Neaviperla forcipata Ricker

Neaviperla forcipata Ricker

(Figs. 1-18)

Alloperla forcipata Neave 1929:160. Holotype ♂ (Canadian National Insect Collection), Lake Edith, Jasper National Park, Canada

Alloperla (Neaviperla) forcipata: Ricker, 1943:142.

Neaviperla forcipata: Illies, 1966:448.

Suwallia forcipata: Alexander & Stewart, 1999:202. Suwallia forcipata: Stewart & Stark, 2002:280.

Suwallia forcipata: Stark et al. 2012:11.

Suwallia forcipata: Stewart and Stark in Merritt et al. 2008

Suwallia forcipata: DeWalt et al. 2014

Material examined. CANADA: Alberta, Minnewanka Creek, passing through lower bank head, Banff National Park, 22 VIII 1969, C.M. Yarmoloy, 1♂, 1♀ (BYUC); Stoney Creek, Banff National Park, 22 VIII 1969, C.M. Yarmoloy, 2♀ (BYUC). British Columbia, Babine River, Kisgegas, 21 VIII 1952, W.E. Ricker, 3♂, 6♀ (CNCI); Chilliwack River, Vedder Crossing, 16-IX-4-X-1937, W.E. Ricker, 31♂, 31♀ (CNCI); Chilliwack River, Chilliwack River Provincial Park, 26 VIII 1991, J. Myack, 6♂, 3♀ (CSUC); Frost Creek, Cultus Lake, 22 X 1937, W.E. Ricker, 4♂, 40♀ (CNCI); creek near Garabaldi Station, 27 IX 1964, W.E. Ricker, 1♀ (CNCI); creek at mile 20, N Hazelton, 20 VIII 1952, W.E. Ricker, 4♂ (CNCI); Kitimat River, Hwy 37, 30 km N Kitimat, 1 I 2000, W.N. Mendel, 1♀ (BYUC); Kleanza Creek, near Terrace, 18 VIII 2008, W.M. Jessop, 1♂ (BYUC); Upper Pitt River, Alvin, 23 IX 1938, W.E. Ricker, 1♂, 6♀ (CNCI); Ritchie Creek, Cassiar Hwy, mi 128, 10 IX 1985, R.L. Botorff, 1♂, 7♀ (RLBC); Skeena River, 14 IX 2005, T. Bansak, 2♀ (UMBS); Telegraph Creek, Trapper Lake Horse Camp, 22 VIII 1959, J. Ricker, 1♂ (CNCI); Waukwash River inlet, Owikeno Lake, 12 X 1956, D. Foskett, 1♀ (CNCI). Yukon, Blackstone River, km 141, Demster Hwy, 11 VIII 1980, R.J. Cannings, 1♂ (SMDV); Canol Road, km 154, 61° 34’N 133°05’W, 28 VII 1997, B. Marshall, 1♂ (ROME); Rose River, Rose Lake, 61° 35’N 133° 05’W, 28 VII 1981, C.S. Guppy, 1♀ (SMDV); 7 km E Rancheria, 60° 04’N 130° 29’W, 5 VIII 1981, C.S. Guppy, 1♂ (SMDV); Starr Creek, 61° 46’N 131° 51’W, 3 VIII 1981, C.S. Guppy 1♂ (SMDV). UNITED STATES: Alaska, Athel Creek, Glenn Hwy, MP 264.5, 11 VIII 1954, C.P. Alexander, 1♀ (USNM); Barr Creek, Seward Peninsula, 70 mi N Nome, 8 VIII 1982, R.L. Newell, 2 larvae, (BYUC); Bear River, Bear Lake, Alaska Peninsula, 4 VIII 1958, J.D. Fiske, 5 larvae (USNM); Same locality, 11 VIII 1958, J.D. Fiske, 1 larva (USNM); creek near Black Lake, Hwy 98, 2 mi N Skagway, 59° 30’N 135°16’W, 21 VIII 2002, Baumann, Huntzinger & Clark, 1♂ (BYUC); Cold Bay, 24 VII 1958, C.H. Lindroth, 2♂, 1♀ (USNM); Granite Creek, Anchorage, 8 IX 1966, K.M. Sommerman, 4♂, 21♀ (USNM); Indian River, near Sitka, 14 IX 2009, G.M. Smith, 1♂, 1♀ (BYUC); Same locality, 13 IX 2010, G.M. Smith, 3♀ (BYUC); Indian River, Sitka National Historic Park, 28 VII 2010, G.M. Smith, 5♂, 5♀ (BYUC); Indian River, Tongass National Forest, 13 VIII 2010, G.M. Smith, 16♂, 10♀ (BYUC); Kwethluk River, Yukon Delta, 17 VIII 2011, Z.J. Crete, 