The genus Tetralicia Harrison (Hemiptera: Sternorrhyncha: Aleyrodidae) in California, U. S. A., with the description of five new species and a redescription of Tetralicia granulata Sampson & Drews, 1941 Author Ellenrieder, Natalia Von 0000-0002-1159-2019 Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento, CA 95832 - 1448, U. S. A. & natalia. von. ellenrieder @ cdfa. ca. gov; https: // orcid. org / 0000 - 0002 - 1159 - 2019 natalia.von.ellenrieder@cdfa.ca.gov Author Gill, Raymond J. 0009-0005-9047-2742 Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento, CA 95832 - 1448, U. S. A. & Plant Pest Diagnostics Center, California Department of Food & Agriculture, 3294 Meadowview Road, Sacramento, CA 95832 - 1448, U. S. A. & rayromagill @ gmail. com; https: // orcid. org / 0009 - 0005 - 9047 - 2742 * Corresponding author rayromagill@gmail.com text Zootaxa 2024 2024-10-22 5527 1 1 129 http://dx.doi.org/10.11646/zootaxa.5527.1.1 journal article 305014 10.11646/zootaxa.5527.1.1 0d9d4ebd-5dd8-473c-80b9-19960053d8b4 1175-5334 14021823 771D2E7B-4025-45BF-B328-6EC8A8851ECD Tetralicia ceanothi Sampson, 1945 Figs 5, 6 , 50–68 Tetralicia ceanothi Sampson, 1945: 59 ; Valencia & Evans 2024: 215 . Aleuropleurocelus ceanothi : Drews & Sampson 1956: 281 ; Mound & Halsey 1978: 59 ; Evans 2007: 170 ; Dubey, 2021: 2 . Material examined . 440 puparia: U.S.A. , California ; 404 slide mounted: 1, W.M. Yothers coll. [ USNM ]; 1, on Arctostaphylos viscida , T. Bridges coll. [ USNM ]; 2, Southern California , on oak, xii.1908 , R.S. Woglum coll. [ USNM ]; 12, Sierra Madre, xii.1907 , R.S. Woglum coll. [ USNM ]; Imperial County: 5, Ocotillo, on manzanita, 27.iv.1972 , Flock, Paddock & Pineda coll. [ CSCA ]; 3, Ocotillo ( 32.743778 N , 116.002915 W ), on Larrea tridentata , 19.ii.2023 , J. Bailey coll. [ CSCA ]; 4, same data but Flock & Paddock coll. [ CSCA ]; 4, same data but on Eriogonum sp. , 23.iii.1968 , R.A. Flock coll., CDFA PDR 68019-19 [ CSCA ]; 8, 15 mi W of El Centro, on Rattlesnake weed ( Euphorbia sp. ), 17.iv.1985 , R.J. Gill coll. [ CSCA ]; 3, El Centro, on Euphorbia sp. , 28.v.1993 , G. Curtner coll., CDFA PDR 843632A [ CSCA ]; 3, Glamis, on Euphorbia sp. , 2.ii.1966 , R.A. Flock coll., CDFA PDR 66I14-40 [ CSCA ]; 1, Indian Wells, on Trixis californica , R.D. Goeden coll., CDFA PDR TC 85-5A [ CSCA ]; San Diego County: 11, Lake Sutherland, on manzanita, 5.iii. 1983 , R.J. Gill coll. [ CSCA ]; 7, Alpine, 1.iii.1984 , R. Dowell coll. [ CSCA ]; 25, San Felipe, Junction of Highways 52 & 522, on manzanita, 18.iii.1986 , R.J. Gill coll. [ CSCA ]; 6, San Diego, on Eriogonum , 8.ii.1973 , E. Paddock & F. Swall coll. [ CSCA ]; 1, Julian, Desert View Park ( 33.049617 N , 116.569002 W ), 30.vii.2022 , on Ceanothus foliolosus , J. Bailey coll. [ CSCA ]; Orange County: 11, Rancho Capistrano, on manzanita, 4.ix.1983 , M. Rose coll. [ CSCA ]; 5, same data but 1 mile N [ CSCA ]; Riverside County: 4, Riverside, on Ceanothus , 28.xi.1950 , L.D. Anderson coll. [ USNM ]; 3, La Quinta, on Larrea tridentata , xi.1951 , R.C. Dickson coll. [ USNM ]; 9, Twin Pine