Torrenticola trimaculata Fisher, 2015

Fisher, J. R., Fisher, D. M., Nelson, W. A., O'Neill, J. C., Skvarla, M. J., Ochoa, R., Bauchan, G. R., Radwell, A. J. & Dowling, A. P. G., 2015, Torrenticola trimaculata n. sp. (Parasitengona: Torrenticolidae), a three-spotted water mite from eastern North America: taxonomic history, species delimitation, and survey of external morphology, Acarologia 55 (1), pp. 71-116 : 89-109

Torrenticola trimaculata Fisher n. sp.

(http://species-id.net/wiki/ Torrenticola _trimacula ta)

LSID — urn:lsid:zoobank.org:act:BE021914-89AE-4DE7-A0DC-18CF80BA78AF

IMAGO GENERAL FEATURES. Color variable across two distinct color morphs that do not coexist within a stream ( Fig. 2 , 9 ). For both morphs, the integument is yellowish, with a central red structural coloration on the dorsal plate. Both morphs also have pigmentation in the form of three dark spots on the dorsal plate that do not extend beyond the area of primary sclerotization; this pigmentation is destroyed during clearing ( Fig. 3 ). This adequately describes the less pigmented morph (Morph-2: Fig. 2B, D, F ), although some specimens express light pigmentation on the genital plates ( Fig. 2B ) and hind coxae (males) ( Fig. 2D ), and size and shape of the dorsal spots varies considerably ( Fig. 9B, D ). The more pigmented morph (Morph-1: Fig. 2A, C, E ) is marked by the following: 1) area of primary sclerotization on the venter is darkly pigmented; 2) the three dorsal spots are larger; 3) entire gnathosoma, including pedipalps, is pigmented; 4) dorsal pigmentation can extend to the anteriomedial platelets; and 5) legs are pinkish-orange. Although noticeable variability is present in the size and shape of dorsal spots, it is much less extreme in Morph-1 ( Fig. 9A, C ). As with Morph-2, Morph- 1 exhibits considerable variability in size and shape of dorsal spots, as well as the extent and vibrancy of all pigmentation.

Sexes are somewhat dimorphic ( Fig. 10 ). Sexual dimorphism consistent with most/all Torrenticola include the following: 1) body slightly smaller in males and consistently ovoid; females larger and vary from round to ovoid; 2) genital fields comparatively small and rectangular in males; female genital plates are larger and trapezoidal; 3) medial suture of males long–comparable to the length of the genital field; female medial suture short–about as long as wide. Additional sexually dimorphic characters that are not shared by most Torrenticola include the following: 1) hind coxae not extending anteriorly beyond hind leg insertions in males but do in females; 2) hind coxae not extending far posteriorly beyond the genital field; female hind coxae extend beyond the genital field by approximately half the length of the genital field; and 3) rostrum directed normally (forward) in females, but is directed downwards in males.

FEMALE (n=49) (holotypic measurements in parentheses when available) with characters described in generic diagnosis and general features, with following specifications.

Gnathosoma ( Fig. 11-14 ) — Subcapitulum [250 – 341 (313) ventral length; 185 – 255 (239) dorsal length; 110 – 169 (142) tall] posterior edge nearly vertical, ventral bend depth slight [4 – 20 (5)], and with short rostrum [95 – 133 (129) long] that is directed forwards. Two pairs of adoral setae rim the rostral opening ( Fig. 11-12 ). Chelicerae [230 – 329 (310) long; 15 – 31 (17) high] unmodified with strongly curved fangs [33 – 74 (56) long]. Each fang with lateral and medial teeth presumably used to anchor to prey after puncturing ( Fig. 12 B-D). Pedipalps [248 – 338 (311) long] with dentate ventrodistal projections medially on femora and mediocentrally on genua ( Fig. 11 , 13 ). These projections vary across individuals in thickness, length, and shape. Further, their appearance can vary according to their position in a given slide preparation, sometimes even appearing tuburculate/edentate. Regardless, they are never lamellate as in the T. serratipalpis -group identified by Goldschmidt (2007). Trochanters [24 – 38 (35) long; 28 – 43 (33) wide] with one dorso-distal fringed spatulate seta (fss). Femora [80 – 123 (109) long; 48 – 68 (60) wide] with one long simple seta (lss) associated with the ventral projection and six dorsal setae as follows: proximally one short simple grooved seta (sgs); two central fss, and three distal fss (two medial; one lateral). Genua [55 – 85 (77) long; 40 – 56 (51) wide] shorter than femora with one lss associated with the ventral projection, one short sgs laterally, and four dorsal setae as follows: one central lss, and three setae distally as follows: one sgs medially, one lss medially, and one lss laterally. Tibiae [73 – 124 (107) long; 19 – 37 (34) wide] subequal in length to femora, with two short, spiny tubercles midventrally that are edentate and associated with 3-4 lss ( Fig. 11 , 13E ). Mid-dorsally, there are two sss (one proximo-lateral; one disto-medial). Distally, there is one lss doro-centrally; two lss dorso-medially; two lss dorso-laterally; one lss laterally; and one large, grooved, spine-like seta dorso-medially ( Fig. 11 , 14 A-B). Tarsi [20 – 27 (25) long; 12 – 16 (16) wide] are accompanied by four tarsal claws, with the bottom two paired ( Fig. 14B ), thus appearing as three claws in most slide preparations. Ventrally, there are 2-3 short bifurcating setae (sbs) and dorsally there are three lss ( Fig. 14 A-B).

