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 10.11646/zootaxa.5527.1.1 1175-5334 14021823 771D2E7B-4025-45BF-B328-6EC8A8851ECD Key to puparia of California species 1. TMS reaching apparent margin ( Figs 50 , 51 , 69 , 72 , 82 , 83 , 127 , 128 , 134 , 135 , 168 –170, 175–177, 187–189, 201–203)... 2 1'. TMS not reaching apparent margin ( Figs 29 , 36, 38, 43, 44, 89–91, 97, 98, 104, 105, 107, 108, 118–120, 159–161, 186–188)................................................................................................... 11 2(1). Longitudinal molting suture smooth, not lined with tubercles; dorsal disc of head with tubercles on entire median area ( Figs 142–145 , 187–189 )................................................................................... 3 2'. Longitudinal molting suture lined with tubercles; dorsal disc of head with tubercles entirely absent from median area or limited to area over mouthparts ( Figs 50 , 51 , 69 , 70 , 82 , 83 , 127 , 128 , 134 , 135 , 168 –170, 175–177, 201–203)................. 4 3(2'). Mature pupae devoid of dorsal wax; eyespots and Ce setae present ( Figs 142–145 ); marginal glandular teeth quadrangular with rounded tips and not differentiated at level of tracheal and caudal openings ( Fig. 147 ); VO lacking petal-shaped projections ( Fig. 149 ) [on Quercus ]................................................................ T. mexicana (in part) 3'. Mature pupae with one dorsolateral wax band on each side divided into cephalic, thoracic and abdominal sections ( Fig. 27 ); eyespots and Ce setae absent ( Figs 187–189 ); marginal teeth subtriangular and slightly differentiated (wider) at level of tracheal and caudal openings, accompanied by granulations on venter (gr; Figs 193, 194 , 196, 199 ); VO with four petal-shaped lateroapical projections (PePr; Figs 195–197, 199, 200 ) [on Hymenoclea salsola ]........................... T. salsolae 4(2') Mature pupae with amorphous lateral wax directed upwards from true margin covering deflexed portion of submargin laterally ( Fig. 8 ); puparium more elevated, with deflexed submargin as wide as 0.7–1.0 of body radius; dorsomedial area of A1–A7 with short longitudinal striations ( Fig. 86 ); venter medially to leg bases smooth ( Fig. 84 ) [on Eriogonum ]......... T. eriogonum 4'. Mature pupae with lateral wax not directed upwards from true margin covering deflexed portion of submargin laterally (e. g., Figs 19 , 23 , 26 ); puparium less elevated, with deflexed submargin as wide as 0.4–0.7 of body radius; dorsomedial area of A1–A7 lacking longitudinal striations ( Figs 56–58 , 75–77 , 132 , 137 , 169 , 182 , 205 ); venter medially to leg bases with spinulae (sp; Figs 62 , 73 , 130 , 138 , 171, 180, 204).................................................................. 5 5(4'). Puparium elliptical-oblanceolate with a pronounced caudal protuberance ( Figs 18, 19 , 23, 24 , 34, 35, 168, 169); VO located on a promontory (best seen in dorsolateral view, Fig. 141 ); with two membranous ventral sacs medially to bases of mesothoracic legs ( Figs 138 , 171)................................................................................... 6 5'. Puparium elliptical to oval with caudal protuberance moderate to only insinuated ( Figs 4–6 , 16 , 17 , 25, 26 , 50, 51, 69, 127, 176, 201, 202); VO not located on a promontory; usually with four membranous ventral sacs medially to bases of mesothoracic legs ( Figs 62 , 130 , 180 , 204 )................................................................................ 7 6(5). Puparium markedly oblanceolate, distinctly narrowed caudally (width at level of anterior margin of operculum 80–120; maximum width/width at level of anterior margin of operculum 2.7–4.4); dorsal disc on submedian area of thorax entirely covered with tubercle-shaped ornamentations ( Fig. 136 ); dorsal disc on submedian area of abdomen with tubercle-shaped ornamentations across both anterior and posterior areas of A1–A6 ( Figs 137, 139, 141 ); caudal setae ( Figs 134 , 135 ) as long as 2.8–3.4 times the length of VO ring [commonly on Lantana , but also on Hibiscus , Eupatorium , and Morus ]..... T. lantanae 6'. Puparium less markedly oblanceolate, less narrowed caudally (width at level of anterior margin of operculum 130–205; maximum width/width at level of anterior margin of operculum 2.0–2.6); dorsal disc on submedian area of thorax devoid of tubercle-shaped ornamentations (Fig. 170); dorsal disc on submedian area of A1–A6 with tubercle-shaped ornamentations only across anterior area ( Fig. 169 ); caudal setae ( Fig. 169 ) as long as 0.4–0.6 times the length of VO ring [on Rhamnus ]............................................................................................... T. oblanceolata 7(5'). Dorsal submargin lacking crescent shaped crenulations or tubercles ( Figs 69 , 71 , 127 , 128 )........................... 8 7'. Dorsal submargin with crescent shaped crenulations or tubercles arranged in transverse rows ( Figs 51 , 52 , 57–60 , 178 , 201– 203, 205 )........................................................................................... 