Revision of torrent mites (Parasitengona, Torrenticolidae, Torrenticola) of the United States and Canada: 90 descriptions, molecular phylogenetics, and a key to species Author Fisher, J. Ray Author Fisher, Danielle M. Author Skvarla, Michael J. Author Nelson, Whitney A. Author Dowling, Ashley P. G. text ZooKeys 2017 701 1 496 http://dx.doi.org/10.3897/zookeys.701.13261 journal article http://dx.doi.org/10.3897/zookeys.701.13261 1313-2970-701-1 23BDD7CE1C7E4D2092A8ED47267579FD 23BDD7CE1C7E4D2092A8ED47267579FD Torrenticola trimaculata Fisher, 2015 Torrenticola trimaculata Fisher et al. (2015) : 71, 89. Material examined. HOLOTYPE (♀): USA, Arkansas, Madison County, Withrow Springs State Park, War Eagle Creek ( 36°8'59.3"N , 93°44'26.94"W ), 27 Jul 2011, by IM Smith, IMS110034. PARATYPES (49 ♀; 37 ♂): Arkansas, USA: 1 ♂ (ALLOTYPE) from Madison County, Withrow Springs State Park, War Eagle Creek ( 36°8'59.3"N , 93°44'26.94"W ), 27 Jul 2011, by IM Smith, IMS110034 * 2 ♀ and 3 ♂ from Madison County, Withrow Springs State Park, War Eagle Creek ( 36°8'59.3"N , 93°44'26.94"W ), 27 Jul 2011, by IM Smith, IMS110034 * 1 ♀ from Marion County, Crooked Creek ex. Northern hogsucker ( Hypentelium nigricans ) ( 36°15'9.9"N , 94°26'25.8"W ), 22 Jul 2014, by CT McAllister * 3 ♀ and 2 ♂ from Montgomery County, Ouachita National Forest, Ouachita River ( 34°34'53.20"N , 93°53'0.16"W ), 5 Oct 2007, by AJ Radwell, & HW Robison, AJR070300A * 8 ♀ and 5 ♂ from Montgomery County, Ouachita National Forest, South Fork of Ouachita River, 29 Jul 2011, by AJ Radwell, & B Crump, AJR110302 * 2♀ and 1 ♂ from Montgomery County, Ouachita National Forest, Ouachita River, 27 Aug 2011, by AJ Radwell, AJR110307 * 4 ♀ and 4 ♂ from Montgomery County, Ouachita National Forest, South Fork of Ouachita River, 29 Jul 2011, by IM Smith, IMS110040 * 1 ♀ from Montgomery County, Caddo River, 29 Jul 2011, by IM Smith, IMS110037 * 1 ♂ from Newton County, Ozark National Forest, Mill Creek ( 36°3'42.12"N , 93°8'7.62"W ), 20 Jun 2012, by TD Edwards, TDE 12-0620-010 * 2 ♀ and 2 ♂ from Newton County, Ozark National Forest, Little Buffalo River, 2 Sep 2012, by TD Edwards, TDE 12-0902-003 * 1 ♂ from Newton County, Buffalo National River, Whiteley Creek ( 35°59'28.14"N , 93°23'57.24"W ), 23 May 2012, by TD Edwards, TDE 12-0523-002 * Illinois, USA: 2 ♀ and 1 ♂ from Union County, Clear Creek ( 37°33'N , 89°23'W ), 13 Sep 1991, by IM Smith, IMS910036A * Indiana, USA: 1 ♀ from Wayne County ( 39°51'13"N , 85°8'4"W ), 24 Jul 2014, by MJ Skvarla, MS 14-0731-001 * Georgia, USA: 1 ♀ from Chattooga County, Johns Creek ( 34°34'N , 80°5'W ), 4 Jul 1990, by IM Smith, IMS900076 * Kentucky, USA: 1 ♀ and 2 ♂ from McCreary County, Rock Creek ( 36°42'N , 84°36'W ), 8 Jul 1990, by IM Smith, IMS900082B * Michigan, USA: 2 ♀ and 2 ♂ from Barry County, Thornapple River ( 42°39'N , 85°17'W ), 29 Jul 1959, by DR Cook, DRC590034 * Missouri, USA: 2 ♀ and 1 ♂ from Crawford County, Huzzah Creek, 23 Jul 2011, by IM Smith, IMS110029 * New York, USA: 3 ♀ and 1 ♂ from St. Lawrence County, Canton ( 44°35'N , 75°10'W ), 15 May 1986, by BP Smith, BPS860508 * 1 ♀ from USA, New York, Delaware Co., Roscoe ( 41°55'N , 74°54'W ), 11 June 1988, by PW Schefter and R MacCulloch, IMS880110 * Nova Scotia, Canada: 1 ♀ from Victoria County, Baddeck River ( 44°52'N , 61°5'W ), 18 Jul 1981, by IM Smith, IMS810082 * Ontario, Canada: 4 ♀ and 2 ♂ from Grey County, Saugeen River (44°10'N, 80°49'W), 9 Jun 1989, by IM Smith, IMS890028A * 1 ♀ from Madoc ( 44°30'N , 77°28'W ), 4 May 1980, by IM Smith, IMS800003A * 1 ♂ from Renfrew County, Madawaska River ( 45°21'N , 76°40'W ), 25 May 1980, by IM Smith, IMS800012 * 1 ♀ and 1 ♂ from Lanark County, Mississippi River ( 45°3'N , 76°23'W ), 6 Oct 1983, by IM Smith and CJ Hill, IMS830093A * Texas, USA: 1 ♀ from Gillespie County, Fredericksburg, Pedernales River ( 30°14'42"N , 98°54'50"W ), 30 May 1998, by IM Smith, IMS980029 * Virginia, USA: 1 ♀ and 1 ♂ from Scott County, North Fork of Holston River ( 36°39'N , 82°28'W ), 7 Jul 1990, by IM Smith, IMS0900080 * 2 ♀ and 4 ♂ from Alleghany County, Potts Creek ( 37°44'N , 80°2'W ), 13 Jul 1990, by IM Smith, IMS900091B * 1 ♀ and 1 ♂ from Bath County, Jackson River ( 38°8'N , 79°46'W ), 16 Jul 1990, by IM Smith, IMS900100 * West Virginia, USA: 2 ♀ from Pendleton County, North Fork of South Branch of Potomac River ( 39°0'N , 79°22'W ), 17 Jul 1990, by IM Smith, IMS900104. Type deposition. Holotype (♀), allotype (♂), and other paratypes (44 ♀; 32 ♂) deposited in the CNC; other paratypes (4 ♀; 4 ♂) deposited in ACUA. Diagnosis. Torrenticola trimaculata are similar to other members of the Tricolor Complex ( T. bittikoferae , T. hoosieri , T. larvata , T. pearsoni , T. olliei , T. sierrensis , T. tricolor , T. unimaculata , T. cardia , T. kringi , T. dimorpha , and T. mohawk ) in having a short, conical rostrum. T. trimaculata can be differentiated from all Torrenticola , including other members of the Tricolor Complex, by having a distinct dorsal pattern. Individuals are reported in two distinct morphs primarily based upon ventral coloration. T. trimaculata are most similar to other members of the Tricolor Complex that have bold patterning ( T. larvata , T. unimaculata , T. tricolor , T. cardia , T. kringi , and T. mohawk ). T. trimaculata can be further differentiated from T. larvata by being rounder (dorsum length/width = 1.2-1.37 in T. trimaculata , 1.41-1.57 in T. larvata ); Dgl-4 closer to the muscle scars (dorsal width/distance between Dgl-4 = 1.49-1.69 in T. trimaculata , 1.18-1.35 in T. larvata ); and a stockier rostrum (length/width = 1.91-2.22 in T. trimaculata , 2.32-2.53 in T. larvata ). T. trimaculata can be further differentiated from T tricolor by having shorter genital field (♀ = 157.5-185 in T. trimaculata , 187.5-210 in T. tricolor ; ♂ = 120-140 in T. trimaculata , 145-170 in T. tricolor ) and a stockier rostrum (length/width ♀ = 1.91-2.1 in T. trimaculata , 2.14-2.39 in T. tricolor ; ♂ = 2.05-2.22 in T. trimaculata , 2.37-2.5 in T. tricolor ). T. trimaculata can be further differentiated from T. unimaculata by Dgl-4 closer to the muscles scars (dorsal width/distance between Dgl-4 = 1.49-1.69 in T. trimaculata , 1.23-1.41 in T. unimaculata ) and females with shorter medial suture (♀ = 17.5-27.5 in T. trimaculata , 40-47.5 in T. unimaculata ). T. trimaculata can be further differentiated from T. cardia by having Dgl-4 further from the edge of the dorsum (dorsum width/distance between Dgl-4 = 1.5-1.7 in T. trimaculata , 1.15-1.4 in T. cardia ) and having a stockier rostrum (length/width ♀ = 1.191-2.10 in T. trimaculata , 2.24-2.50 in T. cardia ; ♂ = 2.05-2.22 in T. trimaculata , 2.27-2.47 in T. cardia ). T. trimaculata can be differentiated from T. kringi by having a stockier