Search

Idagona westcotti Buckett & Gardner 1967 Idagona westcotti Buckett & Gardner 1967, p. 120; Shear, 1972, p. 270, 2007, p. 4. New records: IDAHO: Blaine Co., Bear Trap Cave, 18.2 mi N of Minidoka on Arco-Minidoka Road, Craters of the Moon National Monument, 42.9840°N, - 113.351°W, elev. 1445 m., 1 July 2007, f. Lincoln Co., Diamond Cave, 43.1750°N, - 114.3568°W, elev. 1430 m., March 2017, R. Winton, ff; Pot O’ Gold Cave, 43.09959°N, - 114.39556°W, elev. 1390 m., 5 September 2017, R. Winton, m f; Giant Arch Cave, 43.10027°N, - 114.36399°W, elev. 1400 m., 28 March 2017, R. Winton, mm ff. Power Co., Kids Cave, 43.10632°N, - 113.06565°W, elev. 1560 m., 22 March 2017, R. Winton, mm ff. Notes: For illustrations and further details on this species, see Shear (2007). Idagona westcotti will likely be found in many other lava tubes in central Idaho. The species may disperse through the porous lava to colonize tubes. While pale in color and with somewhat reduced numbers of ocelli, populations are not conspicuously adapted for subterranean life., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 75, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Buckett, J. S. & Gardner, M. R. (1967) A new family of cavernicolous millipedes with the description of a new genus and species from Idaho (Diplopoda: Chordeumida: Chordeumidea). The Michigan Entomologist, 1, 117 - 126.","Shear, W. A. (1972) Studies in the millipede order Chordeumida (Diplopoda): a revision of the family Cleidogonidae and a reclassification of the order in the New World. Bulletin of the Museum of Comparative Zoology, 144, 151 - 352."]}

Bifurcatella bifurcata (Shear, 1974), new combination Figs. 204, 205 New records: Bollmanella bifurcata Shear 1974, p. 143. OREGON: Umatilla Co., Umatilla National Forest, FS- 65 0.4 mi E of Tiger Creek Road, elev. 1520 m., 45.9442°N, - 118.0094°W, 24 October 2003, W. Leonard, mm ff. Union Co., Little Indian Creek, 8 mi N of Cove, ca. 45.4134°N, - 117.7988°W (estimated coordinates taken from 8 air mi N of Cove along FS-6205), 10 September 1977, R. K. Johnson, mm ff (2 vials, FSCA). Notes: This species was originally described (Shear 1974) from 2 mi west of Joseph, along Hurricane Creek, Wallowa Co., Oregon, from specimens collected 23 November 1968. The new records are consistent with a range encompassing northeastern Oregon. In Fig. 204, the mesal branch of the anterior gonopod has been broken; it is normally about half the length of the lateral branch. Descriptions of new species of Bifurcatella, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 53-55, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (1974) The milliped genus Bollmanella (Diplopoda, Chordeumida, Conotylidae). Psyche, 81, 134 - 146. https: // doi. org / 10.1155 / 1974 / 51907"]}

Calityla, new genus Type species: Calityla siskiyou , n. sp. Diagnosis: Related to Taiyutyla, but differing in the posterior gonopod coxites, which lack the characteristic Tshaped branch and other modifications but instead have a single posterior fimbriate process, which may itself be branched; there may be a deep pit or pore at the base of this branch. A deep pit or pore on the anterior gonopods is lacking. The terminus of the coxite is often narrowed and with 2–3 acute, curved branches at the tip. Femoral knobs may appear on legpairs 3–7, 4–7, 6–7 or 7 only, or be completely absent (see species accounts below). Etymology: Cali- for California, where all the know species occur, with - tyla, a combining stem frequently used in the family. Gender is feminine. Distribution: Northern California. Each of the five new species of Calityla are only known from their type localities. Descriptions of new species of Calityla

Karagama, new genus Type species: Karagama ladybird , n. sp. Diagnosis: Differs from Taiyutyla and the new genus Vancouvereuma in the very simple, acuminate, curved coxites of the posterior gonopods, which lack a T-shaped branch but do have narrow fimbriate regions posteriorly, near a suppressed pore (Figs. 107, 108, 110). The anterior gonopods (Figs. 105, 109) are two-branched, similar to those of Bifurcatella, n. gen. (see below) but the branches occupy distinctly anterior and posterior positions, rather than lateral and mesal, as in Bifurcatella species. Etymology: The genus name is an arbitrary combination of letters that should be treated as feminine in gender. Included species: Besides the type, Karagama lupus (Shear, 2004), n. comb., and Karagama loftinae (Shear & Krejca, 2011), n. comb. are also included in the genus. Notes: This genus is somewhat heterogeneous, particularly with regard to the anterior gonopod, which in the type species has the posterior branch tightly appressed to the anterior; while in the other two, it is well separated. Geography may present another problem, since the type species is from northern California (Redwood National Park) and the other two included species are from the southern Sierra Nevada and Vancouver Island. Nevertheless for the present, relying only on morphology, they seem to belong together. The name of K. lupus does not change gender since the specific epithet is a noun in apposition rather than an adjective.

Lophomus shelleyi, new species Figs. 258–261 Types: Male holotype and female paratype from MONTANA: Ravalli Co., Woods Creek, Helena-Lewis and Clark National Forest, 47.4613°N, - 112.8371°W, elev. 1721 m., collected 6 October 2006 by R. Killackey, m f; male paratype from Granite Co., Butte Cabin Creek, Lolo National Forest, 46.5181°N, - 113.7601°W, elev. 1310 m., collected 2 October 2006 by P. Hendricks. Diagnosis: Distinguished from L. laxus by details of the gonopods. The sternal processes of the anterior gonopods are slightly longer in L. shelleyi, and the tips of the gonopods are divided into three parts, with the median the longest; in L. laxus the gonopods are divided into two parts at their tips, with the lateral the longest. A strong lateral hook is present distally on the posterior gonopod coxites of L. laxus but is much smaller in L. shelleyi. Etymology: The species name recognizes the many and varied contributions to myriapodology by our colleague the late Rowland M. Shelley (1942–2018), who also transmitted to us the Montana material reported on here. Description: Male paratype: Length, 14.0 mm. Twenty-seven to 29 ocelli in triangular patch. Metazonital shoulders developed into small, but distinctly flattened paranota; outermost segmental setae long, slightly curved, acute, median two segmental setae shorter, straight, blunt-tipped. Color light brown heavily mottled with darker purplish brown. Legpairs one and two reduced. Legpairs three through seven enlarged, pairs four and five the largest; small, acute, backward-facing basal knobs present on femora four and five. Anterior gonopods (Fig. 258) with anterior sternal processes nearly as large as gonopods; gonopods slightly sigmoid, ending in three processes, mesal process longest, lateral processes much shorter, curved posteriorly. Posterior gonopod coxite (Fig. 259) broad, tipped with large mesal hook, lateral hooked process much smaller; anterior flagellum greatly exceeding coxite in length, composed of many separate, spiral, flattened filaments (Fig. 260) with many tiny, retrorse cuticuar teeth at their tips. Coxae 10 with prominent glands, enlarged. Prefemur 11 with strong basal process, processes also on trochanters and coxae 11 (Fig. 261). Female similar to male in nonsexual characters. Distribution: Known only from the two type localities. Notes: The compound nature of the anterior gonopod flagellum has not been noted before. The flagellum is made up of 10–12 individual, flattened, spiral filaments tightly appressed to one another, but evidently the drying associated with preparing the gonopods for SEM examination caused them to separate. Small, retrorse cuticular thorns cover the acute tips of these filaments.

Brunsonia benewah, new species Figs. 183–189 Types: Male holotype and two male paratypes from IDAHO: Benewah Co., East Fork Charlie Creek, Idaho Panhandle National Forests, 3.25 mi S, 2.75 mi E of Emida, elev. 945 m, ca. 47.0674°N, - 116.5373°W, collected 17 September 1978, by A. L. Johnson (FSCA). Diagnosis: Unlike any other Brunsonia species, B. benewah has low, minutely toothed distal processes on the prefemora, not the femora, of the sixth and seventh legpair. Otherwise it is close to the preceding species in having femoral knobs on legpairs four and five, with the one on five being large and oblong, and fairly complicated posterior gonopod coxites. See the above species for further distinctions. Etymology: The name is a noun in apposition after Benewah County, Idaho. Description: Male paratype: Length, 9.5 mm. Seventeen ocelli in triangular eyepatch. Metazonites with low shoulders on all trunk rings from three to 25. Color light brown, lightly marked darker purplish brown, markings more distinct anteriorly. Legpairs one and two reduced, three to seven enlarged, mesal knobs present only on femora four and five, mesal and large on both, as in B. selwayana (see above). Anterior gonopods (Figs. 182, 183) with low, triangular mesobasal projections, distinct acute tooth lateral about midlength. Posterior gonopod coxites (Fig. 184) complex, with a large, distally attenuate laterobasal branch; fimbriate area reduced, subtended by cuticular teeth; terminal process single, helically curved. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Females not collected. Distribution: Known only from the type locality. Notes: We had too little material to prepare for SEM study. This species and the one above are probably peripheral to the genus as a whole, based on the rather large and well-developed anterior gonopods, but seem to us best placed here. We are hopeful that at some time in the future, genetic evidence may clarify the relative positions of the species now placed in this complex genus., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 47, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Subfamily Lophominae Loomis & Schmitt 1971 Lophomus Loomis & Schmitt 1971, p. 125. Notes: Species of this genus are distinctive in that the anterior gonopods have a pair of anterior sternal processes that are covered in fine cuticular projections. The posterior gonopod coxites have an articulated flagellum that is composed of many smaller filaments appressed and twisted together. New records of previously described species, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 72-73, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Loomis, H. F. & Schmitt, R. (1971) The ecology, distribution and taxonomy of the millipeds of Montana west of the continental divide. Northwest Science, 45, 107 - 131."]}

Taiyutyla clatsop Shear, 1976 Taiyutyla clatsop Shear1976: 6. Type locality: Oregon, Clatsop Co., 5 mi N, 7 mi W of Elsie, estimated coordinates taken from Saddle Mountain Road at 5 horiz mi N and 7 W of Elsie, 45.9339°N, - 123.7263°W. Male holotype and female paratypes (AMNH). New record: OREGON: Curry Co., Jerry’s Flat Road at Quosatana Creek, 9.5 mi E, 4.6 mi N of Gold Beach, Rogue River-Siskiyou National Forest, estimated coordinates taken from intersect of Quosatana Creek and Agnes Road, elev. 30 m., 42.4890°N, - 124.2331°W, 26 March 1976, A. K. Johnson, m (FSCA). Notes: This species is small for a member of Taiyutyla, and some of the gonopod features resemble species of Bifurcatella, n. gen. We retain it in Taiyutyla due to the pregonopodal leg modifications and its similarity to other Taiyutyla species described as new below., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 11, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (1976) The milliped family Conotylidae (Diplopoda, Chordeumida): revision of the genus Taiyutyla, with notes on recently proposed taxa. American Museum Novitates, 2600, 1 - 22."]}

Bollmanella bombus, new species Figs. 111–114 Types: Male holotype and 2 female paratypes from IDAHO: Idaho Co., Idaho Panhandle National Forests, Bumblebee Creek, elev. 685 m., 47.6325°N, - 116.2965°W, collected 6 November 2004, by W. Leonard, C. Richart, J. Baugh from the litter and woody debris of a mixed forest including Tsuga heterophylla, Populus trichocarpa, Abies grandis, Pinus monticola, and Rubus parviflorus. Gonopods of the holotype male are on SEM stub WS30-15, deposited with the specimen (CAS). Diagnosis: Distinct from other Bollmanella species in the two thin branches of the posterior gonopod coxites and a basal, bulbous projection from the posterior gonopod prefemur. This species also occurs at a considerable distance from the coastal Oregon and Washington distribution of the other known species, and is one of the smallest conotylids yet described. Etymology: The species epithet, a noun in apposition, is the Latin generic name of some bumblebee species, and refers to the type locality, Bumblebee Creek. Description: Male holotype: Length, 4.5 mm. Ten or 12 ocelli in three rows, the rows diminishing in length ventrally. Color uniform pale tan. Segmental shoulders moderately developed; metazonital setae small, spatulate. Pregonopodal legpairs three and four much enlarged but lacking knobs. Anterior gonopods (Figs. 111, 113) short, broad, curved at tip, with short process posteriorly in distal third. Posterior gonopods with setose swelling basal, posterior on prefemora; coxites (Figs. 112, 114) two-branched, posterior branch with few small distal teeth or fimbriae, anterior branch with distal hook. Legpair 10 with coxae swollen, glandular; legpair 11 with dorsally directed, small prefemoral lobes. Female 5 mm long, nonsexual characters as in male. Distribution: Known only from the type locality. Notes: This species, at 4.5 mm extended length in the males, is one of the smallest of conotylids. Several immature specimens were part of the type collection, interestingly their metazonital setae were longer and acute, rather than short and spatulate as in the adults. Possibly they represent another, unrecognized, syntopic species.

