Tetralicia guajavae 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 : 27-30

Tetralicia guajavae von Ellenrieder & Gill , n.sp.

Figs 11, 12 , 97–103, 219–230, 255–257

Aleurotrachelus cecropiae Carapia-Ruiz 2020b: 5 (in part: material from Mexico).

Tetralicia cecropiae : Valencia & Evans 2024: 212 , in part: fig. 4F from Carapia-Ruiz 2020b.

Etymology. This species is named after the specific epithet of its preferred host, Psidium guajava .

Type material. 145 puparia: U.S.A., California; 45 slide mounted: Orange County: 1 holotype (circled in red on slide), Garden Grove , on Psidium sp. , 9.x.2015, Lyon, Marty, de Nijs & Cairo coll., CDFA PDR 570P06364047 [ CSCA]; 3 paratypes, same data as holotype [ CSCA]; 4 paratypes, Santa Ana , in nursery, on Psidium guajava , 30.iv.2007, J. Rivas coll. [ CSCA]; 22 paratypes, same data but 12.v.2010, J.N. Nisson coll., CDFA PDR 1544518A [ CSCA]; Los Angeles County: 10 paratypes, Compton , in nursery, on guava, 14.xi.2018, C. Foy coll., CDFA PDR 190P06620744 [ CDFA]; 1 paratype, Sylmar , in nursery, on Psidium sp. , 8.viii.2013, Matsumoto coll., CDFA 190P06058724 [ CSCA]; 4 paratypes, interception of unknown origin, on Psidium guajava , 28.vi.2014, Licea & Chavez coll., CDFA DR0P06177819 [ CSCA] .

1 adult male paratype, 1 adult female paratype, 5 first instar nymphs paratypes, 2 second instar nymphs paratypes, 10 third instar nymphs paratypes: Orange County: Santa Ana, in nursery, on Psidium guajava , 12.v.2010, J.N. Nisson coll. [ CSCA] .

Additional material examined: 100 + dry mounted puparia: U.S.A., California, Orange County, Santa Ana , in nursery, on Psidium guajava , 30.iv.2007, J. Rivas coll. [ CSCA] .

Hosts. Myrtaceae : guava ( Psidium guajava ) in California; Carapia-Ruiz (2020b) reported this species (as T. cecropiae ) from Psidium guajava and Curatella americana (Dilleneaceae) in Mexico.

Description.

Field characteristics. Puparium oblanceolate with pronounced caudal protuberance, black with pale brown venter, with a narrow ring of lateral wax along true margin ( Figs 11, 12 ). Adult color in life unknown; nymphal instars 1–3 pale yellow.

Slide-mounted characters. TMS extending onto submargin but not reaching apparent margin, not lined with tubercles ( Figs 97 , 98 ); longitudinal molting suture lined with tubercles from T2/T3 suture to head submargin ( Fig. 97 ); eyespots absent; Ce setae absent; T2 and T3 setae present, T3 setae arising well behind anterior margin of metanotum ( Figs 97 , 98 ); medial area with depressions lined with small granulations on each side of head (two), T2 (two) and T3 (one), and abdominal segments (one) ( Figs 97–99 ); with paired pores and microsetae on each side of submedial area of head (2), T2 (1), T3 (1), A1 (1), A2 (0), A3 (1), A4 (0–1), A5 (1), A6 (0), A7 (1), and A8 (1); dorsal disc smooth except for small granulations across intersegmental areas and along lateral areas of head, thorax, and abdomen ( Figs 97–99 ); with paired pores and microsetae along sides of dorsal disc ( Fig. 99 ); dorsal submargin with small granulations turning into transverse rows of larger paired crenulations externally ( Figs 97–99 ); deflexed submargin lacking microsetae and pores and with granulations arranged in transverse bands ( Figs 100, 101 ); marginal glandular teeth subquadrangular, with tips smooth or dentate; VO oval, inset from posterior margin by more than its own length, and located on a promontory visible in oblique or lateral view ( Figs 102, 103 ); operculum cordate, its dorsal surface with longitudinal and transverse ridges and wavy striations in between, with microspinulae across distal third to fourth; lingula concealed by operculum; VO ring very narrow around lateral and posterior sections of VO, with anterior portion wide and rectangular ( Fig. 102 ), with dorsal setae of A8 arising from sides anteriorly to level of anterior margin of operculum ( Fig. 102 ); bases of caudal setae moderately separated, arising just outside level of lateral margins of VO ring ( Figs 102, 103 ); with two membranous ventral sacs medially to bases of mesothoracic legs ( Fig. 101 ); tegument of venter smooth except for spinulae on a wide band along medial bases of legs and margins of head and abdomen ( Figs 100, 101 ).

