Torrenticola ellipsoidalis (Marshall, 1943)

Fisher, J. Ray, Fisher, Danielle M., Skvarla, Michael J., Nelson, Whitney A. & Dowling, Ashley P. G., 2017, Revision of torrent mites (Parasitengona, Torrenticolidae, Torrenticola) of the United States and Canada: 90 descriptions, molecular phylogenetics, and a key to species, ZooKeys 701, pp. 1-496 : 1

Torrenticola ellipsoidalis (Marshall, 1943)

Atractides ellipsoidalis : Marshall 1943: 308.

T. ellipsoidalis : Mitchell 1954: 40 • Habeeb 1955: 2 • Viets 1956: 241 • Crowell 1961: 330 • Habeeb 1967: 3 • Conroy 1968: 28 • Habeeb 1974: 1 • Conroy and Scudder 1975: 307 • Quaglia and Conroy 1984: 89 • Viets 1987: 759.

Torrenticola rectiforma Habeeb, 1974: 1.

Material examined.

Type series. HOLOTYPE (♀): from USA, California, Nevada County, north of Lake Tahoe, Martis Creek, Jun 1933, by PR Needham, RM330010.

PARATYPES (1 ♀; 0 ♂): California, USA: 1 ♀ from Nevada County, north of Lake Tahoe, Martis Creek, Jun 1933, by PR Needham, RM330010.