Brunsonia chelanomagna, new species Figs. 180, 181 Types: Male holotype, five male and 11 female paratypes from WASHINGTON: Chelan Co., Lake Chelan, Wenatchee National Forest, Fields Point, elev. 360 m., 47.9699°N, - 120.2160°W, collected 26 October 2007, by W. Leonard, J. Baugh. Diagnosis: Distinct from the syntopic B. chelanoparva in the distinctly pyramidal anterior gonopods, the posterior gonopod coxites lacking process a, and in having femoral knobs on legpairs 3 and 4, as opposed to 4 and 5. Etymology: The species name is a combination of Lake Chelan, the type locality, and Latin magna, noting the fact that it is the larger of two species of Brunsonia syntopic there. Description: Male holotype: Length, 15–15.5 mm. Nineteen ocelli in triangular patch. Metazonites with poorly developed shoulders on all trunk segments, segmental setae short, acute, curved. Color pale tan, with transverse bands of purplish brown on metazonites, pale spot surrounding outermost segmental setae, similar but lighter bands on prozonites. Legpairs three to seven encrassate as usual, femoral knobs present on legpairs three and four. Anterior gonopods (Fig. 180) broad at base, quickly tapering to acute tip, tip posteriorly curved, dorsolateral pit large, deep, channel to tip of gonopod trending first lateral, then posterior, ending on short, subapical process. Posterior gonopod coxites (Fig. 181) with finger-like projection at tip, strongly curved posteriolaterally, process a evidently lacking, channeled process large, prominent, with large pore basally, rugose or warty area small, restricted. Posterior gonopod prefemur clavate. Legpair 10 enlarged, with glands, legpair 11 prefemora with dorsally directed, curved processes. Female 15.0–16.0 mm long, nonsexual characters similar to those of male. Distribution: Known only from the type locality. Notes: The anterior gonopods are closer in form to those of B. pulchra, but less complex at the tips. As noted above, this species is syntopic with B. chelanoparva at the type locality., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 45, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Brunsonia chelanoparva, new species Figs. 168–172 Types: Male holotype, five male paratypes and nine female paratypes from WASHINGTON: Chelan Co., Lake Chelan, Wenatchee National Forest, Fields Point, elev. 360 m., 47.9699°N, - 120.2160°W, collected 26 October 2007, by W. Leonard, J. Baugh. Diagnosis: The anterior gonopods are roughly similar to those of B. albertana, but more acute at the tip and with a short, anterior branch, while the posterior gonopod coxites are less complex, relatively smaller than that species, and with a very prominent warty area on the main branch. Etymology: The species name is a combination of Lake Chelan, the type locality, and Latin parvus, noting the fact that it is the smaller of two species of Brunsonia syntopic there. Description: Male holotype: Length, 12.5 mm. 21–23 ocelli in triangular patch. Metazonites with well-developed shoulders on all trunk segments, on midbody segments shoulders slightly below midline dorsum, with angular posterior corners, segmental setae acute, curved, moderately long. Color pale to medium brown, with well-defined dark purplish brown spots on segmental shoulders, lateral prozonites, less well-defined spots on either side of midline. Legpairs three to seven slightly encrassate compared to postgonopodal legs, femoral knobs on legpairs 4 and 5 only. Anterior gonopods (Figs. 168, 170, 171) very similar to those of B. albertana, but apically more acute, anterolateral rugose area less so. Channel from pore to gonopod tip occupied by long, filamentous pseudoflagellum (Fig. 171). Posterior gonopod coxites (Figs. 169, 172) massive, channeled branch somewhat reduced, basal process displaced toward body of coxa, other branches apparently absent. Posterior gonopod prefemora distinctly clavate. Legpair 10 enlarged, with glands, legpair 11 prefemora with dorsally directed stout processes. Female 12.5–13.0 mm long, nonsexual characters similar to male. Distribution: Known only from the type locality. Notes: The filamentous structure occupying the posterior groove of the anterior gonopod is not known from any other species. It is hollow or possibly channeled, with a pore opening at the tip, which is slightly flared. The species and the following were in a mixed collection and when sorted out by size proved to be two different species. It is unusual to have two species of the same genus syntopic; perhaps each has distinct microhabitat preferences that were not recorded on the collection label., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 44-45, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Calityla, new genus Type species: Calityla siskiyou , n. sp. Diagnosis: Related to Taiyutyla, but differing in the posterior gonopod coxites, which lack the characteristic Tshaped branch and other modifications but instead have a single posterior fimbriate process, which may itself be branched; there may be a deep pit or pore at the base of this branch. A deep pit or pore on the anterior gonopods is lacking. The terminus of the coxite is often narrowed and with 2–3 acute, curved branches at the tip. Femoral knobs may appear on legpairs 3–7, 4–7, 6–7 or 7 only, or be completely absent (see species accounts below). Etymology: Cali- for California, where all the know species occur, with - tyla, a combining stem frequently used in the family. Gender is feminine. Distribution: Northern California. Each of the five new species of Calityla are only known from their type localities. Descriptions of new species of Calityla, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 17, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Bifurcatella hoh, new species Figs. 210, 211, 214, 215 Types: Male holotype, female paratype from WASHINGTON: Jefferson Co., Alder Creek at Hoh River Road, elev. 125 m., 47.8301°N, - 124.2272°W, collected 14 December 2003, by W. Leonard, C. Richart. Diagnosis: The unique feature of this species is the thin mesobasal process (mbp, Fig. 215) of the posterior gonopod coxite, found in no others; further, the apical process of the coxite is sharply curved anteromesally. The mesal branch of the anterior gonopod is more than 2/3 the length of the lateral branch and is straight and blunt. Etymology: Named for the Hoh River of Olympic National Park; a noun in apposition. Description: Male holotype: Length, 11.0 mm. Ocelli 22 in triangular patch. Metazonites with moderately well-developed shoulders, segmental setae acute, curved. Color pale to medium tan, marked darker purplish brown, mostly on dorsum. Legpairs one and two reduced, three to seven enlarged, three to five with swollen femora, femora of pair seven slender; large femoral knobs on pairs five and six. Anterior gonopods (Figs. 210, 214) with lateral branches robust, apically expanded and decurved, terminus cap-like; mesal branches straight, cylindrical, blunt, two-thirds to three-quarters length of lateral branches. Posterior gonopod coxites (Figs. 211, 215) with evenly curved and tapered pseudoflagellar process, fimbriate branch small, subtending shallow, rimmed pit with pore, mesobasally arising thin process. Terminal processes both strongly recurved. Tenth coxae and eleventh prefemora as usual. Female: Length, 11.5 mm, nonsexual characters as in male. Distribution: WASHINGTON: Jefferson Co., Olympic National Park, near Hoh Rain Forest Visitor Center, elev. 180 m., 47.8597°N, - 123.9413°W, 14 December 2003, W. Leonard, C. Richart, mm; Clallam Co., Olympic National Park, Heart of the Hills Campground, elev. 545 m., 48.0355°N, - 123.4263°W, 18 November 2003, W. Leonard, m f.

Karagama, new genus Type species: Karagama ladybird , n. sp. Diagnosis: Differs from Taiyutyla and the new genus Vancouvereuma in the very simple, acuminate, curved coxites of the posterior gonopods, which lack a T-shaped branch but do have narrow fimbriate regions posteriorly, near a suppressed pore (Figs. 107, 108, 110). The anterior gonopods (Figs. 105, 109) are two-branched, similar to those of Bifurcatella, n. gen. (see below) but the branches occupy distinctly anterior and posterior positions, rather than lateral and mesal, as in Bifurcatella species. Etymology: The genus name is an arbitrary combination of letters that should be treated as feminine in gender. Included species: Besides the type, Karagama lupus (Shear, 2004), n. comb., and Karagama loftinae (Shear & Krejca, 2011), n. comb. are also included in the genus. Notes: This genus is somewhat heterogeneous, particularly with regard to the anterior gonopod, which in the type species has the posterior branch tightly appressed to the anterior; while in the other two, it is well separated. Geography may present another problem, since the type species is from northern California (Redwood National Park) and the other two included species are from the southern Sierra Nevada and Vancouver Island. Nevertheless for the present, relying only on morphology, they seem to belong together. The name of K. lupus does not change gender since the specific epithet is a noun in apposition rather than an adjective., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 28, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (2004) Two new species in the millipede genus Taiyutyla from British Columbia, Canada, and Washington, USA. Myriapodologica, 8, 13 - 20.","Shear, W. A. & Krejca, J. K. (2011) Cave millipeds of the United States. IX. A new species of the genus Taiyutyla (Diplopoda, Chordeumatida, Conotylidae) from caves in Sequoia and Yosemite National Parks, California, USA. Journal of Cave and Karst Studies, 73, 93 - 98. https: // doi. org / 10.4311 / jcks 2010 lsc 157"]}

Brunsonia Loomis & Schmitt, 1971 Brunsonia Loomis & Schmitt, 1971:122. Type species Brunsonia albertana (Chamberlin, 1920), by original designation of its junior synonym Brunsonia complexipes Loomis & Schmitt 1971. Conotyla, Shear, 1971 (“Atrolineata group,” not Conotyla Cook & Collins 1895): 80; 1976:5. Diagnosis: Moderate-sized conotylids (10–15 mm long) distinct from other genera of the subfamily Conotylinae in the mostly reduced anterior gonopods coupled with the complex coxites of the posterior gonopods. Notes: The genus includes the previously described species Brunsonia albertana (Chamberlin, 1920) and B. atrolineata (Bollman, 1893). The latter species is known from western British Columbia, Canada; Shear (1971) failed to give specific collection data but mentioned Yoho, Glacier, and Mt. Robson National Parks. We have no new records of this species. Brunsonia albertana is a senior synonym of the designated type species, B. complexipes (Shear 1971), and occurs in southern Alberta and northwestern Montana (Loomis & Schmitt 1971). Again, Shear (1971) does not provide detailed collection records but mentions Banff and Jasper National Parks. Records given below extend the distribution to western Washington and Oregon in the United States. The four new species described herein are all from the United States and represent considerable extensions of the generic range. The gonopods of Brunsonia species show variations on a common theme. The anterior gonopods are small in some species, but nearly the size of the posterior gonopod coxites in at least one ( B. pulchra, n. sp. ), in which they also approach the form of the anterior gonopods of Taiyutyla species. However, the posterior gonopod coxites are far more complicated in this species than in almost any other conotylid, fitting the Brunsonia plan. Except for B. digitata, n. sp. , the anterior gonopods have an anteriolateral lobe set with small nodules, and all species have a deep pit or pore posteriolaterally in the anterior gonopods, with a narrow channel running from it to the tip of the gonopod; in B. chelanoparva, n. sp. , this channel seems also to be occupied by a hair-like, grooved filament. We think this pit or pore is homologous with pores and gland openings that appear on the posterior gonopod coxites and tenth legpair, and which are also present in other species of conotylids. The posterior gonopod coxites bear multiple features. The body of the coxa is globular, even swollen on the posterior side, and with one or two fimbriate branches on the lateral side. The coxite itself is massive, with various terminalia differing in each species; in B. albertana and B. chelanoparva, n. sp. , there is a region set with irregular, distinct nodules. A long process (process a) extends laterally, and just dorsal to it is another process (c) with a pore at its base and a long channel leading from the pore along the process. Process b may be multiple, and may or may not parallel a, but when present it is shorter than a and is often “fuzzy” and set with many small cuticular projections. Modifications of the femora of the male pregonopodal legs range from none to knobs on pairs three to six. New records of a previously described species, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 35, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Loomis, H. F. & Schmitt, R. (1971) The ecology, distribution and taxonomy of the millipeds of Montana west of the continental divide. Northwest Science, 45, 107 - 131.","Chamberlin, R. V. (1920) Some records of Canadian myriopods. The Canadian Entomologist, 52 (4), 94 - 95.","Shear, W. A. (1971) The millipede family Conotylidae in North America, with a description of the new family Adritylidae (Diplopoda: Chordeumida). Bulletin of the Museum of Comparative Zoology, 141, 55 - 96.","Cook, O. F. & Collins, G. (1895) The Craspedosomatidae of North America. Annals of the New York Academy of Sciences, 9, 1 - 100. https: // doi. org / 10.1111 / j. 1749 - 6632.1896. tb 55430. x","Bollman, C. H. (1893) The Myriapoda of the United States. United States National Museum Bulletin, 43, 1 - 210."]}

