Tetralicia lantanae von Ellenrieder & Gill, sp. nov.

Figs 18–20, 135–141, 243–254, 258–260

Aleuropleurocelus granulata: Evans 2007: 170; Stocks 2016: 7; Sanchez-Flores et al. 2020: 278–280 (misidentification in part: records of specimens on Lantana from U.S.A. and Mexico).

Tetralicia granulata: Valencia & Evans 2024: 217 (misidentification in part: characterization, key, fig. 5L from Sanchez-Flores et al. 2020).

Etymology. This species is named after the generic epithet of its preferred host, Lantana .

Type material. 189 slide mounted puparia: U.S.A., California, Riverside County: Holotype (circled with red on slide), Riverside, on Lantana, 17.xii.1991, T. Viztnum coll., CDFA PDR 976224 [CSCA]; 34 paratypes, same data as holotype [CSCA]; 18 paratypes, University of California Riverside, on Lantana, 30.i.1992, E. Reeves coll. [CSCA]; 5 paratypes, same data [FSCA]; San Diego County: 22 paratypes, San Diego, on Lantana, 22.x.1992, D. Kellum coll., CDFA PDR 1015430 [CSCA]; Orange County: 11 paratypes, Mission Viejo, on Lantana, 28.vii.1988, V. Oppenheimer coll., CDFA PDR 861275 [CSCA]; 32 paratypes, Anaheim, on Lantana, 4.xii.1990, M. Beach coll., CDFA PDR 929986 [CSCA]; San Bernardino County: 1 paratype (specimen circled with red on slide), [CSCA]; Santa Barbara County: 6 paratypes, Santa Barbara, on Hibiscus, 19.iii.1993, J. Davidson coll., CDFA PDR 965797 [CSCA]. Arizona, Maricopa County: 7 paratypes, Mesa, on Morus, ix.1995, C. Pickett coll. [CSCA]; Pima County: 25 paratypes, Tucson, on Lantana, viii.1991, D. Cross coll. [CSCA]; 4 paratypes, interception from Arizona at Blythe, on house plants, 18.ix.1988, D.A. El Nawasrah coll. [CSCA]. Texas, Hidalgo County: 2 paratypes, Weslaco, on Lantana camara, 23.iii.1992, C. Mooman coll. [FSCA]; 3 paratypes, Weslaco, on Lantana horrida, 26.ix.1992, W.A. Jones coll. [CSCA]; 10 paratypes, same data but on Eupatorium coelestinum [CSCA]; Florida, Lake County: 4 paratypes, on Lantana, 26.xi.1979, L.J. Chambliss [FSCA]; 1 paratype, same data [CSCA]; Pinellas County: 1 paratype, Pinellas Park, 7651 US 19 N, on Lantana camara, 2.iii.2016, M. Spearman coll., E-2016-711-1 [FSCA]; Martin County: 2 paratypes, Hobe Sound, on Lantana sp., 6.iii.1979, E.W. Campbell coll. [FSCA] .

57 slide mounted paratype adults: 10 males, 14 females, San Diego County, San Diego, on Lantana, 22.x.1992, D. Kellum coll. [CSCA]; 9 males, 24 females, Anaheim, on Lantana, 4.xii.1990, M. Beach coll. [CSCA] .

11 slide mounted paratype nymphal instars 1–3: 3 first instar, 3 second instar, 6 third instar: University of California Riverside, on Lantana, 30.i.1992, E. Reeves coll. [CSCA] .

Additional material examined: 7 slide mounted puparia: U.S.A., Florida, Miami-Dade County: 6, Miami, on Lantana sp., 27.vi.1979, R. Larkin coll. [FSCA]; Palm Beach County: 1, Greenacres 5013, Solar Point Drive, on Lantana camara, 26.vi.2008, L. Smith coll., E-2008-4251-1 [FSCA]. 600 + dry mounted puparia, U.S.A., California: 200 +, Orange County, Mission Viejo, on Lantana, 28.vii.1988, V. Oppenheimer coll., CDFA PDR 861275 [CSCA]; 400 +, Riverside County, University of California Riverside, on Lantana, 30.i.1992, E. Reeves coll. [CSCA].

Hosts. Common on Lantana sp. ( Verbenaceae) and found also on Asteraceae: Eupatorium colestinum, Malvaceae: Hibiscus sp., and Moraceae: Morus sp.

Description.

Field Characteristics. Puparium markedly oblanceolate, elevated with medial area of dorsum depressed and pronounced caudal protuberance (Figs 18, 19), black, with a narrow ring of white wax around true margin (Fig. 19). Adults yellow; wings white with faint grey marks on forewings: one rounded spot at base, one rounded spot at midlength, and two rounded spots and a longitudinal stripe at distal third, the latter contiguous (Fig. 20) or divided into a basal and a distal spot; nymphal instars 1–3 transparent yellow (Fig. 19).

