Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf NEW SPECIES AND RECORDS OF ANACRONEURIA (PLECOPTERA: PERLIDAE) FROM ECUADOR AND PARAGUAY Bill P. Stark1, Boris C. Kondratieff2 and Brian Gill3 1 Department of Biology, Box 4045, Mississippi College, Clinton, Mississippi 39058, U.S.A. E-mail: stark@mc.edu 2 Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado 80523, U.S.A. E-mail: Boris.Kondratieff@colostate.edu 3 Department of Biology, Colorado State University, Fort Collins, Colorado 80523, U.S.A. E-mail: gillbriana@gmail.com ABSTRACT Five new species of Anacroneuria Klapálek are described from Ecuadorian specimens. New taxa include A. apuela, A. cushueme, A. kayceae, A. malkini, and A. pichincha. New distribution records are also presented for six species including three, A. atrifrons Klapálek, A. chavin Stark & Sivec and A. guambiana Zúñiga & Stark, not previously reported from Ecuador. An additional species represented by a female specimen is described under an informal designation, the larva of A. gaumbiana is described, and a checklist of 54 species reported for Ecuador is presented. The first Paraguayan specimens of A. atrifrons are also reported. Keywords: Anacroneuria, Plecoptera, Perlidae, New species, Ecuador, Paraguay INTRODUCTION Genus Anacroneuria is the dominant stonefly group throughout much of the Neotropics, and this condition certainly applies to Ecuador. Forty six Anacroneuria species are currently recognized as valid for Ecuador, and another five, originally described from the country are regarded as nomen dubium (Stark 2001; Stark & Zuniga 2003; Zuniga et al. 2006); the only other stonefly genus reported for Ecuador is Claudioperla (Turcotte & Harper 1982 a, b), presently considered a monotypic member of family Gripopterygidae (Illies 1963). The present study is based on a small sample of Anacroneuria collected at high elevation sites (>1980 m) by two of the authors (BCK, BG), or on material included in the S.G. Jewett collection, but forwarded to us for study together with specimens from the Monte L. Bean Museum at Brigham Young University, and an additional small series from Paraguay in the C.P. Gillette Museum. Study of these specimens resulted in discovery of five previously undescribed species along with specimens of three species not previously reported from Ecuador, and one species record new for Paraguay. These discoveries bring the Anacroneuria species list for Ecuador to 54 species and the Paraguayan Anacroneuria list to four species (Stark & Baumann 2011). MATERIAL AND METHODS Terminal abdominal segments were clipped and boiled in 10% KOH briefly before study with a Wild M5 dissecting microscope, using the procedures outlined in Stark & Zuniga (2003). Holotype specimens are deposited in the United States Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 78 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf National Museum of Natural History, Washington, D.C. (USNM) or the California Academy of Sciences, San Francisco, California (CASC) as indicated in the text; other specimens are deposited in the C.P. Gillette Museum of Arthropod Diversity, Colorado State University, Fort Collins, Colorado (CSUC) or the Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah (BYUC). Other museum designations used in the text are: Natural History Museum London (NHML) and Museo de Entomología de la Universidad del Valle, Cali, Colombia (MEUV). RESULTS AND DISCUSSION Anacroneuria anchicaya Baena & Zúñiga Anacroneuria anchicaya Baena & Zúñiga in Stark et al., 1999:22. Holotype S (MEUV), Alto Anchicayá, Valle Del Cauca, Colombia Anacroneuria anchicaya: Stark, 2001:4. Ecuador records Material examined. Ecuador: Pastaza, Cushueme, Rio Cushueme, 320 m, 150 km SE Puyo, 15-30 May 1971, B. Malkin, 1S (CASC). Comments. This species was previously reported from Colombia and from Cotopaxi, Napo and Pichincha provinces in Ecuador (Stark et al., 1999; Stark 2001). Anacroneuria apuela Stark & Gill sp. n. (Figs. 1-5) Material examined. Holotype S and 1S paratype, Ecuador, Province Imbabura, Canton Cotacachi, Rio Intag, Apuela, 5020 ft, 00° 21.2'N, 78° 31.0'W, 6 November 2009, S.M. Clark, H.R. Hinkson (Holotype USNM, paratype BYUC). Adult habitus. General color brown patterned with darker brown. Ocellar