Revision of torrent mites (Parasitengona, Torrenticolidae, Torrenticola) of the United States and Canada: 90 descriptions, molecular phylogenetics, and a key to species

Abstract

The descriptive biology of torrent mites (Parasitengona: Torrenticolidae: Torrenticola) of North America (north of Mexico) is investigated using integrative methods. Material examined includes approximately 2,300 specimens from nearly 500 localities across the United States and Canada, and a few collections in Mexico and Central America. Species hypotheses are derived from a phylogenetic analysis of the barcoding region of cytochrome c oxidase subunit 1 (COI) for 476 specimens and supported with morphology and biogeography. Relationships between species are examined with a combined analysis of COI and two expansion regions (D2-3) of the large ribosomal subunit (28S rDNA) for 57 specimens. All previously described species from the US and Canada are examined. Our results indicate the need to synonymize four species: T. mercedensis (Marshall, 1943) is a junior synonym of T. sierrensis (Marshall, 1943); T. rectiforma Habeeb, 1974 is a junior synonym of T. ellipsoidalis (Marshall, 1943); T. neoconnexa Habeeb, 1957 is a junior synonym of T. magnexa Habeeb, 1955; and T. esbelta Cramer, 1992 is a junior synonym of T. boettgeri KO Viets, 1977. We describe 66 new species and re-describe all previously described regional species. Our findings indicate that total diversity of Torrenticola in the United States and Canada comprises 90 species, 57 known from the east and 33 from the west. We organize these species into four species complexes that include 13 identification groups. An additional 13 species do not fit within an identification group. The southern Appalachians are suspected to contain the highest concentration of remaining undescribed diversity. A key is provided to all known species in the US and Canada.

Keywords: Acari; Acariformes; Hydrachnidiae; integrative taxonomy; turbo-taxonomy; water mites.

Figure 1.

Diversity of Torrenticola in Ouachita National Forest, Arkansas. Species from top-left to bottom-right: T. irapalpa sp. n., T. interiorensis sp. n., T. biscutella sp. n. female, T. biscutella sp. n. male, T. pearsoni sp. n., T. trimaculata Fisher & Dowling 2016, T. unimaculata sp. n., T. larvata Cherri et al. 2016, T. solisorta sp. n. Note that, for a given locality, many characters necessary for identification can be seen under low magnification. For example, species in the bottom row can be readily differentiated using color pattern. In contrast, species in the top row have similar color patterns; however, even they can be readily identified: T. irapalpa have dorsal glandularia (Dgl-4) closer together than the other species and T. biscutella have fused anterio-lateral platelets and strong sexual dimorphism (neither character is present in T. interiorensis). Only the identification of T. interiorensis is tentative based on this image, because T. neoanomala (not pictured), which are also found in this locality, can only be differentiated from T. interiorensis by comparing precise measurements of the anterio-lateral platelets, necessitating examining slide-prepared material under higher magnification. Also, note that some species can appear differently than what is depicted in species descriptions. For example, although T. pearsoni sp. n. has a nearly colorless cuticle, it can appear dark due to gut contents, as shown here. Image is a composite of stacked images from a Samsung GS7 and edited in Adobe Photoshop.

Figure 2.

Summary of all sampling localities. 327 red dots represent material examined; 14 blue diamonds represent published accounts where material was not available for direct examination. Each of the 341 localities represent a generalized locality, not an individual collection event, as multiple events often occured in close proximity. In total, this map represents nearly 500 collection events. Note the absence of material from the Great Plains (e.g., Oklahoma, Kansas, Nebraska, and the Dakotas) and Alaska; these are areas that were sampled for water mites, but Torrenticola were not found.

Figure 3.

Terminology for coloration: blue – bluish-purple – purple-pink – reddish-purple – orange. Species depicted from top-left to bottom-right: T. gorti; T. erectirostra; T. biscutella; T. neoanomala; T. nigroalba; T. gorti; T. skvarlai; T. magnexa, and T. delicatexa.