Slide-mounted characters. TMS ending at apparent margin, not lined with tubercles medially; longitudinal molting suture lined with tubercles from TMS to submargin (Figs 134, 135); eyespots absent; Ce setae absent; T2 and T3 setae present, T3 setae arising well behind anterior margin of metanotum; medial area of head with some tubercles on a posterolateral patch (Figs 134, 135), medial area of thorax almost entirely covered with large tubercles distributed uniformly (Fig. 136), except in a few specimens with a small bare area posterior to setal bases; medial area of abdomen with large tubercles adjacent to anterior and posterior margins of abdominal segments, forming a complete anterior row on each A1–A7, a second anterior row incomplete medially on A2–A5, a complete posterior row on A1–A3 or A1–A4, and a posterior row incomplete medially on A4–A7 or A5–A7 (Figs 134, 135, 137); anterolateral depressions on abdomen inconspicuous, usually with pores and microsetae on each side of submedial area of head (2), T1 (2), T2 (2), T3 (2), A1 (2), A2 (0), A3 (2), A4 (0), A5 (0–2), A6 (0–2), A7 (2), and A8 (2); lateral areas of dorsal disc with large tubercles except for head (Figs 134, 135), with pores between dorsal disc and submargin; dorsal submargin with transverse double rows of tubercles turning into crescent-shaped imbrications towards apparent margin, not reaching it (Figs 134, 135, 137, 141), fewer and much smaller on head (Figs 134, 135); deflexed submargin lacking microsetae and pores, covered with large granulations distributed uniformly in wide transverse bands (Fig. 138); marginal glandular teeth subquadragular with tips serrate (Fig. 138); VO subcordate, inset from posterior margin by more than its own length; operculum ornamented with longitudinal ridges and with a patch of distal microspinulae; VO ring subquadrate, wide anteriorly and narrow posteriorly (Figs 139, 140), forming an elevated promontory (Fig. 141), with rugose surface (Figs 140, 141); dorsal A8 setae not detected; with two membranous ventral sacs medially to bases of mesothoracic legs (Fig. 138); bases of caudal setae close together, within level of operculum lateral margins (Figs 134, 135, 139, 140); venter smooth except for groups of spinulae medially to leg bases (Fig. 138).

Adults. Female (Fig. 243) with 2 pairs and male (Fig. 245) with 4 pairs of wax plates on abdomen. In both sexes upper and lower compound eyes joined by two or three ommatidia (Figs 247, 248) and antennae 7 segmented (Figs 247, 248), 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 249, 252) with a tibial comb of 15–17 setae in male, 15–16 setae in female; metatibial brush with 2 or 3 setae in male, 2 setae in female. Mesotibial brushes with 2 and/or 3 setae in male and female. Female cement gland tube-like and capitate (Fig. 244). Male parameres ending on a single tooth along distal ninth and bearing three short subapical teeth (Fig. 246).

Nymphal instars 1–3. All with a central longitudinal rachis along abdomen (Figs 258–260). First instar (Fig. 258): 6 pairs of submarginal setae on cephalothorax and 1 on posterior abdomen (mostly broken off in available specimens). 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 259, 260): same as first except for VO located on a promontory and operculum subcordate; venter of tracheal and caudal openings with granulations in third instar (Fig. 260).

Measurements. Puparium length: 594 ± 41 [652]; maximum width (at level of T2/T3 suture): 343 ± 25 [373]; length/maximum width: 1.7 ± 0.1 [1.7]; width at level of anterior margin of operculum: 99 ± 9 [97]; maximum width/width at anterior margin of operculum: 3.5 ± 0.5 [3.9]; deflexed submargin/body radius: 0.6 ± 0.05 [0.6]; Ce setae: absent [absent]; T2 setae: 16 ± 2 [missing]; T3 setae: 18 ± 3 [missing]; dorsal A8 setae: none visible; caudal setae:129 ± 18 [139]; anterior marginal setae: 14 ± 5 [none visible]; posterior marginal setae 16 ± 4 [19]; ventral A8 setae: 25 ± 2 [29]; VO ring length: 43 ± 2 [41]; VO ring width: 37 ± 3 [36]; VO ring length/width: 1.2 ± 0.1 [1.1]; caudal seta/VO ring length: 3 ± 0.5 [3.4]; caudal seta/operculum length: 6 ± 1 [6.3]; VO length: 25 ± 2 [25]; VO width: 24 ± 1 [23]; VO length/width: 1.1 ± 0.1 [1.1]; operculum length: 22 ± 1 [22]; operculum/VO length: 0.9 ± 0.1 [0.9] (see Table 1 for ranges).

Adults. Male: body length (including parameres): 899 ± 62 [805–980]; rostral length: 214 ± 15 [194–235]; hind tibia length: 283 ± 15 [267–306]; third antennal segment length: 112 ± 4 [107–117]; paramere length: 101 ± 4 [97–107]; aedeagus length: 96 ± 4 [92–102]. Female: body length (including ovipositor): 961 ± 66 [815–1038]; rostral length: 249 ± 16 [218–267]; hind tibia length: 303 ± 13 [281–325]; third antennal segment length: 116 ± 2 [114–119]; ovipositor length: 142 ± 9 [129–155].

