Torrenticola multiforma Habeeb, 1974

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 multiforma Habeeb, 1974

Torrenticola multiforma Habeeb, 1974: 4.

Material examined.

LECTOTYPE (1 ♀): from USA, California, Humboldt County, Prairie Creek Redwoods State Park, Prairie Creek, 12 Jul 1964, by Habeeb, HH640021.

PARALECTOTYPE (1 ♂): from USA, California, Humboldt County, Prairie Creek Redwoods State Park, Prairie Creek, 12 Jul 1964, by Habeeb, HH640021.

OTHER MATERIAL (95 ♀; 84 ♂): Arizona, USA: 1 ♀ from Cochise County, Chiricahua Mountains, Turkey Creek at Sycamore campground, east of Sunizona, 15 Jul 1987, by IM Smith, IMS870091 • 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 • 2 ♀ and 2 ♂ 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 • 3 ♀ and 2 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Belarko, Atnarko River, 24-26 Jul 1983, by IM Smith, IMS830049B & IMS830049C • 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Hotnarko River, at end of Atnarko tote road, 27 Jul 1983, by IM Smith, IMS830052 • 5 ♀ and 6 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Hotnarko River, at end of Atnarko tote road, 31 Jul 1983, by IM Smith, IMS830059A & IMS830059B • 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Youngs Creek Picnic Area, Atnarko Slough, 24-27 Jul 1983, by IM Smith, IMS830048A • 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Youngs Creek Picnic Area, tributary of Atnarko Slough, 2 Aug 1983, by IM Smith, IMS830062C • 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 ♂ from McClinchy River, beside Highway 20, west of Kleena Kleene, 5 Aug 1983, by IM Smith, IMS830068A • 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 • 5 ♀ from Vancouver Island, Caycuse, Nixon Creek, 8 Jul 1976, by IM Smith, IMS760197 & IMS760198 • 3 ♀ and 1 ♂ 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, 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 1 ♂ from Vancouver Island, Malahat, Goldstream Provincial Park, Goldstream River, 26 Jun 1979, by IM Smith, IMS790028A • 1 ♀ and 2 ♂ from Vancouver Island, North Shore Road, 3.2 kilometers south of Youbou, 4 Jul 1976, by IM Smith, IMS760190 • 3 ♂ 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-fed pool beside South Shore Road, 2.3 kilometers south of Caycuse, 26 Jul 1979, by IM Smith, IMS790052 • 1 ♀ from Vancouver Island, spring run beside South Shore Road, 2.3 kilometers north of Lake Cowichan, 6 Jun 1979, by IM Smith, IMS790007 • California, USA: 1 ♀ and 2 ♂ from Del Norte County, Six Rivers National Forest, Middle Fork Smith River (41°51'20"N, 123°53'10"W), 15 Aug 2013, by JR Fisher, JRF 13-0815-002 • 1 ♀ and 2 ♂ from Mendocino County, Cottaneva Creek, beside Route 1, 21.8 kilometers southwest of Route 101, 5 Aug 1987, by IM Smith, IMS870129A • 4 ♀ and 4 ♂ 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 ♀ and 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 ♀ and 1 ♂ from Mono County, Humboldt-Toiyabe National Forest, West Walker River (38°21'59"N, 119°28'55"W), 31 Aug 2013, by JR