Tetralicia nigrans (Bemis, 1904)

Figs 21, 159–167

Aleyrodes nigrans Bemis, 1904: 522 .

Tetraleurodes nigrans: Quaintance & Baker 1914: 618.

Tetralicia nigrans: Baker 1937: 281; Valencia & Evans 2024: 220.

Aleuropleurocelus nigrans: Drews & Sampson 1956: 281; Mound & Halsey 1978: 59; Sanchez-Flores et al. 2017: 606.

Material examined. 232 puparia: U.S.A., California: 1, no data [USNM]; 2, on chaparral, R.S. Woglum coll. [USNM]; San Diego County: 3, Escondido, on apple, 2.x.1975, Giesing & Bixby coll., CDFA PDR 7557-34 [CSCA]; 8, Fallbrook, on native shrub, 16.i.1975, Giesing/Metcalf coll. [CSCA]; 8, Sweetwater Dam, on Malosma laurina, 13.ix.1911, P.H. Timberlake coll. [USNM]; Orange County: 1, Trabuco Canyon, on toyon, 4.ii.1933, T. E. Mehead coll. [CSCA]; 1, same data but on manzanita, #12 [ex W.W. Sampson Coll., CSCA]; 3, same data but on Pear, 16.ix.1965, Johnson & Byers coll. [USNM]; 1, Silverado Canyon, on Rhamnus sp., i.1954 [ex W.W. Sampson Coll., CSCA]; 3, Irvine Park, on toyon, 7.i.1934, E.H.P. coll. [CSCA]; Riverside County: 5, Reche Canyon, on toyon, #15 [ex W.W. Sampson Coll., CSCA]; Los Angeles County: 1, San Clemente Island, on Heteromeles sp., 25.viii.1894, E.A. Mearns coll. [USNM]; 1, Los Angeles, on Prunus, xii.1907, R.S. Woglum coll. [USNM]; 3, Glendale, on Rhus, v.1908, R.S. Woglum coll. [USNM]; 4, Mount Wilson, on Rhus, v.1908, R.S. Woglum coll. [USNM];16, Whittier, on Heteromeles arbutifolia, 31.iii.1912, P.H. Timberlake coll. [USNM]; San Bernardino County: 2, Ontario, on Rhamnus crocea, 16.ii.1919, M. Moles coll. [USNM]; 1, same data but on Quercus chrysolepis [USNM]; San Bernardino Mountains, on Rhamnus, #18 [ex W.W. Sampson Coll., CSCA]; 6, San Bernardino, on Rhus, prob. ovata, 10.iii.1960, P. Prescott coll. [CSCA]; 5, same data but on Aronia arbutifolia [USNM]; 1, Mountain Home Village, Mill Creek, 4000 feet, on manzanita, 21.iii.1986, R.J. Gill coll. [CSCA]; Santa Barbara County: 4, Franchese Peak, on Garrya sp., 16.ii.1960, G. Beevor coll. [USNM]; 2, Santa Barbara, Mission District, on Garrya sp., 25.iii.1961, G. Reever coll. [CSCA]; 2, Santa Barbara, on toyon, 21.iii.1986, J. Karl coll. [CSCA]; San Benito County: 1, San Benito Mountain National Area, on Garrya sp., 13.iv.2004, R.L. Zuparko coll. [CSCA]; Tulare County: 14, Exeter, on toyon, 18.iv.1974, Thomas & Akana coll. [CSCA]; 7, Tulare, on ash leaves, ix.1976, E. Russell coll. [CSCA]; Santa Cruz County: 1, Boulder Creek, on Arctostaphylos nummularia, 15.x.1891, V. Bailey coll. [USNM]; Santa Clara County: 5 syntypes, Santa Clara, on Ceanothus integerrimus, 17.iv. 15. v.1901, F.E. Bemis coll. [USNM]; 4 syntypes, Stanford University Campus, on Heteromelas arbutifolia, 2.iv.1901, F.E. Bemis coll. [7098, USNM] ; 6, same data but 24.iv.1902, J.E.D. coll. [CSCA]; 3, same data but 9.iv.1901 [BME]; 1, same data but on Prunus ilicifolia, 25.iv.1901 [BME]; 1, same data but on Rhamnus californica, 26.iv.1945, G.F. Ferris coll. [USNM]; 3, Stanford, on Aronia arbutifolia, 7.iv.1907, L. Swan coll. [USNM]; 1, same data but 6.vi.1951 [USNM]; Mariposa County: 8, on Berberis dictyota, 15.viii.1953, C.R. Quick coll. [USNM]; Napa County: 1, on madrone, 25.iv.1999, C. Ko & R.J. Gill coll. [CSCA]; 1, Santa Helena, on Arbutus menziesii, 17.iii.1994, Campell & R.J. Gill coll. [CSCA]; Nevada County: 9, Nevada City, on manzanita, 5.ii.1931 [CSCA]; Mendocino County: 1, Fort Bragg, on Arctostaphylos nummularia, v.1900 [USNM]; 7, on Rhamnus californicus, 2.iv.1901, Stanford University Entomology Lab. coll. [BME]; 2, Redwood Valley, on Fraxinus sp., 12.ii.1993, J. Xerogeanes coll. [CSCA]; Plumas County: 3, Feather River Canyon, on manzanita, R.S. Woglum coll. [USNM]; Siskiyou County: 6, Tulelake, on mahonia, 5.ix.1981, M. Lundon coll. [CSCA]. Oregon, Jackson Co.: 60, Medford, on Oregon grape, 26.ii.1947 [CSCA, ex Sampson Coll.]. Arizona, Pima County: 3, Tucson, Desert Museum, 20.v.2003, J. Dooley coll. [CSCA]. Nevada, Clark County: 1, Red Rock Canyon National Conservation Area, Lost Creek Canyon, 19.viii.2008, J. Dooley coll. [CSCA].