region covered with moderately dark brown pigment extending to central frons, becoming darker forward of poorly defined Mline; lappets and antennae brown (Fig. 1). Pronotum pale along median suture and along lateral margins; disk predominantly brown with scattered pale rugosities. Wing membrane pale brown, veins darker except costa pale. Femora dark over apical half and along dorsal and ventral margins for most of length; tibiae dark along outer margin and in basal and apical fourth. Male. Forewing length 10 mm. Apical section of aedeagus gradually narrowed from shoulders, then slightly widened subapically and narrowed again to slightly notched tip (Fig. 3). Dorsal keel consists of two narrowly separated and broken lines which diverge sharply and reach lateral margins near subapical wide point (Fig. 5). Ventral aedeagal apex covered by a large pair of membranous lobes. Hooks large, subchelate and narrowed to acute tips. Lateral aspect of apical section shaped somewhat like a bird's head (Fig. 4). Hammer thimble shaped (Fig. 2). Female. Unknown. Larva. Unknown. Etymology. The species name, used as a noun in apposition, is based on the type locality. Diagnosis. This species is a member of the A. aymara Stark & Sivec, 1998 species group discussed by Zuniga & Stark (2002). The group also includes A. chaima Stark, 1999, A. cotacachi Stark, 2001, A. farallonensis Rojas & Baena, 1993 (see Zuniga & Stark 2002), A.perija Stark, 1999, A. portilla Stark & Rojas, 1999 (in Stark et al. 1999), A. ricki Zuniga & Stark, 2002, A. segnini Stark & Maldonado, 2002 (in Maldonado et al. 2002), and A. toni Zuniga & Stark, 2002, and appears to be widely distributed in the northern and central Andes. The new species is most similar to A. toni in structure of the dorsal aedeagal keel, sharing with that species the long, apically divergent pair of keel lines. However, in A. apuela these lines are broken, and appear as a series of small bumps separated by narrow gaps. In addition, the ventral membranous lobes are much larger than in A. toni (see Zuniga & Stark 2002, Figs. 17-19). Anacroneuria atrifrons Klapalek Anacroneuria atrifrons Klapalek 1922:89. Holotype S (NHML). Amazon River, Sao Paulo de Olivenga, Brazil Anacroneuria montera Stark & Sivec 1998:41. Holotype S (USNM). Iquitos, Loreto, Peru. Syn. Froehlich, 2008:129. Anacroneuria montera: Stark et al., 2001:121. Colombian records Material examined. Ecuador: Pastaza, Rio Cushueme, Cushueme, 150 km SE Puyo, 320 m, 15-30 May 1971, B. Malkin, 1S (CASC). Same site, 23 July 1971, B. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 79 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf 1 , , (, ¡" '■J i v 1 (/ III , -?/',", / ,( 1 't'/l, , iy' • r / 11 'V '< " ' "-i ■ t i ' ' , I I - t . ■ I , I , ' I i , l i. ' ' I ' I / I / ' I t * 1 '• ' ',' / i' V"y ' vj1^;:-' //M.v; ■, , 'i«> -' 11 < ■ •/ i i «i;!'.' IV t . v 1 ' ' I / '' Í / /1 7 8 9 10 Figs. 6-10. Anacroneuria cushueme male structures. 6. Head and pronotum. 7. Abdominal sternum 9. 8. Aedeagus, ventral. 9. Aedeagus, lateral. 10. Aedeagus, dorsal. Anacroneuria azul Rojas & Baena Anacroneuria azul Rojas & Baena in Stark et al., 1999:23. Holotype S (MEUV), Rio Azul, Valle del Cauca, Colombia Anacroneuria azul: Stark, 2001:7. Ecuador records Material examined. Ecuador: Pichincha, Santo Domingo de Los Colorados, 500 m, 3-8 April 1971, B. Malkin, 1 $ (CASC). Comments. Stark (2001) previously reported this species from three sites in the general vicinity of this collection in Pichincha. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 81 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Anacroneuria carchi Stark Anacroneuria carchi Stark 2001:11. Holotype $ (CMNH), Chical, Carchi, Ecuador Material examined. Ecuador: Esmeraldas, Cantón San Lorenzo, Rio Lita, 1830 ft, 00° 51.64'N, 78° 27.16'W, 2 December 2008, R.C. Mower, 2$ (BYUC). Comments. This species was known previously from the holotype (Stark 2001). Anacroneuria chavin Stark & Sivec Anacroneuria chavin Stark & Sivec 1998:16. Holotype $ (CASC). 