Dorsum ( Fig. 15-19 ) — [560 – 765 (723) long; 415 – 596 (543) wide] round to ovoid; armored with a central dorsal plate that is divided into an area of primary sclerotization [435 – 530 (477) long; 365 – 521 (440) wide] and an area of secondary sclerotization posteriorly [extends dorsal plate length by 87 – 176 for a total dorsal plate length of 522 – 706 (653)]. Specimens that have recently emerged from the imagochrysalis (i.e., teneral) have not yet developed the area of secondary sclerotization ( Fig. 6 ). The dorsal plate is bordered by ten platelets: two anterio-medials [109 – 178 (130) long; 53 – 86 (69) wide]; two anterio-laterals [135 – 220 (178) long; 65 – 101 (86) wide]; and a posterior ring of six smaller platelets in a striated membranous fold (partially visible in Fig. 18 ). Dgl-4 slightly lateral to Dgl-5 and usually in the area of secondary sclerotization, but occasionally near edge of primary sclerotization.

Eyes are apparently paired and located within sclerotized capsules on the margin of the anteriomedial platelets and dorsal covering of the gnathosoma ( Fig. 16 , 17 , 19 , 21 ).

Venter ( Fig. 20-25 ) — [615 – 870 (830) long; 487

– 668 (601) wide] round, fully sclerotized, and di-

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vided into primary and secondary areas of sclerotization. Gnathosomal bay [119 – 300 (126) long; 68 – 98 (74) wide] not narrow (length/width <3; 1.9 average). Cx-1 narrowed to blunt tip, bearing Cxgl-4 ventro-apically ( Fig. 19 , 20 , 22 A-B, 23). Medial length of Cx-II + Cx-III short, barely longer than wide [10 – 42 (28)]. Genital plates large [153 – 210 (177) long; 133 – 185 (157) wide] and trapezoidal, extending anteriorly beyond level of Leg IV. Each genital plate rimmed in small setae ranging from simple to slightly barbulate ( Fig. 24C ). Additional measurements as follows: Cx-I total length 190 – 307 (281); Cx-III width 319 – 410 (399); Cx-I medial length 109 – 155 (155); genital field to excretory pore 140 – 240 (202); genital field to cauda 128 – 342 (341). Ovipositor morphology unknown.

Legs — Podomere measurements as follows. Leg I (422 – 536 total length): trochanter 37 – 75, basifemur 74 – 114, telofemur 75 – 119 (88), genu 97 – 120 (116), tibia 105 – 135 (125), tarsus 94 – 123 (106). Leg II [450 – 541 total length (541)]: trochanter 35 – 75 (47), basifemur 70 – 110 (88), telofemur 67 –

(a-m p); dorsal glandularia (Dgl); dorsal furrow (df); dorsal plate (dp); eye capsule (ec) lateral glandularia (Lgl); muscle scars (ms);

pre-ocularial setae (pr); post-ocularial setae (po); and area of primary (1°) and secondary (2°) sclerotization. 119 (91), genu 92 – 128 (101), tibia 107 – 148 (118), tarsus 103 – 153 (130). Leg III [511 – 631 total length (596)]: trochanter 44 – 83 (44), basifemur 70 – 115 (87), telofemur 66 – 98 (74), genu 105 – 143 (122), tibia 123 – 168 (145), tarsus 121 – 175 (150). Leg IV [726 – 867 total length (846)]: trochanter 90 – 138 (111), basifemur 108 – 170 (127), telofemur 103-145 (111), genu 139 – 190 (172), tibia 159 – 213 (184), tarsus 155 – 210 (187).