Figure 4.

Dorsal measurements: A dorsal length B dorsal width C distance between dorsal glandularia, Dgl-4 D anterio-lateral platelet length E anterio-lateral platelet width F anterio-medial platelet length G anterio-medial platelet width. T. multiforma Habeeb, 1974 depicted.

Figure 5.

Gnathosomal measurements. Palpomeres – A trochanter length B femur length C genu length D tibia length E tibia width F tarsus length. Chelicerae – G cheliceral base length H fang length. Subcapitulum – I ventral length J dorsal length K rostrum length L rostrum width M height. T. multiforma Habeeb, 1974 depicted.

Figure 6.

Venter measurements: A ventral length B ventral width C anterior venter D coxa-I total length E coxa-I medial length F medial suture length G coxa-III width H gnathosomal bay length I gnathosomal bay width J genital field length K genital field width. T. multiforma Habeeb, 1974 depicted.

Figure 7.

Bayesian inferred cladogram for combined analysis (28S+COI) of North American Torrenticola: species complexes and identification groups. Four well-supported complexes are organized into 13 groups that are helpful for identification. Note that 1) not all identification groups are monophyletic (in quotation marks); 2) not all species are placed in a complex; and 3) some species are not placed within a group, reflecting their identification difficulty. Numbers are DNA identification numbers. Dots denote posterior probably >95%.

Figure 8.

Bayesian inferred phylogram for combined analysis (28S+COI) of North American Torrenticola: distributions of 57 species. Pink represents lineages distributed in eastern North America and blue represents lineages distributed in western North America (gray branches represent lineages distributed in both regions). Species marked with an asterisk are primarily eastern but have distributions extending into the west. Note that larger lineages contain species distributed in either the east or west, but not both. Numbers are DNA identification numbers. Scale bar indicates 0.1 substitutions per site. Dots denote posterior probably >95%.

Figure 9.

Bayesian inferred circular phylograms of North American Torrenticola, COI only. Scale bars indicate 1.0 substitutions per site. A Overall tree depicting the four complexes recovered in combined analysis. Dots denote posterior probability >95% B Species distributions; branch colors correspond to regions of North America depicted on the right (eastern, pink; western teal). Asterisks denote eastern species that extend into the west. Note that lineages tend to contain species distributed in either the east or west, but not both. Detail of this figure is presented in Figures 10–18.

Figure 10.

Raptor Complex (part I). These species are eastern, although T. irapalpa extends further west. Note that two species (T. skvarlai and T. bondi) do not fit into identification groups. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Raptor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Raptor Complex (in part) from B. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state or territory abbreviation.

Figure 11.

Raptor Complex (part II). Species of the Raptor Complex are eastern, but the species that do not fit into the complex (T. rala; T. sharkeyi; T. raptoroides; & T. skvarlai) are western. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Raptor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Raptor Complex (in part) from B. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state or territory abbreviation.

Figure 12.

Miniforma Complex (part I). All species are western. Note that the phylogenetic structure in the Miniforma Group corresponds to biographic regions (Rockies, Pacific Ranges, southwest). A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Miniforma Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Miniforma Complex (in part) from B C Detail of Miniforma Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 13.

Miniforma Complex (part II). All species are western. One species (T. regalis) does not fit within a species group. Note that T. ellipsoidalis and T. multiforma are widespread in the west (C). A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Miniforma Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Miniforma Complex (in part) from B C Detail of Miniforma Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 14.

Rusetria Complex (part I). All species are eastern, but note the distribution of T. sellersorum extends into New Mexico and Saskatchewan. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Rusetria Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Rusetria Complex (in part) from B C Detail of Rusetria Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 15.

Rusetria Complex (part II). All species are eastern. Relationships among species groups remain unresolved, except for three species (T. dunni; T. pollani; and T. shubini), which form a well-supported clade in all analyses. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Rusetria Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Rusetria Complex (in part) from B C Detail of Rusetria Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 16.