Adults. Female ( Fig. 204 ) with 2 pairs and male ( Fig. 206 ) with 4 pairs of wax plates on abdomen. Upper and lower compound eyes joined by three ommatidia. Antennae 7 segmented in both sexes ( Figs 223, 224 ), with an elongate third segment with two primary round sensoria near the apex, fifth and seventh segments each with 1 round primary sensorium located at apex on fifth and at about two thirds of its length on seventh, and third, sixth, and seventh segments with an elongated rod-like sensorium each. Hind tibiae ( Figs 225, 228 ) with a tibial comb of 12 or 13 setae in male, 12–14 setae in female; metatibial brush with 2 setae in male, 3 setae in female. Mesotibial brushes with 2 and 3 setae in male, 3 setae in female. Female cement gland not discernible ( Fig. 220 ). Male parameres very narrow along distal fourth, ending on a single long apical tooth, lacking subapical teeth ( Fig. 222 ).

Nymphal instars 1–3. All with a central longitudinal rachis along abdomen ( Figs 255–257 ). First instar ( Fig. 255 ): with 6 or 7 pairs of long submarginal setae along sides of cephalothorax and 1 pair at 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 ( Figs 256, 257 ): same as first except for absence of long submarginal setae, VO located on a promontory, and operculum subcordate.

Measurements (values of holotype in square brackets). Puparium length: 622 ± 50 [679]; maximum width (between level of T2/T3 suture and A1): 376 ± 40 [432]; length/maximum width: 1.7 ± 0.1 [1.6]; width at level of anterior margin of operculum: 154 ± 19 [175]; maximum width/width at anterior margin of operculum: 2.4 ± 0.2 [2.5]; deflexed submargin/body radius: 0.55 ± 0.05 [deflexed submargin not measurable]; Ce setae: absent; T2 setae: 13 ± 2 [missing]; T3 setae: 15 ± 2 [13]; dorsal A8 setae: 12 ± 2 [15]; caudal setae: 75 ± 8 [84]; anterior marginal setae: 9 ± 1 [not visible]; posterior marginal setae: 14 ± 1 [not visible]; ventral A8 setae: 16 ± 5 [15]; VO ring length: 47 ± 7 [49]; VO ring width: 39 ± 4 [41]; VO ring length/width: 1.2 ± 0.2 [1.2]; caudal seta/VO ring length: 1.6 ± 0.3 [1.7]; caudal seta/operculum length: 3 ± 0.3 [3.1]; VO length: 29 ± 3 [29]; VO width: 30 ± 3 [34]; VO length/width: 1 ± 0.1 [0.9]; operculum length: 26 ± 3 [27]; operculum/VO length: 0.9 ± 0.05 [0.9] (see Table 1 for ranges).

Adults. Male: body length (including parameres): 766; rostral length: 165; hind tibia length: 252; third antennal segment length: 97; paramere length: 92; aedeagus length: 82. Female: body length (including ovipositor): 815; rostral length: 179; hind tibia length: 291; third antennal segment length: 107; ovipositor length: 121.

Nymphal instars 1–3. First instar: length: 234 ± 2 [232–237]; tibiotarsal length: 12; submarginal setae: 21 ± 5 [17–24]; T2 setae: 22; metathoracic setae: 24; dorsal A8 setae: 4; caudal setae: 5. Second instar: length: 283 ± 3 [281–285]; T2 setae: 52; metathoracic setae: 54; dorsal A8 setae: 12; caudal setae: 21 ± 1 [20–22]. Third instar: length: 414 ± 24 [376–441]; anterior setae: 8; posterior setae: 12; T2 setae: 19; T3 setae: 29; dorsal A8 setae: 9 ± 2 [7–10]; caudal setae: 56 ± 7 [49–68].

Similar species. Tetralicia cecropiae , T. lantanae , and T. oblanceolata share an oblanceolate puparium and VO located on a pronounced promontory.

