See Fisher et al. (2015) for diagnosis.

For larval diagnosis see Smith (1982). Adults differ from torrenticolines in having three pairs of acetabula (six in Torrenticolinae); condyles present over the insertions of leg-IV; long posterio-dorsal subcapitular apodemes (also long in Monatractides); a ridge extending anteriorly from the leg-IV socket; and a ring of platelets closely affiliated with the central dorsal plate, i.e., not hidden within a dorsal furrow as in torrenticolines (Fig. 1). They are further characterized by having a single anterio-medial dorsal platelet and pedipalps without ventral projections, although some torrenticolines also have these characters. Testudacarinae can be further diagnosed by the following combination of characters. Medial dorsal plate exhibiting secondary and occasionally tertiary sclerotization. Dorsal platelets variable in size, shape, and coloration. Anterio-medial platelet smaller than anterio-lateral platelets and trapeziform (rounded to rectangular). Anterio-lateral platelets long with anterior bulge and posterior tapering. Seven pairs of lateral platelets present. Lateral-platelet-2, -4, and -6 large and elongate and -1, -3, -5, and -7 smaller and rounded. Lateral-platelet-3 highly variable and positioned either anterior or lateral to lateral-platelet-4. Lateral-platelet-4 highly variable in shape mostly depending on lateral-platelet-3 position. Dorso-glandularia-2 and post-ocularial setae located together on anterio-lateral platelet. Dorso-glandularia-3, -5, and -6 located on lateral-platelet-1, -5, and -7 respectively. Dorso-glandularia-4 located on the large medial dorsal plate. Latero-glandularia-4 located on lateral-platelet-3. Ventro-glandularia-3 posterior to coxae-IV (on coxae-IV in other torrenticolids). Coxo-glandularia-4 located at tip of coxae-I (as in Monatractides and many Torrenticola). Pedipalp, femur, and genu with plumose setae ventrally. Also similar to Monatractides, posterio-dorsal subcapitular apodemes are long. Rostrum short.

Testudacarines have been reported on many occasions outside of their original descriptions. Furthermore, the Canadian National Collection in Ottawa, Canada includes thousands of testudacarines collected from across most of North America (Smith et al. 2010). In Asia there have only been a handful of additional reports (Walter 1929, Pešić and Smit 2007, Jin et al. 2010, Morimoto 2012). This is not completely due to a lack of torrenticolid work in Asia, for an extensive list see Walter et al. (2009, pg. 256) and Fisher et al. (2015). Extensive work has also been done on water mites in Europe, Africa, and Australia without any reports of testudacarines. Therefore, testudacarines are currently thought to be widely distributed throughout most of North America (with southern limits in Mexico and northern limits around the 60^th parallel), and sparsely distributed in parts of Asia.

The three pairs of acetabula, coxae-IV condyles, and “generalized” pedipalps are plesiomorhphic states that clearly show testudacarines as retaining ancestral torrenticolid characteristics (Wiles 1997a). Wiles (1997a) and other authors suggest latero-glandularia-3 is present on the dorsum of testudacarines. However, we suggest that this is latero-glandularia-4 due to its posterior-most positioning. We also detail sexually dimorphic characters (Fig. 7). Although Habeeb (1954) first noted differences between the sexes of T. vulgaris, he did not present these distinctions in their wider context as overall conditions of Testudacarinae. Sexual dimorphism present in Testudacarinae include: 1) female dorso-glandularia-4 positioned closer to the muscle scars; 2) dorsal secondary sclerotization always present in females and usually absent in males (very small if present in males); 3) female coxae-II+III midline short; 4) genital field almost entirely enveloped by coxal field in females but only around half of male genital field within coxal field; 5) females idiosoma larger and rounder (males around 80% of female size) with less of the ventral shield composed of coxal field; and 6) excretory pore well separated from ventral line of secondary sclerotization in females, and is either in direct contact with or nearly so in males.

Figure 7. 

Testudacarine sexual dimorphism: female dorsal shield (A) and ventral shield (C) differing from male (B, D) by the following characters: 1) dorso-glandularia-4 positioned far closer to muscle scares; 2) area of secondary sclerotization always present (males rarely present; very small if present); 3) with shorter coxae-II+III midline; 4) genital field enveloped by coxal field; 5) larger and rounder body (males around 80% of female size); 6) excretory pore well separated from ventral line of secondary sclerotization.

Debsacarus oribatoides (Habeeb, 1961).

Debsacarus differ from all other Testudacarinae in having four-segmented pedipalps (instead of five) and projections on the anterio-tips of coxae-I. With the exception of Testudacarus hyporhynchus, Debsacarus differ from all other Testudacarinae in having an elongate gnathosoma and an extremely narrow gnathosomal bay that is covered dorsally and ends anterior to the leg-I insertion ventrally.

Known from only two counties (Los Angeles and Monterey) in California.

Lectotype (1♀): California, USA: 1♀ from Los Angeles County, Coldbrook Guard Station, North Fork of San Gabriel River, 25 June 1961, by H Habeeb, HH610024; Paralectotype (1♂): California, USA: 1♂ from Los Angeles County, Coldbrook Guard Station, North Fork of San Gabriel River, 25 June 1961, by H Habeeb, HH610024.

Other (10♀, 8♂): California, USA: 1♂ from Monterey County, Salmon Falls Creek, beside Route 1 12.5 km south of Gorda (35°48'56.00"N, 121°21'30.00"W), 2 June 2010, by IM Smith, IMS100045; 5♀ and 3♂ from Monterey County, Los Padres National Forest, Lucia beside Ferguson-Nacimiento Road 5.6 km east of Route 1 (36°0'3.00"N, 121°28'31.00"W), 3 June 2010, by IM Smith, IMS100048; 1♀ and 3♂ from Monterey County, Los Padres National Forest, Lucia beside Nacimiento-Ferguson Road 11.3 km west of Nacimiento Campground (36°1'N, 121°27'W), 30 July 1987, by IM Smith, IMS8700119; 1♀ and 1♂ from Monterey County, Los Padres National Forest, Salmon Creek, beside Route 1 south of Gorda (35°49'N, 121°22'W), 29 July 1987, by IM Smith, IMS870118; 1♀ from Monterey County, Limekiln State Park, Hare Canyon Creek, near campground (36°0'41.00"N, 121°31'1.00"W), 6 September 2013, by JR Fisher, JRF13-0906-001; 1♀ from Monterey County, Salmon Creek, beside Route 1 south of Gorda (35°49'N, 121°22'W), 28 July 1987, by IM Smith, IMS870114A; 1♀ from Los Angeles County, Angeles National Forest, North Fork of San Gabriel River, off Route 39 (34°16'16.00"N, 117°50'46.00"W), 8 September 2013, by JR Fisher, JRF13-0908-001.

Lectotype (1♀), and paralectotype (1♂) deposited at CNC.

Female (n=11) with characteristics of the genus with following specifications.

Gnathosoma (Fig. 9) — Subcapitulum [260–290 ventral length; 125–145 dorsal length; 73–84 tall] elongate with long rostrum. Chelicerae [195–220 long] noticeably straight with short, almost straight fangs [28–33 long]. Pedipalp [217–234 long] highly modified: lanceolate and with four segments. Trochanter [7–9 long; 38–40 wide] shortened. Femur [39–44 long; 30–34 wide]. Fused genu and tibia [41–47 long; 25–28 wide]. Tarsus [17–20 long; 12–15 wide].

Dorsum (Fig. 10) — [574–741 long; 471–561 wide] round to ovoid. Dorsal plate [465–586 long; 391–451 wide]. Primary sclerotization [436–510 long] grey-blue. Dorso-glandularia-4 [163–194 apart] in line with and lateral to [29-48] muscle scars. Platelets extremely robust and colorless. All three anterior platelets similar in size and noticeably rectangular. Anterio-medial platelet [173–209 long; 74–128 wide] large trapezoid with slightly rounded anterior margin. Anterio-lateral platelet [185–207 long; 97–127 wide] without noticeable bulge or posterior narrowing. Lateral platelets as follows: lateral-1 [38–50 long; 25–38 wide]; lateral-2 [143–172 long; 40–66 wide]; lateral-3 [39–64 long; 16–32 wide]; lateral-4 [107–132 long; 28–51 wide]; lateral-5 [51–78 long; 28–48 wide]; lateral-6 [92–128 long; 25–55 wide]; lateral-7 [49–101 long; 22–50 wide].

Figure 8. 

Debsacarus oribatoides molecular phylogeny: 28S and COI Bayesian analysis showing strong support single distinct clade (●: >95% posterior probability); clade exhibits <.6% divergence in COI within and >15% divergence between any other clade (not pictured); continuation of (E) lineage from Fig. 6.

Figure 9. 

Debsacarus oribatoides gnathosoma (generalized).

Figure 10. 

Debsacarus oribatoides female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 10) — [779–929 long; 510–610 wide] round to ovoid and colorless. Primary sclerotization [668–756 long. Gnathosomal bay [33–45 dorsal length; 128–148 ventral length; 33–38 wide] very narrow; dorsal bay length extremely short giving the bay a “covered” appearance and ventral bay base ending anterior to the leg-I insertion. Coxal field [520–567 long; 325–353 wide]. Coxa-I [292–334 long; 160–186 midlength] long and with characteristic secondary growth attached at the anterior tips. Coxa-II + III [137–153 distance to top of coxa-II; 210–237 distance to top of coxa-III; 379–424 distance to bottom of coxa-III; 228–274 total length]. Coxa-IV [355–400 distance to top; 155–173 total length]. Genital field [362–409 distance to top; 556–601 distance to bottom; 185–208 total length; 155–175 width; 221–274 distance from gnathosomal bay; 59–101 distance from coxa-I; 163–227 distance to excretory pore; 215–366 distance to caudad] large. Eggs [200 long; 1–2 eggs]. Distance to excretory pore [727–809].

Legs — colorless. Total leg and podomere lengths as follows: Leg-I [459–505 total; trochanter 54–62; basifemur 81–91; telofemur 63–68; genu 81–91; tibia 86–100; tarsus 84–95]. Leg-II [516–554 total; trochanter 62–65; basifemur 85–100; telofemur 63–71; genu 84–96; tibia 100–114; tarsus 106–115]. Leg-III [593–644 total; trochanter 63–69; basifemur 97–105; telofemur 70–78; genu 104–118; tibia 125–143; tarsus 130–142]. Leg-IV [779–862 total; trochanter 84–96; basifemur 118–127; telofemur 115–129; genu 141–166; tibia 160–181; tarsus 148–170].

Male (n=9) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma (Fig. 9) — Subcapitulum [229–266 ventral length; 120–132 dorsal length; 64–78 tall]. Chelicerae [175–200 long]. Fangs [25–26 long]. Pedipalp [206–219 long]. Trochanter [7–9 long; 35–38 wide]. Femur [36–40 long; 30–32 wide]. Fused genu and tibia [43–45 long; width 23–26 wide]. Tarsus [16–17 long; 13–15 wide].