67 mi E Tingo Maria, Yurac, Peru Material examined. Ecuador: El Oro, 9 mi S Santa Rosa, 23 January 1955, E.I. Schlinger, E.S. Ross, 11$ (CASC). Comments. Forewing lengths for these males range from 8-9 mm and the holotype forewing length is 10.5 mm. This difference and subtle differences in the aedeagal apex suggest these specimens may represent a sibling species in this complex. Specifically, the narrow dark markings along the posterior shoulder margins are less distinct for the Ecuadorian specimens and the lateral margins of the aedeagal apex are not constricted at the shoulder as they are in the holotype. A larger sample from the type locality in Peru is needed to more fully evaluate the status of these specimens. Anacroneuria cushueme Stark & Gill sp. n. (Figs. 6-10) Material examined. Holotype $ and 1$ paratype, Ecuador, Pastaza, Rio Cushueme, Cushueme, 150 km SE Puyo, 320 m, 15-30 May 1971, B. Malkin (CASC). Adult habitus. General color pale yellow patterned with pale brown. Head pattern indistinct, almost entirely yellow with only slightly darkened lappets (Fig. 6). Pronotum with sublateral pale brown bands; area adjacent to median suture and anterolateral margins pale. Femora pale brown, tibiae slightly darker at apex and base. Wings transparent, veins pale amber. Male. Forewing length 9 mm. Apical section of aedeagus with median, projecting lobe narrowed slightly from base and truncate at tip (Fig. 8). Shoulders strongly sclerotized, bulging laterad; body of aedeagus wider proximal to shoulder bases. Dorsal keel consists of two long, divergent ridges and two pairs of shorter ones near base of longer median keel lines (Fig. 10). Hooks relatively wide and acute. Lateral lobes on apical section project away from aedeagal body, one pair lunate and the other almost quadrate (Fig. 9). Hammer thimble shaped (Fig. 7). Female. Unknown. Larva. Unknown. Etymology. The species name, used as a noun in apposition, is based on the type locality. Diagnosis. Based on aedeagal morphology, this species is related to the A. yameo Stark & Sivec 1998 group of species, which also includes A. pakitza Stark & Sivec 1998, A. pinza Stark 1995, and A. zunigae Stark 2001. The new species bears a greater similarity in aedeagal structure to A. yameo and A. pakitza but in these species the shoulder region bulges toward the aedeagal apex creating an apically trilobed structure; this is quite distinct in the dorsal aspect and differs from the comparable structures in A. cushueme which bulge more laterad. In A. cushueme the mesal projection of the aedeagal apex is truncate at the tip and broad at the base which also differs from both related species. Anacroneuria guambiana Zuniga & Stark (Figs. 11-13) Material examined. Ecuador, Napo Province, 6.38 km W Baeza, Hwy E-20, 1,987 m, 25 January 2012, B.C. Kondratieff, B. Gill 1$, 1?, 1 larva, 1 exuvium (CSUC). Napo Province, 8.68 km NW Baeza, Hwy E-20, 2,134 m, 24 January 2012, B.C. Kondratieff, B. Gill, 1$, 1 exuvium (CSUC). Larva. Body length pre-emergent female 16-17 mm. General color brown, patterned with dark brown. Head brown with pale areas at frontoclypeus, M-line, lateral to ocelli and over most of occiput (Fig. 11); Mline incomplete adjacent to pale mesal spot. Pronotum brown but with pale leg shaped areas adjacent to median suture, and with a few additional pale rugosities. Lateral pronotal setal fringe incomplete anterolaterally. Femora pale brown but darker along ventral and apical margins; fine setal Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 82 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf 11 13 Figs. 11-13. Anacroneuria guambiana larval structures. 11. Head, pronotum and terminal abdominal segments. 12. Left fore femur. 13. Cercal segments 5-7, 16-18, 31-33, lateral. fringe well developed; basal patch of short thick setae extensive along dorsal margin; transverse setal row consists of ca. 7 long, thick setae; scattered thick setae sparse beyond transverse row and with ca. 16 on lower half of femoral surface. Ventral margin with a few thick setae and interspersed fine short setae (Fig. 12). Abdomen dark brown but apical segments with a small median pale spot (Fig. 11), terga without apparent intercalary setae. Cerci without well-developed fine setal fringe (Fig. 13), only a few fine setae occur along dorsal margin of apical segments; cercal segments bear apical whorls of short, thick setae, except on some basal segments. Comments. The male specimen from the first site listed