Diagnosis. Puparium can be distinguished from other oblanceolate species in California (characters for T. lantanae and T. oblanceolata in square brackets) by its TMS not reaching apparent margin ( Figs 97 , 98 ) [reaching apparent margin; Figs 134 , 135 , 168 –170], lacking large tubercles on submedian abdomen ( Figs 97 , 99 ) [with large tubercles at least along anterior margin of abdominal segments; Figs 134 , 135 , 137 , 169 ], and mediolateral abdominal depressions conspicuous and lined with small granulations ( Figs 97 , 99 ) [depressions inconspicuous; Figs 134 , 135 , 137 , 169 ]. It can be recognized from T. cecropiae (characters for T. cecropiae in square brackets, based on pictures of syntypes at USNM kindly sent by G. Evans and I. Stocks) by TMS extending halfway onto submargin ( Figs 97 , 98 ) [ending at submargin without extending onto it], dorsal submargin with granulations turning into transverse rows of paired crenulations externally [dorsal submargin lacking granulations or crenulations], spinulae present along medial bases of legs but absent from legs' surfaces ( Figs 100, 101 ) [spinulae entirely covering legs], absence of a sclerotized arch on A7 ( Figs 97 , 103 ) [with a sclerotized arch on posterior margin of A7], setae on T2, T3, and on dorsum of A8 shorter (seta length/VO length T2: 0.3–0.5; T3: 0.4–0.6, A8: 0.3–0.5) and caudal setae longer (caudal seta/VO length 2.3–3.1) compared to length of VO [longer and shorter respectively; seta length/VO length T2: 0.8, T3: 1.0, A8: 0.7, caudal: 1.4], and VO located on a less pronounced promontory [on a higher promontory]). Bondar (1923) described and illustrated the lateral wax ring on true margin as extending dorsally along submargin and beyond in T. cecropiae ; in our dry mounted specimens of T. guajavae it only extends partially along ventral most section of submargin ( Fig. 12 ).

Among the known adults of Tetralicia from California, T. abnormis has two smoky spots near midlength and three on distal portion of forewings ( Quaintance 1900: 18), whereas in addition to those markings T. lantanae has also a basal spot, and T. nigrans has wings 'somewhat dusky at distal ends' ( Bemis 1904: 523). The color of live adults of T. guajavae is unknown, and that of T. hoelmeri was not described ( Polaszek & Gill 2011), and since the faint markings of species with marked forewings are usually not discernible in slide-mounted specimens (pers. obs.), it is not possible to know if the forewings in these two species are unmarked (like they would appear to be in slide-mounted specimens) or marked. Tetralicia abnormis and T. guajavae appear to be the smallest and T. hoelmeri the largest ( Table 2 ). The upper and lower sections of the compound eyes are connected by three ommatidia in T. guajavae and two to three ommatidia in T. lantanae ( Figs 247, 248 ), but only by one in T. hoelmeri ( Figs 235, 236 ). The adult male of T. guajavae can also be recognized from those of T. lantanae and T. hoelmeri by parameres ending on a single slightly curved long distal tooth, as long as about a fourth of parameres length, lacking subapical teeth ( Fig. 222 ). In T. hoelmeri the distal tooth is about as long as a sixth of parameres length and is more markedly curved medially ( Fig. 234 ), whereas in T. lantanae it is as long as a ninth of parameres length and it is preceded by three subapical teeth ( Fig. 246 ). Both T. guajavae and T. lantanae have metatibial and mesotibial brushes of 2 or 3 setae, but these are not defined in T. hoelmeri . The female of T. hoelmeri ( Fig. 232 ) differs further from those of T. guajavae ( Fig. 220 ) and T. lantanae ( Fig. 243 ) by its much longer ovipositor (see Table 2 ). Based on the absence of tibial brushes and the long ovipositor, the adults of T. hoelmeri resemble those of T. ericae ( Valencia & Evans 2024: 208, fig. 3). Bemis' (1904) description of the adults of T. nigrans was very brief, but their size is comparable to those of T. lantanae (see Table 2 ), from which they differ by the wing pattern as described above.

First instar nymphs of T. guajavae and T. lantanae differ from those of T. hoelmeri and T. hyptisemoryi by their shorter submarginal setae (see Table 3 ), and second and third instar nymphs, as well as third instar nymph of T. salsolae , by the absence of submarginal papillae and marginal button-like structures ( Figs 256, 257, 259, 260 , 268 ) that are present in T. hoelmeri and T. hyptisemoryi ( Figs 262–264 , 267 ). The third instar nymph of T. salsolae can be recognized from those of T. guajavae and T. lantanae by its longer T2 and T3 setae and shorter caudal setae, much larger size (see Table 3 ) and presence of four petal-shaped lateroapical projections on VO ( Fig. 270 ), which are lacking in the latter two species ( Figs 257, 260 ). All nymphal instars 1–3 of T. guajavae can be recognized from those of T. lantanae by their larger size (see Table 3 ).

Remarks. Based on the photograph included with the record from Mexico ( Carapia-Ruiz 2020b: 6, fig. 9) we believe that the record of T. cecropiae from Oaxaca corresponds to T. guajavae , and T. cecropiae is so far only known from Brazil.