Ranch Road, Banning-Idyllwild Highway, on manzanita, 15.iii. 1986 , R.J. Gill coll. [ CSCA ]; 14, El Cariso Fire Station, Ortega Highway, on manzanita, 19.iii.1986 , R.J. Gill [ CSCA ]; 6, Joshua Tree National Monument, on Monardella robinsonii , 4.ix.1979 , K. Stolte coll. [ CSCA ]; Los Angeles County: 7, Los Angeles, on Ceanothus cyaneus , 12.xi.1943 , R.H. Smith coll. [ USNM ]; 7, same data but on Ceanothus thyrsiflorus [ USNM ]; 3, 2008 La Salle Ave., Los Angeles, on Ceanothus sp. , 24.iv.1933 , F. Wilson coll. [ BME ]; 11, Arcadia, on Ceanothus thrysiflorus , 17.xii.1968 , H.G. Walker coll. [ CSCA ]; 1, same data but 15.vii.1969 [ CSCA ]; 3, same data but on Acacia melanoxylon , 3.ix.1969 [ CSCA ]; 20, same data but on Ceanothus cyaneus , 3.xi.1970 [ CSCA ]; 4, Sylmar, on manzanita, 11.vi.1976 , Kemper coll., CDFA PDR 76F11-42 [ CSCA ]; 3, Monrovia, 152 m , on Eriogonum fasciculatum , 20.viii.1915 , Phillipson coll., CDA Herbarium 0022584 [ CSCA ]; 2, 3 mi S of Gorman, Hwy 99, on Eriogonum fasciculatum , 3.vii.1953 , A.C. Browne coll., CDA Herbarium 0022588 [ CSCA ]; 5, Mount Wilson [ USNM ]; 3, same data but 20.ii.1910 , H.M. Russell coll. [ USNM ]; 3, same data but 4000–5886 feet [ USNM ]; 2, San Gabriel Mountains near Camp Rincon, on Arctostaphylos sp. , 4.vii.1911 , P.H. Timberlake coll. [ USNM ]; 3, Santa Ana Botanic Garden, Claremont, on Arctostaphylos pajaroensis , 11.xii.1957 , Laird coll. [ USNM ]; 3, same data but on Ceanothus sp. , Johnson coll. [ USNM ]; San Bernardino County: 4, E Highland, on outer ends of leaves of Eriogonum sp. , 11.xii.1957 , Laird coll. [ USNM ]; 25, Mountain Home Village, Mill Creek, 1219 m . a.s.l., on manzanita, 21.iii.1986 , R.J. Gill [ CSCA ]; Tulare County: 15, Three Rivers, on manzanita, 24.vii.1982 , Celis & O’Neil coll. [ CSCA ]; 2, Porterville, on Eriodictyon californicum , 7.iii.1962 , H.H. Keifer coll. [ USNM ]; Inyo County : 2 paratypes ?, Bishop , on Ceanothus cuneatus , 29.iii.1940 , N. Stahler coll. [ex W.W. Sampson Coll., CSCA ] ; Alameda County: 2, Pleasanton, on Arctostaphylos sp. , 23.xi.1971 , V. Chensy & D. Wilston coll., CDFA 71K30-10 [ CSCA ]; San Joaquin County: 2, Comanche Dam, on Arctostaphylos sp. , 28.xii.1978 , Todd & Shepard coll., CDFA 78AB-13 [ CSCA ]; Mariposa County: 6, near Yosemite, on Arctostaphylos patula , 9.v.1914 , T.D.A. Cockerell coll. [ USNM ]; Amador County: 7, W of Drytown on Highway 16, on Ceanothus sp. , 30.iii.1976 , E. Paddock & R.J. Gill coll. [ CSCA ]; Napa County: 1, St. Helena , on Arctostaphylos sp. , 17.iii.1994 , B. Campbell & R.J. Gill coll. [ CSCA ]; Berryessa, on madrone, 24.iv.1994 , C. Ko & R.J. Gill coll. [ CSCA ]; 2, same data but Fringe [ CSCA ]; Yolo County: 2, Monticello Dam, on Ceanothus cuneatus , 22.vi.1979 , T. Kono & R. Harris coll. [ CSCA ]; Lake County: 2, on Arctostaphylos tomentosa , 1888, Brandegec coll. [ USNM ]; 3, Kelseyville, on Arctostaphylos stanfordiana , 1.iv.1907 , H.P. Chandler coll. [ USNM ]; 4, same data but on Arctostaphylos manzanita , 25.iii.1930 , L. Benson coll. [ USNM ]; 8, Upper Lake, on manzanita, C. Townsend