Dorsum (Fig. 11) — [534–590 long; 416–478 wide]. Dorsal plate [421–477 long; 332–380 wide] without secondary sclerotization. Dorso-glandularia-4 [157–188 apart] equally anterior to [25–55] and lateral to [31–53] muscle scars. Anterio-medial platelet [151–200 long; 90–108 wide]. Anterio-lateral platelet [169–186 long; 97–118 wide]. Lateral platelets as follows: lateral-1 [33–46 long; 22–31 wide]; lateral-2 [134–155 long; 48–55 wide]; lateral-3 [29–51 long; 17–24 wide]; lateral-4 [88–113 long; 31–40 wide]; lateral-5 [41–49 long; 22–35 wide]; lateral-6 [82–101 long; 27–42 wide]; lateral-7 [36–59 long; 20–38 wide].

Figure 11. 

Debsacarus oribatoides male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 11) — [686–773 long; 449–515 wide]. Primary sclerotization [637–705 long]. Gnathosomal bay [21–36 dorsal length; 118–132 ventral length; 27–42 wide]. Coxal field [474–532 long; 307–332 wide]. Coxa-I [272–294 long; 143–169 midlength]. Coxa-II + III [123–138 distance to top of coxa-II; 192–208 distance to top of coxa-III; 377–417 distance to bottom of coxa-III; 253–279 total length]. Coxa-IV [327–368 length to top; 136–164 total length]. Genital field [388–435 distance to top; 536–598 distance to bottom; 148–165 total length; 126–150 width; 270–307 distance from gnathosomal bay; 108–144 distance from coxa-I; 88–107 distance to excretory pore; 143–179 distance to caudad]. Genital skeleton [210–265 long; 95–115 wide]. Distance to excretory pore [637–705].

Legs — total leg and podomere lengths as follows: Leg-I [447–476 total; trochanter 59–63; basifemur 80–88; telofemur 61–68; genu 79–88; tibia 85–95; tarsus 80–92]. Leg-II [479–526 total; trochanter 54–67; basifemur 82–89; telofemur 60–72; genu 80–90; tibia 94–105; tarsus 105–114]. Leg-III [544–624 total; trochanter 56–66; basifemur 79–102; telofemur 65–74; genu 95–110; tibia 119–138; tarsus 120–137]. Leg-IV [743–857 total; trochanter 85–110; basifemur 107–125; telofemur 113–130; genu 134–160; tibia 152–177; tarsus 145–158].

Same as genus.

Same as genus.

Debsacarus oribatoides show at least 15% COI divergence from all other Testudacarinae and less than .6% divergence from one another (Fig. 8). Additionally, Habeeb (1961) describes a protrusible maxillary tube, however, we find no evidence in the additional specimens examined that the maxillary tube or subcapitulum is any more protrusible than what is commonly found in other Testudacarus, and certainly is not protrusible like in Pseudotorrenticola. Habeeb (1961) did not designate types, however, he described the species from the only two specimens available. From those two specimens, we have designated a lectotype (♀) and paralectotype (♂).

Testudacarus tripeltatus Walter, 1928

Members of this genus, unlike Debsacarus, lack projections on the anterior tips of coxae-I and have five-segmented pedipalps (instead of four). Furthermore, with the exception of Testudacarus hyporhynchus, they differ from Debsacarus in having a rounded gnathosoma (rather than elongate) and a wide gnathosomal bay that is uncovered dorsally and ventrally ends posterior to the leg-I insertion.

Same as subfamily.

Holotype (1♀): North Carolina, USA: 1♀ from Haywood, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road at Hannah Hoglen Cemetery site (35°38'29.00"N, 83°3'22.00"W), 22 September 2010, by IM Smith, IMS100154 (Specimen 141898 – DNA#1493); Paratypes (9♀, 10♂): North Carolina, USA: (allotype) 1♂ from Haywood, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road at Hannah Hoglen Cemetery site (35°38'29.00"N, 83°3'22.00"W), 22 September 2010, by IM Smith, IMS100154 (Specimen 146756 – DNA#2171); 1♀ and 2♂ from Haywood, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road at Hannah Hoglen Cemetery site (35°38'29.00"N, 83°3'22.00"W), 22 September 2010, by IM Smith, IMS100154; 2♀ and 1♂ from Haywood County, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road near bridge 1.7km north of road to campground (35°38'45.00"N, 83°4'32.00"W), 20 September 2010, by IM Smith, IMS100150; 2♂ from Macon County, Rainbow Springs, beside Forest Road 67 4.4 km south of Standing Indian Campground (35°3'6.00"N, 83°30'45.00"W), 1 July 2006, by IM Smith, IMS060040; 2♂ from Yancey County, Pisgah National Forest, South Toe River, Lost Cove beside Toe River Road (Forest Road 472) 0.4km east of Forest Road 2074 (35°45'0.00"N, 82°12'53.00"W), 9 September 2007, IM Smith, IMS070059; 1♀ from Yancey County, Pisgah National Forest, South Toe River, Lost Cove Picnic Area beside Toe River Road (Forest Road 472) 2.8 km east of Route 80 (35°45'13.00"N, 82°12'42.00"W), 27 September 2009, by IM Smith, IMS090127; Tennessee, USA: 1♂ from Monroe, beside Forest Route #35 2.3km northeast of road from Route 165 to Miller Chapel Baptist Church (35°21'47.00"N, 84°9'47.00"W), 12 September 2009, by IM Smith, IMS090112; 3♀ and 1♂ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'47.00"N, 83°31'52.00"W), 7 September 2009, by IM Smith, IMS090101; 1♀ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'48.00"N, 83°31'53.00"W), 3 September 2009, by IM Smith, IMS090095; Georgia, USA: 1♀ from Floyd County, beside road from Everrett Springs to Villanow 1.4 km south of The Pocket Recreation Area, 4 July 1990, by IM Smith, IMS900077.

(1♀, 2♂): North Carolina, USA: 1♀ from Haywood County, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road near bridge 1.7km north of road to campground (35°38'45.00"N, 83°4'32.00"W), 20 September 2010, by IM Smith, IMS100150; 1♂ from Haywood County, Great Smoky Mountains National Park, tributary of Hemphill Creek, Appalachian Highlands Science Learning Center near Ferguson Cabin site, (35°34'56.00"N, 83°4'30.00"W), 21 September 2010, by IM Smith, IMS100153; Tennessee, USA: 1♀ from Sevier County, Great Smoky Mountains National Park, Catron Branch, Elkmont Road off Little River Road (35°39'51.00"N, 83°35'19.00"W), 24 September 2010, IMS100156.

Holotype (1♀), allotype (1♂), and eight paratypes (4♀, 4♂) deposited at CNC; ten paratypes (5♀, 5♂) at ACUA.

These mites differ from all others in the complex in having large medial pores on the dorsal plate surrounded by a ring of smaller pores (all pores uniform in other species). Males also have a “bleached” or colorless area posterior to the coxal plate that is colored in other members of the complex.

Female (n=10) with characteristics of genus with following specifications.

Gnathosoma — Subcapitulum [165–175 ventral length; 99–106 dorsal length; 90–100 tall] ovoid with short rostrum. Chelicerae [139–150 long] unmodified with lightly curved fangs [29–32 long]. Pedipalp [192–205 long] unmodified. Trochanter [25–28 long; 29–32 wide]. Femur [54–57 long; 37–40 wide]. Genu [40–46 long; 29–33 wide]. Tibia [51–55 long; 20–23 wide]. Tarsus [19–21 long; 10–11 wide].

Dorsum (Fig. 24) — [591–669 long; 445–504 wide] round to ovoid. Dorsal plate [485–556 long; 375–424 wide] with noticeable pore variation: medial pores large surrounded by smaller distal pores. Primary sclerotization [425–470 long] violet to blue. Dorso-glandularia-4 [124–175 apart] lateral to [19–43] and around the anterior tips of the muscle scars. Platelets violet to blue or colorless. Anterio-medial platelet [146–168 long; 81–101 wide] colorless rounded trapezoid noticeably smaller than anterio-lateral platelets. Anterio-lateral platelet [170–197 long; 89–102 wide] with violet to blue restricted to posterior half or third of the platelet. Lateral platelets as follows: lateral-1 [53–69 long; 46–57 wide]; lateral-2 [125–140 long; 35–52 wide]; lateral-3 [39–53 long; 20–27 wide]; lateral-4 [96–115 long; 32–43 wide]; lateral-5 [50–62 long; 29–39 wide]; lateral-6 [81–96 long; 29–43 wide]; lateral-7 [61–77 long; 27–33 wide].

Figure 24. 

Testudacarus hitchensi female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 24) — [765–870; 482–553 wide] round to ovoid. Primary sclerotization [631–717 long] violet to blue. Gnathosomal bay [71–90 dorsal length; 149–170 ventral length; 53–62 wide]. Coxal field [482–543 long; 325–409 wide]. Coxa-I [256–289 long; 99–126 midlength]. Coxa-II + III [118–140 distance to top of coxa-II; 187–215 distance to top of coxa-III; 347–401 distance to bottom of coxa-III; 224–264 total length]. Coxa-IV [333–375 distance to top; 139–168 total length]. Genital field [329–382 distance to top; 493–542 distance to bottom; 158–172 total length; 125–150 width; 178–212 distance from gnathosomal bay; 69–100 distance from coxa-I; 175–234 distance to excretory pore; 272–349 distance to caudad]. Eggs [150–168 long; 1–4 eggs]. Distance to excretory pore [688–777].

Legs — orange and restricted violet to blue. Total leg and podomere lengths as follows: Leg-I [473–524 total; trochanter 60–62; basifemur 83–93; telofemur 65–76; genu 86–95; tibia 92–105; tarsus 83–95]. Leg-II [501–552 total; trochanter 54–63; basifemur 83–93; telofemur 65–72; genu 88–99; tibia 101–111; tarsus 102–115]. Leg-III [586–635 total; trochanter 61–65; basifemur 89–100; telofemur 70–80; genu 105–113; tibia 122–137; tarsus 132–144]. Leg-IV [805–876 total; trochanter 93–109; basifemur 115–132; telofemur 115–125; genu 151–167; tibia 167–180; tarsus 158–177].

Male (n=10) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [150–160 ventral length; 95–106 dorsal length; 86–95 tall]. Chelicerae 127–139 long]. Fangs [26–29 long]. Pedipalp [180–195long]. Trochanter [25–27 long; 27–30 wide]. Femur [50–55 long; 34–37 wide]. Genu [38–41 long; width 26–29 wide]. Tibia [49–52 long; 19–22 wide]. Tarsus [17–20 long; 9–11 wide].

Dorsum (Fig. 25) — [491–567 long; 387–436 wide]. Dorsal plate [404–474 long; 326–375 wide]. Dorso-glandularia-4 [116–152 apart] far anterior to [53–75] and lateral to [13–32] muscle scars. Anterio-medial platelet [137–152 long; 71–91 wide]. Anterio-lateral platelets [163–184 long; 74–88 wide]. Lateral platelets as follows: lateral-1 [45–54 long; 37–44 wide]; lateral-2 [101–120 long; 34–41 wide]; lateral-3 [39–50 long; 19–32 wide]; lateral-4 [74–110 long; 30–35 wide]; lateral-5 [46–58 long; 25–33 wide]; lateral-6 [53–75 long; 27–34 wide]; lateral-7 [46–62 long; 24–33 wide].