MALE (n=37) (allotypic measurements in parentheses when available) similar to female, except with sexually dimorphic characters discussed above, and with following specifications.

Gnathosoma — Subcapitulum [225 – 289 (249) ventral length; 168 – 215 (195) dorsal length; 93 – 134 (108) tall] posterior edge nearly vertical, ventral bend depth slight [5 – 15 (6)], and with short rostrum [84 – 128 (102) long] that is directed slightly downwards. Chelicerae [200 – 285 (251) long; 13 – 22 (15) high] unmodified with strongly curved fangs [37 – 58 (49) long]. Pedipalps [222 – 290 (263) long] with ventral projections and chaetotaxy as in female. Podomere measurements as follows: trochanters 23 – 34 (30) long and 24 – 37 (29) wide; femora 73 – 101 (91) long and 33 – 59 (47) wide; genua 57 – 81 (69) long and 36 – 52 (45) wide; tibiae 52 – 106 (87) long and 21 – 35 (30) wide; tarsi 18 – 34 (31) long and 9 – 15 (141) wide.

Dorsum — [520 – 650 (590) long; 366 – 495 (430) wide] ovoid to narrow. Dorsal plate with area of primary sclerotization [369 – 517 (429) long; 315 – 422 (367) wide] and an area of secondary sclerotization posteriorly [extends dorsal plate length by 107 – 148 (115) for a total dorsal plate length of 476 – 570 (545)]. Anterior platelets as follows: anteriomedials 96 – 120 (100) long and 49 – 75 (61) wide; anterio-laterals 148 – 190 (161) long and 65 – 90 (74) wide.

Venter — [589 – 800 (715) long; 432 – 605 (568) wide] ovoid to narrow. Gnathosomal bay [70 – 125 (100) long; 49 – 85 (76) wide] not narrow (length/width <3; 1.6 average). Medial length of Cx-II + Cx-III long [58 – 123 (86)]. Genital plates small [105 – 150 (131) long; 74 – 125 (95) wide] and rectangular, not extending anteriorly beyond level of Leg IV. Additional measurements as follows: Cx-I total length 212 – 297 (261); Cx-III width 387 – 371 (349); Cx-I medial length 126 – 167 (152); genital field to excretory pore 73 – 140 (128); genital field to cauda 150 – 242 (220). Genital skeleton ( Fig. 25 ) apically short, broad, and tapering abruptly. Cella proximalis large, with reduced processus proximalia; branchia proximalia well-developed, but branchia distalia only moderately developed.

Legs — Podomere measurements as follows. Leg I (410 – 515 total length): trochanter 40 – 70, basifemur 60 – 103, telofemur 74 – 96, genu 81 – 114 (90), tibia 85 – 125 (116), tarsus 88 – 110. Leg II [394 – 517 total length (484)]: trochanter 28 – 78 (71), basifemur 56 – 103 (85), telofemur 61-84 (69), genu 85 – 110 (90), tibia 93 – 135 (116), tarsus 101 – 133 (121). Leg III [431 – 578 total length (543)]: trochanter 31 – 73 (56), basifemur 55 – 103 (74), telofemur 64 – 89 (83), genu 90 – 125 (103), tibia 110 – 150 (129), tarsus 112 – 158 (133). Leg IV [678-805 total length (787)]: trochanter 89 – 128 (110), basifemur 93 – 155 (119), telofemur 95 – 128 (117), genu 138 – 170 (153), tibia 151 – 190 (177), tarsus 143 – 188 (173).

IMMATURES — unknown.

Etymology — Torrenticola (torrens -, L. a torrent; - colo, L. inhabitant) translates to "torrent dwellers" and refers to the lotic habitat of most species. The specific epithet trimaculata (tres -, L. three; - maculo, L. spotted) refers to the three dorsal spots of adults.

Habitat — Rocky and sandy areas (especially riffles) of healthy streams.