coll., CDFA PDR 76J2A-22 [ CSCA ]; El Dorado County: 2, Shingle Springs, on Eriodictyon , 4.iv.1963 , Simonds coll. [ USNM ]; 7, T. 12 N. R.12 E. Int. Diablo, El Dorado National Forest, on Arcostaphylos patula , 1.ix.1938 , Clarende & Quick coll. [ USNM ]; Placer County: 24, Alta, on Arctostaphylos sp. , 24.iii.1992 , L. Connolly coll. [ CSCA ]; 12, Auburn, on manzanita, 1.viii.1986 , D. Mitani coll., CDFA PDR 86H5-147 [ CSCA ]; 1, 3 mi W Omo Ranch, 28.iv.1968 , on Arctostaphylos sp. , R.F. Wiley coll. [ CSCA ]; 6, Georgetown, on Arctostaphylos sp. , 15.iii.1966 , R. Blomstrom coll. [ CSCA ]; 10, Smith Flat, on Ceanothus sp. , 3.iv.1968 , W.W. Wiard coll. [ CSCA ]; Butte County: 5, Chico, on Ceanothus sp. , 22.iii.1954 , H.J. Crawford coll. [ USNM ]; 2, 2 mile S of Forest Ranch, on Arctostaphylos sp. , 22.ii.1928 , A.A. Heller coll. [ USNM ]; 5, Plumas National Forest, on Arctostaphylos patula , 13.ix.1951 , C.R. Quick coll. [ USNM ]; Plumas County: 5, Feather River Canyon, on manzanita, R.S. Woglum Coll. [ USNM ]; Shasta County: 3, Redding, on Arctostaphylos sp. , 25.ix.1973 , Paddock & Gilbert coll. [ CSCA ]; 2, Shingletown, on Arctostaphylos sp. , 6.ii.1959 , I. Green coll. [ USNM ]; Colusa County: 5, Colusa, on manzanita, 9.ii.1973 , D. Brown coll., CDFA 73B14-3 [ CSCA ]; Oregon, Jackson County: 2, Medford, on madrone, 22.v.1952 , H. Sempert coll. [ USNM ]. 38 dry mounted: California, San Bernardino County: 2, Mountain Home Village, Mill Creek, 1219 m . a.s.l., on manzanita, 21.iii.1986 , R.J. Gill coll. [ CSCA ]; El Dorado County: 26, Georgetown, on Arctostaphylos sp. , 15.iii.1966 , R. Blomstrom coll. [ CSCA ]; 10, Smith Flat, on Ceanothus sp. , 3.iv.1968 , W.W. Wiard coll. [ CSCA ]. Hosts . Asteraceae :American threefold ( Trixis californica ); Boraginaceae :yerba santa ( Eriodictyon californicum , Eriodictyon sp. ); Ericaceae : madrone ( Arbutus menziesii ), manzanita ( Arctostaphylos manzanita , A. pajaroensis , A. patula , A. stanfordiana , A. tomentosa , A. viscida , Arctostaphylos sp. ); Euphorbiaceae : spurge ( Euphorbia sp. ); Fabaceae : black acacia ( Acacia melanoxylon ), oak ( Quercus ; Fagaceae ); Lamiaceae : Robison’s monardella ( Monardella robisonii ); Polygonaceae : wild buckwheat ( Eriogonum fasciculatum , Eriogonum sp. ); Rhamnaceae : California lilac ( Ceanothus cuneatus , C. cyaneus , C. foliolosus , C. thyrsiflorus , Ceanothus sp. ); Zygophyllaceae : creosote bush ( Larrea tridentata ). Characterization . Field characteristics . Pupal case oval, jet black, true margin with a lateral ring of amorphous white wax( Figs 5, 6 ). Slide-mounted characters ( Figs 50–68 ). TMS ending at apparent margin, lined with tubercles medially; longitudinal molting suture lined with tubercles from TMS to level of mouthparts or head submargin ( Figs 50–52 ); eyespots absent; Ce setae absent; T2 and T3 setae present, T3 setae arising well behind anterior margin of metanotum ( Figs 53–55 ); medial area with anterolateral depressions on abdomen moderately developed ( Figs 56–58 ), usually with a row of large tubercles