1♂ (ZJCC); Kwethluk River, 22 VIII 2012, Z.J. Crete, 11♂, 9♀ (ZJCC); Same locality, 1 IX 2012, Z.J. Crete, 3♂, 18♀ (ZJCC); Montana Creek, 3 mi N Auke Bay, 21 VIII 2011, R.L. Bottorff, 8♂, 6♀ (RLBC); Montana Creek, near rifle range, Juneau, 10 VIII 1998, J. Hudson, 29♂, 26♀ (BYUC); Same locality, 30 VIII 2011, J. Hudson, 1♀ (BYUC); Mount McKinley Park, 17 VIII 1958, C.H. Lindroth, 1♀ (USNM); Orzenoi Bay, opposite Shumigan Island, 26 VII 1957, J.D. Fiske, 10 larvae (USNM); Same locality, 23 VIII 1957 J.D. Fiske, 10 larvae (USNM); Nome, Seward Peninsula, 13 VIII 1958, C.H. Lindroth, 1♂, 5♀ (USNM); Palmer Creek, Anchorage, 9 IX 1966, K.M. Sommerman, 4♀ (USNM); Paxson, 19 VIII 1958, C.M. Lindroth, 5♂, 9♀, 1 larva (USNM); Pullen Creek, Skagway, 10 VIII 2002, K.T. Huntzinger, 21♂, 4♀ (BYUC); same locality, 18 VIII 2002, Baumann, Huntzinger & Clark, 45♂, 35♀ (BYUC); Reid Creek, Reid Falls, Skagway, 19 VIII 2002, Baumann, Huntzinger & Clark, 2♂, 4♀ (BYUC); Russian River, 0.1 mi above confluence Kenai River, 12 IX 2002, M. Floyd, 3♂, 3♀ (MUIC); Salmon Creek, above jct. Gastineau Channel, Juneau, 27-30 VIII 2002, S.M. Clark, 2♂, 5♀ (BYUC); stream entering Gastineau Channel ¼ mi S Salmon Creek, Juneau, 29 VIII 2002, S.M. Clark, 1♂ (BYUC); Skagway River, Skagway, 59° 28’N 135° 18’W, 18 VIII 2002, R.W. Baumann & S.M. Clark, 107♂, 12♀ (BYUC); Same locality, 21 VIII 2002, Baumann, Huntzinger & Clark, 17♂, 3♀ (BYUC); Steep Creek, near Mendenhall Lake, 15 km N Juneau, 12 VIII 1998, J. Hudson, 11♂, 15♀ (BYUC); Tributary, Surprise Lake, Aniakchak National Monument, Alaska Peninsula, 56° 56’N 158° 06’W, Date?, Collector?, 1♀ (SMDV); Taiya River, Chilkoot Trailhead, Dyea, 19-23 VIII 2002, Baumann, Clark & Huntzinger, 68♂, 19♀ (BYUC); ponds near White Pass, above Skagway, 19 VIII 2002, Baumann, Clark & Huntzinger, 24♂, 6♀ (BYUC). Montana, Flathead Co., Avalanche Creek, Avalanche Campground, Glacier National Park, 10 IX 1968, R.W. Baumann & R.L. Newell, 2♀ (BYUC); South Fork Cut Bank Creek, Schildt Road, 8 VIII 1986, B.C. Kondratieff, 3♂ 5♀ (CSUC); Logan Creek, jct. Mac Donald Creek, Glacier National Park, 10 IX 1968, R.W. Baumann & R.L. Newell, 2♂ (BYUC); Mac Donald Creek, above Mac Donald Lake, Glacier National Park, 10 IX 1968, R.W. Baumann & R.L. Newell, 3♂, 5♀ (BYUC); Park Creek, Lake Isabell Trail, 8 VIII 1986, B.C. Kondratieff, 8♂, 3♀ (CSUC). Washington, Jefferson Co., Hoh River, Hoh River Campground, Olympic National Park, 12-IX-2013, J.J. Lee & R.A. Lee, 10♂, 14♀, 57 larvae (BYUC, JJLC); Whatcom Co., Canyon Creek, Mosquito Lake Road, 4 mi E Deming, 12 IX 1993, Kraft & Minakawa, 7♂, 22♀ (BYUC); Devils Creek, Ross Lake, near Newhalem, 2 X 1993, Kraft, Busse & Minakawa, 2♀ (BYUC); Lightning Creek, Ross Lake, near Newhalem, 2 X 1993, Kraft, Moody, Fritz & Kraft, 3♂ 7♀ (BYUC); Middle Fork Nooksack River, 8 mi E Van Zandt, 1 IX 1993, Kraft, Sammeth & Minakawa, 1♂, 1♀, 44 larvae (BYUC); North Fork Nooksack River, 9 X 1966, K. Vander Mey, 1♂, 3♀ (BYUC).

