Tetralicia hoelmeri von Ellenrieder & Gill, 2024

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 : 31-35

Tetralicia hoelmeri von Ellenrieder & Gill , sp. nov.

Figs 13, 14 , 104–117, 231–242, 261–264

Aleuropleurocelus hyptisemoryi Gill in Polaszek & Gill 2011: 52 (in part: Figs 2, 3 ; description of nymphal instars 1–3 and adults).

Etymology. Named after Kim A. Hoelmer, one of the collectors of this species, who provided several whitefly species on native host plants resulting from his Southern California desert survey of native whiteflies and their parasitoids to CSCA.

Type material. 75 slide mounted puparia: U.S.A., California, Riverside County: Holotype (circled with red on slide), Agua Caliente Indian Reservation, Palm Springs , on Condea emoryi , 27.ii.1984, E. Reeves coll. [ CSCA]; 54 paratypes, same data as holotype [ CSCA]; 12 paratypes (paratypes of T. hyptisemoryi ), Living Desert , Palm Desert , on desert lavender, 10.vi.1997, K.A. Hoelmer coll. [ CSCA]; Imperial County: 5 paratypes, Imperial National Wildlife Refuge , on Condea emoryi , 1943, F.B. McMurry coll., CDA Herbarium 0021354 [ CSCA]; San Bernardino County: 3 paratypes, Vidal Junction , on Condea emoryi , 5.x.1972, Davis et al. coll. [ CSCA].

Adults: 3 male paratypes, 3 female paratypes (paratypes of T. hyptisemoryi ), Riverside County, Living Desert , Palm Desert, on desert lavender, 10.vi.1997, K.A. Hoelmer coll., #20 [ CSCA] .

Nymphal instars 1–3: 8 third instar nymphs, 7 of which have overlapping exuviae of second and first instars (all folded in half except for 1), paratypes (paratypes of T. hyptisemoryi ), Riverside County, Living Desert , Palm Desert, on desert lavender, 10.vi.1997, K.A. Hoelmer coll., #20 [ CSCA] .

Additional material examined. 92 dry mounted puparia: U.S.A., California, Riverside County: 80, Agua Caliente Indian Reservation, Palm Springs , on Condea emoryi , 27.ii.1984, E. Reeves coll. [ CSCA]; 12, Living Desert , Palm Desert, on desert lavender, 10.vi.1997, K.A. Hoelmer coll. [ CSCA] .

Hosts. Condea emoryi ( Lamiaceae ).

Description.

Field characteristics. Pupal case elliptical to oval, black, with a narrow ring of lateral wax ( Fig. 14 ) not visible dorsally, VO located on a promontory, five to seven lobate to strongly conical projections on each side of T3 and abdomen along apparent margin pointing laterally, and three to five (usually four) dorsolateral conical papillae along submargin of abdomen ( Fig. 13 ). Adult color in life unknown; nymphal instars 1–3 transparent yellow ( Fig. 13 ).