along margins of T2/T3 suture and across anteromedial margin of each A1–A6, with microsetae and pores on each side of submedial area of head (3–4), T2 (2–4), T3 (0–2), A1 (0–4), A2 (0–2), A3 (2), A4 (0–2), A5 (0–2), A6 (0–2), A7 (0–2), and A8 (2); lateral areas of dorsal disc with variable number of large tubercles, sometimes restricted to level of abdominal sutures or faintly marked, with microsetae and pores between sides of dorsal disc and submargin; dorsal submargin with a row of microsetae (not visible in excessively bleached specimens) and transverse double rows of crescent-shaped imbrications ( Fig. 59 ), sometimes faintly marked ( Fig. 60 ) or not visible; deflexed submargin with rows of microsetae and pores, and with granulations along its entire width ( Figs 61, 63, 64 ); marginal glandular teeth subquadragular with tips smoothly rounded, toothed or serrate; VO subcordate to oval, inset from posterior margin by about its own length ( Figs 65–68 ); operculum cordate, its dorsal surface with a few longitudinal ridges and with microspinulae across distal third to fourth; lingula concealed by operculum; VO ring oval and wide ( Figs 65–68 ), with dorsal setae of A8 arising on its anterior margin anterior to operculum anterior margin ( Figs 65–68 ) [note that sclerotized ring in Fig. 66 was pushed forward when flattened during mounting, distorting ring shape and relative position of dorsal A8 setae]; with four membranous ventral sacs medially to bases of mesothoracic and metathoracic legs ( Fig. 62 ), although in a few cases posterior pair is not visible; bases of caudal setae close together, within level of operculum lateral margins ( Fig. 63, 64 ); venter smooth except for groups of spinulae medially to leg bases ( Fig. 62 ). Measurements . Puparium length: 730 ± 85; maximum width (between level of T2/T3 suture and A1): 457 ± 50; length/maximum width: 1.6 ± 0.1; width at level of anterior margin of operculum: 253 ± 42; maximum width/width at anterior margin of operculum: 1.8 ± 0.2; deflexed submargin/body radius: 0.5 ± 0.1; Ce setae: absent; T2 setae: 12 ± 11; T3 setae: 13 ± 11; dorsal A8 setae: 12 ± 3; caudal setae 72 ± 12; anterior marginal setae: 11 ± 1; posterior marginal setae 18 ± 3; ventral A8 setae: 16 ± 4; VO ring length: 64 ± 8; VO ring width: 58 ± 6; VO ring length/ width:1.1 ± 0.1; caudal seta/VO ring length: 1.2 ± 0.3; caudal seta/operculum length: 1.9 ± 0.4; VO length: 43 ± 6; VO width: 38 ± 4; VO length/width: 1.1 ± 0.1; operculum length: 38 ± 3; operculum/VO length: 0.9 ± 0.1 (see Table 1 for ranges). Similar species . Slide mounted puparia are very similar to T. ornata and T. sierrae , sharing the presence of large tubercles on lateral area of dorsal disc and crescent-shaped imbrications on submargin. It further shares the presence of tubercle-shaped ornamentations across anteromedial margins A1–A6 and granulations on deflexed margin with T. ornata . TABLE 1 . Measurements ranges (in µm) of puparia of Tetralicia spp. treated in this contribution. L: length; Op: Operculum; TMS: transverse molting suture; Width: W.