Distribution — Eastern North America. Given the breadth of material examined from the western US, it can be confidently concluded T. trimaculata is absent west of the 100 th meridian. The species is probably unable to cross the Great Plains, which should be considered the western-most bor- der. Further, it seems to be absent from the southeastern coastal plains. The species is most common in the Appalachian Mountains and Interior Highlands (Ozark and Ouachita Mountains), where it can be the dominant mite in a stream. Morph-1, the more pigmented morph ( Fig. 2A, C, E ), is known only from the Interior Highlands. It is the dominant morph in the Ozark Mountains, but is less common than Morph- 2 in the Ouachita Mountains. Morph- 2, the less-pigmented morph ( Fig. 2B, D, F ), is the only morph east of the Mississippi River. Morphs do not coexist within a given stream.

Common name — Three-spotted torrent mite.

Remarks — Torrenticola trimaculata are easily differentiated from other Torrenticola in eastern North America by the three distinct spots on the dorsal plate ( Fig. 1 , 2 , 9 ). Additionally, the following characters are important in combination: anterior platelets not fused with dorsal plate; rostrum short (less than maximum depth of subcapitulum) and sexually dimorphic (angled downward in males); ventral bend of female subcapitulum slight; pedipalpal femora and genua bear ventral dentate projections that are not lamellate; hind coxae extend posteriorly beyond the genital plates in females (in line with genital plates in males); and medial suture of females short. Torrenticola trimaculata sp. nov. exhibit considerable variability across many character states, such as coloration, especially within Morph- 2 ( Fig. 9 ).

We identified four anomalies that are worth reporting. Occasionally, individuals of Morph-1 were found with reddish coloration of the gnathosoma and surrounding sclerites ( Fig. 26A ). Although size and shape of the dorsal spots varied considerably, two extremes were identified: 1) spots enlarged so much so that the posterior spots merged into a contiguous U-shape, only found in Morph- 1 males ( Fig. 26B ); and 2) spots so much reduced that the posterior spots were merely comma-shaped, found

Fisher J.R. et al.

in both sexes of Morph-2 ( Fig. 26C ). Finally, one individual had an under-developed anterio-medial platelet ( Fig. 26 D-E).

Samples often contained at least a few mites with epibionts. Several suctorian ciliates (Ciliophora: Suctorea) are known epibionts on aquatic arthropods, including mites ( Dovgal and Peši´c 2007, 2012, Dovgal et al. 2008), and our samples occasionally contained small numbers of mites covered in unidentified suctorians. Additionally, unidentified bacteria were surprisingly abundant on the integument surface, especially within depressions ( Fig. 22 B-D, 27C).

Commonly, mites were covered with epiphytic diatoms identified by Andy Alverson, a diatom spe- cialist at the University of Arkansas, as Cocconeis placentula Ehrenberg, 1838 ( Fig. 27 A-B). Cocconeis are common epibionts well-known for adhering to plants and algae (e.g., Sand-Jensen 1977, Ferreira and Seeliger 1985, Hardwick et al. 1992, Siqueiros- Beltrones et al. 2002). However, few records exist of these diatoms adhering to animals; but see Siqueiros-Beltrones et al. (2001) for a record of C. notata Petit, 1877 living inside the body of a hydrozoan that itself was epizootic on giant kelp, Macrocystis pyrifera (L.) CA Agardh. The present report represents the first record of C. placentula as epiphytic on water mites.

Type series — HOLOTYPE (♀): USA, Arkansas, Madison Co., Withrow Springs State Park, War Eagle Creek (36°8’59.3" N, 93°44’26.94" W), 27 Jul 2011,

by IM Smith, IMS110034.

ALLOTYPE (♂): 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 (48♀; 36♂): Arkansas, USA : 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 and HW Robison, AJR070300A 8♀ and 5♂ from Montgomery County, Ouachita National Forest , South Fork of Ouachita River, 29 Jul 2011, by AJ Radwell and 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, IMS910036 A 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, IMS900082 B Michigan, USA : 2♀ and 2♂ from Barry County, Thornapple River (42°39’ N, 85°17’ W), 29 Jul 1959, by DR Cook, DRC 590034 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 , IMS890028 A 1♀ from Madoc (44°30’ N, 77°28’ W), 4 May 1980, by IM Smith , IMS800003 A 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 , IMS830093 A 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 , IMS900091 B 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 50 (30♀; 20♂) paratypes deposited at the CNC; 4♀ and 4♂ paratypes deposited at the ACUA; 4♀ and 4♂ paratypes deposited at the OSUAC; 4♀ and 4♂ paratypes deposited at the GMNH. The holotype and allotype are slide mounted in Hoyer’s medium; paratypes are a mixture of Hoyer’s and glycerin jelly slide mounts.