Slide-mounted characters (values of holotype in square brackets). Puparium apparent margin festooned with 5–7 [6] (usually 6) very low lobate ( Fig. 104 ) to strongly conical projections ( Fig. 105 ) extending from T3 to caudal abdomen; TMS extending over submargin almost to but not reaching apparent margin, not lined with tubercles ( Figs 104 , 105 , 107, 108 ); longitudinal molting suture not lined with tubercles; eyespots absent; Ce setae absent; T2 and T3 setae long and flattened, arising from bulbous bases (107, 108, 112); T3 setae arising just behind anterior margin of metanotum (107, 108, 112); medial area with large tubercles on head lateral to mouth, along T1/T2 and T2/T3 sutures ( Figs 106 , 112 ), with large tubercles in a row along anterior area of A1–A7, and smaller in a row along posterior area of each segment; abdominal depressions indistinct ( Figs 112, 114, 115 ); with pairs of pores and porettes on each side of submedial area of head (1–2), T2 (0–1), T3 (0), A1 (0), A2 (0), A3 (0–1), A4 (1), A5 (0–1), A6 (0–1), A7 (0), and A8 (0); dorsal disc with large tubercles and pairs of pores and porettes along sides ( Figs 109 , 114, 116 ); dorsal submargin with tubercles along inner portion, with irregular darkened depressions, with pores, and with 3–5 [4] (usually 4) conical papillae on each side on abdomen ( Figs 104–109 , 114–116 ), some of which with a marginal disc pore; deflexed submargin with a row of pores, with dark granulations extending over entire width lacking a particular pattern ( Fig. 113 ); marginal glandular teeth subquadrangular with tips smooth; VO oval, inset from posterior margin by more than its own length or about its own length; operculum cordate, its dorsal surface with a few longitudinal ridges and with microspinulae across distal third to fourth; lingula concealed by operculum; VO ring subtriangular, anteriorly wide and open ( Figs 116, 117 ), with dorsal setae of A8 arising from bulbous bases anterior to level of anterior margin of operculum ( Figs 116, 117 ); bases of caudal setae widely separated, at level of lateral margins of VO ring ( Figs 104 , 105 , 116 ); with five membranous ventral sacs medially to bases of mesothoracic and metathoracic legs and on center ( Fig. 110 ); tegument of venter smooth except for spinulae along margins and medially to leg bases ( Fig. 111 ).

Adults. Female ( Fig. 231 ) with 2 pairs and male ( Fig. 233 ) with 4 pairs of wax plates on abdomen. Upper and lower compound eyes connected by one ommatidium ( Figs 235, 236 ). Antennae 7 segmented ( Figs 235, 236 ), with an elongate third segment with two primary round sensoria near the apex, fifth, sixth, and seventh segments each with 1 round primary sensorium located at apex on fifth and sixth, and at about two thirds of its length on seventh, and sixth and seventh segments with an elongated rod-like sensorium each. Hind tibiae ( Figs 237, 240 ) with a tibial comb of 12–15 setae in male, 15–18 setae in female; metatibial and mesotibial brushes not defined. Female cement gland not discernible ( Fig. 232 ). Male parameres narrowing along distal sixth ending on a single tooth markedly curved medially, lacking subapical teeth ( Fig. 234 ).

Nymphal instars 1–3. First instar ( Fig. 261 ): with 6 pairs of long submarginal setae along sides of cephalothorax and 1 pair posterior to level of VO. VO oval, wider than long, with anterior margin straight; operculum oval occupying most of VO; lingula head bulbous and covered with microsetae; other characters not discernible. Second and third instars: lacking long submarginal setae, with button-like structures with a central pore along margins of cephalothorax and abdomen (25–32 on third instar; exact number not discernible on second instar) which are set close together, separated from each other by a space subequal to width of each structure ( Figs 263, 264 ), and with conical submarginal abdominal papillae (4–6 on third instar; exact number not discernible on second instar), VO located on a promontory, and operculum subcordate ( Figs 262–264 ).

Measurements (values of holotype in square brackets). Puparium length: 649 ± 75 [723]; maximum width (between level of T2/T3 suture and A3): 489 ± 63 [543]; length/maximum width: 1.3 ± 0.1 [1.3]; width at level of anterior margin of operculum: 283 ± 74 [359]; maximum width/width at anterior margin of operculum: 1.8 ± 0.3 [1.5]; deflexed submargin/body radius: 108 ± 18 [0.5]; Ce setae: absent [absent]; T2 setae: 33 ± 6 [45]; T3 setae: 34 ± 3 [36]; dorsal A8 setae: 93 ± 17 [102]; caudal setae: 60 ± 10 [70]; anterior marginal setae: 10 [none visible]; posterior marginal setae: 13 ± 4 [non visible]; ventral A8 setae: 32 [none visible]; VO ring length: 64 ± 6 [68]; VO ring width: 76 ± 8 [87]; VO ring length/width: 0.8 ± 0.05 [0.8]; caudal seta/VO ring length: 0.9 ± 0.1 [1]; caudal seta/operculum length: 1.7 ± 0.2 [1.8]; VO length: 39 ± 4 [43];VO width: 40 ± 5 [41]; VO length/width: 1 ± 0.1 [1]; operculum length: 35 ± 4 [39]; operculum/VO length: 0.9 ± 0.05 [0.9] (see Table 1 for ranges).