Figure 25. 

Testudacarus hitchensi male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 25) — [641–718 long; 418–481 wide]. Primary sclerotization [593–671 long]. Gnathosomal bay [62–89 dorsal length; 131–164 ventral length; 45–67 wide]. Coxal field [441–500 long; 309–340 wide]. Coxa-I [233–276 long; 95–114 midlength]. Coxa-II + III [105–128 distance to top of coxa-II; 171–202 distance to top of coxa-III; 357–409 distance to bottom of coxa-III; 245–288 total length]. Coxa-IV [304–355 length to top; 127–159 total length]. Genital field [378–440 distance to top; 524–598 distance to bottom; 143–157 total length; 115–131 width; 239–284 distance from gnathosomal bay; 143–173 distance from coxa-I; 55–91 distance to excretory pore; 110–153 distance to caudad]. Genital skeleton [190–207 long; 115–126 wide]. Distance to excretory pore [593–671].

Legs — total leg and podomere lengths as follows: Leg-I [444–508 total; trochanter 55–62; basifemur 75–89; telofemur 63–73; genu 80–91; tibia 85–99; tarsus 84–96]. Leg-II [474–533 total; trochanter 60–64; basifemur 77–90; telofemur 61–71; genu 82–93; tibia 92–106; tarsus 99–113]. Leg-III [537–598 total; trochanter 57–64; basifemur 80–92; telofemur 65–73; genu 96–108; tibia 113–128; tarsus 121–137]. Leg-IV [721–778 total; trochanter 88–99; basifemur 96–117; telofemur 102–113; genu 135–151; tibia 147–168; tarsus 142–156].

Specific epithet hitchensi after the late Christopher Eric Hitchens, the English author, journalist, and literary critic. As Sam Harris’ wife, Annaka, said: “Nothing Hitchens does is ever boring.” Hitchens has inspired thousands of free-thinkers to remain clever and engaged in our attempts to understand the world around us.

Eastern United States east of the Mississippi River with southern limits in Florida.

As it is likely that this species represents a cryptic species complex, measurements were only included from specimens exhibiting less than 2% COI divergence within the clade (those highlighted in red in Fig. 23 were excluded). This was done so measurements would remain useful if more species were diagnosed in the future.

Holotype (1♀): North Carolina, USA: 1♀ from Haywood County, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road near bridge 1.7km north of road to campground (35°38'45.00"N, 83°4'32.00"W), 20 September 2010, by IM Smith, IMS100150 (Specimen 146752 – DNA#2166); Paratypes (12♀, 10♂): North Carolina, USA: (allotype) 1♂ from Haywood County, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road near bridge 1.7km north of road to campground (35°38'45.00"N, 83°4'32.00"W), 20 September 2010, by IM Smith, IMS100150 (Specimen 146750 – DNA#2164); Tennessee, USA: 1♂ from Sevier County, Great Smoky Mountains National Park, Crosby Creek, Cosby Recreation Area beside Cosby Campground Road 0.3km from Route 321 (35°46'54.00"N, 83°13'2.00"W), 16 September 2010, by IM Smith, IMS100140; 2♀ and 2♂ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'47.00"N, 83°31'52.00"W), 7 September 2009, by IM Smith, IMS090101; 1♀ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'48.00"N, 83°31'53.00"W), 3 September 2009, by IM Smith, IMS090095; 1♂ from Blount County, Great Smoky Mountains National Park, Cades Cove, near parking lot for Abrams Falls Trail (35°35'26.00"N, 83°51'10.00"W), 17 September 2010, by IM Smith, IMS100143; 2♀ from Sevier Co, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'47.00"N, 83°31'51.00"W), 10 September 2010, by IM Smith, IMS100125; North Carolina, USA: 2♀ and 2♂ from Haywood County, Great Smoky Mountains National Park, Cataloochee Creek, beside Cataloochee Road 0.3km north of Cataloochee Campground (35°38'1.00"N, 83°5'2.00"W), 6 September 2009, IMS090097; 2♀ and 1♂ from Haywood County, Great Smoky Mountains National Park, Big Creek, Waterville Big Creek Picnic Area (35°44'59.00"N, 83°6'42.00"W), 16 September 2010, by IM Smith, IMS100138; 1♀ and 1♂ from Haywood County, Great Smoky Mountains National Park, Cataloochee Creek, beside Mount Sterling Road near bridge 1.7km north of road to campground (35°38'45.00"N, 83°4'32.00"W), 20 September 2010, by IM Smith, IMS100150; 1♂ from Yancey County, Pisgah National Forest, South Toe River, Lost Cove beside Toe River Road (Forest Road 472) 0.4km east of Forest Road 2074 (35°45'0.00"N, 82°12'53.00"W), 9 September 2007, IM Smith, IMS070059; 1♀ from Haywood County, Great Smoky Mountains National Park, Rough Fork Creek, beside road to Nellie 0.3 km west of Pretty Hollow Gap Trailhead (35°37'31.00"N, 83°6'46.00"W), 20 September 2010, by IM Smith, IMS100148; Pennsylvania, USA: 1♀ from Fayette County, State Game Lands #51, Dunbar Creek, off Furnace Hill Road east of Dunbar (39°57'50.00"N, 79°35'8.70"W), 10 August 2014, MJ Skvarla, MS14-0810-001.

Holotype (1♀), allotype (1♂) and ten paratypes (5♀, 5♂) deposited at Canadian National Collection; eleven paratypes (7♀, 4♂) at ACUA.

These mites have violet to blue coloration over the majority of their anterio-lateral platelets while the rest of the complex have coloration restricted to the posterior half of the platelet.

Female (n=13) with characteristics of the genus with following specifications.

Gnathosoma — Subcapitulum [143–165 ventral length; 90–105 dorsal length; 84–95 tall] ovoid with short rostrum. Chelicerae [119–136 long] unmodified with lightly curved fangs [24–32 long]. Pedipalp [167–191 long] unmodified. Trochanter [23–30 long; 29–31 wide]. Femur [47–53 long; 33–38 wide]. Genu [37–42 long; 27–30 wide]. Tibia [40–53 long; 17–22 wide]. Tarsus [15–20 long; 9–12 wide].

Dorsum (Fig. 26) — [527–617 long; 420–482 wide] ovoid. Dorsal plate [375–495 long; 355–515 wide]. Primary sclerotization [358–419 long] violet to blue. Dorso-glandularia-4 [113–167 apart] lateral to [16–42] and around the anterior tips of the muscle scars. Platelets violet to blue or clear. Anterio-medial platelet [112–144 long; 70–94 wide] colorless rounded trapezoid noticeably smaller than anterio-lateral platelets. Anterio-lateral platelets [156–183 long; 74–86 wide] mostly violet to blue with anterio-most corner generally colorless; anterio-medial corner often with orange spot. Lateral platelets as follows: lateral-1 [38–50 long; 35–44 wide]; lateral-2 [103–133 long; 30–40 wide]; lateral-3 [29–45 long; 16–30 wide]; lateral-4 [90–119 long; 26–37 wide]; lateral-5 [47–64 long; 22–34 wide]; lateral-6 [65–90 long; 27–34 wide]; lateral-7 [56–69 long; 24–36 wide].

Figure 26. 

Testudacarus harrisi female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 26) — [668–786 long; 453–509 wide] round to ovoid. Primary sclerotization violet to blue. Gnathosomal bay [61–78 dorsal length; 128–156 ventral length; 45–56 wide]. Coxal field [418–480 long; 281–363 wide]. Coxa-I [213–254 long; 85–105 midlength]. Coxa-II + III [109–131 distance to top of coxa-II; 170–195 distance to top of coxa-III; 295–356 distance to bottom of coxa-III; 186–234 total length]. Coxa-IV [291–335 distance to top; 125–154 total length]. Genital field [284–335 distance to top; 426–489 distance to bottom; 139–154 total length; 114–127 width; 152–184 distance from gnathosomal bay; 65–82 distance from coxa-I; 167–207 distance to excretory pore; 241–307 distance to caudad]. Distance to excretory pore [593–693].

Legs — violet to blue and orange. Total leg and podomere lengths as follows: Leg-I [449–485 total; trochanter 54–62; basifemur 77–83; telofemur 62–70; genu 80–90; tibia 89–99; tarsus 80–90]. Leg-II [471–510 total; trochanter 54–60; basifemur 74–84; telofemur 61–66; genu 82–94; tibia 98–107; tarsus 87–109]. Leg-III [548–612 total; trochanter 55–64; basifemur 79–91; telofemur 66–74; genu 96–114; tibia 116–137; tarsus 119–141]. Leg-IV [737–825 total; trochanter 79–99; basifemur 103–123; telofemur 103–121; genu 138–154; tibia 154–169; tarsus 147–167].

Male (n=10) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [133–144 ventral length; 83–90 dorsal length; 72–84 tall]. Chelicerae [110–120 long]. Fangs [25–30 long]. Pedipalp [168–183 long]. Trochanter [22–25 long; 25–29 wide]. Femur [45–50 long; 32–36 wide]. Genu [36–40 long; width 24–30 wide]. Tibia [44–52 long; 18–20 wide]. Tarsus [16–20 long; 8–11 wide].

Dorsum (Fig. 27) — [418–488 long; 312–380 wide]. Dorsal plate [340–402 long; 271–322 wide]. Dorso-glandularia-4 [89–129 apart] far anterior to [31–71] and lateral to [12–24] muscle scars. Anterio-medial platelet [111–132 long; 63–80 wide]. Anterio-lateral platelets [141–164 long; 63–79 wide]. Lateral platelets as follows: lateral-1 [30–38 long; 29–32 wide]; lateral-2 [85–96 long; 24–33 wide]; lateral-3 [30–40 long; 15–25 wide]; lateral-4 [61–78 long; 21–32 wide]; lateral-5 [35–44 long; 18–27 wide]; lateral-6 [38–56 long; 19–27 wide]; lateral-7 [39–50 long; 17–29 wide].

Figure 27. 

Testudacarus harrisi male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 27) — [544–612 long; 346–399 wide]. Primary sclerotization [504–578 long]. Gnathosomal bay [49–64 dorsal length; 119–133 ventral length; 48–54 wide]. Coxal field [387–443 long; 272–316 wide]. Coxa-I [210–229 long; 88–96 midlength]. Coxa-II + III [97–112 distance to top of coxa-II; 156–173 distance to top of coxa-III; 312–346 distance to bottom of coxa-III; 215–240 total length]. Coxa-IV [267–297 length to top; 120–154 total length]. Genital field [329–369 distance to top; 451–501 distance to bottom; 121–132 total length; 97–104 width; 208–238 distance from gnathosomal bay; 119–143 distance from coxa-I; 53–79 distance to excretory pore; 93–111 distance to caudad]. Genital skeleton [150–167 long; 77–92 wide]. Distance to excretory pore [504–578].