Figures

FIGURE 2: Torrenticola trimaculata n. sp. morphs (A-D compound light micrographs; E-F stereomicrographs): A – Morph I female, note large dorsal spots, pigmented gnathosoma and venter (within area of primary sclerotization), and orange legs; B – Morph II female, note small dorsal spots, and colorless gnathosoma, legs, and venter (except for genital plate); C – Morph I male (note same coloration as female); D – Morph II male (note same coloration as female but with hind coxae pigmented); E-F – Dorsal habitus of Morph I & II, respectively.

FIGURE 9: Torrenticola trimaculata n. sp. variation (dorsal shield compound light micrographs): A – Morph I females, note only slight variably in body shape and spots; B – Morph II females, note high variability in body shape, size and shape of spots, and shape of red marking; C – Morph I males, note only slight variability in body shape body and spots; D – Morph II males, note high variability in body shape and spots. Overall light/dark appearance is a result of exposure differences across multiple cameras, not real-life variation.

FIGURE 3: Torrenticola trimaculata n. sp. color loss in Hoyer’s medium (slide preparation of holotype with separated dorsum and venter): A – prior to warming in Hoyer’s medium, note dorsal spots and ventral coloration; B – same specimen after warming in Hoyer’s medium, note pigmentation (dark color) is cleared, but structural (red) coloration is retained.

FIGURE 10: Torrenticola trimaculata n. sp. sexual dimorphism: A – female gnathosoma, note forward-pointing rostrum and deeper ventral bend; B – male gnathosoma, note down-pointing rostrum and thus shallower ventral bend; C-D – female and male venters, respectively: i) female with shorter coxa II+III medial length; ii) female with larger, rounder body; iii) female genital plates larger, pentagonal (males are rectangular), and extending anteriorly beyond Leg IV insertion; iv) female coxae IV extend posteriorly well beyond genital plates.

FIGURE 11: Torrenticola trimaculata n. sp. female gnathosoma: adoral setae (ad); bifurcating short setae (bss), fringed spatulate setae (fss), long simple setae (lss), oral opening (o), posterio-dorsal apodeme (p-d a), posterio-ventral apodeme (p-v a), simple grooved setae (sgs).

FIGURE 12: Torrenticola trimaculata n. sp. rostral opening and fangs (LT-SEM): A – frontal aspect of rostrum showing opening for fangs surrounded by adoral setae (ad); B – lateral view of rostrum with both fangs partially extended; C – lateral view of fangs; D – frontal view depicting extended right fang (left fang just emerging from rostrum), note lateral and medial teeth of right fang probably used for anchoring into prey; E – dorsal view of rostral opening with fangs retracted.

FIGURE 13: Torrenticola trimaculata n. sp. pedipalp (LT-SEM): A-B – medial (A) and lateral (B) view of genu depicting medial placement of disto-ventral dentate projections; C-D – lateral (C) and inner (D) detail of femoral projection; E – lateral detail of mid-ventral tibial spines. Fringed spatulate setae (fss); long simple setae (lss); short grooved setae (sgs).

FIGURE 14: Torrenticola trimaculata n. sp. pedipalp setae (LT-SEM): A – dorsal view of right tarsus; B – latero-medial view of right tarsus, note dual ventral claws; C – dorso-lateral view of pedipalps. Fringed spatulate setae (fss); grooved spine-like seta (gss); short grooved setae (sgs); short bifurcating setae (sbs); long simple setae (lss)

FIGURE 15: Torrenticola trimaculata n. sp. dorsal plates: anterio-lateral platelet (a-l p); anterio-medial platelet (a-m p); dorsal glandularia (Dgl); dorsal plate (dp); muscle scars (ms); post-ocularial setae (po); and area of primary (1°) and secondary (2°) sclerotization.