Biology. A species recently introduced in the state, probably from Mexico, found in large numbers in Southern California between 2007 and 2010 probably due to its parasitoids not being originally introduced with it. The scarcity of recent findings might indicate that it is now under biological control.

Distribution. Found outdoors in Orange County and occasionally in Southern California nurseries; recorded from Mexico (as T. cecropiae ; Carapia-Ruiz 2020b).

Figures

FIGURE 97. Tetralicia guajavae sp. nov., Holotype: Habitus.

FIGURE 134. Tetralicia lantanae sp. nov., Holotype: Habitus.

FIGURE 135. Tetralicia lantanae sp. nov., Paratype puparium, Santa Barbara Co., Santa Barbara, on Hibiscus: Habitus.

FIGURE 168. Tetralicia oblanceolata, Neotype: Habitus.

FIGURE 169. Tetralicia oblanceolata, Orange Co., Silverado Canyon, on Rhamnus ilicifolia: Habitus.

FIGURES 9–12. Puparia of Tetralicia in situ: 9, 10: T. fouquieriasplendens, Riverside Co., Palm Desert, Living Desert Museum, on Fouquieira digetii, 9: dorsal view, 10: dorsolateral view. 11, 12: T. guajavae, Orange Co., Santa Ana, on Psidium guajava, 11: dorsal view, 12: ventral view. Figs 9–12 by N. von Ellenrieder & R.W. Garrison.

FIGURES 98–103. Tetralicia guajavae sp. nov.: 98–100, 102: Holotype; 101: Santa Ana, on Psidium guajava; 103: Los Angeles Co., Compton, on guava. 98: Dorsal submargin and disc of thorax; 99: Dorsal submargin and disc of A1–A5; 100, 101: Detail deflexed submargin; 102, 103: Detail caudal end, 102: dorsal view, 103: dorsolateral view.

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 223–230. Tetralicia guajavae sp. nov., adults, Paratypes, Orange Co., Santa Ana, in nursery, on Psidium guajava: 223: female antenna; 224: male antenna; 225: female metatibia; 226: female mesotibia; 227: female protibia; 228: male metatibia; 229: male mesotibia; 230: male protibia.

FIGURES 219, 220. Tetralicia guajavae sp. nov., adult female, Paratype, Orange Co., Santa Ana, on Psidium guajava: 219: habitus; 220: ovipositor, ventral view.

FIGURES 221, 222. Tetralicia guajavae sp. nov., adult male, Paratype, Orange Co., Santa Ana, on Psidium guajava: 221: habitus; 222: A10, ventrolateral view.

FIGURES 255–260. Nymphs: 255–257: Tetralicia guajavae sp. nov., Orange Co., Santa Ana, on Psidium guajava; 258–260: Tetralicia lantanae sp. nov., paratypes, Riverside Co., University of California Riverside, on Lantana: 255, 258: first instar nymph; 256, 259: second instar nymph; 257, 260: third instar nymph.

FIGURES 136–141. Tetralicia lantanae sp. nov.: 136–139: Holotype; 140: Paratype, San Diego Co., San Diego, on Lantana; 141: Paratype, Orange Co., Anaheim, on Lantana. 136: Dorsal disc of thorax; 137: Dorsal disc and submargin of A1–A6; 138: Venter of T2 and T3; 139: Caudal end of abdomen, dorsal view; 140: Detail VO; 141: Caudal end of abdomen, dorsolateral view.

FIGURES 247–254. Tetralicia lantanae sp. nov., adults, Paratypes, Riverside Co., University of California Riverside, on Lantana: 247: female compound eye and antenna; 248: male compound eye and antenna; 249: female metatibia; 250: female mesotibia; 251: female protibia; 252: male metatibia; 253: male mesotibia; 254: male protibia.

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 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 245, 246. Tetralicia lantanae sp. nov., adult male, Paratype, Riverside Co., University of California Riverside, on Lantana: 245: habitus; 246: S10, ventrolateral view.

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 243, 244. Tetralicia lantanae sp. nov., adult female, Paratype, Riverside Co., University of California Riverside, on Lantana, 30.i.1992, E. Reeves coll.: 243: habitus; 244: ovipositor, ventral view.

FIGURES 268–271. Tetralicia salsolae, third instar nymph, Paratype, Imperial Co., Ocotillo, on Ambrosia salsola: 268: Habitus; 269: Detail thoracic setae; 270: Detail tracheal opening and legs; 271: Caudal end.

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 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).