Bifurcatella germania, new species Figs. 220, 221, 224, 225 Types: Male holotype from WASHINGTON: Cowlitz Co., Germany Creek Road 5.5 mi N of Stella at SR-4, elev. 140 m., 46.2593°N, - 123.1344°W, collected 22 November 2003, by W. Leonard, C. Richart; male paratype from Germany Creek Road 5.3 mi N of Stella at SR-4, elev. 140 m., 46.2565°N, - 123.1356°W, collected 11 November 2004, by W. Leonard and C. Richart. Diagnosis: The posterior gonopod coxites are similar to those of the preceding species but the anterior gonopods are quite different, lacking the midlength shelf and sharp taper, and with three large subterminal teeth. Etymology: This species takes its adjectival name from Germany Creek, where the holotype and paratype were collected. Description: Male holotype: Length, 9.0 mm. Nineteen ocelli in triangular group. Metazonites with well-developed shoulders, segmental setae long, curved, acute. Color as described for B. pacifica. Legpairs one and two reduced, pairs three to five enlarged, sixth pair somewhat smaller, seventh pair approximately same size as postgonopodal legs; prefemora three to five with conspicuous knobs. Anterior gonopods (Figs. 220, 222) with mesal branches less than one-third size of lateral branches, broadly flattened in lateral view; lateral branches curved, with large, wing-like expansion anteriorly, bearing three triangular subterminal teeth. Posterior gonopod coxites (Figs. 221, 223) with pseudoflagellar branch tightly reflexed, not strongly tapering, two large fimbriate branches, lateralmost embraces pseudoflagellum, mesal branch subtends small pore. Tenth coxae and eleventh prefemur as usual. Females not collected. Distribution: Known only from the two localities along Germany Creek, which are about 340 meters apart., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 60, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Brunsonia benewah, new species Figs. 183–189 Types: Male holotype and two male paratypes from IDAHO: Benewah Co., East Fork Charlie Creek, Idaho Panhandle National Forests, 3.25 mi S, 2.75 mi E of Emida, elev. 945 m, ca. 47.0674°N, - 116.5373°W, collected 17 September 1978, by A. L. Johnson (FSCA). Diagnosis: Unlike any other Brunsonia species, B. benewah has low, minutely toothed distal processes on the prefemora, not the femora, of the sixth and seventh legpair. Otherwise it is close to the preceding species in having femoral knobs on legpairs four and five, with the one on five being large and oblong, and fairly complicated posterior gonopod coxites. See the above species for further distinctions. Etymology: The name is a noun in apposition after Benewah County, Idaho. Description: Male paratype: Length, 9.5 mm. Seventeen ocelli in triangular eyepatch. Metazonites with low shoulders on all trunk rings from three to 25. Color light brown, lightly marked darker purplish brown, markings more distinct anteriorly. Legpairs one and two reduced, three to seven enlarged, mesal knobs present only on femora four and five, mesal and large on both, as in B. selwayana (see above). Anterior gonopods (Figs. 182, 183) with low, triangular mesobasal projections, distinct acute tooth lateral about midlength. Posterior gonopod coxites (Fig. 184) complex, with a large, distally attenuate laterobasal branch; fimbriate area reduced, subtended by cuticular teeth; terminal process single, helically curved. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Females not collected. Distribution: Known only from the type locality. Notes: We had too little material to prepare for SEM study. This species and the one above are probably peripheral to the genus as a whole, based on the rather large and well-developed anterior gonopods, but seem to us best placed here. We are hopeful that at some time in the future, genetic evidence may clarify the relative positions of the species now placed in this complex genus.

Ovaskella ovaskae, new species Figs. 86–91 Types: Male holotype and male paratype from BRITISH COLUMBIA: SGang Gwaay, (Anthony Island), Haida Gwaii (Queen Charlotte Islands), SGang Gwaay UNESCO World Heritage Site and National Historic Site, elev. 15 m., 52.0980°N, - 131.2164°W, 15 September 2004, K. Ovaska, L. Sopuk. Diagnosis: Distinct from the other species of the genus in bearing obvious paranota on most metazonites, and lacking femoral knobs on any pregonopodal legs. The paranotal character is shared by two other British Columbia species, Vancouvereuma shawi (Shear), 2004 and Karagama lupus (Shear), 2004. Vancouvereuma shawi, however, has gonopods similar to Taiyutyla jonesi, and K. lupus has bifid gonopods with subequal branches; both of these species have more complicated posterior gonopod coxites than O. ovaskae. Etymology: We name this species for Kristiina Ovaska, Victoria, British Columbia, the collector. Description: Paratype male from SGang Gwaay: Length, 12.0 mm. 21 ocelli in triangular eyepatch. Metazonites with well-developed paranota, in place of rounded shoulders, on metazonites 6–25; two outermost segmental setae on anterior and posterior corners of paranotum. Color dark brown to chestnut brown. Legpairs one and two reduced, three to seven enlarged, mesal knobs lacking on all pregonopodal legs, but femora three to five more strongly bowed than six and seven. Anterior gonopods (Figs. 86, 89) slender, with anterior midlength knob bearing coarse teeth, apical process bifurcate with subterminal anterior lamella. Posterior gonopod coxites (Figs. 87, 88, 90, 91) with basal portion of prefemur set off by distinct groove, coxite much simplified, lacking fimbriate branches or areas. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Female 12.0 mm long, similar to male in nonsexual characters. Distribution: Only known from the type locality. Note: The former Queen Charlotte Islands, more than 150 mountainous islands off the west coast of British Columbia, are now known as Haida Gwaii. These islands are protected as the Gwaii Haanas National Park Reserve. SGang Gwaay, or Ninstints, is a village abandoned by the Haida nation in 1880 and is now recognized by UNESCO as a World Heritage Site. The region may have escaped glaciation during the Wisconsinan, and hosts a remarkable array of endemic species, such that it is sometimes called the “Canadian Galápagos,” but its soil and litter fauna has been little explored.

Bollmanella washingtonensis, new species Figs. 115–117 Types: Male holotype, male paratype and two female paratypes from WASHINGTON: Mason Co., W Beerbower Road, at W Schafer State Park Road, elev. 45 m., 47.1001°N, - 123.4706°W, collected 31 December 2004, by W. Leonard. Diagnosis: Unlike any other species of Bollmanella, B. washingtonensis has pregonopodal legpairs three to seven enlarged, and pairs five through seven with femoral knobs. Etymology: The species name notes that this species is endemic to Washington. Description: Male paratype: Length, 10.5 mm. 18–20 ocelli in rounded, triangular eyepatch. Color pale tan, mottled light purplish brown. Segmental shoulders prominent, metazonital setae long, acute. Pregonopodal legpairs three to seven strongly encrassate, legpairs five, six and seven with femoral knobs. Anterior gonopods (Fig. 115) slender, widely divergent, prominently curved near midlength, with acute subapical process as in the foregoing species. Posterior gonopods (Fig. 117) with relatively short telopodites, coxites (Figs. 116, 117) two-branched, posterior branch shorter, tapering, curved laterad, anteromesal branch distally bifurcate, one terminal four times broader than other, slightly curved. Legpairs 10 and 11 as usual for genus. Female 11 mm long, nonsexual characters as in male. Distribution: WASHINGTON: Grays Harbor Co., Middle Satsop Road 2 mi N of Brady near Satsop River, elev. 20 m., 47.0245°N, - 123.5196°W, under maple leaves, 3 January 2003, W. Leonard et al., mm; 10 mi N of US- 12, East band of Wynoochee River, ca. 47.0920°N, - 123.6798°W (estimated coordinates measured 10.0 mi N of US- 12 along Wynoochee Valley Rd), under maple leaves, 3 January 2003, W. Leonard et al., m f; Canyon River, 6 mi W, 1.25 mi N of Matlock, ca. 47.2585°N, - 123.5271°W, in maple and fern litter, 17 January 2003, W. Leonard et al., mm ff; Garrard Road at Weyerhaeuser D-line, elev. 45 m., ca. 46.8041°N, - 123.3033°W, 7 February 2005, W. Leonard, mm ff. Thurston Co., Capitol State Forest, 2 mi S of Rock Candy Mountain, ca. 46.9809°N, - 123.1090°W (estimated coordinates taken from 2 air mi S of Rock Candy Mountain summit), 10 December 2002, W. Leonard et al., mm; Summit Lake, elev. 138 m., 47.0526°N, - 123.1304°W, 18 December 2004, W. Leonard, m f; McAllister Springs S of Steilacoom Rd SE, elev. 50 m., 47.0472°N, - 122.7286°W, 7 February 2004, W. Leonard, C. Richart, from the litter and woody debris of a mixed forest including Acer macrophyllum, Alnus rubra, Pseudotsuga menziesii, Thuja plicata, Polystichum munitum, and Mahonia nervosa, mm f; St. Martin’s College Campus, Woodland Creek, elev. 92 m., 47.0416°N, - 122.8134°W, 12 February 2003, W. Leonard, mm ff. King Co., I-90 at Snoqualmie River, Twin Falls/Iron Horse trailhead, elev. 370 m., 47.4410°N, - 121.6687°W, 25 February 2004, W. Leonard, C. Richart, m f; Along Holder Creek, SR-18 1.5 mi N of the Issaquah exit near Hobart, elev. 310 m., 47.4522°N, - 121.9535°W, 25 February 2004, W. Leonard, C. Richart, m f; Ollalie State Park, elev. 380 m., 47.4364°N, - 121.6594°W, 25 February 2004, W. Leonard, C. Richart, m f; FR-9034 near Granite Mountain Trailhead, Mt. Baker-Snoqualmie National Forest, elev. 580 m., 47.3977°N, -121.4855, 27 October 2007, W. Leonard, mm f; same as previous but 26 March 2004, W. Leonard, C. Richart, m; same as previous but 25 October 2003, m. Cowlitz Co., SR-504 2.1 mi E of Toutle, elev. 152 m., 46.3483°N, - 122.7066°W, 1 March 2004, W. Leonard, C. Richart, from the litter and woody debris of a mixed forest including Alnus rubra, Acer macrophyllum, and Polystichum munitum, m f. Jefferson Co., Falls View Campground, Olympic National Forest, US-101 7.8 mi NW of Brinnon, elev. 150 m., 47.7893°N, - 122.9255°W, 22 February 2003, W. Leonard, m f. Pacific Co., Ellsworth Creek Nature Society Preserve, old growth forest, 14 March 2003, elev. 25 m., 46.3981°N, - 123.8898°W, W. Leonard et al., m ff; same as previous but 23 November 2003, W. Leonard et al., mm; Trap Creek Road/B-line 1.1 mi S of SR-6, elev. 58 m., 46.5432°N, - 123.6151°W, 19 November 2005, W. Leonard, C. Richart, mm ff; same as previous but 1.9 mi S of SR-6, elev. 65 m, 46.5401°N, - 123.6296°W. Notes: This widespread species could be considered as a Taiyutyla, since the pregonopodal leg modifications do not conform to the other species of Bollmanella. Nevertheless, the form of the gonopods indicates that it belongs in the latter genus., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 32-34, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Bifurcatella, new genus Type species: Bollmanella bifurcata Shear, 1974 Diagnosis: This genus is distinguished from other conotylines by two features of the gonopods in combination. The anterior gonopods are composed of two branches which occupy lateral and mesal positions. Except for one species, the lateral branch is the longest and usually has a more or less complicated tip, while the mesal branch may be short, of medium length, or long and is always simple. The posterior gonopod coxites carry a flagellar or pseudoflagellar basal branch, which, unlike other genera, arises laterally on the coxite, is strongly sinuous and more or less fimbriate towards its tip. Etymology: The genus name is derived from the species epithet of the type species, referring to the anterior gonopods. Notes: Besides the type, Bifurcatella unca (Shear, 1974), new combination, and the following new species. With regard to the form of the gonopods, the species of this genus conform well to a common pattern. The anterior gonopods are upright, with mesal and lateral branches. The lateral branch (L, Fig. 206) may be considered the main body of the gonopod and is either acute, flared or branched at its tip. The mesal branch (M, Fig. 206) ranges from only about a sixth the length of the lateral to longer than the lateral and is always simple and cylindrical. The posterior gonopod coxites have a characteristic flagellar or pseudoflagellar (we are not sure if it is movable or not) process (pf, Fig. 207) that arises basolaterally and curves around the posterior side of the coxite, describing various curves. This process is robust at its base but tapers to a very fine point and is often fimbriate over at least part of its distal surface. The pseudoflagellar process is usually enclosed in a fimbriate sheath (sh, Fig. 205, 207). On the mesal side of the coxite about midway in its length a fimbriate process (fp, Fig. 207) is seen, which subtends a pore. The main body of the coxite is decurved near its tip and divided into two or three processes. The modifications of the male pregonopodal legs are various. We cannot be sure that the big pseudoflagellar branch of the posterior gonopod coxites of Bifurcatella is homologous to the T-shaped branch found in nearly the same position in Taiyutyla, but it is tempting to think so, and to infer that the two genera may share a common ancestral form. Further evidence lies in the resemblance of the T-shaped branch of T. napa to the pseudoflagellar branch in Bifurcatella species. New records of a previously described species, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 53, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (1974) The milliped genus Bollmanella (Diplopoda, Chordeumida, Conotylidae). Psyche, 81, 134 - 146. https: // doi. org / 10.1155 / 1974 / 51907"]}