Legs — total leg and podomere lengths as follows: Leg-I [419–451 total; trochanter 45–56; basifemur 70–77; telofemur 58–68; genu 74–82; tibia 81–90; tarsus 79–84]. Leg-II [429–472 total; trochanter 47–52; basifemur 69–77; telofemur 56–63; genu 76–86; tibia 86–98; tarsus 93–99]. Leg-III [491–540 total; trochanter 49–53; basifemur 70–86; telofemur 59–66; genu 89–98; tibia 107–120; tarsus 114–124]. Leg-IV [665–739 total; trochanter 74–90; basifemur 95–109; telofemur 95–108; genu 128–138; tibia 139–150; tarsus 131–145].

Specific epithet after Samuel Benjamin Harris, the American author, philosopher, and co-founder of Project Reason. Sam Harris, more than any speaker and author, has challenged my (JCO) views and assumptions and kept me on my toes.

Eastern United States east of the Mississippi river, with southern limits in Florida.

Holotype (1♀): Pennsylvania, USA: 1♀ from Fayette County, Ohiopyle State Park, Laurel Run, fishing access #2 off T798 (Meadow Run Rd) Ohiopyle State Park (39°50'58.00"N, 79°30'51.00"W), 10 August 2014, by MJ Skvarla, MS14-0810-005 (Specimen 143645 – DNA#2071); Paratypes (8♀, 7♂): Mississippi, USA: (allotype) 1♂ from Tishomingo County, Tishomingo State Park, Rocky Quarry Branch, beside road just outside park entrance (34°36'43.00"N, 88°12'4.00"W), 20 September 2009, by IM Smith, IMS090115 (Specimen 146784 – DNA#2202); 3♀ and 4♂ from Tishomingo County, Tishomingo State Park, Rocky Quarry Branch, beside road just outside park entrance (34°36'43.00"N, 88°12'4.00"W), 20 September 2009, by IM Smith, IMS090115; 2♀ and 2♂ from Tishomingo County, Tishomingo State Park, Rocky Quarry Branch, (34°36’” N, 88°11'W), 18 September 1991, by IM Smith, IMS910049; Pennsylvania, USA: 2♀ from Fayette County, State Game Lands #51, Dunbar Creek, off Furnace Hill Road east of Dunbar (39°57'50.00"N, 79°35'8.70"W), 10 August 2014, MJ Skvarla, MS14-0810-001; Alabama, USA: 1♀ from DeKalb County, Desoto State Park, beside Trail Y (Yellow) (34°29'N, 85°32'W), 26 September 1992, by IM Smith, IMS920053A.

Holotype (1♀), allotype (1♂), and six paratypes (3♀, 3♂) deposited at CNC; eight paratypes (5♀, 3♂) at ACUA.

Both Testudacarus dennetti and T. dawkinsi have dorsal plates with uniform pores (unlike T. hitchensi) and anterio-lateral platelets with color restricted to the posterior half (unlike T. harrisi). However, they can be distinguished based on size. Testudacarus dennetti females and males have dorsal lengths less than 575 and 450 µm, respectively. Testudacarus dawkinsi females and males have dorsal lengths greater than 600 and 475 µm, respectively.

Female (n=9) with characteristics of the genus with following specifications.

Gnathosoma — Subcapitulum [139–152 ventral length; 85–97 dorsal length; 85–91 tall] ovoid with short rostrum. Chelicerae [117–126 long] unmodified with lightly curved fangs [24–28 long]. Pedipalp [168–189 long] unmodified. Trochanter [23–27 long; 28–31 wide]. Femur [42–52 long; 33–35 wide]. Genu [35–41 long; 25–32 wide]. Tibia [45–52 long; 17–22 wide]. Tarsus [18–20 long; 8–12 wide].

Dorsum (Fig. 28) — [473–558 long; 368–429 wide] round to ovoid. Dorsal plate [348–459 long; 353–442 wide]. Primary sclerotization [319–400 long]. Dorso-glandularia-4 [121–150 apart] lateral to [16–41] and around the anterior tips of muscle scars. Platelets violet to blue or colorless. Anterio-medial platelet [115–128 long; 65–83 wide] colorless rounded trapezoid noticeably smaller than anterio-lateral platelets. Anterio-lateral platelets [150–171 long; 68–78 wide] with violet to blue restricted to posterior half or third of the platelet. Lateral platelets as follows: lateral-1 [36–48 long; 29–44 wide]; lateral-2 [96–129 long; 24–37 wide]; lateral-3 [26–44 long; 14–27 wide]; lateral-4 [68–95 long; 19–39 wide]; lateral-5 [39–56 long; 13–32 wide]; lateral-6 [65–81 long; 16–34 wide]; lateral-7 [42–69 long; 15–30 wide].

Figure 28. 

Testudacarus dennetti female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 28) — [627–738 long; 411–474 wide] round to ovoid. Primary sclerotization [534–600 long] violet to blue. Gnathosomal bay [61–70 dorsal length; 125–142 ventral length; 48–57 wide]. Coxal field [406–438 long; 286–320 wide]. Coxa-I [216–236 long; 89–103 midlength]. Coxa-II + III [103–116 distance to top of coxa-II; 164–171 distance to top of coxa-III; 298–321 distance to bottom of coxa-III; 195–208 total length]. Coxa-IV [281–301 distance to top; 125–142 total length]. Genital field [279–304 distance to top; 416–455 distance to bottom; 137–151 total length; 110–123 width; 149–170 distance from gnathosomal bay; 54–75 distance from coxa-I; 160–185 distance to excretory pore; 211–295 distance to caudad]. Distance to excretory pore [581–640].

Legs — violet to blue and orange. Total leg and podomere lengths as follows: Leg-I [431–463 total; trochanter 48–58; basifemur 70–78; telofemur 59–65; genu 77–84; tibia 85–93; tarsus 81–88]. Leg-II [455–487 total; trochanter 51–56; basifemur 72–79; telofemur 57–64; genu 80–88; tibia 92–102; tarsus 98–109]. Leg-III [538–572 total; trochanter 54–59; basifemur 73–83; telofemur 62–67; genu 95–106; tibia 114–127; tarsus 128–134]. Leg-IV [641–768 total; trochanter 84–89; basifemur 96–115; telofemur 102–110; genu 137–144; tibia 147–163; tarsus 142–158].

Male (n=7) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [125–134 ventral length; 80–86 dorsal length; 74–83 tall]. Chelicerae [100–115 long]. Fangs [24–28 long]. Pedipalp [164–179 long]. Trochanter [22–24 long; 26–29 wide]. Femur [44–50 long; 30–35 wide]. Genu [36–43 long; width 25–28 wide]. Tibia [44–49 long; 17–20 wide]. Tarsus [17–19 long; 9–10 wide].

Dorsum (Fig. 29) — [408–440 long; 333–351 wide]. Dorsal plate [333–370 long; 268–305 wide]. Dorso-glandularia-4 [98–131 apart] lateral to [15–32] and far anterior to [46–62] muscle scars. Anterio-medial platelet [104–123 long; 60–66 wide]. Anterio-lateral platelets [133–153 long; 59–69 wide]. Lateral platelets as follows: lateral-1 [29–35 long; 25–31 wide]; lateral-2 [80–101 long; 24–32 wide]; lateral-3 [27–35 long; 18–21 wide]; lateral-4 [56–78 long; 21–28 wide]; lateral-5 [32–42 long; 20–25 wide]; lateral-6 [46–54 long; 23–25 wide]; lateral-7 [30–47 long; 19–23 wide].

Figure 29. 

Testudacarus dennetti male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 29) — [537–570 long; 352–370 wide]. Primary sclerotization [498–536 long]. Gnathosomal bay [55–61 dorsal length; 110–129 ventral length; 42–53 wide]. Coxal field [378–417 long; 275–292 wide]. Coxa-I [195–219 long; 80–95 midlength]. Coxa-II + III [84–103 distance to top of coxa-II; 141–165 distance to top of coxa-III; 299–327 distance to bottom of coxa-III; 215–236 total length]. Coxa-IV [261–284 length to top; 117–133 total length]. Genital field [322–341 distance to top; 443–471 distance to bottom; 121–130 total length; 96–103 width; 207–222 distance from gnathosomal bay; 120–136 distance from coxa-I; 54–66 distance to excretory pore; 85–100 distance to caudad]. Genital skeleton [152–169 long; 80–95 wide]. Distance to excretory pore [498–536].

Legs — total leg and podomere lengths as follows: Leg-I [414–434 total; trochanter 47–54; basifemur 67–73; telofemur 55–62; genu 72–79; tibia 81–85; tarsus 79–85]. Leg-II [432–450 total; trochanter 48–54; basifemur 66–72; telofemur 54–61; genu 73–80; tibia 88–91; tarsus 96–99]. Leg-III [478–525 total; trochanter 49–58; basifemur 66–76; telofemur 58–64; genu 83–93; tibia 102–114; tarsus 114–126]. Leg-IV [658–685 total; trochanter 76–86; basifemur 85–103; telofemur 90–100; genu 124–130; tibia 140–141; tarsus 133–140].

Specific epithet dennetti after Daniel Clement Dennett III, the American philosopher, writer, and cognitive scientist. Dennett’s work has been the focus of many late night debates in close social circles just as he adds the necessary philosophical spice to the New Athiests.

Eastern United States east of the Mississippi River, with southern limits in Florida.

Holotype (1♀): New York, USA: 1♀ from Franklin County, Little Aldo Creek, Little Aldo Creek trail from Keese Mill Rd (44°25'32.00"N, 74°20'43.00"W), 19 July 2013, by AJ Radwell and C Milewski, AJR13-0719-205 (Specimen 141897 – DNA#1501); Paratypes (5♀, 9♂): Tennessee, USA: (allotype) 1♂ from Sevier County, Great Smoky Mountains National Park, Crosby Creek, Cosby Recreation Area beside Cosby Campground Road 0.3km from Route 321 (35°46'54.00"N, 83°13'2.00"W), 16 September 2010, by IM Smith, IMS100140 (Specimen 146744 – DNA#2156); 1♂ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'47.00"N, 83°31'51.00"W), 10 September 2010, by IM Smith, IMS100125; 2♀ and 1♂ from Sevier County, Great Smoky Mountains National Park, Crosby Creek, Cosby Recreation Area beside Cosby Campground Road 0.3km from Route 321 (35°46'54.00"N, 83°13'2.00"W), 16 September 2010, by IM Smith, IMS100140; 1♂ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'47.00"N, 83°31'52.00"W), 24 September 2010, by IM Smith, IMS100158; 1♂ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'48.00"N, 83°31'53.00"W), 3 September 2009, by IM Smith, IMS090095; 1♀ and 1♂ from Sevier County, Great Smoky Mountains National Park, Bullhead Branch, Sugarlands Nature Trail off Route 441/71 (35°40'47.00"N, 83°31'52.00"W), 7 September 2009, by IM Smith, IMS090101; 1♀ and 2♂ from Sevier County, Great Smoky Mountains National Park, Cosby Creek, beside road to Cosby Campground at Gabes Mountain Trailhead (35°45'27.00"N, 83°12'36.00"W), 19 September, by IM Smith, IMS050093A; Virginia, USA: 1♂ from Alleghany County, Simpson Creek, Longdale Furnace beside Route 850 2.2 km northeast of I-64 overpass (37°49'41.00"N, 79°39'30.00"W), 14 August 2008, by IM Smith, IMS080044; North Carolina, USA: 1♀ from Macon County, Rainbow Springs, beside Forest Road 67 4.4 km south of Standing Indian Campground (35°3'6.00"N, 83°30'45.00"W), 1 July 2006, by IM Smith, IMS060040.