above varies slightly from typical A. guambiana specimens in having the apical aedeagal section with convergent lateral margins and in having the aedeagal shoulders approaching the apical section at an obtuse angle. Larger samples are needed to resolve whether this specimen represents a sibling member of the A. guambiana complex. The larva is generally similar to that of A. paleta Stark 1995 (Maldonado et al. 2002), but that species has a complete M-line and more extensive dark pigment on the occiput. These specimens are the first representatives of this species to be reported for Ecuador. The first site listed in the materials section above (Figs. 32-33) is located in the lower montane cloud forest at 1,987 m, and is composed mainly of alternating step-pools. The substrate ranges from depositional fines to large boulders, however, size extremes of sand and boulders dominate. The stream channel has numerous debris dams and downed trees from adjacent riparian zones. Steep walls surrounding the stream are covered with plants that Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 83 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf 16 17 18 Figs. 14-18. Anacroneuria kayceae male structures. 14. Head and pronotum. 15. Abdominal sternum 9. 16. Aedeagus, ventral. 17. Aedeagus, lateral. 18. Aedeagus, dorsal. confine the channel and create approximately 71% canopy cover. The channel is 1.56 m wide and depth averaged 0.13 m. Discharge measured during the collection at this site was 37.1 L s-1. Interestingly, the substrate was partially covered at the time of the collection by filamentous algae not typically seen in the region. The second stream (Fig. 34) is characterized by stretches of cobble filled riffle between cascades formed by large boulders. The channel is braided and narrow at various points, and is relatively free of debris dams or downed trees. Channel width is 2.32 m and depth averages 0.19 m. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 84 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Discharge was measured at 105.3 L s-1 during collecting. Canopy cover varies from open to closed along the channel. Adjacent riparian areas are composed of vegetation typical of lower montane cloud forest. Anacroneuria jewetti Stark Anacroneuria jewetti Stark 2001:19. Holotype S (USNM). San Francisco de Borja, Napo, Ecuador Anacroneuria jewetti: Zúñiga et al., 2006:55. Female description Material examined. Ecuador: Imbabura Province, Cantón Cotacachi, Rio Intag, Apuela, 5020 ft, 00° 21.2'N, 78° 31.0'W, 6 November 2009, S.M. Clark, H.R. Hinkson 8^ (BYUC). Comments. These specimens are the first to be reported from Imbabura Province. Anacroneuria kayceae Kondratieff & Gill sp. n. (Figs. 14-18) Material examined. Holotype S, Ecuador, Napo, 6.38 km W Baeza, Hwy E-20, 1,987 m, 0° 27.123'S, 77° 56.468'W, 25 January 2012, B.C. Kondratieff, B. Gill (USNM). Adult habitus. General color pale yellow, white in alcohol. Head with slightly darker, but obscure markings over ocelli, lappets and posterolateral areas of occiput (Fig. 14). Pronotum pale along lateral margins and median suture, but with irregular, dark brown, sublateral bands. Ocelli narrowly separated; wings transparent, veins pale. Legs, antennae and palpi pale. Male. Forewing length 17 mm. Hammer almost cylindrical, but slightly wider at base than apex (Fig. 15). Aedeagal apex truncate, lateral margins slightly expanded and appearing swollen at base (Figs. 16, 18); hooks strongly sclerotized at base with a distinct dark line outlining inner basal margin, and bearing a strong ventral keel from near midlength to near tips. Dorsal keel consists of two close-set, low ridges, extending for about half of apical area length (Fig. 18). In lateral aspect much of aedeagal apex membranous (Fig. 17). Female. Unknown. Larva. Unknown. Etymology. We honor Dr. Kayce Anderson, Colorado State University, for her dedication to ecological studies in Ecuador, and her appreciation of Neotropical aquatic biology. Diagnosis. The aedeagus of this species is similar to that of the Peruvian-Bolivian species, A. pachacuti Stark & Sivec 1998, particularly in lateral aspect. However, in that species the apical section of the aedeagus is offset by prominent basolateral notches