Bifurcatella pacifica, new species Figs. 218, 219, 222, 223 Types: Male holotype and male and female paratypes from WASHINGTON: Pacific Co., Ellsworth Creek Preserve, elev. 50 m., 46.3981°N, - 123.8898°W, collected 23 November 2003 by W. Leonard et al. in old growth forest. Diagnosis: Like the following species, B. pacifica has the pseudoflagellar branch of the posterior gonopod coxite tightly doubled on itself, and the fimbriate branch is large. However, the anterior gonopods are quite different; in B. pacifica the lateral branch has a distinct shoulder distal of which the branch narrows to a simple tip, while in B. germania n. sp. (see below) the anterior gonopod lateral branch tapers more or less evenly and has three triangular teeth subtending the tip. Etymology: The species is named for Pacific County, Washington. Description: Male holotype: Length, 10.00 mm. Eighteen to 20 ocelli in triangular group. Metazonites with well-developed shoulders, segmental setae curved, acute. Color pale tan to chestnut, marked purplish brown on metazonites and prozonites such that dorsally two parallel rows of dark spots appear, additional rows on both lateral surfaces of rings. Legpairs one and two reduced, three through seven enlarged, fourth pair with low femoral knobs, fifth pair with large, acute, distal femoral knobs. Anterior gonopods (Figs. 218, 222) with mesal branches, blunt, cylindrical, approximately one-third length of lateral branches; lateral branches thickest at midlength with distinct constriction distal to posterior shelf, distal part curved, acute at tip, with subdistal ridged region. Posterior gonopod coxites (Figs. 219, 223) with main branch narrow, ending bluntly, two strong fimbriate branches embrace tightly folded, rapidly tapering pseudoflagellar branch. Tenth coxae conspicuously enlarged, glandular; eleventh prefemora with usual process. Female 10.00 mm long, nonsexual characters as in male. Distribution: WASHINGTON: Pacific Co., Walberg Road 1.0 mi N of Chinook Valley Rd, or 0.4 mi W of Ilwaco Water Park, elev. 30 m., 46.3235°N, - 123.9417°W, 2 January 2005, W. Leonard, C. Richart, from the litter and woody debris of a mixed forest including Alnus rubra, Tsuga heterophylla, Polystichum munitum, and Rubus spectabilus, mm., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 58-59, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Taiyutyla napa Shear, 1971 Fig. 22 Taiyutyla napa Shear 1971: 87. Type locality: California, Napa Co., Mt. St. Helena, male holotype and two female paratypes (AMNH). New record: California, Napa Co., Mt. St. Helena, estimated coordinates taken from the Mt. St. Helena Trail Head along SR-29, elev. 715 m., 38.6523°N, - 122.5998°W, 13 March 1962, J. S. Buckett, M. E. Irwin, m f (EMEC). Notes: These specimens are topotypical. The scanning electron micrograph (Fig. 22) shows many details not seen using optical microscopes and not reported in the original description (Shear 1971). Figure 22 illustrates, in posterior view, the anterior (ag) and posterior (pg) gonopods. The ventral part of the T-shaped branch (T) of the posterior gonopod coxite is evidently absent, so the coxite somewhat resembles those of species of Bifurcatella, n. gen., but the anterior gonopod does not. For the time being, this species is better accommodated in Taiyutyla. Note the secretion (sec) emerging from a pore in the anterior gonopod., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 11, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (1971) The millipede family Conotylidae in North America, with a description of the new family Adritylidae (Diplopoda: Chordeumida). Bulletin of the Museum of Comparative Zoology, 141, 55 - 96."]}

Bifurcatella hobo, new species Figs. 234–239 Types: Male holotype, male and female paratypes from IDAHO: Shoshone Co., Hobo Cedar Grove Botanical Area, Idaho Panhandle National Forests, elev. 1360 m, 47.0880°N, - 116.1152°W, collected 16 October 2009, by W. Leonard, C. Richart, from the litter and woody debris of an old-growth forest including Tsuga heterophylla, Thuja plicata, and Abies grandis. Diagnosis: As with the preceding species, B. hobo, in keeping with their geographic separation from the remainder of Bifurcatella species, differs from the Washington forms. The anterior gonopods recall those of Karagama ladybird from northern California in that one or more small branches are tightly appressed to the posterior side. However, the posterior gonopod coxites are very different and have the typical Bifurcatella pseudoflagellar branch; the fimbriate branch is significantly reduced. Etymology: The species name is a noun in apposition referring to the type locality, a preserve noted for the diversity of its plant life. Description: Male holotype: Length, 12.0 mm. Twenty to 22 ocelli in triangular group. Shoulders of metazonites weakly developed, segmental setae long, curved, acute. Color almost uniform pale tan, lightly marked darker. Legpairs one and two reduced, pairs three to seven enlarged, three to five much the largest; femora five with very large, oblong knobs distally acute, femora four with smaller knobs. Anterior gonopods (Figs. 234, 236, 237) with low, broad mesal branches, lateral branches curved, with thin, acute apical process; in posterior view with two or three posterior branches tightly appressed to main body of gonopod. Posterior gonopod coxites (Figs. 235, 238, 239) simpler than in other species of Bifurcatella, with subapical teeth; pseudoflagellar branch sharply angled at base, not sinuate; fimbriate branch small. Coxae 10 and eleventh prefemora as usual. Female 12 mm long, nonsexual characters as in male. Distribution: IDAHO: Latah Co., E Fork of Meadow Creek Road (FS-1443) at SR-6, tributary of Mannering Creek, Idaho Panhandle National Forests, elev. 915 m, 47.0350°N, - 116.6729°W, 17 October 2009, W. Leonard, C. Richart, from litter and stream-side woody debris in a mixed forest including Thuja plicata, Abies grandis, and Tsuga heterophylla, m. Benewah Co., SR-6 0.6 mi S of Dennis Mountain Road and 2.1 mi N of the Latah County line, Idaho Panhandle National Forests, headwaters of Mannering Creek, elev. 1030 m., 47.0639°N, - 116.6751°W, 17 October 2009, W. Leonard, C. Richart, m ff. Shoshone Co., same as type locality but 17 October 2009, elev. 1290 m., 47.0880°N, - 116.1152°W, m ff. Notes: The Mannering Creek specimens are considerably darker than the holotype, some of them appearing almost solidly dark purplish brown., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 64, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Bifurcatella pacifica, new species Figs. 218, 219, 222, 223 Types: Male holotype and male and female paratypes from WASHINGTON: Pacific Co., Ellsworth Creek Preserve, elev. 50 m., 46.3981°N, - 123.8898°W, collected 23 November 2003 by W. Leonard et al. in old growth forest. Diagnosis: Like the following species, B. pacifica has the pseudoflagellar branch of the posterior gonopod coxite tightly doubled on itself, and the fimbriate branch is large. However, the anterior gonopods are quite different; in B. pacifica the lateral branch has a distinct shoulder distal of which the branch narrows to a simple tip, while in B. germania n. sp. (see below) the anterior gonopod lateral branch tapers more or less evenly and has three triangular teeth subtending the tip. Etymology: The species is named for Pacific County, Washington. Description: Male holotype: Length, 10.00 mm. Eighteen to 20 ocelli in triangular group. Metazonites with well-developed shoulders, segmental setae curved, acute. Color pale tan to chestnut, marked purplish brown on metazonites and prozonites such that dorsally two parallel rows of dark spots appear, additional rows on both lateral surfaces of rings. Legpairs one and two reduced, three through seven enlarged, fourth pair with low femoral knobs, fifth pair with large, acute, distal femoral knobs. Anterior gonopods (Figs. 218, 222) with mesal branches, blunt, cylindrical, approximately one-third length of lateral branches; lateral branches thickest at midlength with distinct constriction distal to posterior shelf, distal part curved, acute at tip, with subdistal ridged region. Posterior gonopod coxites (Figs. 219, 223) with main branch narrow, ending bluntly, two strong fimbriate branches embrace tightly folded, rapidly tapering pseudoflagellar branch. Tenth coxae conspicuously enlarged, glandular; eleventh prefemora with usual process. Female 10.00 mm long, nonsexual characters as in male. Distribution: WASHINGTON: Pacific Co., Walberg Road 1.0 mi N of Chinook Valley Rd, or 0.4 mi W of Ilwaco Water Park, elev. 30 m., 46.3235°N, - 123.9417°W, 2 January 2005, W. Leonard, C. Richart, from the litter and woody debris of a mixed forest including Alnus rubra, Tsuga heterophylla, Polystichum munitum, and Rubus spectabilus, mm.

Loomisella, new genus Type species: Loomisiella evergreen , n. sp. Diagnosis: Loomisiella, n. gen. , differs from other conotylids in the form of the large anterior gonopods, widely separated at the base, then curving sharply inward and posterior. The telopodites of the posterior gonopods are reduced in size; the posterior gonopod coxites are as broad as long, anteriorly excavated and with several processes arising from the anterior surface. Both known species are small, 5 mm or less in length. Etymology: The genus name honors Harold F. Loomis (1896–1976), who not only named more than 500 species of millipedes, primarily from Central America and the Caribbean, but also served 27 years as Director of the U. S. Plant Introduction Station in Miami, Florida. The name should be treated as feminine in gender. Notes: Other included species are, in addition to the type, Loomisiella pylei, n. sp. This genus includes some of the smallest known conotylids, both species being 5 mm or less in length at maturity. Descriptions of new species of Loomisiella