Hosts. Bemis (1904) described this species from several chaparral plants, including Anacardiaceae: Malosma laurina, Rhus trilobata, R. prob. ovata; Boraginaceae: Eriodictyon californicum; Caprifoliaceae: Lonicera involucrata, Symphoricarpos racemosus; Ericaceae: Arbutus menziesii, Arctostaphyllos manzanita; Fabaceae: Quercus agrifolia; Lauraceae: Umbellularia californica; Ranunculaceae: Clematis lingusticifolia; Rhamnaceae: Ceanothus integerrimus, Rhamnus californica; Rosaceae: Heteromeles arbutifolia, Prunus ilicifolia . We also have records on Berberidaceae: Berberis dictyota, Mahonia aquifolium; Ericaceae: Arctostaphylos nummularia; Fabaceae: Quercus chrysolepis; Garryaceae: Garrya sp.; Oleaceae: Fraxinus sp.; Rhamnaceae: Rhamnus crocea; Rosaceae: apple, Aronia arbutifolia, pear.

Characterization.

Field Characteristics. Pupal case elliptical, black (Fig. 21). Often without visible wax formation although some specimens, particularly in southern California, may have a small amount of amorphous white lateral wax (R.J. Gill pers. obs.). Eggs dark yellow to dusky brown, elliptical with apex pointed and a short pedicel; nymphal instars 1–3 and adults yellow with wings somewhat dusky at distal end (Bemis 1904).

Slide-mounted characters. TMS extending over submargin but not reaching apparent margin, not lined with tubercles (Figs 159–161); longitudinal molting suture lined with tubercles from T2/T3 suture to head submargin (Figs 161, 162); eyespots absent; Ce setae absent; T2 and T3 setae present, T3 setae arising just behind anterior margin of metanotum (Figs 159–161); medial area with depressions lacking granulations on each side of head (two), T2 (three) and T3 (one), and abdominal segments (one) (Figs 159–162); with paired pores and microsetae on each side of submedial area of head (2–6), T2 (1–2), T3 (0–2), A1 (0–2), A2 (0–1), A3 (0–2), A4 (0–2), A5 (0–4), A6 (0–3), A7 (0–3), and A8 (0–2); dorsal disc with small tubercles along lateral areas (Figs 159, 160, 165), with anterior margin of A1–A7 sclerotized (Fig. 165); with some pairs of pores along sides of dorsal disc (Fig. 161); dorsal submargin with small tubercles (Fig. 162); deflexed submargin with a row of pores and transverse rows of small granulations usually arranged in 7–9 compact clusters extending for about 3/4 of its width (Fig. 164); marginal glandular teeth subquadrangular with tips bidentate; VO subcordate, inset from posterior margin by more than its own length; operculum cordate, its dorsal surface with short longitudinal and transverse ridges, with microspinulae across distal third to fourth; lingula concealed by operculum; VO ring cordate and open anteriorly, very narrow around lateral and posterior sections of VO, with anterior portion wide and subrectangular, lacking longitudinal crests and not separated from remainder of ring (Figs 166, 167), with dorsal setae of A8 arising from sides anterior to anterior margin of operculum (Figs 166, 167); bases of caudal setae close together, within level of operculum lateral margins (Fig. 166); with two membranous ventral sacs medially to bases of mesothoracic legs (Fig. 163); venter with groups of spinulae medially to legs (Fig. 163).

Measurements. Puparium length: 743 ± 76; maximum width (at level of TMS): 486 ± 57; length/maximum width: 1.5 ± 0.1; width at level of anterior margin of operculum: 238 ± 42; maximum width/width at anterior margin of operculum: 2.1 ± 0.2; deflexed submargin/body radius: 128 ± 25; Ce setae: absent; T2 setae: 48 ± 12; T3 setae: 63 ± 17; dorsal A8 setae: 24 ± 2; caudal setae: 68 ± 17; anterior marginal setae: 9 ± 1; posterior marginal setae: 21 ± 3; ventral A8 setae: 22 ± 2; ring length: 53 ± 5; ring width: 56 ± 8; ring length/width: 1 ± 0.1; caudal seta/ring length: 1.3 ± 0.4; caudal seta/operculum length: 2.5 ± 0.7; VO length: 29 ± 3; VO width: 37 ± 6; VO length/width: 0.8 ± 0.1; operculum length: 27 ± 3; operculum/VO length: 0.9 ± 0.5 (see Table 1 for ranges).

Similar species. Tetralicia acaudata .

Diagnosis. See under T. acaudata .

Biology. Eggs always found on an upright position on the underside of the leaf. Early nymphal instars present from April to June and in September, and puparia abundant year-long except on deciduous host plants (Bemis 1904).

Distribution. This is the most widespread species in California, found along the full extension of the State (we have records from Alameda, Contra Costa, Los Angeles, Mariposa, Mendocino, Monterrey, Napa, Nevada, Orange, Plumas, Riverside, San Bernardino, San Benito, San Diego, San Mateo, Santa Barbara, Santa Clara, Santa Cruz, Siskiyou, and Tulare Counties). It is also known from Arizona, Oregon (this paper), and Nevada (Dooley et al. 2010) in the U.S.A., and Mexico (Sanchez-Flores et al. 2017), and it has been intercepted from El Salvador and Haiti (Valencia & Evans 2024).