Holotype (1♀), allotype (1♂), and six paratypes (3♀, 3♂) deposited at CNC; seven paratypes (2♀, 5♂) at ACUA.

Both Testudacarus dennetti and T. dawkinsi have dorsal plates with uniform pores (unlike T. hitchensi) and anterio-lateral platelets with color restricted to the posterior half (unlike T. harrisi). However, they can be distinguished based on size. Testudacarus dennetti females and males have dorsal lengths less than 575 and 450 µm, respectively. Testudacarus dawkinsi females and males have dorsal lengths greater than 600 and 475 µm, respectively.

Female (n=6) with characteristics of the genus with following specifications.

Gnathosoma — Subcapitulum [160–168 ventral length; 102–105 dorsal length; 91–95 tall] ovoid with short rostrum. Chelicerae [136–141 long] unmodified with lightly curved fangs [29–30 long]. Pedipalp [188–193 long] unmodified. Trochanter [26–29 long; 28–32 wide]. Femur [50–54 long; 35–37 wide]. Genu [39–41 long; 30–33 wide]. Tibia [51–52 long; 19–22 wide]. Tarsus [19–20(–21) long; 9–11 wide].

Dorsum (Fig. 30) — [615–640 long; 475–501 wide] round to ovoid. Dorsal plate [497–528 long; 402–421 wide]. Primary sclerotization [421–453 long] violet to blue. Dorso-glandularia-4 [136–171 apart] lateral to [23–48] and around the anterior tips of muscle scars. Platelets violet to blue or colorless. Anterio-medial platelet [132–153 long; 80–102 wide] colorless rounded trapezoid noticeably smaller than anterio-lateral platelets. Anterio-lateral platelets [170–179 long; 81–91 wide] with violet to blue restricted to posterior half or third of the platelet. Lateral platelets as follows: lateral-1 [52–56 long; 44–49 wide]; lateral-2 [117–138 long; 31–46 wide]; lateral-3 [29–46 long; 20–26 wide]; lateral-4 [95–129 long; 33–38 wide]; lateral-5 [57–68 long; 32–36 wide]; lateral-6 [79–99 long; 32–43 wide]; lateral-7 [62–76 long; 32–39 wide].

Figure 30. 

Testudacarus dawkinsi female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 30) — [790–798; 510–534 wide] round to ovoid, fully sclerotized, and with anterior area of primary sclerotization [620–654 long] and posterior area of secondary sclerotization, violet to blue. Gnathosomal bay [77–84 dorsal length; 148–152 ventral length; 51–66 wide]. Coxal field [473–495 long; 330–368 wide]. Coxa-I [250–266 long; 100–114 midlength]. Coxa-II + III [119–125 distance to top of coxa-II; 188–195 distance to top of coxa-III; 350–370 distance to bottom of coxa-III; 229–245 total length]. Coxa-IV [325–343 distance to top; 144–155 total length]. Genital field [329–343 distance to top; 490–501 distance to bottom; 150–162 total length; 122–137 width; 181–194 distance from gnathosomal bay; 69–90 distance from coxa-I; 191–208 distance to excretory pore; 293–304 distance to caudad]. Distance to excretory pore [682–707].

Legs — violet to blue. Total leg and podomere lengths as follows: Leg-I [487–500 total; trochanter 57–63; basifemur 84–86; telofemur 67–73; genu 90–94; tibia 94–99; tarsus 87–94]. Leg-II [532–548 total; trochanter 58–65; basifemur 84–89; telofemur 67–72; genu 94–99; tibia 107–113; tarsus 110–116]. Leg-III [599–629 total; trochanter 63–68; basifemur 88–97; telofemur 73–76; genu 107–117; tibia 128–138; tarsus 134–140]. Leg-IV [830–861 total; trochanter 83–104; basifemur 113–130; telofemur 119–130; genu 156–164; tibia 172–181; tarsus 164–175].

Male (n=9) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [143–156 ventral length; 89–97 dorsal length; 81–91 tall]. Chelicerae [113–129 long]. Fangs [26–30 long]. Pedipalp [180–195 long]. Trochanter [24–30 long; 26–29 wide]. Femur [50–53 long; 33–36 wide]. Genu [38–43 long; width 27–29 wide]. Tibia [47–52 long; 19–22 wide]. Tarsus [17–20 long; 9–10 wide].

Dorsum (Fig. 31) — [491–540 long; 368–421 wide]. Dorsal plate [401–443 long; 322–364 wide]. Dorso-glandularia-4 [101–143 apart] lateral to [6–35] and far anterior to [43–81] muscle scars. Anterio-medial platelet [121–144 long; 73–83 wide]. Anterio-lateral platelets [156–173 long; 74–80 wide]. Lateral platelets as follows: lateral-1 [37–46 long; 33–43 wide]; lateral-2 [95–115 long; 30–43 wide]; lateral-3 [33–48 long; 19–23 wide]; lateral-4 [75–95 long; 27–32 wide]; lateral-5 [40–50 long; 22–28 wide]; lateral-6 [58–69 long; 24–33 wide]; lateral-7 [42–56 long; 23–27 wide].

Figure 31. 

Testudacarus dawkinsi male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 31) — [618–683 long; 395–454 wide]. Primary sclerotization [583–641 long]. Gnathosomal bay [59–78 dorsal length; 139–153 ventral length; 50–61 wide]. Coxal field [434–484 long; 305–329 wide]. Coxa-I [227–256 long; 86–106 midlength]. Coxa-II + III [106–122 distance to top of coxa-II; 171–194 distance to top of coxa-III; 346–388 distance to bottom of coxa-III; 240–269 total length]. Coxa-IV [303–334 length to top; 130–154 total length]. Genital field [367–419 distance to top; 502–556 distance to bottom; 135–146 total length; 108–120 width; 224–267 distance from gnathosomal bay; 135–171 distance from coxa-I; 70–93 distance to excretory pore; 105–134 distance to caudad]. Genital skeleton [170–173 long; 85–105 wide]. Distance to excretory pore [583–641].

Legs — total leg and podomere lengths as follows: Leg-I [452–497 total; trochanter 52–63; basifemur 78–87; telofemur 63–72; genu 84–92; tibia 89–99; tarsus 84–93]. Leg-II [486–519 total; trochanter 53–61; basifemur 77–85; telofemur 59–70; genu 87–91; tibia 102–107; tarsus 101–112]. Leg-III [551–588 total; trochanter 54–60; basifemur 81–90; telofemur 66–73; genu 100–107; tibia 118–129; tarsus 126–134]. Leg-IV [752–796 total; trochanter 85–94; basifemur 99–120; telofemur 107–117; genu 143–146; tibia 158–163; tarsus 148–158].

Eastern United States east of the Mississippi River, with southern limits in Florida.

Specific epithet dawkinsi after Clinton Richard Dawkins, the English evolutionary biologist and writer. Dawkins has proven repeatedly that one can change the world as a biologist by day and keep going as a free-thinker by night.

Holotype (1♀): California, USA: 1♀ from Santa Cruz County, Waddell Creek, 29-30 June 1933, by PR Needham, RM330008

Other (9♀, 8♂): Oregon, USA: 1♀ and 1♂ from Lincoln County, Siuslaw National Forest, Lord Creek, (44°14'24.00"N, 123°46'11.00"W), 8 August 2013, by JC O’Neill and WA Nelson, JNOW13-0808-002; 3♀ and 4♂ from Lane County, Cape Perpetua, Cape Perpetua Campground (44°16'51.00"N, 124°5'38.00"W), 15 September 2004, by IM Smith, IMS040077; 1♀ and 1♂ from Lane County, Rock Creek, Rock Creek Campground off Route 101 between Heceta Head and Yachats (44°11'6.00"N, 124°6'34.00"W), 14 September 2004, by IM Smith, IMS040076; 1♀ from Lane County, Cape Creek, Cape Perpetua, Cape Perpetua Campground (44°16'51.00"N, 124°5'38.00"W), 24 June 2010, by IM Smith, IMS100083; 1♀ and 1♂ from Curry County, Port Orford, beside road from Humbug Mountain State Park to McGribble Campground (Forest Road 5002) 5.3 km from Route 101 (42°42'11.00"N, 124°23'54.00"W), 25 June 1976, by IM Smith, IMS760161; 1♀ from Curry County, Port Orford, beside road from Humbug Mountain State Pk to McGribble Campground (Forest Road 5002) 4.6 km from Route 101 (42°42'3.00"N, 124°24'21.00"W), 17 June 2010, by IM Smith , IMS100070; 1♀ from Curry County, Siskiyou National Forest, North Fork of Foster Creek, beside Road #33 between Powers and Agness (42°39'N, 124°4'W), 2 July 1983, IMS 830019; Washington, USA: 1♂ from Kittitas County, Wenatchee National Forest, Squawk Creek, (47°16'51.00"N, 120°41'53.00"W), 31 July 2013, by JC O’Neill, WA Nelson, JNOW13-0731-002.

Holotype (1♀) deposited at CNC.

Resembling most Testudacarus smithi, these mites differ by shape, color, and several other characters. Most notably, T. americanus are elliptical and colorless to peach and have a small cheliceral fang (<33 µm) while T. smithi are rounded and are grey to colorless with large cheliceral fangs (>40 µm).

Female (n=10) with characteristics of the genus with following specifications.

Gnathosoma — Subcapitulum [180–199 ventral length; 108–121 dorsal length; 110–124 tall] ovoid with short rostrum and colorless. Chelicerae [148–173 long] unmodified with lightly curved fangs [30–33 long]. Pedipalp [209–236 long] unmodified. Trochanter [29–37 long; 31–33 wide]. Femur [54–59 long; 38–45 wide]. Genu [47–60 long; 32–36 wide]. Tibia [53–59 long; 21–24 wide]. Tarsus [19–23 long; 9–13 wide].

Dorsum (Fig. 33) — [826–890 long; 570–688 wide] ovoid to oblong and colorless with a peach tint. Dorsal plate [696–765 long; 497–561 wide]. Primary sclerotization [626–710 long]. Dorso-glandularia-4 [221–260 apart] anterior to [0–35] and lateral to [50–67] muscle scars. Anterio-medial platelet [198–241 long; 85–105 wide] broad, thin, very slightly rounded trapezoid similar in size to anterio-lateral platelets. Anterio-lateral platelets [219–253 long; 103–130 wide]. Lateral platelets as follows: lateral-1 [562–78 long; 48–59 wide]; lateral-2 [175–191 long; 41–59 wide]; lateral-3 [50–81 long; 22–45 wide]; lateral-4 [137–180 long; 40–56 wide]; lateral-5 [62–78 long; 40–55 wide]; lateral-6 [140–153 long; 39–56 wide]; lateral-7 [76–102 long; 36–52 wide].

Figure 33. 