OTHER MATERIAL (94 ♀; 94 ♂): Alberta, Canada: 1 ♀ and 2 ♂ from Waterton Lakes National Park, Cameron Creek, beside Akamina Parkway (49°5'N, 113°52'W), 2 Aug 1985, by IM Smith, IMS850133 • 2 ♀ and 2 ♂ from Waterton Lakes National Park, Cameron Creek, beside Akamina Parkway, west of Rowe Creek, 27 Jun 1980, by IM Smith, IMS800086A • 1 ♀ and 2 ♂ from Waterton Lakes National Park, Cameron Creek, beside Akamina Parkway, 12-15 Jun 1980, by IM Smith, IMS800069A & IMS800069B • 2 ♀ and 2 ♂ from Waterton Lakes National Park, Little Prairie Picnic Area, Cameron Creek, 5-9 Jun 1980, by IM Smith, IMS800053B • British Columbia, Canada: 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Atnarko River, at campground, 28 Jul 1983, by IM Smith & AB Smith, IMS830054A • 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Atnarko Slough, beside Highway 20, west of Youngs Creek Picnic Area, 4 Aug 1983, by IM Smith, IMS830064 • 1 ♀ from Bella Coola Valley, Tweedsmuir Provincial Park, Belarko, Atnarko River, 24-26 Jul 1983, by IM Smith, IMS830049B • 1 ♀ and 2 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Hotnarko River, at end of Atnarko tote road, 27 Jul 1983, by IM Smith, IMS830052 • 2 ♀ and 3 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Hotnarko River, at end of Atnarko tote road, 31 Jul 1983, by IM Smith, IMS830059A & IMS830059B • 3 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Youngs Creek Picnic Area, Atnarko Slough, 24-27 Jul 1983, by IM Smith, IMS830048A • 2 ♀ and 1 ♂ from Fernie, Lizard Creek, beside Highway 3, 1.8 km west of Fernie Mountain Provincial Park, 16 Aug 2012, by IM Smith, IMS120073 • 1 ♀ and 1 ♂ from Tweedsmuir Provincial Park, Youngs Creek, beside Highway 20, between Heckman Pass & Bella Coola Valley, 5 Aug 1983, by IM Smith, IMS830065 • 2 ♀ and 2 ♂ from Vancouver Island, Caycuse, Nixon Creek, 8 Jul 1976, by IM Smith, IMS760197 & IMS760198 • 1 ♀ and 2 ♂ from Vancouver Island, beside Highway 4, 35.6 kilometers east of Pacific Rim Road, 9 Jul 1976, by IM Smith, IMS760206 • 1 ♀ and 1 ♂ from Vancouver Island, Lake Cowichan, Cowichan River, above Skutz Falls, 9 Jul 1979, by IM Smith, IMS790035 • 1 ♀ and 1 ♂ from Vancouver Island, Lake Cowichan, Skutz Falls, Skutz Creek, near Cowichan River, 9 Jul 1979, by IM Smith, IMS790036A & IMS790036B • 1 ♀ and 2 ♂ from Vancouver Island, Lake Cowichan, South Shore Road, north of Mesachie Lake, Robertson River, 4 Jul 1976, by IM Smith, IMS760183A • 1 ♀ from Vancouver Island, Lake Cowichan, South Shore Road, north of Mesachie Lake, tributary of Robertson River, 4-10 Jul 1976, by IM Smith, IMS760182 • 1 ♀ and 2 ♂ from Vancouver Island, Lost Shoe Creek, beside Highway 4, 1.3 kilometers east of Pacific Rim Road, 9 Jul 1976, by IM Smith, IMS760202 • 1 ♀ and 2 ♂ from Vancouver Island, Malahat, Goldstream Provincial Park, Goldstream River, 26 Jun 1979, by IM Smith, IMS790028A • 1 ♂ from Vancouver Island, spring run beside North Shore Road, 1.7 kilometers north of Lake Cowichan, 7 Jun 1979, by IM Smith, IMS790008A • 3 ♀ and 2 ♂ from Vancouver Island, North Shore Road, 3.2 kilometers south of Youbou, 4 Jul 1976, by IM Smith, IMS760190 • 1 ♀ from Vancouver Island, Youbou, Shaw Creek, North Shore Road, 4.3 kilometers south of north end of Cowichan Lake, 8 Jul 1976, by IM Smith, IMS760196 • 1 ♀ from Vancouver Island, spring run beside South Shore Road, 2.3 kilometers north of Lake Cowichan, 6 Jun 1979, by IM Smith, IMS790007 • 1 ♀ and 1 ♂ from Vancouver Island, Ucluelet, beside Highway 4, 16.6 kilometers east of Pacific Rim Road, 18-19 Jul 1979, by IM Smith, IMS790047 • California, USA: 1 ♀ and 1 ♂ from Humboldt County, Prairie Creek State Park, Prairie Creek, 12 Jul 1964, by H Habeeb, HH640021 • 1 ♀ from Humboldt County, Trinidad, stream beside Patrick Point Road, near Bishop Pine Lodge, 7 Aug 1987, by IM Smith, IMS870134 • 1 ♀ from Inyo County, Inyo National Forest, Bishop Creek (37°17'23"N, 118°33'14"W), 2 Sep 2013, by JR Fisher, JRF 13-0902-003 • 1 ♂ from Mendocino County, Cottaneva Creek, beside Route 1, 21.8 kilometers southwest of Route 101, 5 Aug 1987, by IM Smith, IMS870129A • 2 ♀ and 3 ♂ from Mendocino County, small stream at beach access road, off Route 1, 2.6 kilometers south of Westport, 5 Aug 1987, by IM Smith, IMS870128A • 1 ♀ from Mono County, Humboldt-Toiyabe National Forest, Leavitt Creek (38°18'40"N, 119°34'49"W), 31 Aug 2013, by JR Fisher, JRF 13-0831-004 • 1 ♂ from Monterey County, Los Padres National Forest, Salmon Creek (35°48'57"N, 121°21'29"W), 6 Sep 2013, by JR Fisher, JRF 13-0906-003 • 2 ♀ and 2 ♂ from Monterey County, Nacimiento River, beside Nacimiento-Ferguson Road at Nacimiento campground, 30 Jul 1987, by IM Smith, IMS870120A • 2 ♀ and 2 ♂ from Monterey County, Salmon Creek, beside Route 1, south of Gorda, 28 Jul 1987, by IM Smith, IMS870114A • 1 ♀ from Nevada County, beside Route 89, north of Hobart Mills, 13 Jun 1976, by IM Smith, IMS760109 • 1 ♀ and 2 ♂ from San Bernardino County, Claremont, Mt Baldy, 3.5 kilometers above Mt Baldy Village, 24 Jul 1987, by IM Smith, IMS870107 • 1 ♀ from San Bernardino