Adults. Male: body length (including parameres): 1203 ± 145 [1038–1310]; rostral length: 280 ± 7 [272–286]; hind tibia length: 297 ± 9 [286–303]; third antennal segment length: 143 ± 7 [138–148]; paramere length: 124 ± 3 [121–126]; aedeagus length: 121. Female: body length (including ovipositor): 1267 ± 62 [1222–1339]; rostral length: 318 ± 7 [310–325]; hind tibia length: 362 ± 3 [359–364]; third antennal segment length: 155; ovipositor length: 379 ± 8 [373–388].

Nymphal instars 1–3. First instar: length: 191 ± 12 [177–211]; submarginal setae: 45 ± 3 [36–46]; Ce setae: 57 ± 10 [44–73]; T2 setae: 70 ± 10 [56–83]; T3 setae: 54 ± 7 [49–66]; dorsal A8 setae: 66 ± 10 [51–75]; caudal setae: 29 ± 5 [22–34]. Second instar: length: 235 ± 10 [224–243]; T2 setae: 38 ± 4 [36–44]; T3 setae: 37 ± 5 [32–44]; dorsal A8 setae: 43 ± 6 [36–53]; caudal setae: 33 ± 6 [24–43]. Third instar: length: 365 ± 20 [335–398]; ventral A8 setae: 13 ± 2 [12–15]; dorsal A8 setae: 60 ± 8 [49–70]; caudal setae: 46 ± 7 [39–56].

Similar species. It shares a smooth longitudinal molting suture lacking tubercles with T. fouquierasplendens , T. hyptisemoryi , and T. salsolae , and the presence of lateral projections along apparent margin, VO ring open anteriorly, and all T2, T3, and dorsal A8 setae originating from bulbous bases with T. fouquierasplendens and T. hyptisemoryi . Tetralicia hoelmeri and T. hyptisemoryi share the presence of dorsal A8 setae longer than caudal setae (in puparia and earlier nymphs) with puparia of T. agrifoliae . Both species are the only known members of this genus with submarginal papillae (in puparia and second and third instar nymphs) and marginal button-like structures (in second and third instar nymphs). The latter resemble the gland-tubercles described by Nakahara (1995) for the puparia of the species of Tetraleurodes Cockerell in the acaciae group.

Diagnosis. Tetralicia hoelmeri can be recognized from T. hyptisemoryi (characters for the latter species in square brackets) by the lower number of conical papillae on submargin ( Figs 104 , 105 ), three to five—usually four—on each side of abdomen only [six to eight on each side of thorax and abdomen—usually seven—and usually also a medial one on head; Figs 118 , 119 ], TMS ending almost at apparent margin ( Figs 107, 108 ) [barely extending onto submargin; Fig. 120 ], dorsum extensively covered with large tubercles (104, 105) [devoid of large tubercles; Figs 118 , 119 ], with paired pores and porettes ( Figs 114, 115 ) on each side of medial disc on mesothorax and A3–A6 [pores and porettes absent from that position; Fig. 122 ], deflexed submargin with dark granulations ( Fig. 113 ) extending over entire width lacking a particular pattern [with transverse rows of one to three clusters of dark granulations adjacent to the margin; Figs 125, 126 ], mediolateral abdominal depressions inconspicuous ( Figs 112, 114, 115 ) [conspicuous; Figs 118 , 119 , 122 ], and venter smooth except for spinulae medially to leg bases ( Fig. 111 ) and along margins [entirely covered with spinulae; Fig. 123 ]. Although their size ranges overlap partially, T. hoelmeri is in general smaller than T. hyptisemoryi (rounded average values followed by range in square brackets; values for T. hyptisemoryi in parenthesis): length 650 [570–760] (740 [665–850]); maximum width 490 [410–550] (540 [485–630]), dorsal A8 setae and caudal setae are relatively longer in T. hoelmeri than in T. hyptisemoryi : dorsal A8 setae length 90 [70–130] (65 [55–90]); caudal setae length: 60 [45–80] (50 [40–55]), and caudal setae are about as long as 1.5 times to 2.5 times the length of operculum in T. hoelmeri and slightly shorter to slightly longer (but less than 1.5 times) the length of operculum in T. hyptisemoryi .