Ovaskella ovaskae, new species Figs. 86–91 Types: Male holotype and male paratype from BRITISH COLUMBIA: SGang Gwaay, (Anthony Island), Haida Gwaii (Queen Charlotte Islands), SGang Gwaay UNESCO World Heritage Site and National Historic Site, elev. 15 m., 52.0980°N, - 131.2164°W, 15 September 2004, K. Ovaska, L. Sopuk. Diagnosis: Distinct from the other species of the genus in bearing obvious paranota on most metazonites, and lacking femoral knobs on any pregonopodal legs. The paranotal character is shared by two other British Columbia species, Vancouvereuma shawi (Shear), 2004 and Karagama lupus (Shear), 2004. Vancouvereuma shawi, however, has gonopods similar to Taiyutyla jonesi, and K. lupus has bifid gonopods with subequal branches; both of these species have more complicated posterior gonopod coxites than O. ovaskae. Etymology: We name this species for Kristiina Ovaska, Victoria, British Columbia, the collector. Description: Paratype male from SGang Gwaay: Length, 12.0 mm. 21 ocelli in triangular eyepatch. Metazonites with well-developed paranota, in place of rounded shoulders, on metazonites 6–25; two outermost segmental setae on anterior and posterior corners of paranotum. Color dark brown to chestnut brown. Legpairs one and two reduced, three to seven enlarged, mesal knobs lacking on all pregonopodal legs, but femora three to five more strongly bowed than six and seven. Anterior gonopods (Figs. 86, 89) slender, with anterior midlength knob bearing coarse teeth, apical process bifurcate with subterminal anterior lamella. Posterior gonopod coxites (Figs. 87, 88, 90, 91) with basal portion of prefemur set off by distinct groove, coxite much simplified, lacking fimbriate branches or areas. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Female 12.0 mm long, similar to male in nonsexual characters. Distribution: Only known from the type locality. Note: The former Queen Charlotte Islands, more than 150 mountainous islands off the west coast of British Columbia, are now known as Haida Gwaii. These islands are protected as the Gwaii Haanas National Park Reserve. SGang Gwaay, or Ninstints, is a village abandoned by the Haida nation in 1880 and is now recognized by UNESCO as a World Heritage Site. The region may have escaped glaciation during the Wisconsinan, and hosts a remarkable array of endemic species, such that it is sometimes called the “Canadian Galápagos,” but its soil and litter fauna has been little explored., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 23-24, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Bifurcatella hoh, new species Figs. 210, 211, 214, 215 Types: Male holotype, female paratype from WASHINGTON: Jefferson Co., Alder Creek at Hoh River Road, elev. 125 m., 47.8301°N, - 124.2272°W, collected 14 December 2003, by W. Leonard, C. Richart. Diagnosis: The unique feature of this species is the thin mesobasal process (mbp, Fig. 215) of the posterior gonopod coxite, found in no others; further, the apical process of the coxite is sharply curved anteromesally. The mesal branch of the anterior gonopod is more than 2/3 the length of the lateral branch and is straight and blunt. Etymology: Named for the Hoh River of Olympic National Park; a noun in apposition. Description: Male holotype: Length, 11.0 mm. Ocelli 22 in triangular patch. Metazonites with moderately well-developed shoulders, segmental setae acute, curved. Color pale to medium tan, marked darker purplish brown, mostly on dorsum. Legpairs one and two reduced, three to seven enlarged, three to five with swollen femora, femora of pair seven slender; large femoral knobs on pairs five and six. Anterior gonopods (Figs. 210, 214) with lateral branches robust, apically expanded and decurved, terminus cap-like; mesal branches straight, cylindrical, blunt, two-thirds to three-quarters length of lateral branches. Posterior gonopod coxites (Figs. 211, 215) with evenly curved and tapered pseudoflagellar process, fimbriate branch small, subtending shallow, rimmed pit with pore, mesobasally arising thin process. Terminal processes both strongly recurved. Tenth coxae and eleventh prefemora as usual. Female: Length, 11.5 mm, nonsexual characters as in male. Distribution: WASHINGTON: Jefferson Co., Olympic National Park, near Hoh Rain Forest Visitor Center, elev. 180 m., 47.8597°N, - 123.9413°W, 14 December 2003, W. Leonard, C. Richart, mm; Clallam Co., Olympic National Park, Heart of the Hills Campground, elev. 545 m., 48.0355°N, - 123.4263°W, 18 November 2003, W. Leonard, m f., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 56-57, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Calityla humboldtensis, new species Figs. 75 –85 Types: Male holotype and female paratype from CALIFORNIA: Humboldt Co., 2.75 mi NNE Orleans, Six Rivers National Forest, estimated coordinates taken from CA-96 2.75 mi NNE of the U.S. Post Office in Orleans, elev. 175 m., 41.3311°N, - 123.5131°W, collected 21 December 1976, by R. K. Johnson (FSCA). Diagnosis: In addition to its remarkably simple gonopods, this species differs from all others in Calityla in having distal knobs on some of the pregonopodal leg prefemora, as well as the femora. Etymology: The species name, an adjective, refers to the type locality in Humboldt County. Description: Male holotype: Length, 9.5 mm. 19 ocelli in triangular eyepatch. Metazonites with moderate shoulders on rings 3–26. Coloration pale tan, lightly marked darker brown. Legpairs one and two reduced, three to seven enlarged, fungiform knobs basal on femora four, five and seven, mesodistal on femora six, knobs of femora seven set with small teeth; prefemora four to seven with small distal knobs, smallest on femora six (Figs. 78–82, 85). Anterior gonopods (Fig. 75) large, broad, with small mesal branches about one-tenth length of main, lateral branches, these with finely corrugated areas just below tips. Posterior gonopod coxites (Figs. 76, 77) with simple, acuminate, distally hooked coxites, anteriorly bearing densely fimbriate process. Legpair 10 coxae of normal size, with small glands (Fig. 83), legpair 11 femora with blunt, dorsally directed knobs (Fig. 84). Female 10 mm long, otherwise as male in nonsexual characters. Distribution: Only known from the type locality. Notes: Probably the extra prefemoral knobs allow the males to have a firmer grip on the females during mating.

Bifurcatella uniclada, new species Figs. 226–229 Types: Male holotype and male and female paratypes from WASHINGTON: Lewis Co., Slide Creek, Weyerhaeuser 4000 Road 0.8 mi SW of Pe-Ell-McDonald Road, elev. 200 m, 46.5241°N, - 123.1934°W, collected 3 January 2005, by W. Leonard. Diagnosis: Similar to the foregoing two species but distinct in that the anterior gonopods lack mesal branches and terminate in a series of finger-like projections from a flattened, palm-like tip. Also, the posterior gonopod coxite pseudoflagellar branch is greatly thickened about midway in its length and then tapers rapidly to a very thin tip. Etymology: The species epithet, an adjective, refers to the lack of a mesal branch on the anterior gonopod. Description: Male holotype: Length, 12.0 mm. Twenty to 22 ocelli in triangular group. Segmental shoulders well-developed, setae long, curved, acute. Color mostly dark purplish brown in continuous bands across metazonites but with conspicuous white to pale tan spots on shoulders. Legpairs one and two reduced, pairs three to seven encrassate, femoral knobs on pairs four and five. Anterior gonopods (Figs. 226, 228) long, thin, curved at tip, with three to five finger-like projections. Posterior gonopod coxites (Figs. 227, 229) robust, broad-tipped, with large basal flange that presses against projection from gonopod prefemur; pseudoflagellar branch strongly sinuate, greatly thickened midlength, abruptly tapering, conspicuously fimbriate near tip; fimbriate branches not as large as in preceding two species. Gonopod femur narrow. Tenth coxae and eleventh prefemora as usual. Female 11.00 mm long, nonsexual characters as in male. Distribution: WASHINGTON: Pacific Co., Trap Creek Road 5.9 mi S of SR-6, Alder Creek drainage of Naselle River, elev. 230 m., 46.4972°N, - 123.6439°W, 19 November 2005, W. Leonard, C. Richart, m; Cement Creek, SR-401 2.5 mi S of Parpala Road in Naselle, elev. 30 m., 46.3341°N, - 123.8002°W, 15 January 2006, W. Leonard, C. Richart, from the litter of a riparian forest including Alnus rubra, Picea sitchensis, Sambucus cf. caerulea, and Polystichum munitum, m; 3 horiz mi S, 2 horiz mi W of Menlo, estimated coordinates and elevation taken from headwater stream adjacent Ehmer Road, elev. ca. 255 m., 46.5769°N, - 123.6892°W, 20 December 2003, W. Leonard et al., m f. Lewis Co., in Lewis County only known from the type locality. Notes: Despite lacking mesal branches on the anterior gonopods, the posterior gonopod coxites clearly mark this species as a Bifurcatella. The basal, posterior flanges and the basal projection from the posterior gonopod prefemur are also distinctive., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 60, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Brunsonia albertana (Chamberlin, 1920) Figs. 121–137 Conotyla albertana Chamberlin, 1920:167. Shear, 1971:82. Brunsonia complexipes Loomis & Schmitt, 1971:122. New records: MONTANA: Missoula Co., Lindbergh Lake Campground, Flathead National Forest, elev. 1320 m., T19 N, R17 W, Sec 19 NENE, ca. 47.4070°N, - 113.7211°W, 25 September 2006, P. Hendricks, mm f?. Saunders Co., East Fork of Bull River, Cabinet Mountains Wilderness, elev. 1000 m., T27 N, R32 W, Sec. 3 SESE, ca. 48.1385°N, - 115.6886°W, 11 October 2006, P. Hendricks, m. Lincoln Co., Gordon Creek at Callahan Creek Road, Kootenai National Forest, elev. 890 m., T31 N, R34 W, Sec. 19WNW, ca. 48.4462°N, - 115.9860°W, 12 October 2006, P. Hendricks, m f. Lake Co., Swan River State Forest, South Fork of Lost Creek, elev. 1350 m., T24 N, R17 W, Sec. 11 SENW, ca. 47.8602°N, - 113.7326°W, 26 September 2006, P. Hendricks, m f. WASHINGTON: Spokane Co., Riverside State Park Bowl & Pitcher, Spokane, elev. 520 m., ca. 47.7002°N, - 117.5004°W, 1 November 2004, W. Leonard, C. Richart, J. Baugh, m. OREGON: Umatilla Co., Umatilla National Forest, Kendall Skyline Road (FS- 64) 0.4 mi N of FS-65, elev. 1725 m., 45.9719°N, - 117.9534°W, W. Leonard, mm ff; Umatilla National Forest, FS- 65 0.4 mi E of Tiger Creek Road, elev. 1520 m., 45.9442°N, - 118.0094°W, 24 October 2003, W. Leonard, f. Notes: The figures given here of Spokane, Washington (Figs. 121–127) and Umatilla County, Oregon (Figs. 128–135) specimens deviate somewhat from the description and illustrations of the species given by Shear (1971). This may represent geographic variation or may be due to viewing the gonopods (particularly the complicated posterior gonopod coxites) from slightly differing angles. Alternatively, genetic studies may eventually reveal that this variation represents distinct species. Illustrations (Figs. 136, 137) of a specimen from Missoula Co., Montana, seem closer to the figures in Shear (1971) and Loomis and Schmitt (1971)., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 35-36, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Chamberlin, R. V. (1920) Some records of Canadian myriopods. The Canadian Entomologist, 52 (4), 94 - 95.","Shear, W. A. (1971) The millipede family Conotylidae in North America, with a description of the new family Adritylidae (Diplopoda: Chordeumida). Bulletin of the Museum of Comparative Zoology, 141, 55 - 96.","Loomis, H. F. & Schmitt, R. (1971) The ecology, distribution and taxonomy of the millipeds of Montana west of the continental divide. Northwest Science, 45, 107 - 131."]}

Taiyutyla Chamberlin 1952 Taiyutyla Chamberlin 1952:1. Type species, Taiyutyla jonesi (Chamberlin, 1951) by original designation of its synonym (see below for full details), T. corvallis Chamberlin, 1952. Hoffman, 1961:270; Shear, 1971:86, 1976:6, 2004:15. Diagnosis: Taiyutyla species may be distinguished from sympatric conotyline genera of western North America by a T-shaped process on the posterior surface of the posterior gonopod coxites (see species descriptions and illustrations below). The stem of the T (which may be very short) connects the branch to the coxite, and the ventral and dorsal limbs of the T-cap may have various modifications. This definition restricts Taiyutyla to the following described and new species: jonesi (Chamberlin, 1951), napa Shear, 1971, francisca Shear, 1971, benedictae Shear, 1976, prefemorata Shear, 1976, variata Shear, 1976, clarki Shear, 1976, T. amicitia n. sp., T. acuphora, n. sp., and T. tillamook n. sp., the last three described below. See Shear (1971, 1972, 1974, 1976, 2004) and Shear and Krejca (2011) for more detailed morphological data. The remaining previously described species and new species described here are grouped in new genera. The anterior gonopods of Taiyutyla species are usually simple and erect, but sometimes with elaborations at the tips. The posterior gonopod coxites carry an anterior part that may be terminally blunt, abruptly acute, or with processes. The posterior part consists of the T-shaped branch, the ventral process of which is covered in fine, short, unbranched cuticular fimbriae. The dorsal process may be either smooth, or bear long, branched fimbriae. Near the point where the T-shaped branch joins the body of the coxite, a pore may be seen, often with exudate preserved, and subtended by one or two fimbriate branches or regions. These branches may serve as a sheath or groove in which the ventral process of the T-shaped branch is carried. The T-shaped branch may be homologous to what we are calling the pseudoflagellar branch of species of Bifurcatella, n. gen. (see below). Distribution: From the San Francisco Bay region north through northern California, including the Sierra Nevada, to Oregon, Washington and British Colombia, Canada. Females of a probable Taiyutyla species yet to be described have been taken in the Alexander Archipelago and continental southern Alaska (Shelley et al. 2009). The inclusion of T. amicitia n. sp. extends the range of the genus east to Idaho. New records of previously described species, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 9, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Chamberlin, R. V. (1952) Two Oregon millipeds of the order Chordeumida. Natural History Miscellanea of the Chicago Academy of Sciences, 113, 1 - 3.","Chamberlin, R. V. (1951) Eleven new western millipeds. Natural History Miscellanea of the Chicago Academy of Sciences, 87, 1 - 12.","Hoffman, R. L. (1961) Systematic and morphological notes on North American conotyloid diplopods. Transactions of the American Entomological Society, 87, 259 - 272.","Shear, W. A. (1971) The millipede family Conotylidae in North America, with a description of the new family Adritylidae (Diplopoda: Chordeumida). Bulletin of the Museum of Comparative Zoology, 141, 55 - 96.","Shear, W. A. (1976) The milliped family Conotylidae (Diplopoda, Chordeumida): revision of the genus Taiyutyla, with notes on recently proposed taxa. American Museum Novitates, 2600, 1 - 22.","Shear, W. A. (2004) Two new species in the millipede genus Taiyutyla from British Columbia, Canada, and Washington, USA. Myriapodologica, 8, 13 - 20.","Shear, W. A. (1972) Studies in the millipede order Chordeumida (Diplopoda): a revision of the family Cleidogonidae and a reclassification of the order in the New World. Bulletin of the Museum of Comparative Zoology, 144, 151 - 352.","Shear, W. A. (1974) The milliped genus Bollmanella (Diplopoda, Chordeumida, Conotylidae). Psyche, 81, 134 - 146. https: // doi. org / 10.1155 / 1974 / 51907","Shear, W. A. & Krejca, J. K. (2011) Cave millipeds of the United States. IX. A new species of the genus Taiyutyla (Diplopoda, Chordeumatida, Conotylidae) from caves in Sequoia and Yosemite National Parks, California, USA. Journal of Cave and Karst Studies, 73, 93 - 98. https: // doi. org / 10.4311 / jcks 2010 lsc 157"]}