and the aedeagal apex is nipple-shaped rather than truncate (see Stark & Sivec 1998, Figs. 124-126). In addition, A. pachacuti is a smaller species (FWL ~11 mm) with a slightly darker habitus. The aedeagus is also similar to that of A. rawlinsi Stark, 2001, but in that species the apical aedeagal section lacks swollen basolateral lobes. Comments. The type locality (Figs. 32-33) is described under A. guambiana. Anacroneuria malkini Stark & Kondratieff sp. n. (Figs. 19-23) Material examined. Holotype Ecuador, Pastaza, Rio Cushueme, Cushueme, 150 km SE Puyo, 320 m, 15-30 May 1971, B. Malkin (CASC). Adult habitus. General color yellow-brown patterned with darker pigment. Head mostly yellow but with a large dark brown patch covering ocelli, and with dark brown lappets. Antennae dark brown, pronotum brown with scattered pale rugosities and with a pale median band and with areas of lighter pigment along lateral margins (Fig. 19). Femora brown in apical half with narrow black band at knee; basal half of femora pale, tibiae brown. Wings transparent with pale amber tint, veins dark amber with R vein dark brown. Male. Forewing length 8 mm. Aedeagal apex narrowed from shoulders, rounded at tip, and bearing a pair of membranous ventral lobes (Fig. 23). Dorsal keel consists of a pair of divergent, narrow ridges and a small transverse ridge set at the base of the divergent ridges (Fig. 21); lateral aspect of aedeagal apex broad basally, narrowed to an acute tip and directed toward apex (Fig. 22). Aedeagal hooks gradually curved to acute tips. Abdominal sternum 9 bears a thimble shaped hammer (Fig. 20). Female. Unknown. Larva. Unknown. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 85 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf 19 21 22 23 20 Figs. 19-23. Anacroneuria malkini male structures. 19. Head and pronotum. 20. Abdominal sternum 9. 21. Aedeagus, dorsal. 22. Aedeagus, lateral. 23. Aedeagus, ventral. Etymology. The patronym honors the late Borys Malkin who collected the holotype of this and many other species. Mr. Malkin was a famed naturalist and friend of the well-known Plecoptera authority, Stanley G. Jewett, Jr. The specimens were part of the Jewett collection directed to us by R.W. Baumann. Diagnosis. The aedeagus of this species is generally similar to that of A. tzapino Stark 2001, a species also known from Pastaza. The aedeagal apex of that species is slightly notched, the dorsal keel arms are parallel for most of their length, and the apical aedeagal region slants slightly ventrad in lateral aspect (see Stark 2001, Figs. 118-120). In addition, A. tzapino has a more extensive area of brown pigment on the head and the wings are darker, but also have an obscure pale spot at the cord (Stark 2001). Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 86 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf 24 R •■ /■-''■ <','""• i, (AX'' 1 I ■ . \ \ \ t • / ; . ' 1 v I i /' i ' ' / r '1 / < ' / < , 25 26 27 28 Figs. 24-28. Anacroneuria pichincha male structures. 24. Head and pronotum. 25. Abdominal sternum 9. 26. Aedeagus, ventral. 27. Aedeagus, lateral. 28. Aedeagus, dorsal. Anacroneuria pichincha Stark & Kondratieff sp. n. (Figs. 24-28) Material examined. Holotype Ecuador, Pichincha, Santo Domingo de Los Colorados, 500 m, 3-8 April 1971, B. Malkin (CASC). Adult habitus. General color yellow-brown patterned with dark brown. Head with a quadrangular area covering ocellar area forward to about mid-frons; pigmented area with a slight anteromedian notch (Fig. 24); lappets pale brown, antennae brown. Pronotum dark over much of disk but pale along median suture, anterolateral margins and bearing a few small, pale rugosities. Femora pale brown except for narrow, dark knee band; tibiae pale except for small dark bands at base and near apex. Wing membrane dark amber with darker veins; R-vein particularly dark in basal half. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 87 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Male. Forewing length 10.5 mm. Apical section of aedeagus indistinctly trilobed with narrow median lobe projecting well beyond lateral, somewhat bulging shoulders (Figs. 26, 28); entire apical area covered by large ventral membranous lobes; shoulders with short dorsal ridges extending inwards toward bases of median, finger-like lobe. Body of aedeagus constricted near apical third, but with low lateral bulges projecting over bases of hooks; tips of hooks narrowly truncate. Lateral lobes of aedeagal apex project away from aedeagal body in ear-like fashion (Fig. 27). Abdominal sternum 9 bears a thimble shaped hammer (Fig. 25). Female. Unknown. Larva. Unknown. Etymology. The species name, used as a noun in apposition, is based on the type locality. Diagnosis. Aedeagal morphology places this species near A. curiosa Stark 1998, a Mesoamerican species, and A. cosnipata Stark & Sivec 1998, a species known from several Peruvian sites. The latter species bears a broad, dorsomedian, mesa-like structure on the aedeagus, and the median lobe on the aedeagal apex is wider than in the new species (see Stark & Sivec 1998, Figs. 53-55). Neither of these related species has the aedeagus strongly constricted proximal to the apical section, neither have the hooks blunt at the tips and neither shares the distinctive color pattern of A. pichincha. 30 29 31 Figs. 29-31. Anacroneuria EC-1 female structures. 29. Head and pronotum. 30. Egg. 31. Abdominal sterna 8-9. Anacroneuria EC-1 (Figs. 29-31) Material examined. Ecuador: Napo Province, 2.78 km W Papallacta, Hwy E-20, 3296 m, 25 January 2012, B.C. Kondratieff, B. Gill, 1? (CSUC). Adult habitus. General color dark brown to black. Head yellow, with dark brown patches extending forward of each ocellus toward central frons; lappets dark brown, posterolateral occiput and anteromedian frons pale brown (Fig. 29). Antennae and legs dark brown, femora paler in ventral half. Pronotum dark brown over lateral third, and bearing a thin, dark, diagonal line near anterolateral margin. Wing membrane dark brown to black, veins brown except pale R vein. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 88 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Figs. 32-35. Ecuadorian collecting sites. 32. Napo Province, 6.38 km W Baeza, 1,987 m. 33. Same site as previous figure. 34. Napo Province, 8.68 km W Baeza, 2,134 m. 35. Napo Province, 2.78 km W Papallacta, 3,296 m. Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 89 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Male. Unknown. Female. Forewing length 31 mm. Subgenital plate bilobed with truncate posterior margins; lobes separated by a shallow V-shaped notch. Sternum 9 bearing an obscure transverse sclerite, median sclerite covered by dark setae (Fig. 31). Egg. Somewhat thimble shaped with broadly rounded anterior pole almost as wide as collar end (Fig. 30). Collar small and button shaped. Chorionic surface without follicle cell impressions. Comments. Females representing more than 30 Anacroneuria species have been described under temporary designations consisting of a two letter national code followed by a number. Ten have been proposed for Colombia (Stark et al. 1999; Zuniga & Stark, 2002; Zuniga et al. 2007), five for Guyana (Stark 1999; 2000), three for Paraguay (Stark & Baumann 2011), one for Peru (Stark & Sivec 1998), and twelve for Venezuela (Stark 1995; Maldonado et al. 2002). Several additional species have been proposed as Anacroneuria sp., or as Anacroneuria sp. A, etc., without a prefix; six of these (sp. A, B, C, D, E, F) were proposed by Harper (1992) for the Panamanian fauna. Only one of these unassociated females, A. CO-5 (FWL 24-26 mm), has a forewing length approaching that of this species; both species also have the subgenital plate bilobed, however the former species is quite pale in comparison to A. EC-1. Among large Anacroneuria species where sufficient specimens are known to generate a significant sample size [e.g. A. magnirufa Jewett 1958 or A. plutonis (Banks 1914)], male forewing length is approximately 75% of female forewing length (Stark 1998). If this is applied to the current female, the male could be expected to display a forewing length of about 23 mm. The site (Fig. 35) where this female was collected is a high gradient stream characterized by a series of large waterfalls and short stretches of riffles. The channel width is 1.63 m and average depth at the time of collection was 0.12 m. Discharge was 31.3 L s-1 at the time of the collection, and canopy cover was about 47%. Vegetation