The second and third instar nymphs of T. hoelmeri have button-like structures along margins of cephalothorax and abdomen set close together, separated from each other by a space subequal to the width of each structure ( Figs 262–264 ), whereas these structures are separated by a space subequal to two to four times the width of each structure in those nymphs of T. hyptisemoryi ( Figs 266, 267 ). The submarginal papillae are accompanied each by a pore in those nymphs of T. hyptisemoryi ( Fig. 264 ) which is absent in those nymphs of T. hoelmeri ( Fig. 266 ). As in the puparia, they also differ by their relative size, with T. hoelmeri earlier nymphal instars being smaller (first instar 177–211 long, second instar 224–243 long, third instar 335–398 long, vs. 243, 296, and 446 long respectively in T. hyptisemoryi ). And as is also the case in the puparia, T. hoelmeri third instar nymphs have relatively longer dorsal A8 setae (49–70, vs. 39 in T. hyptisemoryi ) and caudal setae (39–56, vs. 29 in T. hyptisemoryi ).

Adults are diagnosed from other known species under T. guajavae .

Remarks. This species is clearly closely related to T. hyptisemoryi , sharing only with it the presence of submarginal papillae. However, numerous differences were observed between them (see diagnosis above), and the combination of characters by which they differ is consistent in all specimens examined. Specimens of this species were included as paratypes of T. hyptisemoryi in Polaszek & Gill (2011), where the presence of tubercles on the dorsal disc and submargin was interpreted as ontogenetic variability, hypothesizing that the tubercles would develop as the pupa aged ( Polaszek & Gill 2011: 52). However, no intermediates were found, with pupae being either entirely smooth ( T. hyptisemoryi ) or extensively covered with tubercles ( T. hoelmeri ), and slide mounting and examination of numerous additional specimens of both species on samples of the CSCA dry collection and CDA herbarium showed that pupae otherwise matching T. hyptisemoryi with pharate adults ready to emerge still have an entirely smooth dorsal disc lacking any tubercles, and puparia matching other characters of T. hoelmeri always have these tubercles irrespective of being young or already eclosed, disproving that hypothesis. Furthermore, both species were found to coexist on the same host plant in two localities (Imperial National Wildlife Refuge and Vidal Junction in Imperial County), indicating that the differences considered here to be interspecific are not due to environmental or geographical variability. The descriptions of nymphal instars 1–3 and adults of T. hyptisemoryi in Polaszek & Gill (2011) correspond to specimens of T. hoelmeri , as do figures 2 and 3. Figure 2 ( Polaszek & Gill 2011: 54) illustrating a puparium of T. hoelmeri from the Living Desert Museum with two submarginal papillae on thorax is inaccurate, since reexamination of that particular specimen (photographed here in Figs 105 , 107 ), as well as of all other puparia of T. hoelmeri in CSCA, shows papillae to be restricted to submargin of abdomen and absent from submargin of thorax in this species.

The projections along apparent margin can be lobate to markedly conical and their development responds most likely to environmental conditions; specimens from Living Desert and Vidal Junction presented projections going from lobate to markedly conical, whereas specimens from Agua Caliente and Imperial Wildlife Refuge presented only lobate projections. Although their position corresponds to the position of the submarginal papillae for the most part, there are usually two additional submarginal projections on each side that are not accompanied by papillae, one on T3 and one on caudal abdomen between the last papilla and the caudal seta.

Carapia-Ruiz et al. (2018a) reported T. hyptisemoryi on Condea albida from Baja California and Baja California Sur in Mexico. However, since the description of T. hyptisemoryi was based on a composite of T. hyptisemoryi and T. hoelmeri , and no images accompany the Mexican records, it is not possible for us to determine if those records correspond to T. hyptisemoryi , to T. hoelmeri , or to both species.