rostrum (length/width = 1.9-2.3 in T. trimaculata , 2.6-3.2 in T. kringi ). T. trimaculata can be differentiated from T. mohawk by having a shorter anterior venter (♀ = 157-180 in T. trimaculata , 187-200 in T. mohawk , ♂ = 230-260 in T. trimaculata , 290-308 in T. mohawk ). Additionally, female T. trimaculata can be differentiated from female T. mohawk by having longer pedipalpal tibiae (♀ = 90-97 in T. trimaculata , 77-88 in T. mohawk ). Re-description (amended from Fisher et al. 2015 ). Female (Figure 256) (n = 6) (holotype measurements in parentheses when available) with characters of the genus with following specifications. Dorsum - (570-725 (725) long; 455-550 (550) wide) circular to ellipsoid with bluish-purple to navy blue coloration in three distinct spots, one anteriorly and two posteriorly, and orange medially. Anterio-medial platelets (125-145 (145) long; 60-72.5 (67.5) wide). Anterio-lateral platelets (162.5-180 (180) long; 80-85 (82.5) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and the dorsum edge (distance between Dgl-4 295-335 (325)). Dorsal plate proportions: dorsum length/width 1.20-1.32 (1.32); dorsal width/distance between Dgl-4 1.49-1.69 1.69); anterio-medial platelet length/width 1.93-2.19 (2.15); anterio-lateral platelet length/width 2.00-2.19 (2.18); anterio-lateral/anterio-medial length 1.23-1.30 (1.24). Gnathosoma - Subcapitulum (250-280 (280) long (ventral); 202-252 (210) long (dorsal); 115-135 (130) tall) colorless or with bluish-purple to navy blue coloration. Rostrum (97.5-110 (107.5) long; 47.5-55 (52.5) wide) short and conical. Chelicerae (261-326 (270) long) with curved fangs (48-62 (55) long). Subcapitular proportions: ventral length/height 2.04-2.17 (2.15); rostrum length/width 1.91-2.10 (2.05). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5-42.5 (42.5) long); femur (97.5-107.5 (103.75) long); genu (65-77.5 (70) long); tibia (90-96.25 (92.5) long; 27.5-30 (27.5) wide); tarsus (22.5-25 (22.5) long). Palpomere proportions: femur/genu 1.39-1.50 (1.48); tibia/femur 0.88-0.92 (0.89); tibia length/width 3.17-3.36 (3.36). Venter - (615-840 (840) long; 533-700 (700) wide) colorless or with variable amount of bluish-purple to navy blue coloration. Gnathosomal bay (130-145 (130) long; 80-90 (80) wide). Cxgl-4 subapical. Medial suture (17.5-27.5 (20) long). Genital plates (157.5-185 (180) long; 152.5-185 (180) wide). Additional measurements: Cx-1 (258-306.75 (260) long (total); 109-154 (145) long (medial)); Cx-3 (328-390 (390) wide); anterior venter (157.5-180 (180) long). Ventral proportions: gnathosomal bay length/width 1.49-1.75 (1.63); anterior venter/genital field length 0.90-1.03 (1.00); anterior venter length/genital field width 1.00-1.14 (1.14); anterior venter/medial suture 6.09-9.29 (9.00). Male (Figure 257) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications. Dorsum - (530-595 (595) long; 390-440 (435) wide) circular to ellipsoid with bluish-purple to navy blue coloration in three distinct spots, one anteriorly and two posteriorly, and orange medially. Anterio-medial platelets (107.5-130 (120) long; 55-67.5 (62.5) wide). Anterio-lateral platelets (70-87.5 (75) long; 70-87.5 (75) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and the dorsum edge (distance between Dgl-4 250-285 (270)). Dorsal plate proportions: dorsum length/width 1.30-1.37 (1.37); dorsal width/distance