Rusetria Complex (part III). All species are western. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Rusetria Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Rusetria Complex (in part) from B. D Detail of Rusetria Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 17.

Tricolor Complex (part I). All species are eastern. These are some of the most recognizable of all Torrenticola, namely the elongate T. projector and a clade of species with diagnostic color patterns (T. trimaculata, T. unimaculata, and T. tricolor). Note that three species (T. pearsoni, T. hoosieri, and T. projector) are repeated in Figure 18. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Tricolor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Tricolor Complex (in part) from B C Detail of Tricolor Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 18.

Tricolor Complex (part II). Most species of the Tricolor Complex are eastern, but one lineage is western: T. sierrensis, widespread; and T. olliei, north Pacific coast. The most apomorphic Torrrenticola (T. dimorpha) is recovered here in all analyses. Note that three species (T. pearsoni, T. hoosieri, and T. projector) are repeated in Figure 17. A Species guide (overview of 28S+COI analysis from Figure 7–8) B Overview of Tricolor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Tricolor Complex (in part) from B C Detail of Tricolor Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 19.

Torrenticola anoplopalpa sp. n. distribution.

Figure 20.

Torrenticola anoplopalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 21.

Torrenticola arktonyx sp. n. distribution.

Figure 22.

Torrenticola arktonyx sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 23.

Torrenticola arktonyx sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 24.

Torrenticola biscutella sp. n. distribution.

Figure 25.

Torrenticola biscutella sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 26.

Torrenticola biscutella sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 27.

Torrenticola bittikoferae distribution.

Figure 28.

Torrenticola bittikoferae male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 29.

Torrenticola boettgeri distribution. Blue star represents type locality (Viets 1977a); blue diamond represents additional published record (Cramer 1992, as T. esbelta comb. n.); and red circles represent new records and material examined.

Figure 30.

Torrenticola boettgeri female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 31.

Torrenticola boettgeri male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 32.

Torrenticola bondi sp. n. distribution.

Figure 33.

Torrenticola bondi sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 34.

Torrenticola bondi sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 35.

Torrenticola caerulea sp. n. distribution.

Figure 36.

Torrenticola caerulea sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 37.

Torrenticola caerulea sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 38.

Torrenticola cardia sp. n. distribution.

Figure 39.

Torrenticola cardia sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 40.

Torrenticola cardia sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 41.

Torrenticola copipalpa sp. n. distribution.

Figure 42.

Torrenticola copipalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 43.

Torrenticola copipalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 44.

Torrenticola daemon sp. n. distribution.

Figure 45.

Torrenticola daemon sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 46.

Torrenticola daemon sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 47.

Torrenticola danielleae sp. n. distribution.

Figure 48.

Torrenticola danielleae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 49.

Torrenticola danielleae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 50.

Torrenticola delicatexa distribution.

Figure 51.

Torrenticola delicatexa female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 52.

Torrenticola delicatexa male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 53.

Torrenticola dentirostra sp. n. distribution.

Figure 54.

Torrenticola dentirostra sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 55.

Torrenticola dentirostra sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 56.

Torrenticola dimorpha sp. n. distribution.

Figure 57.

Torrenticola dimorpha sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 58.

Torrenticola dimorpha sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 59.

Torrenticola dolichodactyla sp. n. distribution.

Figure 60.

Torrenticola dolichodactyla sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 61.

Torrenticola dunni sp. n. distribution.

Figure 62.

Torrenticola dunni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 63.

Torrenticola dunni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 64.

Torrenticola ellipsoidalis distribution. Red dots indicate material examined. Blue crosses indicate additional previous records (Marshall 1943; Quaglia & Conroy 1984).

Figure 65.

Torrenticola ellipsoidalis female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 66.

Torrenticola ellipsoidalis male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 67.

Torrenticola elongata sp. n. distribution.