Bifurcatella germania, new species Figs. 220, 221, 224, 225 Types: Male holotype from WASHINGTON: Cowlitz Co., Germany Creek Road 5.5 mi N of Stella at SR-4, elev. 140 m., 46.2593°N, - 123.1344°W, collected 22 November 2003, by W. Leonard, C. Richart; male paratype from Germany Creek Road 5.3 mi N of Stella at SR-4, elev. 140 m., 46.2565°N, - 123.1356°W, collected 11 November 2004, by W. Leonard and C. Richart. Diagnosis: The posterior gonopod coxites are similar to those of the preceding species but the anterior gonopods are quite different, lacking the midlength shelf and sharp taper, and with three large subterminal teeth. Etymology: This species takes its adjectival name from Germany Creek, where the holotype and paratype were collected. Description: Male holotype: Length, 9.0 mm. Nineteen ocelli in triangular group. Metazonites with well-developed shoulders, segmental setae long, curved, acute. Color as described for B. pacifica. Legpairs one and two reduced, pairs three to five enlarged, sixth pair somewhat smaller, seventh pair approximately same size as postgonopodal legs; prefemora three to five with conspicuous knobs. Anterior gonopods (Figs. 220, 222) with mesal branches less than one-third size of lateral branches, broadly flattened in lateral view; lateral branches curved, with large, wing-like expansion anteriorly, bearing three triangular subterminal teeth. Posterior gonopod coxites (Figs. 221, 223) with pseudoflagellar branch tightly reflexed, not strongly tapering, two large fimbriate branches, lateralmost embraces pseudoflagellum, mesal branch subtends small pore. Tenth coxae and eleventh prefemur as usual. Females not collected. Distribution: Known only from the two localities along Germany Creek, which are about 340 meters apart.

Vancouvereuma shawi (Shear, 2004) Figs. 102–104 Taiyutyla shawi Shear, 2004:16. Type locality: BRITISH COLUMBIA: Vancouver Island, Lower Hanging Sump Cave, 16 km SW of Port McNeil, male holotype, juvenile male paratype (VMNH). New record: WASHINGTON: Grays Harbor Co., unnamed tributary of Wynoochee River, Olympic National Forest, elev. 457 m., 47.4852°N, - 123.5231°W, 28 September 2003, W. Leonard, m f. Notes: Vancouvereuma shawi has also been collected in Thurston Co., Washington (Shear 2004), and females resembling this species have been collected as far north as southern Alaska., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 26-28, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (2004) Two new species in the millipede genus Taiyutyla from British Columbia, Canada, and Washington, USA. Myriapodologica, 8, 13 - 20."]}

Taiyutyla variata Shear, 1976 Taiyutyla variata Shear 1976:13. Type locality: Oregon: Jackson Co., Buckley County Park Rest Area, 3.5 mi S of Ruch on the Upper Applegate River, 2.5 mi from State Highway 238, estimated coordinates taken from Cantrall Buckley County Park which is nearer to Ruch and SR-238 than the written description, 42.2188°N, - 123.0687°W, male holotype and several male paratypes (AMNH). New records: OREGON: Multnomah Co., Towle Butte near Clackamas County line, elev. 170 m., 45.4658°N, - 122.4530°W, 8 November 2016, P. Nosler, 3mm f; Gabbert Butte, 45.473124°N, - 122.43770°W, 30 October 2016, P. Nosler, m. Distribution: Also collected at several localities in Josephine Co., Oregon, and evidently sympatric with T. jonesi in Multnomah Co. Descriptions of new species of Taiyutyla, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 12, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (1976) The milliped family Conotylidae (Diplopoda, Chordeumida): revision of the genus Taiyutyla, with notes on recently proposed taxa. American Museum Novitates, 2600, 1 - 22."]}

Complicatella complicata (Shear, 1974), new combination Bollmanella complicata Shear 1974, p. 146. Figs. 192–193 New Records: WASHINGTON: Thurston Co., Black Lake and Belmore Road at 66 th Avenue, Tumwater, elev. 60 m., 46.9889°N, - 122.9659°W, 19 November 2003, W. Leonard, mm; Hospital Creek, above confluence with Skookumchuck River, 5 horiz mi S, 3 horiz mi E of Vail, elev. 245 m., 46.7732°N, - 122.5855°W, 15 December 2003, W. Leonard, K. McAllister, mm ff; Rainier Road SE, 3.0 mi NW of Military Road SE, Fort Lewis Military Reservation, elev. ca. 100 m., ca. 46.9351°N, - 122.7600°W, 29 November 2003, W. Leonard, m; The Evergreen State College campus, Beach Trail N of Driftwood Road, Olympia, elev. 30 m., 47.0791°N, - 122.9776°W, 26 January 2003, W. Leonard, mm ff; same as previous but 16 March 2003, mm; same as previous but 23 March 2003; same as previous but 22 December 2003, mm ff; Priest Point Park, Olympia, elev. ca. 65 m., ca. 47.07°N, - 122.89°W, 5 January 2003, W. Leonard, mm f; same as previous but 2 February 2003, m; same as previous but 14 November 2004, mm ff; along Rock Candy Mtn. Road, Capitol State Forest, elev. ca. 300 m., 47.0256°N, - 123.0735°W, 18 December 2004, W. Leonard, m. Grays Harbor Co., Porter Creek Campground, Capitol State Forest, elev. 90 m., 46.9779°N, - 123.2565°W, 24 January 2005, W. Leonard, m. Cowlitz Co., Germany Creek Road 5.5 mi N of Stella at SR-4, elev. 110 m., 46.2593°N, - 123.1344°W, 22 November 2003, W. Leonard, C. Richart, mm; same as previous but 8 December 2003, m f; same as previous but 11 November 2004, mm ff. Jefferson Co., along Dosewallips River, Dosewallips Road 7.0 mi W of US-101, Olympic National Forest, elev. 115 m., 47.7326°N, - 123.0201°W, 19 November 2003, W. Leonard, m; Falls View Campground, US-101 7.8 mi NW of Brinnon, elev. 150 m., 47.7893°N, - 122.9255°W, 22 February 2003, W. Leonard, m f. Mason Co., SR-119 6 mi W of US-101 at Hoodsport, elev. 310 m., 47.7326°N, - 123.2097°W, 7 February 2003, W. Leonard, mm. King Co., FR-9034 near Granite Mountain Trailhead, Mt. Baker-Snoqualmie National Forest, elev. 580 m., 47.3977°N, - 121.4855°W, W. Leonard, 27 October 2007, mm f; same as previous but 26 March 2004, W. Leonard, C. Richart, m; same as previous but 25 October 2003. Lewis Co., 604 Roswell Road, Centralia, elev. 65 m., 46.7227°N, - 122.9452°W, 25 January 2004, C. Richart, W. Leonard, mm ff; SE end of Riffle Lake, elev. 250 m., 46.4644°N, - 122.1705°W, 9 April 2004, W. Leonard, C. Richart, m; Slide Creek, Weyerhaeuser 4000 Rd 0.8 mi SW of Pe-Ell-McDonald Rd., elev. 190 m., 46.5250°N, - 123.1913°W, 4 December 2004, W. Leonard, C. Richart, mm ff; same as previous but 3 January 2005, W. Leonard; Stillman Basin, 1.8 mi on 4000 Weyerhaeuser from Pe-Ell McDonald Rd., elev. 290 m., 46.5145°N, - 123.2015°W, 4 December 2004, W. Leonard, C. Richart, from the litter and woody debris of a mixed forest including Thuja plicata, Alnus rubra, Rubus spectabilus, Pseudotsuga menziesii, and Polystichum munitum, m f; North Fork of Newaukum River, below confluence with Middle Fork, elev. 88 m., 46.6046°N, - 122.8481°W, 29 December 2004, W. Leonard, m. Notes: This species was described from 1 mi W of Bayshore and 4 mi N of Shelton, Mason Co., Washington, and heretofore was known only from the type locality. New collecting revealed a much wider distribution, detailed above and in the supplemental KML map, and show this species to be the most widely distributed and most commonly collected conotylid in western Washington. All these populations are linked by identical anterior gonopods (Fig. 192), but the posterior gonopod coxites (i.e., Fig. 193) are so complicated that it is difficult to view them from exactly the same angle and make comparisons. Thus it is not possible really to be sure that all collections belong to the same species. This can be worked out later with genetic data. Descriptions of new species of Complicatella

Calityla trinitaria, new species Figs. 58–66 Types: Male holotype and female paratype from CALIFORNIA: Trinity Co., Indian Valley, parking area for Indian Creek Valley Cave, elev. 700 m., 40.5634°N, - 123.4197°W, collected 27 October 1990, by D. Ubick, W. Rauscher (CAS). Diagnosis: Taiyutyla trinitaria is sympatric, possibly syntopic, with the previous species but may easily be distinguished from it by the far more complicated coxites of the male posterior gonopods, and the femoral knobs of the male pregonopodal legs, which are on legpairs three to seven, while in T. ubicki the knobs are found only on legpairs six and seven. Etymology: The species name is an adjective based on Trinity County. Description: Male holotype: Length, 11.0 mm. 17 ocelli in triangular eyepatch. Metazonites with low, narrow shoulders on rings 4–26. Color pale cream-tan lightly mottled brown. Legpairs one and two reduced, three to seven enlarged, with moderate angular knobs basal on femora three and to seven (Figs. 61–66). Anterior gonopods (Fig. 58) simple, blunt, roughened apicolaterally with small cuticular teeth (Fig. 59). Posterior gonopod coxites bearing large, complex fimbriate branch distal of midlength, distally acuminate, hooked (Fig. 60). Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Female 11.0 mm long, similar to male in nonsexual characters. Distribution: Known only from type locality. Note: This species is clearly quite different from its sympatric congener T. ubicki., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 20, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Bifurcatella bifurcata (Shear, 1974), new combination Figs. 204, 205 New records: Bollmanella bifurcata Shear 1974, p. 143. OREGON: Umatilla Co., Umatilla National Forest, FS- 65 0.4 mi E of Tiger Creek Road, elev. 1520 m., 45.9442°N, - 118.0094°W, 24 October 2003, W. Leonard, mm ff. Union Co., Little Indian Creek, 8 mi N of Cove, ca. 45.4134°N, - 117.7988°W (estimated coordinates taken from 8 air mi N of Cove along FS-6205), 10 September 1977, R. K. Johnson, mm ff (2 vials, FSCA). Notes: This species was originally described (Shear 1974) from 2 mi west of Joseph, along Hurricane Creek, Wallowa Co., Oregon, from specimens collected 23 November 1968. The new records are consistent with a range encompassing northeastern Oregon. In Fig. 204, the mesal branch of the anterior gonopod has been broken; it is normally about half the length of the lateral branch. Descriptions of new species of Bifurcatella

Karagama ladybird, new species Figs. 105–110 Types: Male holotype and female paratype from CALIFORNIA, Humboldt Co., Redwood National Park, Lady Bird Johnson Grove, Bald Hills Road 2.6 mi E of US-101, elev. 445 m., 41.3047°N, - 124.0166°W, collected 21 December 2006, by C. Richart and A. Fusek, from the forest litter of a redwood forest including Sequoia sempervirens, Rhododendron macrophyllum, and Polystichum munitum (CAS). Three male paratypes and one female paratype from Redwood National Park, along Redwood Creek, 40.9946°N, - 123.8605°W, collected 27 November 1976 by R. K. Johnson (FSCA). Diagnosis: The complex anterior gonopods (Fig. 1) with a distinctive posterior branch and fimbriate regions separate this species from all others known so far. Karagama lupus Shear, 2004 has a basal branch, but it is subequal to the main branch and fimbriate regions are lacking. Also, T. lupus has broad paranota on its metazonites. Karagama loftinae (Shear & Krejca, 2011) has the posterior branch of the anterior gonopod with a comb-like distal tip. Etymology: The species name is a noun in apposition referring to the type locality and the late Lady Bird Johnson, former First Lady of the United States, who had a deep interest in the preservation of natural areas. Description: Paratype male from Redwood National Park: Length, 8.0 mm. 22 ocelli in triangular eyepatch. Metazonites with prominent shoulders on rings 6–25. Coloration pale tan, lightly marked darker brown. Legpairs one and two reduced, three to seven enlarged, basal femoral knobs on legpair four, legpair five basally swollen, otherwise pregonopodal femora not modified. Anterior gonopods (Figs. 105, 106, 109) with main (mesal) branch bearing fimbriae from colpocoxal region near base, additional fimbriate region near tip, ending in short, blunt subapical process set with acute, fine cuticular projections. Posterior gonopods with distinct, band-like sternum, coxae partially fused to prefemora, prefemora with anterobasal, short process bearing few setae; posterior gonopod coxites (Figs. 107, 108, 110) simple, long-triangular, with fimbriate lamellae basally, apically hooked, distal row of blunt teeth on lateral side. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with blunt, dorsally directed knobs. Female from Redwood National Park 8.2 mm long, otherwise nonsexual characters as in male. Distribution: Known only from Redwood National Park.