Testudacarus americanus female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 33) — [973–1095 long; 620–731 wide] ovoid to oblong and colorless. Primary sclerotization [828–934 long]. Gnathosomal bay [83–100 dorsal length; 171–196 ventral length; 67–82 wide]. Coxal field [555–615 long; 393–478 wide] noticeably small in relation to the venter when compared to other Testudacarines. Coxa-I [279–326 long; 106–141 midlength]. Coxa-II + III [128–145 distance to top of coxa-II; 211–256 distance to top of coxa-III; 391–444 distance to bottom of coxa-III; 263–304 total length]. Coxa-IV [368–427 distance to top; 170–198 total length]. Genital field [375–438 distance to top; 570–641 distance to bottom; 188–203 total length; 157–170 width; 203–259 distance from gnathosomal bay; 94–118 distance from coxa-I; 282–325 distance to excretory pore; 403–472 distance to caudad]. Eggs [182–200 long; 1–2 eggs]. Distance to excretory pore [880–964].

Legs — colorless. Total leg and podomere lengths as follows: Leg-I [542–604 total; trochanter 68–75; basifemur 91–108; telofemur 75–83; genu 95–111; tibia 108–122; tarsus 101–112]. Leg-II [548–611 total; trochanter 63–72; basifemur 95–108; telofemur 70–83; genu 93–103; tibia 107–123; tarsus 110–127]. Leg-III [595–683 total; trochanter 66–73; basifemur 99–108; telofemur 76–84; genu 103–124; tibia 127–150; tarsus 124–153]. Leg-IV [854–987 total; trochanter 94–112; basifemur 125–152; telofemur 134–154; genu 170–192; tibia 171–216; tarsus 154–181].

Male (n=8) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [164–178 ventral length; 97–109 dorsal length; 96–114 tall]. Chelicerae [132–152 long]. Fangs [28–33 long]. Pedipalp [202–222 long]. Trochanter [28–33 long; 28–33 wide]. Femur [50–58 long; 38–42 wide]. Genu [50–55 long; width 30–35 wide]. Tibia [53–58 long; 20–22 wide]. Tarsus [18–23 long; 11–13 wide].

Dorsum (Fig. 34) — [678–755 long; 475–534 wide]. Dorsal plate [573–645 long; 405–463 wide]. Dorso-glandularia-4 [180–208 apart] lateral to [40–58] and well anterior to [53–90] muscle scars. Anterio-medial platelet [194–220 long; 82–103 wide]. Anterio-lateral platelets [198–227 long; 101–120 wide] without noticeable bump. Lateral platelets as follows: lateral-1 [52–62 long; 35–48 wide]; lateral-2 [125–164 long; 30–45 wide]; lateral-3 [47–68 long; 21–31 wide]; lateral-4 [92–120 long; 31–41 wide]; lateral-5 [55–68 long; 20–39 wide]; lateral-6 [87–125 long; 25–43 wide]; lateral-7 [47–63 long; 26–38 wide].

Figure 34. 

Testudacarus americanus male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 34) — [840–893 long; 516–605 wide]. Primary sclerotization [763–841 long]. Gnathosomal bay [76–93 dorsal length; 150–174 ventral length; 64–89 wide]. Coxal field [520–594 long; 361–402 wide]. Coxa-I [276–291 long; 112–126 midlength]. Coxa-II + III [112–130 distance to top of coxa-II; 190–226 distance to top of coxa-III; 413–452 distance to bottom of coxa-III; 298–324 total length]. Coxa-IV [366–395 length to top; 150–203 total length]. Genital field [448–492 distance to top; 593–638 distance to bottom; 130–159 total length; 120–138 width; 292–323 distance from gnathosomal bay; 168–200 distance from coxa-I; 181–201 distance to excretory pore; 236–280 distance to caudad]. Genital skeleton [163–178 long; 80–88 wide]. Distance to excretory pore [780–833].

Legs — total leg and podomere lengths as follows: Leg-I [501–560 total; trochanter 57–64; basifemur 85–99; telofemur 70–80; genu 90–102; tibia 101–113; tarsus 95–108]. Leg-II [508–567 total; trochanter 58–67; basifemur 88–96; telofemur 67–74; genu 83–99; tibia 101–117; tarsus 105–119]. Leg-III [554–615 total; trochanter 59–63; basifemur 83–98; telofemur 70–76; genu 97–115; tibia 117–138; tarsus 119–132]. Leg-IV [526–882 total; trochanter 79–103; basifemur 116–130; telofemur 121–135; genu 157–175; tibia 171–197; tarsus 149–166].

Western North America within and west of the Rocky Mountains. California (Marshall 1943), Wyoming (Bergstrom 1953), Colorado (Young 1969), Vancouver Island (Smith 1982), Mexico State (Cramer 1992).

Having examined the type material, we suggest that T. americanus galloi Habeeb, 1969 is simply a teneral T. americanus that Habeeb confused for an “a-typical” T. americanus.

Holotype (1♀) Oregon, USA: 1♀ from Douglas County, Rouge River National Forest, Muir Creek, (43°2'53.00"N, 122°21'4.00"W), 12 August 2013, by JC O’Neill and WA Nelson, JNOW13-0812-004 (Specimen 141885 – DNA#1791); Other examined but not measured (1♀): Washington, USA: 1♀ from Snohomish County, Mount Baker National Forest, tributary of South Fork of Sauk River, (48°1'40.00"N, 121°26'24.00"W), 28 July 2013, JC O’Neill and WA Nelson, JNOW13-0728-003

Holotype (1♀) deposited at CNC.

These are the largest known testudacarines and also differ further from their complex in having a smaller, more rounded anterio-medial platelet.

Female (n=1) with characteristics of genus with following specifications.

Gnathosoma — Subcapitulum [245 ventral length; 133 dorsal length; 130 tall] ovoid with short rostrum. Chelicerae [200 long] unmodified with lightly curved fangs [40 long]. Pedipalp [259 long] unmodified. Trochanter [38 long; 43 wide]. Femur [70 long; 50 wide]. Genu [63 long; 40 wide]. Tibia [63 long; 28 wide]. Tarsus [25 long; 15 wide].

Dorsum (Fig. 35) — [918 long; 645 wide] ovoid to oblong. Dorsal plate [758 long; 566 wide]. Primary sclerotization [603 long] light pink to colorless. Dorso-glandularia-4 [232 apart] lateral to [63] and around muscle scar midline. Platelets colorless. Anterio-medial platelet [201 long; 123 wide]. Anterio-lateral platelets [237 long; 134 wide]. Lateral platelets as follows: lateral-1 [65 long; 55 wide]; lateral-2 [173 long; 46 wide]; lateral-3 [67 long; 24 wide]; lateral-4 [173 long; 46 wide]; lateral-5 [91 long; 47 wide]; lateral-6 [149 long; 54 wide]; lateral-7 [93 long; 46 wide].

Figure 35. 

Testudacarus kirkwoodae female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 35) — [1045 long; 853 wide] round to ovoid and colorless. Primary sclerotization [752 long]. Gnathosomal bay [123 dorsal length; 151 ventral length; 69 wide]. Coxal field [551 long; 484 wide] proportionally small compared to venter. Coxa-I [294 long; 144 midlength]. Coxa-II + III [114 distance to top of coxa-II; 202 distance to top of coxa-III; 393 distance to bottom of coxa-III; 279 total length]. Coxa-IV [376 distance to top; 175 total length]. Genital field [373 distance to top; 578 distance to bottom; 205 total length; 69 width; 222 distance from gnathosomal bay; 78 distance from coxa-I; 272 distance to excretory pore; 467 distance to caudad]. Distance to excretory pore [850].

Legs — colorless. Total leg and podomere lengths as follows: Leg-I [620 total; trochanter 70; basifemur 119; telofemur 89; genu 101; tibia 119; tarsus 121]. Leg-II [681 total; trochanter 77; basifemur 113; telofemur 90; genu 117; tibia 145; tarsus 140]. Leg-III [756 total; trochanter 70; basifemur 116; telofemur 93; genu 144; tibia 165; tarsus 167]. Leg-IV [1081 total; trochanter 136; basifemur 148; telofemur 159; genu 207; tibia 224; tarsus 208].

Male (n=0) unknown.

Specific epithet kirkwoodae after my (JCO) mother’s maiden name. It was her bringing me to the leafcutter ant exhibit at our local science center that helped me become interested in biology as a child, and her endless support and advice that helped me finish my education.

As it is likely that this species represents a cryptic species complex (with 4.5% COI divergence between the two available specimens), measurements were included from only one specimen (the other highlighted in red in Fig. 32 was excluded). This was done so measurements would remain useful if more species were diagnosed in the future. Measurments were also not included from the other specimen because it is teneral and badly damaged and would therefore prove poor for any description. While more than a single specimen is certainly desired for new descriptions, the included specimen has unique morphological characters such as its large size (it is larger than any other testudacarine, including species from Asia), and has strong support as a unique clade using COI. Therefore, this single specimen is unique enough that we are comfortable describing it.

Only one specimen know from Douglas County, Oregon.

Holotype (1♀): California, USA: 1♀ Humboldt County, Willow Creek, Willow Creek Campground off Rt. 299 (40°54'17.00"N, 123°42'21.00"W), 14 June 2010, by IM Smith, IMS100065 (Specimen 146762 – DNA#2177); Paratype (1♀, 2♂): California, USA: (allotype) 1♂ Humboldt County, Willow Creek, Willow Creek Campground off Rt. 299 (40°54'17.00"N, 123°42'21.00"W), 14 June 2010, by IM Smith, IMS100065 (Specimen 146763 – DNA#2178); 1♂ Humboldt County, Willow Creek, Willow Creek Campground off Rt. 299 (40°54'17.00"N, 123°42'21.00"W), 14 June 2010, by IM Smith, IMS100065; Oregon, USA: 1♀ from Curry County, Port Orford, beside Elk River Road 9.0 km east of Elk River Fish Hatchery (42°42'22.00"N, 124°20'28.00"W), 22 June 2010, by IM Smith, IMS100080.

Holotype (1♀) and allotype (1♂) deposited at CNC; two paratypes (1♀, 1♂) at ACUA.

These mites differ from the rest of the complex in having a dorsally “covered” gnathosomal bay (short doral gnathosomal bay length) and an elongate gnathosoma with a long rostrum that exteneds below the gnathosoma ventral surface.

Female (n=2) with characteristics of genus with following specifications.

Gnathosoma (Fig. 36) — Subcapitulum [244–250 ventral length; 150–155 dorsal length; 89–97 tall] elongate with long rostrum extending below ventral surface; colorless. Chelicerae [210–220 long] unmodified with lightly curved fangs [32–36 long]. Pedipalp [194–203 long] unmodified. Trochanter [24–25 long; 38–40 wide]. Femur [53–56 long; 42–43 wide]. Genu [44–45 long; 34–35 wide]. Tibia [50–54 long; 22–23 wide]. Tarsus [21–22 long; 9–10 wide].

Figure 36. 

Testudacarus hyporhynchus sp. n. gnathosoma (generalized).