is typical of high altitude Andean cloud forest; however, it has been disturbed by adjacent anthropogenic activities. This stream is one of the few in the Papallacta area where larvae of the gripopterygid Claudioperla cf. tigrina (Klapalek) were found in oxygenated areas with smaller substrate in large numbers. Ecuadorian Anacroneuria Species List Species Provincial Records A. anchicaya Baena & Zúñiga 1999 in Stark et al. 1999 A. angusticollis (Enderlein 1909) A. antizana Stark 2001 A. apuela Stark & Gill sp. n. A. atrifrons Klapálek 1922 A. auca Stark 2001 A. azul Rojas & Baena 1999 in Stark et al. 1999 A. bari Stark 1995 A. cajas Zúñiga & Vimos 2006 in Zúñiga et al. 2006 A. camposi (Banks 1920) A. canelo Stark 2001 A. caraca Stark 1995 A. carchi Stark 2001 A. cayapa Stark 2001 A. chachis Stark & Vásconez 2006 in Zúñiga et al. 2006 A. chavin Stark & Sivec 1998 A. chimborazo Stark 2001 Cotopaxi, Napo, Pastaza, Pichincha Napo, Pastaza, Zamora Chinchipe Napo Imbabura Pastaza Pastaza Pastaza, Pichincha Napo Azuay Pichincha Napo Napo Carchi Pichincha Pastaza, Pichincha El Oro Chimborazo Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 90 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf A. choco Stark & Berosa 2006 in Zúñiga et al. 2006 A. cotacachi Stark 2001 A. cotopaxi Stark 2001 A. cushueme Stark & Gill sp. n. A. guambiana Zúñiga & Stark 1999 in Stark et al. 1999 A. guayaquil Zúñiga & Rojas 1999 in Stark et al. 1999 A. hieroglyphica (Enderlein 1909) A. jewetti Stark 2001 A. jivaro Stark 2001 A. kayceae Kondratieff & Gill sp. n. A. kitchensi Stark 2001 A. kondratieffi Stark 2001 A. major Stark 2001 A. malkini Stark & Kondratieff sp. n. A. mindo Zúñiga & Vásconez 2006 in Zúñiga et al. 2006 A. ohausiana (Enderlein 1909) A. parva Stark 2001 A. pastaza Stark 2001 A. pichincha Stark & Kondratieff sp. n. A. pistacina (Enderlein 1909) A. planada Baena & Rojas 1999 in Stark et al. 1999 A. portilla Stark & Rojas 1999 in Stark et al. 1999 A. puna Stark 2001 A. quijo Stark 2001 A. quilla Stark & Zúñiga 1999 in Stark et al. 1999 A. rawlinsi Stark 2001 A. regleta Stark & Rojas 1999 in Stark et al. 1999 A. ricki Zúñiga & Stark 2002 A. rosita Stark & Rojas 1999 in Stark et al. 1999 A. rugosa Stark 2001 A. schmidti (Enderlein 1909) A. spangleri Stark 2001 A. tena Stark 2001 A. tungurahua Stark 2001 A. tzapino Stark 2001 A. zamora Stark 2001 A. zunigae Stark 2001 Esmeraldas, Pichincha Pichincha Cotopaxi Pastaza Napo Chimborazo Balzapampa (Bolivar Province ?) Azuay, Imbabura, Loja, Morona Santiago, Napo, Pichincha Azuay, Cañar Napo Pastaza Los Rios Azuay Pastaza Pichincha Loja Napo Napo, Pastaza Pichincha Balzapampa (Bolivar Province ?) Napo Loja, Napo, Pichincha Cotopaxi Napo Azuay, Carchi, Zamora Chinchipe Morona Santiago Carchi, Zamora Chinchipe Esmeraldas Pastaza Pichincha Balzapampa (Bolivar Province ?) Los Rios, Pichincha Napo Tungurahua Pastaza Zamora Chinchipe Napo ACKNOWLEDGEMENTS We are grateful to Dr. Richard W. Baumann of Brigham Young University for sending specimens from the Monte L. Bean Museum collection, and for forwarding a small series of specimens from the S.G. Jewett collection for inclusion in this study. Dr. Baumann also provided background information on Borys Malkin. Michael Kippenhan, McMinnville, Oregon is thanked for providing specimens from Paraguay. We also thank Dr. Claudio Froehlich for sharing information on location of the Anacroneuria atrifrons holotype. Lori Discoe, Fort Collins, Colorado, produced the line illustrations of the head and prothorax of the new taxa. The field work in Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 91 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Ecuador was supported by NSF Award 1046408 Dimensions: "Collaborative Research: An integrative traits-based approach to predicting variation in vulnerability of tropical and temperate stream biodiversity to climate change." We thank Drs. N. LeRoy Poff, Colorado State University, Alexander S. Flecker, Cornell University, and Andrea C. Encalada, Universidad San Francisco de Quito, Ecuador, for field assistance and other support. REFERENCES Banks, N. 1914. New neuropteroid insects, native and exotic. Proceedings of the Academy of Natural Sciences of Philadelphia, 66:608-632. Banks, N. 1920. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology, 64:314-325. Enderlein, G. 1909. Klassification der Plecopteren sowie Diagnosen neuer Gattungen und Arten. Zoologischer Anzeiger, 34:385-419. Froehlich, C.G. 2008. Old species of Neotropical Plecoptera. Pp. 125-132 in Hauer, F.R., J.A. Stanford & R.L. Newell [editors]. International advances in the ecology, zoogeography and systematic of mayflies and stoneflies. University of California Publications in Entomology, Volume 128. University of California Press, Berkeley, California. 424 pp. Harper, P.P. 1992. Stoneflies of Panama (Plecoptera. Pp. 114-121 in Quintero, D. & A. Aiello [editors]. Insects of Panama and Mesoamerica, selected studies. Oxford University Press, Oxford. Illies, J. 1963. Revision der südamerikanischen Gripopterygidae (Plecoptera). Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 36:145-248. Jewett, S.G. 1958. Stoneflies of the genus Anacroneuria from Mexico and Central America (Plecoptera). The American Midland Naturalist, 60-159-175. Klapâlek, F. 1922. Plécoptères nouveaux. Quatrième partie. Annales de la Societé Entomologique de Belgique, 62:89-95. Maldonado, V., B.P. Stark, & C. Cressa. 2002. Descriptions and records of Anacroneuria from Venezuela (Plecoptera: Perlidae). Aquatic Insects, 24:219-236. Rojas, A.M. & M.L. Baena. 1993. Anacroneuria farallonensis (Plecoptera: Perlidae) una nueva especie para Colombia. Boletin del Museo de Entomologia de la Universidad del Valle, 1:23-28. Stark, B.P. 1995. New species and records of Anacroneuria (Klapâlek) from Venezuela. Spixiana, 18:211-249. Stark, B.P. 1998. The Anacroneuria of Costa Rica and Panama (Insecta: Plecoptera: Perlidae). Proceedings of the Biological Society of Washington, 111:551-603. Stark, B.P. 1999. Anacroneuria from northeastern South America (Insecta: Plecoptera: Perlidae). Proceedings of the Biological Society of Washington, 112:70-93. Stark, B.P. 2000. Notes on the Anacroneuria (Plecoptera: Perlidae) of Guyana with the description of a new species. Aquatic Insects, 22:305-310. Stark, B.P. 2001. Records and descriptions of Anacroneuria from Ecuador (Plecoptera: Perlidae). Scopolia, 46:1-42. Stark, B.P. & R.W. Baumann. 2011. Records of Anacroneuria (Plecoptera: Perlidae) from Bolivia and Paraguay with descriptions of three new species. Illiesia, 7:182-191. Stark, B.P. & I. Sivec. 1998. Anacroneuria of Peru and Bolivia (Plecoptera: Perlidae). Scopolia, 40:1-64. Stark, B.P. & M.C. Züniga. 2003. The Anacroneuria guambiana complex of South America (Plecoptera: Perlidae). Pp. 229-237 in Gaino, E. [editor]. Research update on Ephemeroptera and Plecoptera. University of Perugia, Perugia, Italy. 488 pp. Stark, B.P., M.C. Züniga, & I. Sivec. 2001. Descriptions of Anacroneuria spp. (Plecoptera: Perlidae) from the upper Rio Amazonas drainage, Colombia and Peru. Acta Entomologica Slovenica, 9:119-122. Stark, B.P., M.C. Züniga, A.M. Rojas, & M.L. Baena. 1999. Colombian Anacroneuria: Descriptions of new and old species (Insecta, Plecoptera, Perlidae). Spixiana, 22:13-46. Turcotte, P. & P.P. Harper. 1982a. Drift patterns in a high Andean stream. Hydrobiologia, 89:141-151. Turcotte, P. & P.P. Harper. 1982b. The macro-invertebrate fauna of a small Andean stream. Freshwater Biology, 12:411-419. Züniga, M.C. & B.P. Stark. 2002. New species and records of Colombian Anacroneuria (Insecta, Plecoptera, Perlidae). Spixiana, 25:209-224. Züniga, M.C., B.P. Stark, W. Cardena, C. Tamaris- Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 92 Stark, B.P., B.C. Kondratieff, and B. Gill. 2012. New species and records of Anacroneuria (Plecoptera: Perlidae) from Ecuador and Paraguay. Illiesia, 8(06):78-93. Available online: http://www2.pms-lj.si/illiesia/papers/Illiesia08-06.pdf Turizo, & O.E. Ortega. 2007. Additions to the Colombian Anacroneuria fauna (Plecoptera: Perlidae) with descriptions of seven new species. Illiesia, 3:127-149. Zúñiga, M.C., B.P. Stark, J.J. Vásconez, F. Berosa, & L.D. Vimos. 2006. Colombian and Ecuadorian Anacroneuria (Plecoptera: Perlidae): seven new species, records and life stages. Studies on Neotropical Fauna and Environment, 41:45-57. Received 2 May 2012, Accepted 16 May 2012, Published 24 May 2012 Illiesia - http://www2.pms-lj.si/illiesia/ Volume 8 - Number 6 - Page 93