Body L Max. body W W at Op anterior margin Body L/ Max. body W Max. body W/ body W at Op ant. margin Cephalothorax L Abdomen L Cephalothorax L/ Abdomen L Deflexed submargin at TMS Body radius at TMS Deflexed submarg./body radius at TMS
abnormis 718–1,164 504–732 223–466 1.2–1.5 1.6–2.5 340–446 373–567 0.7–0.9 78–126 252–364 0.2–0.4
acaudata 757–1,076 524_747 276–485 1.3–1.4 1.5–1.9 349–504 407–572 0.8–0.9 92–165 262–369 0.4
agrifoliae 708–970 495–689 286–446 1.2–1.5 1.5–2.0 330–475 349–495 0.8–1.0 141–233 247–359 0.5–0.7
ceanothi 597–888 388–563 179–320 1.5–1.8 1.5–2.4 267–417 310–485 0.8–0.9 68–160 194–281 0.4–0.7
coachellensis 689–960 388–621 189–383 1.5–1.8 1.6–2.2 315–417 380–543 0.8–0.9 92–165 194–301 0.4–0.6
eriogonum 592–791 403–490 199–306 1.4–1.7 1.6–1.9 335–441 283–349 1.1–1.3 136–228 184–247 0.7–1.0
fouqueirasplendens 791–951 572–728 296–441 1.0–1.4 1.6–1.9 354–427 433–528 0.8 114–150 286–365 0.3–0.5
guajavae 543–679 310–432 121–175 1.6–1.8 2.2–2.8 262–315 291–363 0.8–1.0 87–116 160–218 0.5–0.6
granulata 577–689 325–458 136–184 1.5–1.8 2.1–2.7 291–330 301–378 0.8–1.0 73–112 158–226 0.4–0.6
hoelmeri 572–761 407–553 204–393 1.2–1.4 1.4–2.3 264–359 291–403 0.8–1.0 82–136 194–291 0.4–0.5
hyptisemoryi 664–854 485–629 272–398 1.3–1.4 1.5–1.8 330–410 335–441 0.9–1.0 102–141 243–312 0.4–0.5
laingi 708–873 466–621 238–369 1.4–1.6 1.6–2.2 344–427 361–446 0.8–1.0 102–175 233–310 0.4–0.6
lantanae 514–652 306–378 82–119 1.6–1.9 2.7–4.4 238–320 276–335 0.9–1.0 114–121 146–189 0.6
mexicana 883–980 582-689 369–451 1.4–1.5 1.5–1.6 378–441 500–582 0.7–0.8 83–170 286–344 0.4–0.6
nevadensis 549–689 427–563 233–310 1.2–1.4 1.7_2.0 272–330 296–359 0.9–1.0 61–109 213–281 0.3–0.4
nigrans 608–829 427–602 194–320 1.4–1.8 1.8–2.4 301–388 349–475 0.8–0.9 102–170 213–301 0.4–0.6
oblanceolata 597–722 354–437 136–204 1.5–2.0 2.0–2.6 276–335 320–400 0.8–0.9 82–116 177–218 0.5–0.6
ornata 616–871 409–577 223–310 1.5–1.7 1.6–2.0 301–422 320–456 0.8–1.0 99–155 205–289 0.4–0.6
salsolae 764–994 514–660 252–446 1.4–1.6 1.4–2.0 349–446 405–534 0.8–1.0 102–170 257–330 0.4–0.6
sierrae 640–883 437–640 233–349 1.3–1.5 1.7–1.9 301–407 340–475 0.8–0.9 104–177 218–320 0.5–0.6
......continued on the next page TABLE 1 . Continued. Measurements ranges (in µm) of puparia of Tetralicia spp. treated in this contribution.A8: abdominal segment 8; T2: Mesothoracic; T3: Metathoracic. *: From Carapia et al . 2018.
# apparent Cephalic seta L T2 seta L T3 seta L A8 dorsal Caudal seta L Anterior Posterior A8 ventral
marg. large crenulations per 100 µm seta L marginal seta L marginal seta L seta L
abnormis 0 10–22 15–97 17–107 27–34 24–28 24–29 66–70 19–39
acaudata 0 absent 11–19 11 12–17 12–15 9 11–24 15–17
agrifoliae 0 5–10 7–19 10–22 78–122 41–66 none visible 17–24 22–29
ceanothi 4–7 absent 6–39 5–49 10–17 51–92 10–12 11–38 10_31
coachellensis 0 absent 10–12 10–24 7–13 44–87 9–12 12–22 13–30
eriogonum 5–6 absent missing 10 10 36–73 12–34 19 34
fouqueirasplendens 0 absent 27–66 62–90 16–22 44–72 12 22 15–28
guajavae 5–6 absent 10–15 12–17 10–15 61–85 7–10 13–15 12–24
granulata 5–7 unknown missing missing 27 36–56 none visible 12 10
hoelmeri 0 absent 27–45 32–37 73–129 46–78 10 10–15 32
hyptisemoryi 0 absent 24–61 27–61 56–90 39–57 15 7 24-39
laingi 4–5 absent 11 12 10–15 39–66 12 16–22 7–29
lantanae 5–7 absent 15–19 15–22 none visible 87–141 10–17 12–22 22–29
mexicana 5–9 8–10* 8–10* 8–10* 25* 49–53 10–12* 45* 27–29
nevadensis 0 12–19 12 12 44–49 29–56 16 19–34 19
nigrans 0 absent 36–61 41–84 21–27 39–81 7–10 18–24 19–24
oblanceolata 4–5 absent 5–7 6–10 7–10 24–32 9 9–12 15
ornata 4–6 absent 5 5 10 36–53 7 12–13 12–17
salsolae 5–6 absent 17 9–21 6–12 39–70 missing 15–29 15
sierrae 5–8 absent 5–10 6 9–11 45–70 none visible 22 15–24
......continued on the next page TABLE 1 . Continued. Measurements ranges (in µm) of puparia of Tetralicia spp. treated in this contribution. VO: vasiform orifice.