Nymphal instars 1–3. First instar: length: 207 ± 10 [197–216]; tibiotarsal length: 24 1 [23–24]; submarginal setae: 19. Second instar: length:258; T2 setae: 45 12 [36–53]; T3 setae: 47 12 [39–56]; caudal setae: 50 ± 2 [49–51]. Third instar: length: 389 ± 24 [355–412]; anterior setae: 10; T2 setae: 40 5 [36–44]; T3 setae: 44; dorsal A8 setae: 17; caudal setae: 100 ± 8 [92–114].

Similar species. Puparium shares an oblanceolate shape and dorsal disc of thorax and abdomen extensively covered with large tubercles with T. oblanceolata in California and with T. granulata Sampson & Drews, 1941, T. pseudogranulata Carapia-Ruiz & Sánchez-Flores, 2021, and T. sampsoni Sánchez-Flores et al., 2020 in Mexico.

Diagnosis. Puparium can be recognized from all the species mentioned above except for T. granulata (characters for them in square brackets) by its ornamentation with tubercles usually covering the entire thoracic dorsal disc (Figs 134–136) [with bare patches on the medial area of each segment; Figs 151–154], and further from T. oblanceolata by medial area of A1–A6 with tubercle-shaped ornamentations across both anterior and posterior areas (Figs 134, 135, 137) [only across anterior area of A1–A6; Figs 168, 169]; from T. pseudogranulata by the rounded dorsal tubercles of abdomen and apparent absence of A8 setae [elongate dorsal tubercles and A8 setae 42 long; Carapia Ruiz et al. 2020]; and from T. sampsoni by the presence of spinulae medially to leg bases (Fig. 138) and absence of dorsal wax [lacking spinulae medially to leg bases and with dorsal wax; Sanchez Flores et al. 2020]. It differs from T. granulata by its markedly oblanceolate puparium (Figs 134, 135) [wider apically; Figs 210, 211], caudal setae very long, as long as 3.5 or more times the length of VO (Figs 134, 135) [shorter, as long as 1.5 to 2 times the length of VO; Figs 216–218)], venter with spinulae medially to leg bases (Fig. 138) [venter smooth medially to leg bases, Fig. 215], ornamentation of medial area of abdominal dorsum with tubercles on anterior and posterior rows only (Figs 134, 135, 137) [tubercles covering almost entire length of segments except for small mediolateral patch; Figs 210, 211, 216], dorsal submargin with transverse rows of tubercles separated by bare areas narrower than rows' width, turning into crescent shaped imbrications towards apparent margin (Figs 134, 135, 137) [rows of tubercles separated by bare areas wider than rows' width, with tubercles decreasing in size towards apparent margin; Fig. 212], and setal sockets of caudal setae close together, separated by a distance subequal to width of one to two setal sockets, within level of operculum lateral margins (Figs 134, 135, 139, 140) [setal sockets of caudal setae separated by a distance subequal to the width of three to five setal sockets, located at level of lateral margin of VO ring (Fig. 218) to externally to it (Fig. 217)]. In T. lantanae VO is located on a pronounced promontory (Fig. 141) with a rugose surface (Fig. 140), apparently obscuring the position of A8 dorsal setae, which were not detected in any of the examined specimens. In T. granulata, the VO is not as elevated, and the setal sockets of A8 dorsal setae are visible on its anterior margin (Fig. 216).

Adults and nymphal instars 1–3 are diagnosed under T. guajavae .

Remarks. Examination of the type series of T. granulata (Figs 210–218) deposited at BME allowed us to recognize that the specimens of Tetralicia found on Lantana, identified as T. granulata in the past based on Sampson & Drews' (1941) description which lacks sufficient detail to allow for a reliable diagnosis, represent a different, still undescribed, species. All literature records of T. granulata subsequent to its description (Evans 2007; Stocks 2016; Sanchez-Flores et al. 2020) correspond to T. lantanae . To complement its original description and facilitate its diagnosis we provide a redescription of T. granulata (not known to occur in California; see below the species accounts of Tetralicia species found in California), including characterization, measurements, and illustrations, based on its type series.

In the key in Valencia & Evans (2024: 210, 211) this species (as T. granulata) is considered to have an operculum 'without many short, spinelike-setae along its posterior margin'. Although they are not always readily visible, there are microspinulae covering the distal third to fourth of the dorsal surface of the operculum in all the specimens of T. lantanae that we examined.

Biology. Found in large numbers in Southern California between 1988 and 1993 probably due to its parasitoids not being originally introduced with it. The scarcity of more recent findings might indicate that it is now under biological control.

Distribution. Known to occur in Southern California (Orange, Riverside, San Bernardino, and San Diego Counties); in the U.S.A. also in Arizona (this paper), Florida and Texas (reported as A. granulata, Evans 2007; Stocks 2016; Valencia & Evans 2024), and in Mexico (reported as A. granulata; Sánchez-Flores et al. 2020).