Dorsum (Fig. 37) — [768–849 long; 634–668 wide] round to ovoid and mostly colorless. Dorsal plate [570–578 long; 645–693 wide]. Primary sclerotization [540–583 long] colorless to light pink. Dorso-glandularia-4 [230–252 apart] lateral to [44–61] and just anterior to [0–27] muscle scars. Platelets colorless. Anterio-medial platelet [230–252 long; 103–116 wide]. Anterio-lateral platelets [229–246 long; 117–123 wide]. Lateral platelets as follows: lateral-1 [66–83 long; 56–59 wide]; lateral-2 [156–188 long; 47–51 wide]; lateral-3 [60–84 long; 29–30 wide]; lateral-4 [139–150 long; 35–45 wide]; lateral-5 [79–93 long; 41–42 wide]; lateral-6 [129–144 long; 39–44 wide]; lateral-7 [76–88 long; 35–41 wide].

Figure 37. 

Testudacarus hyporhynchus female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 37) — [1001–1049 long; 700–728 wide] round to ovoid and colorless. Primary sclerotization [814–824 long]. Gnathosomal bay [14–20 dorsal length; 101–102 ventral length; 67–70 wide] short dorsally giving a “covered” appearance and ventrally ending anterior to leg-I insertion. Coxal field [590–616 long; 404–433 wide]. Coxa-I [298–316 long; 196–216 midlength]. Coxa-II + III [127–143 distance to top of coxa-II; 216–264 distance to top of coxa-III; 427–446 distance to bottom of coxa-III; 299–303 total length]. Coxa-IV [399–434 distance to top; 182–191 total length]. Genital field [410–424 distance to top; 627–642 distance to bottom; 217–218 total length; 174–189 width; 308–323 distance from gnathosomal bay; 108–112 distance from coxa-I; 252–270 distance to excretory pore; 374–407 distance to caudad] large. Distance to excretory pore [879–912].

Legs — colorless. Total leg and podomere lengths as follows: Leg-I [566–594 total; trochanter 68–74; basifemur 104–106; telofemur 81–85; genu 108–112; tibia 109–117; tarsus 95–102]. Leg-II [614–645 total; trochanter 75–78; basifemur 102–108; telofemur 77–79; genu 111–116; tibia 124–130; tarsus 125–135]. Leg-III [714–753 total; trochanter 74–79; basifemur 109–119; telofemur 88–94; genu 136–139; tibia 154–160; tarsus 152–161]. Leg-IV [952–961 total; trochanter 105–109; basifemur 142–144; telofemur 138–139; genu 191–192; tibia 199–201; tarsus 175–178].

Male (n=2) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma (Fig. 36) — Subcapitulum [222–239 ventral length; 136–151 dorsal length; 85–89 tall]. Chelicerae [203–218 long]. Fangs [33–34 long]. Pedipalp [195–200 long]. Trochanter [24–25 long; 36–38 wide]. Femur [55–58 long; 42–46 wide]. Genu [45–46 long; width 34–35 wide]. Tibia [50–54 long; 21–23 wide]. Tarsus [17–20 long; 8–9 wide].

Dorsum (Fig. 38) — [667–712 long; 548–582 wide]. Dorsal plate [546–616 long; 470–471 wide]. Dorso-glandularia-4 [192–212 apart] lateral to and well anterior to muscle scars [65–67 anterior to; 40–55 lateral to]. Anterio-medial platelet [244–249 long; 91–100 wide]. Anterio-lateral platelets [225–229 long; 111–116 wide]. Lateral platelets as follows: lateral-1 [49–61 long; 43–60 wide]; lateral-2 [135–152 long; 47–50 wide]; lateral-3 [53–60 long; 24–25 wide]; lateral-4 [135–144 long; 37–41 wide]; lateral-5 [60–75 long; 35–37 wide]; lateral-6 [101–105 long; 32–37 wide]; lateral-7 [59–63 long; 31–33 wide].

Figure 38. 

Testudacarus hyporhynchus male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 38) — [835–898 long; 625–626 wide]. Primary sclerotization [750–791 long]. Gnathosomal bay [16–20 dorsal length; 99–101 ventral length; 67–68 wide]. Coxal field [553–576 long; 403–408 wide]. Coxa-I [296–318 long; 195–218 midlength]. Coxa-II + III [116–127 distance to top of coxa-II; 212–234 distance to top of coxa-III; 430–473 distance to bottom of coxa-III; 314–346 total length]. Coxa-IV [377–413 length to top; 163–176 total length]. Genital field [473–506 distance to top; 636–681 distance to bottom; 163–175 total length; 149–151 width; 372–407 distance from gnathosomal bay; 177–188 distance from coxa-I; 129–134 distance to excretory pore; 200–217 distance to caudad]. Genital skeleton [180–187 long; 117–122 wide]. Distance to excretory pore [765–815].

Legs — total leg and podomere lengths as follows: Leg-I [593–606 total; trochanter 73–79; basifemur 111–114; telofemur 78–83; genu 107–111; tibia 113–118; tarsus 105–106]. Leg-II [635–645 total; trochanter 74–79; basifemur 102–104; telofemur 84–85; genu 115–120; tibia 129–132; tarsus 124–130]. Leg-III [724–726 total; trochanter 77–78; basifemur 109–116; telofemur 87–91; genu 136–137; tibia 155–156; tarsus 152–155]. Leg-IV [905–964 total; trochanter 107–118; basifemur 139–140; telofemur 129–142; genu 183–191; tibia 179–198; tarsus 167–176].

Specific epithet hyporhynchus (hypo-, G. under; rhynchus, G. snout) refers to the long rostrum that extends below the ventral surface of the gnathosoma.

Humbolt County, California and Curry County, Oregon.

Holotype (1♀): British Columbia, Canada: 1♀ from Vancouver Island, spring run, Lake Cowichan, beside North Shore Road 1.7 km north of town (48°49'29.00"N, 124°4'2.00"W), 1 July 2010, by IM Smith, IMS100091 (Specimen 146769 – DNA#2184); Paratypes (10♀, 12♂): British Columbia, Canada: (allotype) 1♂ from Vancouver Island, spring run, Lake Cowichan, beside North Shore Road 1.7 km north of town (48°49'29.00"N, 124°4'2.00"W), 1 July 2010, by IM Smith, IMS100091 (Specimen 146770 – DNA#2185); 2♂ from Vancouver Island, spring run, Lake Cowichan, beside North Shore Road 1.7 km north of town (48°49'29.00"N, 124°4'2.00"W), 1 July 2010, by IM Smith, IMS100091; 3♀ and 2♂ from Vancouver Island, Lake Cowichan, spring run, beside North Shore Road 1.7 km north of town (48°49'29.00"N, 124°4'13.00"W), 11 June 1979, by IM Smith, IMS790013A; 3♀ and 3♂ from Vancouver Island, Lake Cowichan, spring run, beside South Shore Road 2.3 km north of town (48°48'25.00"N, 124°5'13.00"W), 7 July 1976, by IM Smith, IMS760194; 3♀ and 2♂ from Vancouver Island, Port Alberni, beside road to Mount Arrowsmith Ski Area 11.6 km from Highway 4 (49°12'50.00"N, 124°36'18.00"W), 19 September 2004, by IM Smith, IMS040084A; 1♀ and 1♂ from Vancouver Island, Lake Cowichan, spring run, beside South Shore Road 2.3 km north of town (48°48'25.00"N, 124°5'13.00"W), 31 July 1979, by IM Smith, IMS790056; 1♂ from Vancouver Island, Lake Cowichan, spring run, beside South Shore Road 2.3 km north of town (48°48'25.00"N, 124°5'13.00"W), 26 July 1985, by IM Smith, IMS850122A.

Holotype (1♀), allotype (1♂), and eight paratypes (4♀, 4♂) deposited at CNC; thirteen paratypes (6♀, 7♂) at ACUA.

Resembling most Testudacarus americanus, these mites differ by shape, color, and several other characters. Most notably, T. americanus are elliptical and colorless to peach and have a small cheliceral fang (<33 µm) while T. smithi are rounded and are grey to colorless with large cheliceral fangs (>40 µm).

Female (n=10) with characteristics of genus with following specifications.

Gnathosoma — Subcapitulum [217–245 ventral length; 137–145 dorsal length; 125–152 tall] elliptical to ovoid with short rostrum. Chelicerae [190–205 long] unmodified with lightly curved fangs [40–45 long]; fangs characteristically large. Pedipalp [250–272 long] unmodified. Trochanter [37–45 long; 39–46 wide]. Femur [72–80 long; 52–61 wide]. Genu [55–61 long; 41–49 wide]. Tibia [57–66 long; 23–26 wide]. Tarsus [21–26 long; 11–13 wide].

Dorsum (Fig. 39) — [790–864 long; 619–683 wide] round to ovoid. Dorsal plate [643–705 long; 500–549 wide]. Primary sclerotization [541–596 long] grey-violet. Dorso-glandularia-4 [215–246 apart] lateral to [45–72] and just anterior to [0–21] muscle scars. Platelets colorless. Anterio-medial platelet [200–233 long; 103–128 wide] large slightly rounded trapezoid approaching size of anterio-lateral platelets. Anterio-lateral platelets [230–266 long; 125–149 wide]. Lateral platelets as follows: lateral-1 [69–82 long; 47–67 wide]; lateral-2 [127–154 long; 38–54 wide]; lateral-3 [37–65 long; 22–40 wide]; lateral-4 [156–185 long; 31–54 wide]; lateral-5 [80–102 long; 40–60 wide]; lateral-6 [106–158 long; 35–60 wide]; lateral-7 [73–103 long; 36–55 wide].

Figure 39. 

Testudacarus smithi female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 39) — [955–1047; 671–816 wide] round to ovoid and colorless. Primary sclerotization [742–814 long]. Gnathosomal bay [87–130 dorsal length; 164–216 ventral length; 78–105 wide]. Coxal field [578–641 long; 421–488 wide]. Coxa-I [302–361 long; 133–157 midlength]. Coxa-II + III [130–165 distance to top of coxa-II; 223–262 distance to top of coxa-III; 416–476 distance to bottom of coxa-III; 284–317 total length]. Coxa-IV [384–435 distance to top; 184–222 total length]. Genital field [390-455 distance to top; 595–657 distance to bottom; 201–217 total length; 173–184 width; 222–251 distance from gnathosomal bay; 84–102 distance from coxa-I; 229–298 distance to excretory pore; 332–429 distance to caudad]. Eggs [185–200 long; 1–3 eggs]. Distance to excretory pore [853–924].

Legs — colorless. Total leg and podomere lengths as follows: Leg-I [624–660 total; trochanter 74–81; basifemur 110–121; telofemur 84–95; genu 117–125; tibia 123–130; tarsus 108–116]. Leg-II [658–718 total; trochanter 75–87; basifemur 111–123; telofemur 83–95; genu 115–130; tibia 133–149; tarsus 133–145]. Leg-III [755–820 total; trochanter 78–89; basifemur 112–131; telofemur 88–102; genu 139–155; tibia 162–184; tarsus 165–182]. Leg-IV [1017–1058 total; trochanter 117–132; basifemur 139–150; telofemur 140–153; genu 197–210; tibia 212–228; tarsus 194–205].