FIGURE 16: Torrenticola trimaculata n. sp. dorsum (LT-SEM; gnathosoma removed): anterio-lateral platelet (a-l p); anterio-medial platelet

FIGURE 17: Torrenticola trimaculata n. sp. anterio-lateral dorsum (LT-SEM): A – anterio-lateral platelet (a-l p); anterio-medial platelet (a-m p); dorsal glandularia (Dgl); dorsal plate (dp); eye capsule (ec); lateral glandularia (Lgl); lyrifissures (l); pre-ocularial setae (pr); post-ocularial setae (po); B – close-up view of l-2; C – close-up view of eye capsule.

FIGURE 18: Torrenticola trimaculata n. sp. posterio-lateral dorsum (LT-SEM): anterio-lateral platelet (a-l p); dorsal furrow (df); dorsal glandularia (Dgl); dorsal plate (dp); lateral glandularia (Lgl); lyrifissures (l); anterior two posterior-lateral platelets (p-l p) partially visible; and area of primary (1°) and secondary (2°) sclerotization.

FIGURE 19: Torrenticola trimaculata n. sp. frontal view (LT-SEM). Eye capsule (ec); lyrifissure 1 (l-1); latero-glandularia 1 (Lgl-1); and coxal glandularia 4 (Cxgl-4). Note vertically stacked, dual eyes on eye capsule.

FIGURE 6: Torrenticola trimaculata n. sp. primary and secondary sclerotization (Morph-1 female compound light micrographs):

FIGURE 21: Torrenticola trimaculata n. sp. lateral glandularia (LT-SEM; female depicted): coxal glandularia (Cxgl); dorsal furrow (df); dorsal glandularia 1 (Dgl-1); eye capsule (ec); lateral glandularia (Lgl); lyrifissure 1 (l-1); preocularial seta (pr); and areas of primary (1°) and secondary sclerotization (2°).

FIGURE 20: Torrenticola trimaculata n. sp. venter (right legs removed; left leg setae omitted; female depicted): coxae (Cx; fused coxae: Cx II+III); coxal glandularia (Cxgl); excretory pore (ep); latero-glandularia (Lgl); medial length of suture between Cx II+III (Cx II+III mL); ventral glandularia (Vgl); and area of primary (1°) and secondary (2°) sclerotization.

FIGURE 22: Torrenticola trimaculata n. sp. glandularia (LT-SEM): A – ventral view of gnathosoma and surrounding coxae depicting coxal glandularia (Cxgl); B – close-up of Cxgl-4; C – dorsal glandularium 4 (Dgl-4); D – coxal glandularium 2 (Cxgl-2).

FIGURE 23: Torrenticola trimaculata n. sp. venter (LT-SEM; female depicted): coxal glandularia (Cxgl); excretory pore (ep); ventral glandularia (Vgl): and area of primary (1°) and secondary (2°) sclerotization.

FIGURE 24: Torrenticola trimaculata n. sp. ventral posterior (LT-SEM): A – posterior area of secondary sclerotization (2°) depicting vestigial ventral glandularium 1 (Vgl-1), ventral glandularia 2 (Vgl-2), and excretory pore (ep); B – close-up of excretory pore; C – genital plates, note rim of setae surrounding each plate.

FIGURE 25: Torrenticola trimaculata n. sp. male genital skeleton. A – line drawing depicting moderately developed branchia distalia (bd), well-developed branchia proximalia (bp), weak carina anterior (ca), very large cella proximalis (cp), and reduced processus proximalis (pp); B – compound light micrograph.

FIGURE 1: Torrenticola trimaculata n. sp. habitus of types (montaged from iPhone steromicrographs): A – Holotype (female): dorsal and ventral habitus, Morph 1; B – Allotype (male): dorsal and ventral habitus, Morph 1. Coloration is not indicative of sex.

FIGURE 26: Torrenticola trimaculata n. sp. abnormalities (males depicted; A-C are slide preparations with separated gnathosoma, dorsum, and venter; D-E are LT-SEM): A – red color of gnathosoma and surrounding venter, only occurs in Morph 1; B – posterior spots fused, only occurs in Morph 1; C – posterior spots strongly reduced, only occurs in Morph 2; D – left anterio-medial platelets stunted; E – close-up of D depicting eye capsule (ec), anterio-medial platelet (e-m p), and dorsal glandularia 2 (Dgl-2).

FIGURE 27: Torrenticola trimaculata n. sp. commensals (LT-SEM): A – diatoms (Cocconeis placentula Ehrenberg, 1838) covering dorsum; B – close-up of C. placentula covering T. trimaculata dorsum; C – bacteria covering body, especially within depressions.