Fisher, JRF 13-0831-003 • 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 • 1 ♀ and 2 ♂ from Monterey County, Nacimiento River, beside Nacimiento-Ferguson Road at Nacimiento campground, 30 Jul 1987, by IM Smith, IMS870120A • 2 ♀ from Monterey County, Pfeiffer State Park, Big Sur River (36°14'42"N, 121°46'43"W), 4 Sep 2013, by JR Fisher, JRF 13-0904-004 • 2 ♀ and 2 ♂ from Monterey County, Salmon Creek, beside Route 1, south of Gorda, 28 Jul 1987, by IM Smith, IMS870114A • 1 ♀ and 1 ♂ from Nevada County, beside Route 89, north of Hobart Mills, 13 Jun 1976, by IM Smith, IMS760109 • 1 ♀ and 1 ♂ from Nevada County, Tahoe National Forest, Sagehen Creek (39°26'2"N, 120°12'17"W), 26 Aug 2013, by JR Fisher, JRF 13-0826-006 • 1 ♀ from Plumas County, beside Route 89, north of Greenville, 14 Jun 1976, by IM Smith, IMS760113 • 1 ♀ from Siskiyou County, Klamath National Forest, Dead Cow Creek (41°57'16"N, 122°52'23"W), 16 Aug 2013, by JR Fisher, JRF 13-0816-002 • 1 ♀ and 1 ♂ from Tulare County, Stony Creek at Stony Creek Picnic Area, east of Sequoia National Park, 1 Aug 1987, by IM Smith, IMS870124A • 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 Blaine County, Salmon River, beside Route 75 between Obsidian & Galena Summit, 3 Jul 1985, by IM Smith, IMS850067 • 1 ♂ from Blaine County, Sawtooth National Forest, Deer River (43°32'49"N, 114°26'31"W), 29 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0729-001 • 1 ♀ and 1 ♂ from Custer County, Basin Creek campground, Basin Creek, beside Route 75 between Sunbeam & Stanley, 2 Jul 1985, by IM Smith, IMS850066 • 1 ♀ from Custer County, Salmon River at picnic area, beside Route 93 north of Morgan Creek, 2 Jul 1985, by IM Smith, IMS850064 • 1 ♀ and 1 ♂ from Lemhi County, North Fork of Salmon River, beside Route 93, 15 kilometers north of North Fork, 1 Jul 1985, by IM Smith, IMS850062 • 1 ♀ from Lemhi County, Salmon National Forest, Indian Creek (45°24'12"N, 114°10'10"W), 2 Aug 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0802-001 • Nevada, USA: 2 ♀ and 2 ♂ from Elko County, Lamoille, Lamoille Creek near car park, at end of Lamoille Canyon Road, 12 Aug 1987, by IM Smith, IMS870143A • 2 ♀ and 3 ♂ from Humboldt County, Paradise Valley, Dutch John Creek beside road, 8.3 kilometers north of Hinkey Summit, 11 Aug 1987, by IM Smith, IMS870142A • New Mexico, USA: 2 ♀ and 2 ♂ from Catron County, Whitewater Creek, Glenwood Whitewater Picnic Area, 5 May 2012, by IM Smith, IMS120005 • 1 ♀ and 1 ♂ from Catron County, Glenwood Whitewater Creek at Whitewater Creek Picnic Area, 12 Jul 1987, by IM Smith, IMS870084 • 1 ♀ and 1 ♂ from Lincoln County, Eagle Creek, beside Route 532, just above Route 37, 8 Jul 1987, by IM Smith, IMS870077 • 1 ♀ and 1 ♂ from San Miguel County, Pecos, Pecos River, beside Route 63 at Dalton campground, 6 Jul 1987, by IM Smith, IMS870071 • 1 ♀ and 1 ♂ from San Miguel County, stream near Route 63 in Dalton Canyon, 6 Jul 1987, by IM Smith, IMS870073 • 1 ♀ from Sante Fe County, Sante Fe, Tesuque Creek, beside Route 22, south of