Tetralicia ceanothi Sampson, 1945

Ellenrieder, Natalia Von & Gill, Raymond J., 2024, 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, Zootaxa 5527 (1), pp. 1-129 : 17-22

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 .

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).

Figures

FIGURE 50. Tetralicia ceanothi, Paratype?: Habitus.

FIGURE 51. Tetralicia ceanothi, Orange Co., Rancho Capistrano, on manzanita: Habitus.

FIGURE 52. Tetralicia ceanothi, Amador Co., W of Drytown on Highway 16, on Ceanothus: Habitus. Note that puparium is distorted on slide, with deflexed margin folded outwardly so that apparent margin is not marginal.

FIGURES 53–58. Tetralicia ceanothi: 53, 56: Paratype?; 54: Los Angeles Co., Arcadia, on Ceanothus cyaneus; 55: Amador Co., W of Drytown on Highway 16, on Ceanothus; 57: Orange Co., Rancho Capistrano, on manzanita; 58: San Diego Co., San Diego, on Eriogonum. 53: Dorsal disc of thorax; 54, 55: Detail dorsum of T2 and T3; 56–58: Dorsal submargin and dorsal disc of A1–A5.

FIGURES 59–64. Tetralicia ceanothi: 59, 61, 62: Paratype?; 60: San Diego Co., San Felipe, Junction of Highways 52 & 522, on manzanita; 63: Orange Co., Rancho Capistrano, on manzanita; 64: Los Angeles Co., Arcadia, on Ceanothus cyaneus. 59, 60: Detail dorsal submargin; 61: Detail deflexed submargin; 62: Detail venter of thorax; 63, 64: Caudal end, ventral view.

FIGURES 65–68. Tetralicia ceanothi: 65: Paratype?; 66: Orange Co., Rancho Capistrano, on manzanita; 67: San Diego Co., San Felipe, Junction of Highways 52 & 522, on manzanita; 68: San Diego Co., San Diego, on Eriogonum. 65–68: Detail caudal end, dorsal view.

FIGURE 127. Tetralicia laingi, Neotype: Habitus.

FIGURE 176. Tetralicia ornata, Neotype: Habitus.

FIGURE 201. Tetralicia sierrae, Paratype?: Habitus.

FIGURE 202. Tetralicia sierrae, Paratype?: Habitus.

FIGURES 5–8. Puparia of Tetralicia in situ: 5: T. ceanothi, San Bernardino Co., Mountain Home Village, Mill Creek. 6: T. ceanothi, San Joaquin Co., Lake Camanche, on Arctostaphylos; 7: T. coachellensis, Riverside Co., Coachella, on arrowwood; 8: T. eriogonum, San Joaquin Co., 7 mi SW of Tracy, on Eriogonum; Figs 5, 6, 8 by R.J. Gill, 7 by N. von Ellenrieder & R.W. Garrison.

FIGURES 25–28. Puparia of Tetralicia in situ: 25, 26: T. ornata, San Bernardino Co., Mentone, on Eriodictyon californicum, 25: dorsal view, 26: dorsolateral view; 27, 28: T. salsolae sp. nov., Imperial Co., Ocotillo, on Ambrosia salsola, 27: dorsal view, 28: dorsolateral view. Figs 25, 26 by N. von Ellenrieder & R.W. Garrison, 28 by R.J. Gill.

FIGURES 13–16. Puparia of Tetralicia in situ: 13, 14: T. hoelmeri, Riverside Co.,Agua Caliente Indian Reservation, Palm Springs, on native shrub, 13: dorsal view, 14: posterior view; 15: T. hyptisemoryi, Imperial Co., Shell Canyon, on desert lavender, dorsolateral view; 16: T. laingi, Orange Co., Trabuco, on Salvia. Figs 13–16 by N. von Ellenrieder & R.W. Garrison.

FIGURES 17–20. Tetralicia in situ: 17–19: puparia; 20: adult. 17: T. laingi, San Diego Co., highway 76 at E boundary of La Jolla Indian Reservation, on Salvia apiana; 18, 19: T. lantanae sp. nov., Riverside Co., University of California Riverside, on Lantana, 18: dorsal view, 19: dorsolateral view; 20: T. lantanae sp. nov., San Diego Co., Chula Vista, on Lantana, 22.viii.1992, J. Kenyon coll. [no voucher available]. Figs 17, 20 by R.J. Gill, 18, 19 by N. von Ellenrieder & R.W. Garrison.

FIGURES 183–186. Tetralicia ornata, caudal end of puparium, dorsal view. 183, 185: Neotype; 184: San Bernardino Co., Mentone, Eriodictyon; 186: San Bernardino Co., 4 mi E of Mentone.

FIGURES 203–209. Tetralicia sierrae: 203, 205, 207, 208: Paratypes?; 204, 206, 209: Mariposa Co., between Crow and Signal Peaks, on Ceanothus parvifolius. 203: Cephalothoracic dorsal disc and submargin; 204: Venter T2 and T3; 205: Dorsal disc and submargin of abdomen; 206: Detail of deflexed submargin; 207, 208: Caudal end of abdomen, dorsal view; 209: Caudal end of abdomen, ventral view.

FIGURES 128–133. Tetralicia laingi: 128, 129, 132, 133: Neotype; 130, 131: San Diego Co., highway 76 at E boundary of La Jolla Indian Reservation, on Salvia apiana. 128: Dorsal disc and submargin of thorax; 129: Deflexed submargin; 130: Detail venter of T2 and T3; 131: Detail vasiform orifice; 132: Dorsal disc and submargin of abdomen; 133: Caudal end of abdomen, ventral view.

Tables

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

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