Figure 68.

Torrenticola elongata sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 69.

Torrenticola elongata sp. n. male: A dorsal plates, coloration added B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 70.

Torrenticola elusiva sp. n. distribution.

Figure 71.

Torrenticola elusiva sp. n. female: A dorsal plates, coloration added B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 72.

Torrenticola erectirostra sp. n. distribution.

Figure 73.

Torrenticola erectirostra sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 74.

Torrenticola erectirostra sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 75.

Torrenticola feminellai sp. n. distribution.

Figure 76.

Torrenticola feminellai sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 77.

Torrenticola feminellai sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 78.

Torrenticola flangipalpa sp. n. distribution.

Figure 79.

Torrenticola flangipalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 80.

Torrenticola flangipalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 81.

Torrenticola folkertsae sp. n. distribution.

Figure 82.

Torrenticola folkertsae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 83.

Torrenticola folkertsae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 84.

Torrenticola glomerabilis sp. n. distribution.

Figure 85.

Torrenticola glomerabilis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 86.

Torrenticola glomerabilis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 87.

Torrenticola gnoma sp. n. distribution.

Figure 88.

Torrenticola gnoma sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 89.

Torrenticola gnoma sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 90.

Torrenticola gorti sp. n. distribution.

Figure 91.

Torrenticola gorti sp. n. female, Tennessee specimen depicted except for A (top): A dorsal plates, note variation in color between Tennessee specimens (bottom) and elsewhere (top) B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 92.

Torrenticola gorti sp. n. male, Tennessee specimen depicted: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 93.

Torrenticola hoosieri sp. n. distribution.

Figure 94.

Torrenticola hoosieri sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum, note damaged rostrum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 95.

Torrenticola hoosieri sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 96.

Torrenticola indistincta distribution. Red dots indicate material examined. Blue diamonds indicate previously published reports. Green “X” indicates previously published report (Young 1969) that requires verification of identification.

Figure 97.

Torrenticola indistincta female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 98.

Torrenticola indistincta male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 99.

Torrenticola interiorensis sp. n. distribution.

Figure 100.

Torrenticola interiorensis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 101.

Torrenticola interiorensis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 102.

Torrenticola irapalpa sp. n. distribution.

Figure 103.

Torrenticola irapalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 104.

Torrenticola irapalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 105.

Torrenticola ivyae sp. n. distribution.

Figure 106.

Torrenticola ivyae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 107.

Torrenticola ivyae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 108.

Torrenticola karambita sp. n. distribution.

Figure 109.

Torrenticola karambita sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 110.

Torrenticola karambita sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 111.

Torrenticola keesdavidsi distribution. Blue star represents type locality (Cramer 1992); and red circles represent new records and material examined.

Figure 112.

Torrenticola keesdavidsi female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 113.

Torrenticola keesdavidsi male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 114.

Torrenticola kittatinniana distribution.

Figure 115.

Torrenticola kittatinniana female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 116.

Torrenticola kittatinniana male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 117.

Torrenticola kringi sp. n. distribution.

Figure 118.

Torrenticola kringi sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 119.

Torrenticola kringi sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 120.

Torrenticola kurtvietsi distribution. Blue star represents type locality (Cramer 1992); red circles represent new record and material examined.

Figure 121.

Torrenticola kurtvietsi female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 122.

Torrenticola kurtvietsi male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 123.

Torrenticola lamellipalpis distribution. Blue star represents type locality (K.O. Viets 1977b); and red circles represent new records and material examined.

Figure 124.

Torrenticola lamellipalpis female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 125.

Torrenticola lamellipalpis male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 126.

Torrenticola larvata distribution.

Figure 127.

Torrenticola larvata female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 128.

Torrenticola larvata male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 129.

Torrenticola leviathan sp. n. distribution.

Figure 130.

Torrenticola leviathan sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 131.

Torrenticola leviathan sp. n. male (teneral): A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 132.