Tesuque, 5 Jul 1987, by IM Smith, IMS870070 • Montana, USA: 1 ♀ from Lake County, stream beside Route 83, 39.5 kilometers north of Condon, 30 Jun 1985, by IM Smith, IMS850059A • 1 ♀ from Ravalli County, Beaverhead 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 12-0803-004 • 1 ♂ from Ravalli County, Bitterroot National Forest, Piquette Creek (45°51'24"N, 114°11'37"W), 6 Aug 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0806-002 • 1 ♀ and 2 ♂ from Ravalli County, Medicine Springs, Spring Gulch campground, East Fork of Bitterroot River, beside Route 93, 1 Jul 1985, by IM Smith, IMS850060 • Oregon, USA: 1 ♀ from Clackamas County, Rhododendron Pioneer Tollgate campground, Zigzag River, 27 Jun 1976, by IM Smith, IMS760164 • 1 ♀ and 2 ♂ from Coos County, Siskiyou National Forest, Coal Creek, Road 33 between Powers & Agness, 2 Jul 1983, by IM Smith, IMS830015 • 2 ♀ and 2 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, 2 Jul 1983, by IM Smith, IMS830017 • 1 ♀ and 1 ♂ from Curry County, Port Orford, Butler Bar campground, Elk River, 25 Jun 1976, by IM Smith, IMS760162 • 1 ♀ and 6 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, Brush Creek, beside Route 1, 3 Jul 1983, by IM Smith, IMS830020A & IMS830020B • 2 ♀ and 1 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, Brush Creek, 1 Jul 1983, by IM Smith, IMS830012 • 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 • 1 ♀ and 1 ♂ from Curry County, Siskiyou National Forest, road 33 between Powers and Agness, North Fork of Foster Creek, 2 Jul 1983, by IM Smith, IMS830019 • 2 ♀ and 1 ♂ from Curry County, Sixes, Sixes River, beside road at mouth of Edson Creek, 4 Jul 1983, by IM Smith, IMS830021A • 1 ♀ and 1 ♂ from Multnomah County, Columbia River Scenic Highway, Horsetail Falls, 27 Jun 1983, by IM Smith, IMS830005 • Utah, USA: 2 ♀ and 1 ♂ from Summit County, North Fork of Provo River, beside Route 150, west of Provo River Overlook, 14 Aug 1987, by IM Smith, IMS870148 • 1 ♂ from Utah County, Uinta National Forest, Hobble Creek Road, upstream on Right Fork (40°10'9"N, 111°28'36"W), 22 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0722-001 • 1 ♀ and 1 ♂ from Wasatch County, Provo River, beside Route 150 at Upper Provo River Bridge Picnic Area, 14 Aug 1987, by IM Smith, IMS870149 • 2 ♀ and 1 ♂ from Wasatch County, Wasatch-Cache National Forest, Holden Fork (40°47'19"N, 110°53'2"W), 23 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0723-003 • Washington, USA: 2 ♀ from Jefferson County, Rocky Brook (47°43'11"N, 122°56'32"W), 22 Jul 2013, by JC O’Neill & WA Nelson, JNOW 13-0722-002 • 1 ♀ and 1 ♂ from Lewis County, Gifford Pinchot National Forest, Snake Creek (46°38'52"N, 121°43'8"W), 23 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0723-006 • Wyoming, USA: 2 ♀ and 1 ♂ from Carbon County, Medicine Bow Mountains, Medicine Bow River, west of Arlington, 30 Jul 2012, by IM Smith, IMS120045 • 1 ♀ and 1 ♂ from Fremont County, Wind River Mountains, Sinks Canyon, Popo Agie River, 1 Aug 2012, by IM Smith, IMS120049.