County, Claremont, Mt Baldy, San Antonio Falls, above Monker Flats, 24 Jul 1987, by IM Smith, IMS870105 • 3 ♀ and 2 ♂ from San Bernardino County, Claremont, Mt Baldy, stream below San Antonio Falls, above Monker Flats, 24 Jul 1987, by IM Smith, IMS870106 • 2 ♀ and 2 ♂ from Shasta County, Battle Creek, beside Route 44, 5.6 kilometers west of Viola, 10 Aug 1987, by IM Smith, IMS870139A • 1 ♀ and 1 ♂ from Trinity County, small cascading trickle beside Route 36, 5.2 kilometers west of Forest Glen Station, 6 Aug 1987, by IM Smith, IMS870132 • 3 ♂ from Tulare County, Stony Creek at Stony Creek Picnic Area, east of Sequoia National Park, 1 Aug 1987, by IM Smith, IMS870124A • 1 ♀ and 1 ♂ from Ventura County, Ojai, North Fork of Ventura River, beside Route 33, just above Wheeler Gorge, 25-26 Jul 1987, by IM Smith, IMS870109A • Idaho, USA: 1 ♀ from Custer County, Challis National Forest, Stanley Creek (44°15'12"N, 115°0'19"W), 30 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0730-005 • 1 ♂ from Custer County, Salmon River (44°12'31"N, 114°55'51"W), 29 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0729-003 • 2 ♀ and 3 ♂ from Lemhi County, North Fork of Salmon River, beside Route 93, 15 kilometers north of North Fork, 1 Jul 1985, by IM Smith, IMS850062 • Montana, USA: 2 ♀ from Flathead County, stream beside Route 487 near north end of Whitefish Lake, 29 Jun 1985, by IM Smith, IMS850057 • 3 ♀ and 4 ♂ from Lake County, stream beside Route 83, 39.5 kilometers north of Condon, 30 Jun 1985, by IM Smith, IMS850059A • 1 ♀ from Missoula County, Lolo National Forest, Lolo Creek (46°46'7"N, 114°27'53"W), 7 Aug 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0807-003 • 1 ♂ from Ravalli County, Bitterroot National Forest, East Fork Bitterroot River (45°51'40"N, 114°1'46"W), 3 Aug 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0803-005 • Oregon, USA: 1 ♀ and 1 ♂ from Benton County, Marys Peak near Philomath, Parker Creek, 27-28 Jun 1983, by IM Smith & AB Smith, IMS830006 • 1 ♀ from Clackamas County, Rhododendron Pioneer Tollgate campground, Zigzag River, 27 Jun 1976, by IM Smith, IMS760164 • 1 ♀ and 2 ♂ from Curry County, Port Orford, Butler Bar campground, Elk River, 25 Jun 1976, by IM Smith, IMS760162 • 1 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, Coquille River, 2 Jul 1983, by IM Smith, IMS830016 • 1 ♀ and 1 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, 2 Jul 1983, by IM Smith, IMS830017 • 2 ♀ and 5 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, Brush Creek, 1 Jul 1983, by IM Smith, IMS830012 • 4 ♀ and 4 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, Brush Creek, beside Route 1, 3 Jul 1983, by IM Smith, IMS830020A & IMS830020B • 1 ♀ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, 1 Jul 1983, by IM Smith, IMS830013 • 1 ♀ from Curry County, Rogue River National Forest, Elk River (42°42'46"N, 124°18'41"W), 13 Aug 2013, by JR Fisher, JRF 13-0813-003 • 2 ♀ and 2 ♂ from Curry County, Siskiyou National Forest, road 33 between Powers and Agness, North Fork of Foster Creek, 2 Jul 1983, by IM Smith, IMS830019 • 1 ♀ from Curry County, Sixes, Sixes River, beside road at mouth of Edson Creek, 4 Jul 1983, by IM Smith, IMS830021A • 1 ♀ from Grant County, Prairie City, Lunch Creek, beside Route 26. east of Dixie Pass, 17-20 Jun 1976, by IM Smith, IMS760125 • 1 ♀ and 1 ♂ from Grant County, Prairie City, Strawberry Forest Camp, Strawberry Creek, 17-20 Jun 1976, by IM Smith, IMS760126 • 2 ♀ from Lane County, Gate Creek (44°8'48"N, 122°34'20"W), 11 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0811-001 • 3 ♀ and 1 ♂ from Marion County, Marion Forks Riverside campground, North Fork of Santiam River, 22 Jun 1976, by IM Smith, IMS760145 • 3 ♀ and 4 ♂ from Multnomah County, Columbia River Scenic Highway, Horsetail Falls, 27 Jun 1983, by IM Smith, IMS830005 • 1 ♂ from Tillamook County, Siuslaw National Forest, Alder Creek (45°9'27"N, 123°47'60"W), 6 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0806-002 • Washington, USA: 2 ♀ from Lewis County, Gifford Pinchot National Forest (46°39'49"N, 121°41'11"W), 23 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0723-005 • 2 ♀ from Mason County, Olympic National Forest, Cabin Creek (47°35'44"N, 123°7'39"W), 22 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0722-004 • 2 ♀ from Snohomish County, Mount Baker National Forest, Marten River (48°4'19"N, 121°36'24"W), 28 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0728-002 • Wyoming, USA: 1 ♂ from Johnson County, Bighorn Mountains, Clear Creek, west of Buffalo Mosier Gulch Picnic Area, 28 Jul 2012, by IM Smith, IMS120041 • 1 ♀ and 1 ♂ from Johnson County, Crazy Woman Creek, beside Route 16, 14.9 kilometers west of road to South Fork campground, 18 Aug 1987, by IM Smith, IMS870158A • 1 ♀ and 2 ♂ from Washakie County, Ten Sleep, Ten Sleep Creek, beside Route 16, 4 kilometers west of Route 435, 18 Aug 1987, by IM Smith, IMS870157 • 2 ♀ and 1 ♂ from Washakie County, Ten Sleep Creek, Ten Sleep Wigwam Rearing Station, 26 Jul 2012, by IM Smith, IMS120044.