The following characters illustrate the distinctive differences which serve to separate Neaviperla as a genus in the family Chloroperlidae and to add characters useful in diagnosis.

Adult. A distinct sclerotized line is present on the meso and meta thorax that intersects the center of the large U-shaped mark at the base of the segment. This appears as a W in contrast to the U commonly seen in Suwallia and makes it easy to separate adults quickly. Male. Five basal segments of cerci modified into clasper-like structures (Figs. 1-4). First segment very large, curved outward and heavily sclerotized (Fig.4), with large inward directed, triangular–shaped process at base (Figs. 4, 7). Segments 2-5 smaller, forming stout inward pointing lobes, sharp apically, and bearing many large setae, which cover irregular grooves (Figs. 4, 7). Ninth segment enlarged, about length of segments 7 and 8, tergum covered by large plate-like structure (Figs. 3, 8), base broadly rounded, tapered to narrow pointed tip, which extends anteriorly to segment 8 (Figs. 3, 7). Lateral margins of tergum 10 formed into inward curved lobes, bluntly truncate apically, with irregular grooves, bearing few long hairs near base, with narrow patch of shorter hairs near apex (Fig. 6).

Aedeagus. Stout and capped by dorsal medial hump, when recurved, composed of densely compacted hair-like structures. Aedeagal base with pair of sclerotized hook-like processes that are visible through the ventral cuticle of sternum 9. Processes composed of numerous flat sclerotized tubercles that join together to form a comma-shaped structure (Fig. 9).

Female. Cerci with first segment elongate, much longer than remaining segments and slightly concave inwardly (Fig. 10).

Larvae. Head and prothorax heavily covered with long hairs (Figs. 11, 12). Basal cercal segment of mature larvae long, with whorl of stout setae that is not associated with segment margins (Fig. 15).

Egg. Collar with two layers and chorion smooth.

Surdick (1985) constructed a phylogenetic tree that considers the modified cerci of Neaviperla as the apomorphic state and also mentions the unique modifications of the ninth tergum. She considers the hemitergal process of Suwallia as an apomorphic character with the poorly developed processes of Neaviperla as plesiomorphic. Alexander and Stewart (1999) do not address Surdick’s phylogenetic tree but instead place Neaviperla basally in a clade with four Palearctic species: Suwallia jezoensis (Kohno), S. kerzhneri Zhiltzova and Zwick, S. talalajensis Zhiltzova and S. teleckojensis (Samal) based on egg structure, color pattern and hemitergal process shape. However, the exact characters are not given and no phylogeny is included for this clade. The egg of S. kerzhneri is unknown. Neaviperla forcipata is the only species mentioned in Alexander and Stewart (1999) whose egg has a collar with two layers and a smooth chorion. In addition, based on Alexander and Stewart (1999), Neaviperla is distinct from the four Palearctic species mentioned by having a W-shaped rather than simply a U-shaped meso-metanotal mark. Neaviperla has a brown head pattern that covers the inter ocellar area and extends forward to the labrum, while the mentioned Asian species exhibit coloration primarily between the ocelli, which is stated to be nearly black.

The larvae of Neaviperla and Suwallia are indeed similar but that is not surprising given the fact that the family Chloroperlidae is quite conservative in larval characters. The dorsal surface of the pronotum is covered with hairs in both genera but they tend to be larger and thicker in Neaviperla (Fig. 12). The mesosternum bears few hairs in each and those on Neaviperla are very delicate (Fig. 16). The basal segments of the mature larval cerci show the best diagnostic characters. They are longer in Neaviperla and bear a whorl of stout spines located proximal to the first row of cercal spines (Fig. 15). These spines are not associated with the cercal segment divisions and could be related to the extreme modifications of the adult cerci.