Type deposition.

Types (1 ♀; 1 ♂) deposited in the CNC.

Diagnosis.

Torrenticola multiforma are similar to other members of the Ellipsoidalis Group ( T. ellipsoidalis , 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. multiforma are best differentiated from other members of the Ellipsoidalis Group by having more elongate subcapitular rostra (length/width = 2.5-2.8 in T. multiforma , 1.84-2.27 in other Ellipsoidalis Group).

Re-description.

Female (Figure 159) (n = 6) with characters of the genus with following specifications.

Dorsum - (700-850 (730) long; 500-615 (550) wide) ellipsoid with highly variable coloration, colorless to orange to purple without distinct pattern. Anterio-medial platelets (132.5-150 (145) long; 65-77.5 (65) wide). Anterio-lateral platelets (195-230 (195) long; 77.5-92.5 (82.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 380-465 (400)). Dorsal plate proportions: dorsum length/width 1.31-1.41 (1.33); dorsal width/distance between Dgl-4 1.26-1.38 (1.38); anterio-medial platelet length/width 1.94-2.27 (2.23); anterio-lateral platelet length/width 2.34-2.87 (2.36); anterio-lateral/anterio-medial length 1.34-1.57 (1.34).

Gnathosoma - Subcapitulum (340-385 long (ventral); 255-284 long (dorsal); 145-172.5 tall) colorless. Rostrum (132.5-152.5 long; 50-55 wide). Chelicerae (336-395 long) with curved fangs (62-79 long). Subcapitular proportions: ventral length/height 2.19-2.46; rostrum length/width 2.5-2.8. Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5-45 long); femur (115-127.5 long); genu (75-90 long); tibia (87.5-95 long; 27.5-32.5 wide); tarsus (17.5-20 long). Palpomere proportions: femur/genu 1.42-1.53; tibia/femur 0.71-0.78; tibia length/width 2.77-3.18.

Venter - (840-1010 long; 588-679 wide) colorless. Gnathosomal bay (165-217.5 long; 82.5-100 wide). Cxgl-4 subapical. Medial suture (17.5-32.5 long). Genital plates (210-242.5 long; 197.5-222.5 wide). Additional measurements: Cx-1 (295-380 long (total); 123-170 long (medial)); Cx-3 (359-441 wide); anterior venter (175-212.5 long). Ventral proportions: gnathosomal bay length/width 1.89-2.44; anterior venter/genital field length 0.79-0.92; anterior venter length/genital field width 0.86-1.05; anterior venter/medial suture 5.83-11.14.

Male (Figure 160) (n = 6) with characters of the genus with following specifications.

Dorsum - (665-790 long; 470-580 wide) ellipsoid with highly variable coloration, colorless to orange to purple without distinct pattern. Anterio-medial platelets (125-165 long; 60-77.5 wide). Anterio-lateral platelets (185-222.5 long; 75-87.5 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 365-450). Dorsal plate proportions: dorsum length/width 1.30-1.41; dorsal width/distance between Dgl-4 1.27-1.38; anterio-medial platelet length/width 1.97-2.33; anterio-lateral platelet length/width 2.46-2.67; anterio-lateral/anterio-medial length 1.35-1.48.

Gnathosoma - Subcapitulum (310-380 long (ventral); 236-284) long (dorsal); 137.5-170 tall) colorless. Rostrum (120-150 long; 45-60 wide). Chelicerae (309-382.5 long) with curved fangs (63-75 long). Subcapitular proportions: ventral length/height 2.16-2.34; rostrum length/width 2.50-2.79. Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40-45 long); femur (102.5-122.5 long); genu (70-87.5 long); tibia (80-95 long; 26.25-32.5 wide); tarsus (17.5-20 long). Palpomere proportions: femur/genu 1.40-1.50; tibia/femur 0.73-0.79; tibia length/width 2.91-3.33.

Venter - (805-940 long; 479-653 wide) colorless. Gnathosomal bay (172.5-197.5 long; 75-100 wide). Cxgl-4 subapical. Medial suture (62.5-82.5 long). Genital plates (175-190 long; 140-155 wide). Additional measurements: Cx-1 (308-375 long (total); 127-180 long (medial)); Cx-3 (351-421 wide); anterior venter (225-270 long). Ventral proportions: gnathosomal bay length/width 1.92-2.30; anterior venter/genital field length 1.29-1.46; anterior venter length/genital field width 1.57-1.81; anterior venter/medial suture 3.27-4.00.

Immatures unknown.

Etymology.

Habeeb (1974) did not specify an etymology for the specific epithet ( multiforma ). However, he notes that the males "vary a lot in size." Thus, we suspect the specific epithet refers to the variable body size males of this species (multus, L. many; forma, L. form).

Distribution.

Western (Figure 158). T. multiforma was previously known only from Prairie Creek, California ( Habeeb 1974), but we expand its range to include most of western North America.

Remarks.

Torrenticola multiforma groups other members of the Miniforma Complex with high support and 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) and morphological similarity, it seems apparent that the variability represents a continuum across a large distribution, rather than isolated species.

In all analyses, T. multiforma groups with T. ellipsoidalis 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.

This species hypothesis is supported by phylogenetic affinity, biogeography, high divergence between species (3-15%), and by the morphological characters outlined in the diagnosis.