Loomisiella pylei, new species Figs. 244–246 Types: Male holotype, male and female paratypes from WASHINGTON: Wahakiakum Co., Lower Hendrickson Canyon, 46.3693°N, - 123.6657°W, elev. 30 m, old-growth forest, collected 23 January 2004, by W. Leonard, M. Leonard, C. Richart, R. M. Pyle, and K. Novoselić. Diagnosis: See under preceding species. Etymology: The species name honors Robert Michael Pyle, a notable lepidopterist, author, conservation advocate, and as is this species, denizen of the Willapa Hills. Description: Male: Length, 5.0 mm. Fourteen ocelli in oval-triangular group. Metazonital shoulders poorly developed; segmental setae curved, acute. Legpairs one, two reduced, pairs three, four encrassate, pairs five to seven near same size as postgonopodal legs; femora three, four with basal femoral knobs acute, posteriorly directed.Anterior gonopods (ag, Figs. 244, 245) with large anterior sternum, widely separated, curving lateral to posterior gonopod coxites, apically decurved but without any apical expansion, small lateral branch tightly appressed to gonopod (free in Fig. 245 due to compression on microscope slide). Posterior gonopod telopodites reduced but relatively larger than in L. evergreen; posterior gonopod coxites (pgc, Figs. 244, 245) short, broad, anteriorly cupped, with broad, lamellate process mesally, basal pseudoflagellum originating posteriorly, two processes distolaterally, more lateral process composed of filaments (seen as separate in Fig. 246 due to compression on microscope slide), mesal process curved, apically fimbriate. Coxae 10, prefemora 11 as usual. Female 5.2 mm long, nonsexual characters as in male. Distribution: WASHINGTON: Lewis Co., FS-25 8.5 mi S of Randle, 46.4409°N, - 121.9966°W, elev. 330 m, 21 December 2003, W. Leonard, C. Richart m f; same as previous but 6 December 2003, m. Pacific Co., Cement Creek, SR-401 2.5 mi S of Parpala Road in Naselle, 46.3341°N, - 123.8002°W, elev. 30 m., 15 January 2006, W. Leonard, C. Richart, from the litter of a riparian forest including Alnus rubra, Picea sitchensis, Sambucus cf. caerulea, and Polystichum munitum, m ff; Trap Creek Road 1.7 mi S of SR-6, Trap Creek Basin, 46.5403°N, - 123.6297°W, elev. 60 m., 19 November 2005, W. Leonard, C. Richart, mm ff. Wahkiakum Co., only known from the type locality., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 69, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Taiyutyla amicitia, new species Figs. 27–36 Types: Male holotype, 16 male and 8 female paratypes from IDAHO: Kootenai Co., Beauty Creek, Idaho Panhandle National Forests, 47.6031°N, - 116.6453°W, collected 8 November 2004, by C. Richart, W. Leonard, J. Baugh. Diagnosis: Taiyutyla amicitia is unique in having femoral knobs only on legpairs four and five. Etymology: The species is named to commemorate the friendship that formed over millipede species discovery (L., amicitia, a noun in apposition) between WAS, CHR and William P. Leonard. Description: Paratype male from Beauty Creek: Length, 10.0 mm. Twenty-two ocelli in triangular eyepatch. Metazonites with low shoulders on all trunk rings from three to 25. Color light brown, marked darker purplish brown, markings more distinct anteriorly. Legpairs one and two reduced, three to seven enlarged, mesal knobs present only on femora four and five, mesal and large on both (Figs. 32–36). Anterior gonopods (Figs. 27, 28, 36) simple, with short, triangular process subdistally. Posterior gonopod coxites (Fig. 29, 31) complicated, with T-shaped branch (T, Fig. 29) originating subdistally, a strong, triangular tooth at its base; this branch with two long, thin processes, the more distal one set with small teeth. Terminus of coxite strongly hooked. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Female 14.5 mm long, similar to male in nonsexual characters. Distribution: IDAHO: Shoshone Co., Hobo Cedar Grove Botanical Area, Idaho Panhandle National Forests, elev. 1295 m., 47.0880°N - 116.1152°W, 14 October 2006, W. Leonard, C. Richart, A. Fusek, Habitat: oldgrowth Tsuga heterophylla, Thuja plicata, Abies grandis, under litter and woody debris, m f (CAS). Notes: This species, the first Taiyutyla from Idaho, is syntopic with Brunsonia pulchra, n. sp. (see below) at Beauty Creek, the type locality for both of them. The original collection was mixed, and after the two species were sorted out by size, the gonopod differences became apparent. Taiyutyla amicitia is perhaps the Taiyutyla species closest in gonopod form (particularly the posterior gonopod coxites) to some Bifurcatella, n. gen. species, but the large femoral knobs on legpair five and the somewhat larger body size argue for including it in Taiyutyla.

Corypus cochlearis Loomis & Schmitt 1971 Corypus cochlearis Loomis & Schmitt 1971, p. 123; Shear, 1976, p. 19; Shelley, 1997, p. 123. New record: MONTANA: Sanders Co., Skeleton Creek, 47.4634°N, - 115.8868°W, elev. 973 m., 13 October 2006, P. Hendricks, female presumed this species., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 71, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Loomis, H. F. & Schmitt, R. (1971) The ecology, distribution and taxonomy of the millipeds of Montana west of the continental divide. Northwest Science, 45, 107 - 131.","Shear, W. A. (1976) The milliped family Conotylidae (Diplopoda, Chordeumida): revision of the genus Taiyutyla, with notes on recently proposed taxa. American Museum Novitates, 2600, 1 - 22.","Shelley, R. M. (1997) Corypus kavanaughi, n. sp., a large-bodied chordeumatoid milliped from Idaho, with a review of the genus (Conotylidae). Myriapodologica, 4, 121 - 126."]}

Brunsonia chelanoparva, new species Figs. 168–172 Types: Male holotype, five male paratypes and nine female paratypes from WASHINGTON: Chelan Co., Lake Chelan, Wenatchee National Forest, Fields Point, elev. 360 m., 47.9699°N, - 120.2160°W, collected 26 October 2007, by W. Leonard, J. Baugh. Diagnosis: The anterior gonopods are roughly similar to those of B. albertana, but more acute at the tip and with a short, anterior branch, while the posterior gonopod coxites are less complex, relatively smaller than that species, and with a very prominent warty area on the main branch. Etymology: The species name is a combination of Lake Chelan, the type locality, and Latin parvus, noting the fact that it is the smaller of two species of Brunsonia syntopic there. Description: Male holotype: Length, 12.5 mm. 21–23 ocelli in triangular patch. Metazonites with well-developed shoulders on all trunk segments, on midbody segments shoulders slightly below midline dorsum, with angular posterior corners, segmental setae acute, curved, moderately long. Color pale to medium brown, with well-defined dark purplish brown spots on segmental shoulders, lateral prozonites, less well-defined spots on either side of midline. Legpairs three to seven slightly encrassate compared to postgonopodal legs, femoral knobs on legpairs 4 and 5 only. Anterior gonopods (Figs. 168, 170, 171) very similar to those of B. albertana, but apically more acute, anterolateral rugose area less so. Channel from pore to gonopod tip occupied by long, filamentous pseudoflagellum (Fig. 171). Posterior gonopod coxites (Figs. 169, 172) massive, channeled branch somewhat reduced, basal process displaced toward body of coxa, other branches apparently absent. Posterior gonopod prefemora distinctly clavate. Legpair 10 enlarged, with glands, legpair 11 prefemora with dorsally directed stout processes. Female 12.5–13.0 mm long, nonsexual characters similar to male. Distribution: Known only from the type locality. Notes: The filamentous structure occupying the posterior groove of the anterior gonopod is not known from any other species. It is hollow or possibly channeled, with a pore opening at the tip, which is slightly flared. The species and the following were in a mixed collection and when sorted out by size proved to be two different species. It is unusual to have two species of the same genus syntopic; perhaps each has distinct microhabitat preferences that were not recorded on the collection label.

Taiyutyla acuphora, new species Figs. 19–21, 25, 26 Types: Male holotype and female paratype from Washington, Lewis Co., Stillman Basin, Weyerhaeuser 4000 Road 1.8 mi SW of Pe-Ell McDonald Road, elev. 282 m., 46.5145°N, - 123.2015°W, collected 4 December 2004, by W. Leonard, C. Richart from the litter and woody debris of a mixed forest including Thuja plicata, Alnus rubra, Rubus spectabilus, Pseudotsuga menziesii, and Polystichum munitum. Diagnosis: The anterior gonopods have a unique, anterior wing-like extension, while the posterior gonopod coxites appear squared off, but have a distal, acute terminal process and femoral knobs are borne on male legpairs 47. No other species of Taiyutyla has this combination of characters. Etymology: The species epithet means “needle bearer” and refers to the acute terminal process of the posterior gonopod coxite. Description:Male holotype.Length, 12.5 mm.Twenty to twenty-two ocelli in triangular eyepatch.Metazonites with well-developed shoulders, segmental setae long,acute,curved.Color light brown,irregularly marked brownish purple. Legpairs one and two reduced, three to seven moderately enlarged, podomeres not swollen, femoral knobs on pairs 4–7. Anterior gonopods (Figs. 19, 20, 25) distinctly clavate, distally expanded anterior into broad wing, bearing small, distal posterior process. Posterior gonopod coxites (Figs. 21, 26) short, stout, apically squared off, but with long, curved, acute process arising from lateral angle apically; ventral arm of T-shaped branch (T, Fig. 26) with fine fimbriae, embraced by fimbriate region, distinct pore near base of this region (with coagulated secretion in some specimens). Tenth coxae of normal size, with glands; eleventh prefemora with long, curved distal processes. Female 13.5 mm long. Nonsexual characters as in male. Distribution: WASHINGTON: Grays Harbor Co., Middle Satsop Road 2 mi N of Brady near Satsop River, elev. 15 m., 47.0245°N, - 123.5196°W, 3 January 2003, W. Leonard et al., under maple leaves, mm ff. Notes: This is the first Taiyutyla species to be recorded from western Washington., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 12-13, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Subfamily Idagoninae Buckett & Gardner 1967 Idagona Buckett & Gardner 1967, p. 119. New records of a previously described species, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 75, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Buckett, J. S. & Gardner, M. R. (1967) A new family of cavernicolous millipedes with the description of a new genus and species from Idaho (Diplopoda: Chordeumida: Chordeumidea). The Michigan Entomologist, 1, 117 - 126."]}

