holotype included Alloperla nanina Banks, 1911 , A. usa Ricker, 1952 , Amphinemura wui (Claassen, 1936) , Isoperla dewalti Verdone & Kondratieff, 2017 , Leuctra grandis Banks, 1906 , L. sibleyi Claassen, 1923 , Sweltsa lateralis (Banks, 1911) , S. mediana (Banks, 1911) , S. urticae ( Ricker, 1952 ) , Tallaperla anna (Needham & Smith, 1916) , and T. laurie ( Ricker, 1952 ) .