VO ring L VO ring W VO ring L/W VO L VO W VO L/W Operculum L Operculum/ VO L Caudal seta/ VO ring L Caudal seta/ operculum L
abnormis 58–75 68–92 0.8–0.9 33–45 34–46 0.8–1.1 27–34 0.7–0.9 0.3–0.4 0.7–1.0
acaudata 56–78 51–74 1.0–1.2 32–46 34–51 0.7–1.1 27–38 0.7–1.0 0.2–0.3 0.4–0.5
agrifoliae 63–78 73–165 0.5–0.9 36–45 39–51 0.9–1.1 33–44 0.8–1.0 0.6–0.9 1.0–1.7
ceanothi 51–79 51–70 1.0–1.3 35–55 32–49 0.8–1.5 33–43 0.7–1.0 0.8–1.7 1.5–2.7
coachellensis 46–78 49–78 0.8–1.2 38–44 36–41 0.9–1.2 35–46 0.9–1.1 0.8–1.1 0.9–1.4
eriogonum 44–53 41–61 0.9–1.1 29–36 28–41 0.8–1.2 27–34 0.9–1.0 0.8 1.4
fouqueirasplendens 69–90 56–78 1.0–1.3 44–56 38–61 0.8–1.2 41–51 0.9–1.0 0.6–0.9 1.0–1.4
guajavae 30–55 32–44 0.9–1.7 22–34 24–34 0.9–1.1 19–29 0.8–0.9 1.3–2.2 2.5–3.5
granulata 41–49 36–53 0.9–1.2 27–33 27–29 1.0–1.2 23–27 0.8–0.9 0.8–1.2 1.5–2.1
hoelmeri 56–73 66–90 0.8–0.9 34–43 32–49 0.9–1.1 29–43 0.8–1.0 0.8–1.1 1.4–2.3
hyptisemoryi 58–83 68–90 0.9–1.0 43–53 39–61 0.8–1.1 34–49 0.8–0.9 0.6–0.8 0.9–1.3
laingi 56–75 56–72 0.9–1.2 39–56 34–46 0.9–1.2 34–46 0.9 0.6–0.9 1.0–1.4
lantanae 39–46 32–41 1.1–1.4 22–28 23–27 0.9–1.1 21–24 0.8–1.0 2.8–3.4 4.0–6.4
mexicana 49–53 50–56 1.0 39–44 41–44 0.9–1.0 34–39 0.8–0.9 1.0–1.1 1.3–1.4
nevadensis 46–53 63–83 0.6–0.8 29–39 29–34 0.9–1.3 23–34 0.8–0.9 0.6–0.9 1.1–1.9
nigrans 49–66 47–73 0.8–1.1 26–36 32–51 0.7–0.9 24–34 0.9–1.0 0.8–1.6 1.5–3.1
oblanceolata 50–58 41–56 1.0–1.2 30–40 29–44 0.9–1.1 27–36 0.8–1.0 0.4–0.6 0.7–1.2
ornata 56–70 51–70 1.0–1.2 36–51 33–46 0.9–1.2 34–47 0.9–1.0 0.6–0.9 1.0–1.6
salsolae 50–78 55–85 0.9–1.0 32–44 35–41 0.9–1.1 27–40 0.8–0.9 0.7–1.1 1.4–1.8
sierrae 53–61 46–63 0.9–1.2 36–51 34–46 0.8–1.2 34–41 0.8–0.9 0.7–1.2 1.2–1.8
Diagnosis . Mature puparia are devoid of dorsal wax ( Figs 5, 6 ) which is present in mature puparia of T. ornata ( Figs 25, 26 ), and the lateral ring of wax is matted, whereas it is striated in T. laingi ( Figs 16 , 17 ). Microscopically they differ from T. ornata and T. sierrae by puparium elliptical with caudal protuberance almost aligned with apparent margin ( Figs 50 , 51 , 63–68 ) and VO ring orifice oval ( Figs 63 , 65–68 ), whereas puparium is broadly oval with caudal protuberance separated by an obtuse angle from apparent margin ( Figs 176 , 201 , 202 ) and VO ring is subrectangular to subquadrate ( Figs 183–186 , 207, 208 ) in the other two species. It differs further from T. ornata by its relatively longer caudal setae (caudal setae: 51–92; caudal setae/VO ring length: 0.8–1.7; caudal setae/operculum length: 1.5–2.7). In T. ornata caudal setae are shorter (caudal setae: 36–53; caudal setae/VO ring length: 0.6–0.9; caudal setae/operculum length: 1.0–1.6). It differs further from T. sierrae by the entire width of deflexed submargin with granulations ( Fig. 61 ), and usually by the presence of tubercles along anterior margins of A2–A7 ( Figs 50 , 51 , 56, 57 ); in T. sierrae deflexed submargin lacks granulations ( Figs 204, 206 ), and there are sclerotized ridges lacking tubercles along anterior margins of A2–A7 ( Fig. 201 , 202 , 205 ). It differs from T. laingi by ornamentation of dorsal disc with tubercles ( Figs 50 , 51 ), dorsal submargin with crescent shaped crenulations arranged in transverse rows ( Figs 59, 60 ), and puparium elliptical