Torrenticola longitibia sp. n. distribution.

Figure 133.

Torrenticola longitibia sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 134.

Torrenticola magnexa distribution.

Figure 135.

Torrenticola magnexa female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 136.

Torrenticola magnexa male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 137.

Torrenticola malarkeyorum sp. n. distribution.

Figure 138.

Torrenticola malarkeyorum sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 139.

Torrenticola malarkeyorum sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 140.

Torrenticola manni sp. n. distribution.

Figure 141.

Torrenticola manni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 142.

Torrenticola manni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 143.

Torrenticola microbiscutella sp. n. distribution.

Figure 144.

Torrenticola microbiscutella sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 145.

Torrenticola microbiscutella sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 146.

Torrenticola miniforma distribution.

Figure 147.

Torrenticola miniforma female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 148.

Torrenticola miniforma male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 149.

Torrenticola mjolniri sp. n. distribution.

Figure 150.

Torrenticola mjolniri sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 151.

Torrenticola mjolniri sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 152.

Torrenticola mohawk sp. n. distribution.

Figure 153.

Torrenticola mohawk sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 154.

Torrenticola mohawk sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 155.

Torrenticola mulleni sp. n. distribution. Red dots represent material examined. Blue dot represents previously-published record (Young 1969), reported as T. indistincta, but likely representing T. mulleni.

Figure 156.

Torrenticola mulleni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 157.

Torrenticola mulleni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 158.

Torrenticola multiforma distribution.

Figure 159.

Torrenticola multiforma female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 160.

Torrenticola multiforma male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 161.

Torrenticola neoanomala distribution.

Figure 162.

Torrenticola neoanomala female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 163.

Torrenticola neoanomala male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 164.

Torrenticola nigroalba distribution.

Figure 165.

Torrenticola nigroalba female: A dorsal plates, note variation B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 166.

Torrenticola nigroalba male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 167.

Torrenticola nortoni sp. n. distribution.

Figure 168.

Torrenticola nortoni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 169.

Torrenticola nortoni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 170.

Torrenticola occidentalis distribution.

Figure 171.

Torrenticola occidentalis female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 172.

Torrenticola oliveri sp. n. distribution.

Figure 173.

Torrenticola oliveri sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 174.

Torrenticola oliveri sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 175.

Torrenticola olliei sp. n. distribution.

Figure 176.

Torrenticola olliei sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 177.

Torrenticola olliei sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 178.

Torrenticola oregonensis sp. n. distribution.

Figure 179.

Torrenticola oregonensis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 180.

Torrenticola oregonensis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 181.

Torrenticola pacificensis sp. n. distribution.

Figure 182.

Torrenticola pacificensis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 183.

Torrenticola pacificensis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 184.

Torrenticola pearsoni sp. n. distribution.

Figure 185.

Torrenticola pearsoni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 186.

Torrenticola pearsoni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 187.

Torrenticola pendula sp. n. distribution.

Figure 188.

Torrenticola pendula sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 189.

Torrenticola pendula sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 190.

Torrenticola pinocchio sp. n. distribution.

Figure 191.

Torrenticola pinocchio sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 192.

Torrenticola pinocchio sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 193.

Torrenticola pollani sp. n. distribution.

Figure 194.

Torrenticola pollani sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 195.

Torrenticola pollani sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 196.

Torrenticola priapus sp. n. distribution.

Figure 197.

Torrenticola priapus sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 198.

Torrenticola priapus sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 199.

Torrenticola projector distribution.

Figure 200.

Torrenticola projector female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 201.

Torrenticola projector male: A dorsal plates B venter (legs removed) C subcapitulum Dpedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 202.

Torrenticola pulchra sp. n. distribution.

Figure 203.

Torrenticola pulchra sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 204.

Torrenticola pulchra sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 205.

Torrenticola racupalpa sp. n. distribution.

Figure 206.