Type deposition.

Holotype (♀) and paratype (1 ♀) deposited in the CNC.

Diagnosis.

Torrenticola ellipsoidalis are similar to other members of the Ellipsoidalis Group ( T. multiforma , T. occidentalis , and T. leviathan ), in being among the largest Torrenticola in the west (dorsum length ♀ = 700-885; ♂ = 665-850), although T. sierrensis are also large (dorsum length ♀ = 700-880; ♂ = 590-735) but can easily be distinguished from the Ellipsoidalis Group by being circular instead of ellipsoid or rectangular (dorsum length/width = 1.17-1.28 in T. sierrensis , 1.30-1.67 in Ellipsoidalis Group). T. ellipsoidalis can be differentiated from T. multiforma by having stockier subcapitular rostra (length/width = 1.8-2.1 in T. ellipsoidalis , 2.5-2.8 in T. multiforma ). T. ellipsoidalis can be differentiated from T. occidentalis (only known from females) by having a longer medial suture (40-57.5 in T. ellipsoidalis , 20 in T. occidentalis ) and by having stockier anterio-lateral platelets (length/width = 2.00-2.39 in T. ellipsoidalis , 2.54 in T. occidentalis ). T. ellipsoidalis can be differentiated from T. leviathan by having less elongate pedipalpal tibiae (length/width = 2.6-3.3 in T. ellipsoidalis , 3.4-4.2 in T. leviathan ) and stockier anterio-medial platelets (length/width ♀ = 1.43-1.72 in T. ellipsoidalis , 1.94-2.14 in T. leviathan ; ♂ = 1.53-2.00 in T. ellipsoidalis , 2.15 in T. leviathan ).

Re-description.

Female (Figure 65) (n = 8) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum - (765-885 (800) long; 520-605 (540) wide) rectangular and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets (127.5-147.5 137.5) long; 80-97.5 (80) wide). Anterio-lateral platelets (207.5-235 (217.5) long; 90-115 (92.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 390-470 (410)). Dorsal plate proportions: dorsum length/width 1.41-1.64 (1.48); dorsal width/distance between Dgl-4 1.27-1.40 (1.32); anterio-medial platelet length/width 1.43-1.72 (1.72); anterio-lateral platelet length/width 2.00-2.39 (2.35); anterio-lateral/anterio-medial length 1.48-1.77 (1.58).

Gnathosoma - Subcapitulum (285-315 (310) long (ventral); 194-219 (215) long (dorsal); 145-165 (165) tall) colorless. Rostrum (115-127.5 (117.5) long; 57.5-62.5 (60) wide) short and conical. Chelicerae (261-289 long) with curved fangs (61-74 long). Subcapitular proportions: ventral length/height 1.82-2.07 (1.88); rostrum length/width 1.84-2.09 (1.96). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5-42.5 (40) long); femur (101.25-107.5 (102.5) long); genu (70-77.5 (77.5) long); tibia (80-90 (85) long; 26.25-27.5 (27.5) wide); tarsus (17.5-21.25 (20) long). Palpomere proportions: femur/genu 1.32-1.46 (1.32); tibia/femur 0.76-0.85 (0.83); tibia length/width 2.91-3.27 (3.09).