9 8(7). Puparium elliptical ( Fig. 69 ), apparently lacking any lateral wax ( Fig. 7 ); with small tubercles on sides of dorsal disc and minute granulations along sutures (gr; Figs 75–77 ) [on Pluchea and other Asteraceae ]........................ T. coachellensis 8'. Puparium broadly oval ( Fig. 127 ), with a wide marginal fringe of striated shiny wax ( Fig. 17 ); lacking tubercles on sides of dorsal disc and minute granulations along sutures ( Figs 127 , 132 ) [on Salvia and various other plants, including Asteraceae ].............................................................................................. T. laingi 9(7'). Deflexed submargin smooth, lacking granulations ( Fig. 206 ); dorsal disc on anterior margin of A1–A6 with a sclerotized ridge lacking separate distinct tubercles ( Figs 202 , 205 ) [on Ceanothus parvifolius ].............................. T. sierrae 9'. Entire width of deflexed submargin with granulations ( Figs 61 , 179 ); usually with distinct tubercules on dorsal disc across anterior margin of A1–A6 ( Figs 56, 57 , 181, 182 ).......................................................... 10 10(9'). Mature pupae lacking dorsal wax ( Figs 5, 6 ); puparium elliptical with caudal protuberance approximately aligned with apparent margin ( Figs 50 , 51 , 63–68 ); VO oval and surrounded by a relatively wide VO ring ( Figs 63 , 65–68 ); caudal setae about as long as 1.5–3 times the length of operculum ( Figs 51 , 63, 64 ) [on Ceanothus and multiple other hosts including Eriodictyon ]............................................................................................... T. ceanothi 10'. Mature pupae with two pairs of dorsal longitudinal wax bands ( Fig. 25 ); puparium broadly oval with caudal protuberance forming an obtuse angle with apparent margin ( Figs 176 , 183–186 ); VO subrectangular to subquadrate and surrounded by a relatively narrow VO ring ( Figs 183–186 ); caudal setae about as long as 1–1.6 times the length of operculum ( Figs 176 , 185 ) [on Eriodictyon ]................................................................................ T. ornata 11(1'). Longitudinal molting suture lined with tubercles ( Figs 29 , 36, 43, 97, 159, 160).................................. 12 11'. Longitudinal molting suture smooth ( Figs 89 , 90 , 104 , 105 , 118 , 119 ).......................................... 17 12(11) Puparium rounded-oval ( Figs 29 , 43, 151, 152); Ce setae and eyespots present ( Figs 29 , 30, 43, 151–153) [on Quercus in California ]......................................................................................... 13 12'. Puparium oblanceolate, elliptical, or oval with truncate caudal end ( Figs 36 , 97 , 159 , 160 ); Ce setae and eyespots absent ( Figs 36 , 37 , 97 , 159 , 160 ) [on various hosts including Quercus ]................................................... 15 13(12') Longitudinal molting suture lined with tubercles from suture between T2/T3 to submarginal area ( Figs 29 , 30, 36); lateral areas of dorsal disc and medial areas of head and T1 smooth or with small granulations not forming a reticulate pattern ( Figs 29 , 30, 36)....................................................................................... T. abnormis 13'. Longitudinal molting suture lined with tubercles from TMS to level of mouthparts ( Figs 43 , 44 , 151–153 ); lateral areas of dorsal disc and medial areas of head and T1 with large rounded tubercles forming a reticulate pattern ( Figs 43 , 44, 46, 48 , 151–153, 155 )...................................................................................... 14 14(13') Eyespots oval ( Figs 43 , 44 ); TMS U-shaped, with medial section almost straight and distal ends bent at almost 90 degrees ( Figs 43 , 44 ); caudal setae close together, originating at or mesad of levels of operculum lateral margins ( Figs 43 , 49 ). T. agrifoliae 14'. Eyespots slit-like ( Figs 151–153 ); TMS W-shaped, with medial section curved anteriorly and distal ends bent at about 45 degrees ( Figs 151–153 ); caudal setae widely separated, originating laterad of levels of operculum lateral margins ( Figs 151 , 152 , 155, 157, 158 )......................................................................... T. nevadensis 15(12') Puparium oblanceolate (width at anterior margin of operculum 120–175) with a pronounced caudal protuberance ( Figs 97 , 102, 103 ); submedian area of thorax with granulations along sutures and depressions (gr; Fig. 98 ); abdominal depressions lined with granulations (gr; Fig. 99 ); deflexed subdorsum with transverse rows of granulations across its entire width ( Fig. 101 ) [on Psidium guajava in California ].................................................................. T. guajavae 15'. Puparium oval to elliptical (width at anterior margin of operculum 195–485), with caudal protuberance absent or moderate ( Figs 36 , 41, 42 , 159 , 160 , 166, 167 ); submedian area of thorax smooth ( Figs 37 , 161 ); abdominal depressions not lined with granulations ( Figs 36 , 165 ); deflexed subdorsum with transverse rows of small granulations arranged in compact clusters across about half to 3/4 of its width ( Figs 40 , 164 )............................................................... 