between Dgl-4 1.51-1.61 (1.61); anterio-medial platelet length/width 1.92-2.00 (1.92); anterio-lateral platelet length/width 1.91-2.23 (2.23); anterio-lateral/anterio-medial length 1.29-1.46 (1.40). Gnathosoma - Subcapitulum (207.5-242.5 (230) long (ventral); 175-215 (194.75) long (dorsal); 92.5-112.5 (102.5) tall) colorless or with bluish-purple to navy blue coloration. Rostrum (87.5-100 (93.75) long; 40-47.5 (42.5) wide) short and conical. Chelicerae (232-286 (251) long) with curved fangs (40-53 (49) long). Subcapitular proportions: ventral length/height 2.16-2.43 (2.24); rostrum length/width 2.05-2.22 (2.21). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (32.5-40 (32.5) long); femur (85-95 (93.75) long); genu (60-70 (65) long); tibia (75-82.5 (75) long; 25-27.5 (27.5) wide); tarsus (20-25 (22.5) long). Palpomere proportions: femur/genu 1.34-1.44 (1.44); tibia/femur 0.80-0.88 (0.80); tibia length/width 2.73-3.00 (2.73). Venter - (588-719 (715) long; 431-571 (568) wide) colorless or with variable amount of bluish-purple to navy blue coloration. Gnathosomal bay (82.5-112.5 (102.5) long; 62.5-77.5 (77.5) wide). Cxgl-4 subapical. Medial suture (87.5-105 (105) long). Genital plates (120-140 (127.5) long; 92.5-105 (95) wide). Additional measurements: Cx-1 (216-297 (261) long (total); 130-168 (152) long (medial)); Cx-3 (287-372 (349) wide); anterior venter (230-260 (255) long). Ventral proportions: gnathosomal bay length/width 1.32-1.55 (1.32); anterior venter/genital field length 1.79-2.04 (2.00); anterior venter length/genital field width 92.5-105 (95); anterior venter/medial suture 2.43-2.63 (2.43). Immatures unknown. Etymology. Fisher et al. (2015) named the specific epithet ( trimaculata ) in reference to the dorsal coloration of three dark spots (tres, L. three; macula, L. spot). Distribution. Eastern North America (Figure 255). Figure 255. Torrenticola trimaculata distribution. Figure 256. Torrenticola trimaculata female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm . Figure 257. Torrenticola trimaculata male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm . Remarks. Torrenticola trimaculata group with other members of the Tricolor Complex with high support in all analyses. All specimens of this species are less than 2% different in COI sequence from each other and are greater than 7% from sister species. In all analyses, T. trimaculata groups with two other species which also have dorsal spots: T. tricolor and T. unimaculata . This clade represents some of the most distinctive of all Torrenticola . This species hypothesis is supported by low COI divergence within the species (0-2%), high divergence between species (3-15%), and the morphological characters outlined in the diagnosis. Fisher et al. (2015) reported two color morphs based primarily upon presence of ventral coloration. Morph I, with a dark venter, was only known from the Interior Highlands and all eastern specimens were reported as Morph II, with a colorless venter. However, we examined additional specimens from across eastern North America and found colored individuals mixed within samples of uncolored individuals. We remain unsure what controls color in Torrenticola , but with several other species exhibiting great color variation (e.g., T. tricolor , T. gorti ), we do not consider it useful to continue with the "morph" concept in this species.