Venter - (885-1000 (935) long; 605-700 (605) wide) colorless. Gnathosomal bay (157.5-180 (177.5) long; 80-105 (85) wide). Cxgl-4 subapical. Medial suture (40-57.5 (47.5) long). Genital plates (201.25-222.5 (205) long; 167.5-195 (172.5) wide). Additional measurements: Cx-1 (308-337.5 (335) long (total); 122-162.5 (162.5) long (medial)); Cx-3 (393-440 (405) wide); anterior venter (210-237.5 (225) long). Ventral proportions: gnathosomal bay length/width 1.50-2.12 (2.09); anterior venter/genital field length 1.01-1.13 (1.10); anterior venter length/genital field width 1.13-1.30 (1.30); anterior venter/medial suture 3.83-5.94 (4.74).

Male (Figure 66) (n = 6) with characters of the genus with following specifications.

Dorsum - (725-850 long; 450-565 wide) rectangular and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets (122.5-165 long; 72.5-95 wide). Anterio-lateral platelets (195-230 long; 85-107.5 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 350-460). Dorsal plate proportions: dorsum length/width 1.38-1.67; dorsal width/distance between Dgl-4 1.22-1.43; anterio-medial platelet length/width 1.53-2.00; anterio-lateral platelet length/width 2.07-2.36; anterio-lateral/anterio-medial length 1.39-1.67.

Gnathosoma - Subcapitulum (280-290 long (ventral); 196-203.75 long (dorsal); 138.75-155 tall) colorless. Rostrum (102.5-115 long; 52.5-60 wide) short and conical. Chelicerae (263-280 long) with curved fangs (60-74 long). Subcapitular proportions: ventral length/height 1.87-2.04; rostrum length/width 1.86-2.02. Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35-41.25 long); femur 92.5-100 long); genu (65-72.5 long); tibia (72.5-80 long; 25-30 wide); tarsus (17.5-20 long). Palpomere proportions: femur/genu 1.31-1.46; tibia/femur 0.74-0.84; tibia length/width 2.64-3.1.

Venter - (840-980 long; 469-653 wide) colorless. Gnathosomal bay (147.5-177.5 long; 77.5-90 wide). Cxgl-4 subapical. Medial suture (70-90 long). Genital plates (177.5-236.25 long; 131.25-162.5 wide). Additional measurements: Cx-1 (283-345 long (total); 117-167.5 long (medial)); Cx-3 (348-432.5 wide); anterior venter (245-270 long). Ventral proportions: gnathosomal bay length/width 1.69-2.15; anterior venter/genital field length 1.14-1.44; anterior venter length/genital field width 1.66-1.96; anterior venter/medial suture 2.72-3.71.

Immatures unknown.

Etymology.

Marshall (1943) presumably named the specific epithet ( ellipsoidalis ) after the elongate body of this species, as she wrote, "the body is an ellipse."

Distribution.

Western (Figure 64). T. ellipsoidalis was previously recorded from Martis Creek and Gibbon River, Wyoming ( Marshall 1943); and from Torch River, Saskatchewan (Quaglia & Conroy 1984). We expand the range into most of western North America. However, T. ellipsoidalis is not known from the southwest.

Remarks.

Torrenticola ellipsoidalis groups other members of the Miniforma Complex with high support and most specimens are 0-3% different in COI sequence. This is higher sequence variability than in many species hypotheses presented herein. However, given the topology in the COI tree (Figure 10, 16) and morphological similarity, it seems apparent that the variability represents a continuum across a large distribution, rather than isolated species. An exception is that a single specimen (DNA#1930) is 2.9-3.6% different from the rest. This specimen is indistinguishable from other specimens and is collected from the same location. We do not find evidence to propose it as a separate species and therefore include it within T. ellipsoidalis .

In all analyses, T. ellipsoidalis groups with T. multiforma and T. regalis , which are greater than 10% different from each other. Based upon overall similarity, body size, and distribution, we place this species within the Ellipsoidalis Identification Group.

Upon examining the types of T. ellipsoidalis and T. rectiforma , all characters for both species overlap with members of only one clade in our analyses. Furthermore, the main character Habeeb (1974) used to differentiate T. rectiforma from T. ellipsoidalis was body size, which is known to be a highly variable character. Therefore, we synonymize T. rectiforma as the junior synonym of T. ellipsoidalis .