Distribution. Desert areas in Southern California (Imperial, Riverside, and San Bernardino Counties).

Figures

FIGURE 104. Tetralicia hoelmeri sp. nov., Holotype: Habitus.

FIGURE 105. Tetralicia hoelmeri sp. nov., Paratype puparium, Riverside Co., Living Desert, Palm Desert, on desert lavender: Habitus.

FIGURE 118. Tetralicia hyptisemoryi, Holotype: Habitus.

FIGURE 119. Tetralicia hyptisemoryi, Paratype, Riverside Co., Deep Canyon, 25.iii. 2009: Habitus.

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 1–4. Puparia of Tetralicia in situ: 1: T. abnormis, Santa Barbara Co., Los Olivos, on Quercus agrifolia; 2, 3: T. acaudata, 2: Riverside Co., Tin Canyon, on manzanita, 3: San Bernardino Co., Mountain Home Village, Mill Creek Canyon, on manzanita; 4: T. agrifoliae, Riverside Co., Blue Jay Road & Ortega highway, on Quercus agrifolia. Fig. 2, by N. von Ellenrieder & R.W. Garrison, 1, 3, 4 by R.J. Gill.

FIGURES 106–111. Tetralicia hoelmeri sp. nov.: 106, 108, 110: Holotype; 107, 109, 111: Paratype, Riverside Co., Living Desert, Palm Desert, on desert lavender. 106: Cephalothorax; 107: T2–A3 dorsal disc and submargin; 108: Detail submargin T3–A2; 109: Detail submargin A2–A5; 110: Venter cephalothorax; 111: Detail venter of T2 and T3.

FIGURES 112–117. Tetralicia hoelmeri sp. nov.: 112–114, 116, 117: Riverside Co., Agua Caliente Indian Reservation, Palm Springs, on native shrub, 112, 113: Paratype; 114, 116, 117: Holotype; 115: Riverside Co., Living Desert, Palm Desert, on desert lavender. 112: Thorax andA1–A6 dorsal disc; 113: Detail deflexed submargin; 114, 115: Dorsal disc and submargin of abdomen; 116: Caudal end, dorsal view; 117: Detail vasiform orifice.

FIGURES 231, 232. Tetralicia hoelmeri sp. nov., adult female, Paratype, Riverside Co., Living Desert, Palm Desert, on desert lavender: 231: habitus; 232: ovipositor, ventral view.

FIGURES 233, 234. Tetralicia hoelmeri sp. nov., adult male, Paratype, Riverside Co., Living Desert, Palm Desert, on desert lavender: 233: habitus; 234: A10, ventrolateral view.

FIGURES 235–242. Tetralicia hoelmeri sp. nov., adults, Paratypes, Riverside Co., Living Desert, Palm Desert, on desert lavender: 235: female compound eye and antenna; 236: male compound eye and antenna; 237: female metatibia; 238: female mesotibia; 239: female protibia; 240: male metatibia; 241: male mesotibia; 242: male protibia.

FIGURES 261–264. Tetralicia hoelmeri sp. nov., nymphal instars 1–3, Paratypes, Riverside Co., Living Desert, Palm Desert, on desert lavender: 261: first instar nymph; 262: second instar nymph, stacked between first and third instars; 263: third instar nymph, with cephalothorax folded over; 264: third instar nymph, detail of unobstructed abdominal view.

FIGURES 120–126. Tetralicia hyptisemoryi: 120–122, 124: Holotype; 125, 126: Paratype, same data as holotype; 123: Paratype, Riverside Co., Deep Canyon, 25.iii. 2009. 120: Dorsal disc and submargin of T2 and T3; 121: Cephalothoracic venter; 122: Dorsal disc and submargin of abdomen; 123: Venter of abdomen; 124: Caudal end, dorsal view; 125: Detail deflexed submargin of thorax; 126: Detail deflexed submargin of abdomen.

FIGURES 265–267. Tetralicia hyptisemoryi, set of stacked nymphal instars 1–3 exuviae overlaying holotype puparium: 265: first instar nymph; 266: submargin of third instar nymph; 267: Caudal end of second and third instar nymphs (third instar below).