Torrenticola racupalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 207.

Torrenticola racupalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 208.

Torrenticola rala distribution. Red dots represent material examined; blue diamonds represent previous records (Cook 1980); blue star represents holotype locality (Cook 1980).

Figure 209.

Torrenticola rala female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 210.

Torrenticola rala male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 211.

Torrenticola raptor sp. n. distribution.

Figure 212.

Torrenticola raptor sp. n. female: A dorsal plates, note color variation B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 213.

Torrenticola raptor sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 214.

Torrenticola raptoroides sp. n. distribution.

Figure 215.

Torrenticola raptoroides sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 216.

Torrenticola raptoroides sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 217.

Torrenticola reduncarostra sp. n. distribution.

Figure 218.

Torrenticola reduncarostra sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 219.

Torrenticola reduncarostra sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 220.

Torrenticola regalis sp. n. distribution.

Figure 221.

Torrenticola regalis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 222.

Torrenticola regalis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 223.

Torrenticola robisoni sp. n. distribution.

Figure 224.

Torrenticola robisoni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 225.

Torrenticola rockyensis sp. n. distribution.

Figure 226.

Torrenticola rockyensis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 227.

Torrenticola rockyensis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 228.

Torrenticola rufoalba distribution.

Figure 229.

Torrenticola rufoalba female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 230.

Torrenticola rufoalba male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 231.

Torrenticola sellersorum sp. n. distribution.

Figure 232.

Torrenticola sellersorum sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 233.

Torrenticola sellersorum sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 234.

Torrenticola sharkeyi sp. n. distribution.

Figure 235.

Torrenticola sharkeyi sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 236.

Torrenticola sharkeyi sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 237.

Torrenticola shubini sp. n. distribution.

Figure 238.

Torrenticola shubini sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 239.

Torrenticola shubini sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 240.

Torrenticola sierrensis distribution.

Figure 241.

Torrenticola sierrensis female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 242.

Torrenticola sierrensis male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 243.

Torrenticola skvarlai sp. n. distribution.

Figure 244.

Torrenticola skvarlai sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 245.

Torrenticola skvarlai sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 246.

Torrenticola solisorta sp. n. distribution.

Figure 247.

Torrenticola solisorta sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 248.

Torrenticola solisorta sp. n. male: A dorsal plates, note the faint orange coloration, which is a rare character state (most specimens are much brighter), not a male character B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 249.

Torrenticola tahoei distribution.

Figure 250.

Torrenticola tahoei female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 251.

Torrenticola tahoei male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 252.

Torrenticola tricolor distribution.

Figure 253.

Torrenticola tricolor female: A dorsal plates, note color variation B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 254.

Torrenticola tricolor male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

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.

Figure 258.

Torrenticola tysoni sp. n. distribution.

Figure 259.

Torrenticola tysoni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 260.

Torrenticola tysoni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 261.

Torrenticola ululata sp. n. distribution.

Figure 262.

Torrenticola ululata sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 263.

Torrenticola ululata sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 264.

Torrenticola unimaculata sp. n. distribution.

Figure 265.

Torrenticola unimaculata sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 266.

Torrenticola unimaculata sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 267.

Torrenticola ventura distribution.

Figure 268.

Torrenticola ventura female: A dorsal plates, note color variation B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 269.

Torrenticola ventura male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 270.

Torrenticola walteri sp. n. distribution.

Figure 271.

Torrenticola walteri sp. n. female: A dorsal plates, note color variation B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 272.

Torrenticola walteri sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 273.

Torrenticola welbourni sp. n. distribution.

Figure 274.

Torrenticola welbourni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 275.

Torrenticola whitneyae sp. n. distribution.

Figure 276.

Torrenticola whitneyae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 277.

Torrenticola whitneyae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 278.

Torrenticola wiedenmanni distribution.

Figure 279.

Torrenticola wiedenmanni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 280.

Torrenticola wiedenmanni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.