with caudal protuberance not forming an angle with apparent margin ( Figs 50 , 51 , 63–68 ); in T. laingi , dorsal disc is smooth ( Figs 127 , 128, 132 ), dorsal submargin lacks crescent shaped crenulations ( Figs 127 , 128 ), and body is broadly oval with caudal protuberance forming an obtuse angle with sides of puparium ( Figs 127 , 132, 133 ). Remarks . Even though Sampson (1945: 58) indicated that " holotypes and certain paratypes will be deposited in the California Academy of Sciences" in the introduction of his paper describing this and another four species from California , no types of any species described by Sampson were deposited at CAS (Christopher C. Grinter pers. comm .). Instead, the holotype was deposited at BME at some point in the past (Lynn S. Kimsey & Brennen T. Dyer pers. comm .), where the junior author examined and illustrated it in 1985. Unfortunately, it could not be located for inclusion in the current paper. Sampson (1945: 60) did not indicate number of specimens studied under the species description, stating only 'Collected by Nathan Stahler and Thomas Kelly from Ceanothus cuniatus [sic] near Bishop, California , March 29, 1940 '. Two puparia belonging to the portion of W.W. Sampson's collection donated to CSCA have label data matching locality, misspelled host, and date, with collector indicated only as N. Stahler, and we consider that these most likely represent paratypes . In its original description, Sampson (1945: 59) stated "thoracic transverse slit not reaching apparent edge of body", but the transverse suture does reach the apparent margin in the holotype (R. Gill pers. obs. ), paratypes ( Fig. 50 ), and other examined material. We are not certain that all the specimens included here as T. ceanothi are conspecific with the types . The puparia collected on manzanita match the type specimens except for being usually (but not always, see Fig. 57 ) less ornamented along sides of dorsum, and those found on other native hosts ( Eriogonum , Euphorbia , Monardella , Trixis ) seem to also be a match except for tubercles across A1–A6 anterior margins vestigial or absent; we interpret this here as intraspecific variability correlated with the different hosts. However, we believe that more evidence based on adult and nymphal morphology and molecular studies is necessary to confirm their specific identity. As noted under T. abnormis , two morphs were detected based on the thoracic setae. In most specimens in which these setae were preserved ( Figs 53, 54 ) they are hairlike and short (T2 setae 6–34; T3 setae 5–24), but in some others ( Fig. 55 ) they are flattened and longer (T2 setae 39; T3 setae 49). Distribution . Specimens are common on Ceanothus along the western slope of the Sierra Nevada in central California , and on Arctostaphylos and other native hosts throughout the state. We have records from Alameda, Amador, Butte, Colusa, El Dorado, Imperial, Inyo, Los Angeles, Lake, Mariposa, Napa, Orange, Placer, Plumas, Riverside, San Diego, San Bernardino, San Joaquin, Shasta, Tulare, and Yolo Counties. This species was also recorded from Hawaii (host not indicated; Dubey 2021 ), Nevada ( Dooley et al. 2010 ), and Oregon (record in this paper) in the U.S.A. , and from Mexico ( Valencia & Evans 2024 ).