Taiyutyla amicitia, new species Figs. 27–36 Types: Male holotype, 16 male and 8 female paratypes from IDAHO: Kootenai Co., Beauty Creek, Idaho Panhandle National Forests, 47.6031°N, - 116.6453°W, collected 8 November 2004, by C. Richart, W. Leonard, J. Baugh. Diagnosis: Taiyutyla amicitia is unique in having femoral knobs only on legpairs four and five. Etymology: The species is named to commemorate the friendship that formed over millipede species discovery (L., amicitia, a noun in apposition) between WAS, CHR and William P. Leonard. Description: Paratype male from Beauty Creek: Length, 10.0 mm. Twenty-two ocelli in triangular eyepatch. Metazonites with low shoulders on all trunk rings from three to 25. Color light brown, marked darker purplish brown, markings more distinct anteriorly. Legpairs one and two reduced, three to seven enlarged, mesal knobs present only on femora four and five, mesal and large on both (Figs. 32–36). Anterior gonopods (Figs. 27, 28, 36) simple, with short, triangular process subdistally. Posterior gonopod coxites (Fig. 29, 31) complicated, with T-shaped branch (T, Fig. 29) originating subdistally, a strong, triangular tooth at its base; this branch with two long, thin processes, the more distal one set with small teeth. Terminus of coxite strongly hooked. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Female 14.5 mm long, similar to male in nonsexual characters. Distribution: IDAHO: Shoshone Co., Hobo Cedar Grove Botanical Area, Idaho Panhandle National Forests, elev. 1295 m., 47.0880°N - 116.1152°W, 14 October 2006, W. Leonard, C. Richart, A. Fusek, Habitat: oldgrowth Tsuga heterophylla, Thuja plicata, Abies grandis, under litter and woody debris, m f (CAS). Notes: This species, the first Taiyutyla from Idaho, is syntopic with Brunsonia pulchra, n. sp. (see below) at Beauty Creek, the type locality for both of them. The original collection was mixed, and after the two species were sorted out by size, the gonopod differences became apparent. Taiyutyla amicitia is perhaps the Taiyutyla species closest in gonopod form (particularly the posterior gonopod coxites) to some Bifurcatella, n. gen. species, but the large femoral knobs on legpair five and the somewhat larger body size argue for including it in Taiyutyla., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 13, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Bollmanella Chamberlin, 1941 Bollmanella Chamberlin 1941: 12; Shear, 1974: 135. Type species, Bollmanella oregona Chamberlin, 1941. Diagnosis: Small to medium-sized conotylids (4.5–11 mm long). Anterior gonopods unbranched, simple. Coxites of posterior gonopods often with pseudoflagellar branch carried in sheath at least partially fused with main body of coxite. Pregonopodal legs three and four encrassate, with distomesal femoral knobs, or legs three to seven encrassate, five, six and seven with prefemoral knobs ( Bollmanella washingtonensis, n. sp. ). Distinct from the closely related Bifurcatella, n. gen. in the unbranched anterior gonopods, Bifurcatella having the gonopods distinctly twobranched, the median branch usually the shorter, and from Complicatella, n. gen. in having the pseudoflagellum single (multiple in Complicatella); Complicatella also has the sheath of the pseudoflagellum entirely separate from the main body of the coxite; it appears as a basal branch of the main pseudoflagellum. Notes: The eight species of Bollmanella and their known distributions are as follows: Bollmanella oregona Chamberlin, 1941, from Josephine, Douglas and Jackson Counties in Oregon (for a full list of localities, see Shear, 1974); B. reducta Shear, 1974, from near Ashland, Jackson Co., Oregon; B. laminata Shear, 1974, from near Timber, Washington Co., Oregon; B. bella Shear, 1974, from near Allegany on the Douglas/Coos Counties border; B. camassia Shear, 1974, from Camas Valley, Douglas Co., Oregon; B. washingtonensis, n. sp., from Grays Harbor, Thurston, King, Mason, Cowlitz, Jefferson and Pacific Counties, Washington, B. bombus, n. sp. from Idaho Co., Idaho, and B. leonardi, n. sp. from Cowlitz and Thurston Counties, Washington. See descriptions below for detailed localities of the new species. Previously described species of the genus transferred elsewhere: B. unca Shear, 1974, from two localities in Douglas Co., Oregon and B. bifurcata Shear, 1974 from Wallowa Co., Oregon, are moved to Bifurcatella, n. gen. and B. complicata Shear, 1974 from Mason Co., Washington is the type of the new genus Complicatella. This genus seems to be close to Taiyutyla in many respects, and some species in both genera are difficult to place. The type species, B. oregona, had previously been a somewhat aberrant member of the genus in that the main branch of the posterior gonopod coxite is quite narrow, and the sheathing branch is free for much of its length. The three new species described below more closely conform to this plan, despite significant geographic separation from B. oregona. The species removed from the genus seem quite dissimilar to B. oregona and the three new ones; the name Bollmanella has to go with B. oregona, its type. Descriptions of new species of Bollmanella, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 30, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Chamberlin, R. V. (1941) New western millipeds. Bulletin of the University of Utah, Biological Series, 6, 3 - 23.","Shear, W. A. (1974) The milliped genus Bollmanella (Diplopoda, Chordeumida, Conotylidae). Psyche, 81, 134 - 146. https: // doi. org / 10.1155 / 1974 / 51907"]}

Brunsonia wenatchee, new species Figs. 159–167 Types: Male holotype, five male and two female paratypes from WASHINGTON: Kittitas Co., Wenatchee National Forest, North Fork Teanaway River at the Stafford Creek confluence, elev. 835 m., 47.3482°N, - 120.8480°W, collected 14 October 2004, by W. Leonard, J. Baugh (CAS). Diagnosis: No other species of Brunsonia has a long, blade-like lateral branch on the coxite of male posterior gonopods; the distal segment of the male posterior gonopod telopodite is shorter than the proximal, which attaches to the coxa by a narrowed stem. Etymology: The species epithet is a noun in apposition, referring to the type locality in the Wenatchee National Forest, Washington. Description: Male paratype: Length, 10.0 mm. 22 ocelli in triangular eyepatch. Metazonites with moderately well-developed shoulders on all but 28–30. Color tan, marked with dark purplish brown, marking uniform along trunk. Legpairs one and two reduced, three to seven enlarged, mesal knobs present basally on femora four to six, on femora three lower, basal swellings. Knobs of femora four to six set with small, cuticular teeth (Figs. 163–167). Anterior gonopods (Figs. 159, 161) slender, acuminate, terminally hooked, with short, posterior subterminal branch finely toothed. Posterior gonopod coxites (Figs. 160, 162) robust, complex, with lateral blade-like branch, subterminal branch short, ending in tuft of cuticular filaments. Legpair 10 coxae of normal size, with small glands, legpair 11 femora with long, thin, dorsally directed knobs. Female 6.5 mm long, similar to male in nonsexual characters. Distribution: Known only from the type locality. Notes: The anterior gonopod is very unusual for a species of Brunsonia, but the deep pit in the gonopod and the complicated posterior gonopod coxites argue for its placement here. There appears to be a pore enveloped in the posterior, slightly fimbriate branch of the anterior gonopod. We were not able to reliably homologize the processes of the posterior gonopod with the typical pattern (see illustrations of B. albertana, above). More study is required., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 42, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782

Lophomus laxus Loomis & Schmitt 1971 Figs. 255–257 Lophomus laxus Loomis & Schmitt 1971, p. 125; Shear, 1976, p. 17. New records: MONTANA: Granite Co., Welcome Creek, 46.5638°N, - 113.7044°W, elev. 1239 m., 2 October 2006, P. Hendricks, mm; Welcome Creek divide, 46.6438°N, - 113.6949°W, elev. 1553 m., 1 October 2006, P. Hendricks, mm. Notes: The SEMs presented here compliment the drawings in Shear (1971). The two-branched gonopods (Figs. 255, 256) are unique in that the anterior branches, which are set with tiny cuticular projections, seem to arise from the sternal part of the coxosternum, which can be seen in a posterior view (Fig. 256) to be poorly sclerotized. The posterior gonopod coxites (Fig. 257) have an anterior movable flagellum made up of several separate filaments. Descriptions of new species of Lophomus

Vancouvereuma, new genus Type species: Vancouvereuma shawi (Shear, 2004), new combination Diagnosis: Distinct from Taiyutyla, which it otherwise resembles, in the posterior gonopod coxites (Figs. 103, 104), which are broad and stout, lack a T-shaped branch but have a broad, curved, basal posterior process extending between the body of the coxa and the prefemur (B, Fig. 103). The anterior gonopods (Fig. 102) are broad and simple. Etymology: From Vancouver Island and - euma, a combining stem often used in generic names in Chordeumatida. Feminine in gender. Distribution: Vancouver Island, British Columbia, Canada, and northwestern Washington, USA. Notes: Vancouvereuma shawi (Shear, 2004) is the only known species. For a complete description and additional illustrations, see Shear (2004). New records of previously described species, Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on page 26, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Shear, W. A. (2004) Two new species in the millipede genus Taiyutyla from British Columbia, Canada, and Washington, USA. Myriapodologica, 8, 13 - 20."]}

Complicatella neili, new species Figs. 196, 197, 202, 203 Types: Male holotype and male and female paratypes from NEVADA: Pershing Co., Dragon’s Gate Cave, elev. 2660 m., 40.5551°N, - 118.1601°W, collected 19 August 2008, by Neil Marchington. Diagnosis: Unlike the other two species of Complicatella, C. neili has anterior gonopods with an anterior process that is almost as long as the body of the gonopod. The posterior gonopod coxites have the characteristic basally arising, sheathed flagellum but only two pseudoflagella. The ocelli are poorly formed and metazonital shoulders are practically absent. Etymology: The species is named for the collector, Neil Marchington, an indefatigable explorer of northwestern caves. Description: Male holotype: Length, 7.5 mm. Ocelli 13–15 in oblong patch, poorly formed and pigmented (some paratypes have as few as 10 ocelli). Metazonites with no visible shoulders, segmental setae very long, curved, acute. Color white, with some very faint tan shading ventrally on few anterior segments. Legpairs one and two reduced, legpair three enlarged with swollen femur; legpairs four to seven only a little more robust than postgonopodal legs, lacking femoral knobs. Anterior gonopods (Figs. 196, 202) strongly clavate, apically cupped distal to distinct shoulder, with clavate, large, anterior branch. Posterior gonopod coxites (Figs. 197, 201) complex, body of coxa swollen, long, thin flagellum arises mesally near the base, curving sigmoidally into sheathing branch, two long pseudoflagella present, small pore at base of apical expansion of main branch. Coxa 10 not enlarged, with gland. Prefemur 11 with basal process projecting dorsally. Female 8.0 mm long, nonsexual characters as in male. Distribution: Known only from the type locality. Notes: This species is distant in its location from the others of the genus and its placement may be doubtful. We put it in Complicatella based on the similarities of the gonopods with C. complicata. It was totally unexpected to find a troglobiotic conotylid in the circumstances of the type locality, a barren mountaintop in Nevada in a range surrounded by sagebrush desert. However, its appearance there may be somewhat explained by these habitat notes supplied by Neil Marchington (pers. comm. to WAS, 2018): “The Nevada specimens, collected 19 Aug 2008, came from a small cave called Dragon’s Pit in the Humboldt Range of Pershing County, Nevada at 8682 feet elevation. The specimens were clinging to a soft, wet calcite paste on the ceiling of the cave about 60–80 feet from the entrance. The cave is 105 miles NE of Reno or 35 miles SW of Winnemucca. The cave is on a nearly barren limestone peak with scattered sage, gooseberry, rice grass and (strangely) stunted mountain maple. There are stunted juniper growing about 700 feet lower in elevation than the cave. The Humboldt Range was likely nearly an island during the Pleistocene. The projected levels of glacial Lake Lahontan show only a narrow land bridge at the south end of the range connecting to any other land. The confirmed and studied shoreline puts at least 50% of the range as a shoreline area (the west and north sides were for sure).” “The climate history of the region can be summarized fairly easily. About 4 million years ago, the Sierra Nevada Range started to rapidly uplift forming a massive fault block. The peaks started increasingly taking more of the Pacific moisture. This process led to the massive desert regions such as Carson, Great Basin and Mohave. The Ice Age’s lower temps allowed the water that did get through to exceed evaporation in the Great Basin and Mohave. Giant lakes like Lahontan filled most of the basins, and crept over modern passes, forming vast waterways surrounding many mountain ranges entirely. The Humboldt Range was a giant island in this lake. We know from archaeological sites the region had vast marshlands where the natives hunted ducks and other animals. Tule reeds were used in many native artifacts, such as duck decoys, baskets, and sandals. Many of the higher peaks were glaciated, particularly the eastern Sierra Nevada.” “Around 14,000 years ago, the Wisconsin Glaciation was waning. As temperatures rose, the basins gradually started to lose water each year. There were at least pockets of ice age megafauna, such as short-nosed bears, lurking in the mountains as recently as ~8000 years ago, but most of the population was probably gone around 10,000 years ago. The desertification of the area seems to be continuing and possibly accelerating. There are studies monitoring plant communities moving to higher elevations in the region. Most of the Ponderosa groves in Nevada are of 200-year-old trees, with nothing younger present. Even 500 years ago, Ponderosa were growing in lower areas in the Mohave, where only cacti grow today. Douglas Fir cones have been found in 30,000 year old middens throughout areas of Arizona, Nevada, and Utah where there are only cacti or juniper today.” The geological and climatic history of Lake Lahontan is summarized in Morrison (1963). So the clear suggestion is, as with so many other arthropods now limited to caves, that the ancestors of the present population lived on the surface during a moist glacial maximum, and were also able to occupy caves. With the drying and warming starting 140 centuries ago, surface populations were gradually extinguished, leaving only those in the hospitable, buffered environment of the caves. Subsequently the remnant population developed the modest troglobiotic adaptations we see today., Published as part of Shear, William A., Richart, Casey H. & Wong, Victoria L., 2020, The millipede family Conotylidae in northwestern North America, with a complete bibliography of the family (Diplopoda, Chordeumatida, Heterochordeumatidea, Conotyloidea), pp. 1-78 in Zootaxa 4753 (1) on pages 51-53, DOI: 10.11646/zootaxa.4753.1.1, http://zenodo.org/record/3983782, {"references":["Morrison, R. B. (1963) Lake Lahontan: geology of southern Carson Desert, Nevada. Geological Survey Professional Paper, 401, 1 - 156. https: // doi. org / 10.3133 / pp 401"]}