Male (n=12) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [205–232 ventral length; 125–145 dorsal length; 124–133 tall]. Chelicerae [178–202 long]. Fangs [39–42 long]. Pedipalp [250–279 long]. Trochanter [37–42 long; 38–43 wide]. Femur [71–81 long; 52–60 wide]. Genu [55–65 long; width 40–47 wide]. Tibia [59–69 long; 23–25 wide]. Tarsus [21–27 long; 11–15 wide].

Dorsum (Fig. 40) — [682–790 long; 523–626 wide]. Dorsal plate [567–670 long; 440–521 wide] with minute amount of secondary sclerotization. Dorso-glandularia-4 [198–261 apart] roughly equal distance anterior to and lateral to muscle scars [39–92 anterior to; 42–72 lateral to]. Anterio-medial platelet [194–221 long; 99–108 wide]. Anterio–lateral [216–249 long; 116–135 wide]. Lateral platelets as follows: lateral-1 [62–79 long; 45–57 wide]; lateral-2 [114–150 long; 37–52 wide]; lateral-3 [30–67 long; 21–33 wide]; lateral-4 [123–160 long; 30–48 wide]; lateral-5 [67–90 long; 32–48 wide]; lateral-6 [91–121 long; 33–47 wide]; lateral-7 [50–79 long; 29–44 wide].

Figure 40. 

Testudacarus smithi male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 40) — [868–974; 575–730 wide]. Primary sclerotization [728–820 long]. Gnathosomal bay [79–126 dorsal length; 183–206 ventral length; 72–101 wide]. Coxal field [582–657 long; 394–468 wide]. Coxa-I [320–346 long; 131–146 midlength]. Coxa-II + III [132–158 distance to top of coxa-II; 233–264 distance to top of coxa-III; 470–515 distance to bottom of coxa-III; 327–370 total length]. Coxa-IV [378–436 length to top; 182–234 total length]. Genital field [490–545 distance to top; 675–742 distance to bottom; 185–210 total length; 150–166 width; 307–340 distance from gnathosomal bay; 170–200 distance from coxa-I; 90–137 distance to excretory pore; 177–244 distance to caudad]. Genital skeleton [245–272 long; 125–152 wide]. Distance to excretory pore [790–874]. Excretory pore characteristically well separated from line of secondary sclerotization.

Legs — total leg and podomere lengths as follows: Leg-I [617–679 total; trochanter 73–80; basifemur 110–122; telofemur 84–97; genu 116–129; tibia 118–138; tarsus 107–123]. Leg-II [664–743 total; trochanter 74–84; basifemur 110–125; telofemur 85–103; genu 118–137; tibia 134–157; tarsus 131–148]. Leg-III [753–841 total; trochanter 76–83; basifemur 111–133; telofemur 89–106; genu 138–158; tibia 163–186; tarsus 155–186]. Leg-IV [952–1098 total; trochanter 111–131; basifemur 136–154; telofemur 139–161; genu 189–217; tibia 189–238; tarsus 176–210].

Specific epithet smithi after Dr Ian Smith, the Canadian water mite researcher who collected the specimens needed for this description. Dr Smith has advanced water mite research in North America more than anyone else and without him, this work would have been impossible.

British Columbia, Canada.

Holotype (1♀): California, USA: 1♀ from Nevada County, Tahoe National Forest, Bear River, at Sierra Discovery day use area upstream from bridge (39°18'35.00"N, 120°39'56.00"W), 26 August 2013, by JR Fisher, JRF13-0826-001 (Specimen 146725 – DNA# 2135); Paratypes (2♀, 2♂): California, USA: (allotype) 1♂ (allotype)from Nevada County, Tahoe National Forest, Bear River, at Sierra Discovery day use area upstream from bridge (39°18'35.00"N, 120°39'56.00"W), 26 August 2013, by JR Fisher, JRF13-0826-001 (Specimen 146724 – DNA# 2134); 1♀ from Nevada County, Tahoe National Forest, Bear River, at Sierra Discovery day use area upstream from bridge (39°18'35.00"N, 120°39'56.00"W), 26 August 2013, by JR Fisher, JRF13-0826-001; 1♀ and 1♂ from Calaveras County, Calaveras Big Trees State Park, Big Trees River, (38°16'N, 120°16'W), 12 June 1976, by IM Smith, IMS760099; 1♂ from Mendocino County, Navarro River, Paul M. Dimmick Recreation Area beside Route 128 (39°`10'N, 123°38'W), 29 September 1993, by IM Smith, IMS9300026A.

Holotype (1♀) and allotype (1♂) deposited at CNC; four paratypes (2♀, 2♂) at ACUA.

These mites are smaller and more colorful than other species in the complex and therefore resemble most the T. minimus-like mites; however, mites of the T. minimus complex are even smaller and have a smaller (<140 µm, and far less than twice as wide as long), more rounded anterio-medial platelet, while these mites have a larger (>140 µm, and more than or nearly twice as wide as long) more rectangular anterio-medial platelet.

Female (n=3) with characteristics of genus with following specifications.

Gnathosoma — Subcapitulum [176–188 ventral length; 105–107 dorsal length; 100–103 tall] elliptical ovoid with short rostrum and colorless. Chelicerae [145–152 long] unmodified with lightly curved fangs [28–30 long]. Pedipalp [202–212 long] unmodified. Trochanter [30–31 long; 28–30 wide]. Femur [56–58 long; 40–41 wide]. Genu [43–47 long; 33–35 wide]. Tibia [52–55 long; 20–23 wide]. Tarsus [20–22 long; 9–10 wide].

Dorsum (Fig. 41) — [625–680 long; 483–550 wide] ovoid and mostly colorless. Dorsal plate [526–568 long; 410–475 wide]. Primary sclerotization [431–473 long] light pink to colorless. Dorso-glandularia-4 [192–246 apart] lateral to [45–58] and around muscle scar midline. Platelets colorless. Anterio-medial platelet [153–164 long; 83–93 wide] large trapezoid with nearly straight anterior margin. Anterio-lateral platelets [181–211 long; 88–91 wide]. Lateral platelets as follows: lateral-1 [46–52 long; 38–40 wide]; lateral-2 [132–148 long; 33–39 wide]; lateral-3 [50–69 long; 19–26 wide]; lateral-4 [107–112 long; 22–29 wide]; lateral-5 [61–86 long; 27–32 wide]; lateral-6 [112–128 long; 25–34 wide]; lateral-7 [31–77 long; 23–33 wide].

Figure 41. 

Testudacarus rollerae female: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 41) — [786–884 long; 548–644 wide] round to ovoid and colorless. Primary sclerotization [624–709 long]. Gnathosomal bay [81–96 dorsal length; 154–164 ventral length; 56–60 wide]. Coxal field [478–532 long; 335–394 wide]. Coxa-I [261–290 long; 106–126 midlength]. Coxa-II + III [122–137 distance to top of coxa-II; 198–224 distance to top of coxa-III; 363–395 distance to bottom of coxa-III; 237–257 total length]. Coxa-IV [346–385 distance to top; 132–148 total length]. Genital field [347–374 distance to top; 500–539 distance to bottom; 153–165 total length; 130–139 width; 193–210 distance from gnathosomal bay; 79–87 distance from coxa-I; 199–231 distance to excretory pore; 286–345 distance to caudad]. Eggs [165–178 long; 1 egg]. Distance to excretory pore [699–770].

Legs — colorless. Total leg and podomere lengths as follows: Leg-I [503–542 total; trochanter 65–68; basifemur 88–93; telofemur 70–81; genu 93–103; tibia 96–102; tarsus 89–97]. Leg-II [510–577 total; trochanter 52–74; basifemur 82–94; telofemur 66–78; genu 94–102; tibia 103–118; tarsus 106–112]. Leg-III [610–657 total; trochanter 64–71; basifemur 92–99; telofemur 73–82; genu 110–122; tibia 127–146; tarsus 137–141]. Leg-IV [843–914 total; trochanter 96–101; basifemur 116–125; telofemur 117–133; genu 166–185; tibia 181–195; tarsus 168–178].

Male (n=3) similar to female except for sexually dimorphic characters previously discussed and with following specifications.

Gnathosoma — Subcapitulum [160–170 ventral length; 98–109 dorsal length; 81–92 tall]. Chelicerae [132–140 long]. Fangs [27–29 long]. Pedipalp [184–190 long]. Trochanter [23–26 long; 29–30 wide]. Femur [50–51 long; 36–38 wide]. Genu [41–42 long; width 29–30 wide]. Tibia [49–52 long; 19–20 wide]. Tarsus [17–21 long; 8–9 wide].

Dorsum (Fig. 42) — [540–585 long; 412–433 wide]. Dorsal plate [444–487 long; 355–384 wide] with minute secondary sclerotization. Dorso-glandularia-4 [151–167 apart] roughly equal in distance anterior to [32–53] and lateral to [31–43] muscle scars. Anterio-medial platelet [151–158 long; 80–85 wide]. Anterio-lateral platelets [180–188 long; 84–91 wide]. Lateral platelets as follows: lateral-1 [44–49 long; 33–38 wide]; lateral-2 [109–114 long; 31–35 wide]; lateral-3 [50–63 long; 19–22 wide]; lateral-4 [75–91 long; 18–29 wide]; lateral-5 [60–65 long; 22–29 wide]; lateral-6 [66–82 long; 20–33 wide]; lateral-7 [52–61 long; 22–30 wide].

Figure 42. 

Testudacarus rollerae male: (Left) dorsum; (Right) venter. Scale: 100 µm.

Venter (Fig. 42) — [698–740 long; 453–544 wide]. Primary sclerotization [623–655 long]. Gnathosomal bay [71–80 dorsal length; 138–147 ventral length; 54–60 wide]. Coxal field [475–484 long; 325–374 wide]. Coxa-I [253–263 long; 111–117 midlength]. Coxa-II + III [118–131 distance to top of coxa-II; 185–198 distance to top of coxa-III; 382–396 distance to bottom of coxa-III; 251–274 total length]. Coxa-IV [337–356 length to top; 127–139 total length]. Genital field [406–426 distance to top; 547–570 distance to bottom; 142–146 total length; 114–123 width; 263–280 distance from gnathosomal bay; 152–164 distance from coxa-I; 75–91 distance to excretory pore; 148–176 distance to caudad]. Genital skeleton [190–215 long; 110–112 wide]. Distance to excretory pore [623–655].

Legs — total leg and podomere lengths as follows: Leg-I [472 total; trochanter 59–60; basifemur 83–91; telofemur 66–71; genu 88–91; tibia 91–97; tarsus 83–88]. Leg-II [496–515 total; trochanter 61–66; basifemur 84–87; telofemur 65–69; genu 85–97; tibia 96–108; tarsus 102–107]. Leg-III [554–593 total; trochanter 62–69; basifemur 84–89; telofemur 65–74; genu 100–110; tibia 117–126; tarsus 125–136]. Leg-IV [784–822 total; trochanter 80–95; basifemur 109–116; telofemur 114–115; genu 155–162; tibia 164–174; tarsus 155–162].

Specific epithet rollerae after Elizabeth Ashley Roller, my (JCO) life partner.

Reported from only two counties (Mendocino and Nevada) in California.