16 16(15'). Puparium oval with caudal end truncate lacking a caudal protuberance ( Figs 36 , 41, 42 ), with a narrow lateral ring of amorphous wax visible dorsally ( Figs 1, 2 ); lateral area of dorsal disc smooth ( Figs 36–38 ); dorsal submargin smooth ( Fig. 38 ); deflexed submargin with transverse rows of small granulations usually arranged in 1–3 compact clusters extending across about half to 2/3 of its width (gr; Fig. 40 ); VO cordate ( Figs 36 , 41 ); VO ring subcordate with anterior margin well defined and anterior portion delimited from remainder of ring and sculpted with crests ( Figs 36 , 41 ); venter with groups of strong spines (spi) medially to legs ( Fig. 39 ) [on Arctostaphylos ]...................................................... T. acaudata 16'. Puparium elliptical with a caudal protuberance ( Figs 159 , 160 , 16 , 167), with fragmentary lateral wax not visible dorsally ( Fig. 21 ); lateral area of dorsal disc with small tubercles ( Figs 161, 165 ); dorsal submargin with small tubercles ( Fig. 162 ); deflexed submargin with transverse rows of small granulations usually arranged in 7–9 compact clusters extending across about 3/4 of its width (gr; Fig. 164 ); VO subcordate ( Figs 166, 167 ); VO ring cordate and open anteriorly, lacking crests ( Figs 166, 167 ); venter with groups of spinulae (sp) medially to legs ( Fig. 163 ) [on multiple hosts including Arctostaphylos ]...... T. nigrans 17(11'). With eyespots ( Figs 89 , 90 , 143–145 ); dorsal submargin lacking a row of membranous papillae ( Figs 89 , 90 , 142–146 ); with two or four ventral membranous sacs (vs) on thorax ( Fig. 92 )................................................. 18 17'. Lacking eyespots ( Figs 104 , 105 , 118 , 119 ); dorsal submargin with a row of membranous papillae (S pap; Figs 13, 15 , 104, 105, 107–109, 114, 115, 118–120, 122); with five ventral membranous sacs (vs) on thorax ( Figs 110, 111 , 121 ) [on Condea ]... 19 18(17). Apparent margin with an indentation at anterior end and three rounded projections, one each adjacent to lateral ends of TMS and at caudal end ( Figs 9 , 89); Ce setae absent; T2, T3, and dorsal A8 setae originating from bulbous bases ( Figs 89–91, 95 ); dorsal disc and submargin lacking imbrications or granulations, with cobblestone pattern of depressions of variable extension ( Figs 89–91 ); deflexed submargin with narrow band of small granulations adjacent to marginal teeth and with a row of paired pores (pr) and porettes (pt) ( Figs 93, 94 ); venter extensively spinulose ( Figs 92, 93 ) [on Fouquieria and other hosts]........................................................................................ T. fouquieriasplendens 18'. Apparent margin lacking projections ( Figs 142 , 143 ); Ce setae present; T2, T3, and dorsal A8 setae not arising from bulbous bases ( Figs 144, 145, 149 ); sides of dorsal disc with granulations forming a reticulate pattern and dorsal submargin with rows of transverse crests ( Figs 142–146 ); deflexed submargin with a wide band of rounded granulations and a row of single pores (pr; Figs 147 ); venter with spinulae on wide bands along leg bases and around mouthparts (sp; Fig. 147 ) absent from abdomen [on Quercus ]........................................................................ T. mexicana (in part) 19(17'). Dorsal disc with large tubercles on medial area lateral to mouthparts, along thoracic sutures, anterior and posterior margins of A1–A7, and on lateral areas ( Figs 104–107 , 112, 114, 115 ); 3–5 papillae (usually 4) (S pap) on each side of abdominal submargin only ( Figs 104 , 105 , 114–116 ); TMS ending on submargin almost at apparent margin ( Figs 104 , 105 , 107, 108 ); with a row of double pores, a larger one with dark rim and an adjacent minute one (pr & pt), between dorsal disc and submargin ( Figs 114, 116 ); deflexed submargin with dark granulations extending over its entire width lacking a particular pattern ( Fig. 113 ); abdominal anteromedial depressions inconspicuous ( Figs 104 , 105 , 107 , 112, 114, 115 ); VO ring subtriangular ( Figs 115–117 ); venter smooth except for spinulae (sp) along margins and medially to leg bases ( Figs 111 )................... T. hoelmeri 19'. Dorsal disc devoid of large tubercles ( Figs 118–120 ); 6–8 papillae (S pap) on each side of thoracic and abdominal submargin, plus usually a medial one on cephalic submargin ( Figs 118–120, 122 ); TMS barely extending onto submargin ( Figs 118–120 ); with a row of single pores (pr) between dorsal disc and submargin ( Figs 122, 124 ); deflexed submargin with transverse rows of one to three clusters of dark granulations (gr) adjacent to the margin ( Figs 125, 126 ); abdominal anteromedial depressions (de) conspicuous ( Figs 118–122 ); VO ring subcordate ( Figs 119 , 122, 124 ); venter uniformly covered with closely-set spinulae ( Fig. 123 ).................................................................................... T. hyptisemoryi