Poblicia fuliginosa ( Olivier, 1791 )

Poblicia fuliginosa ( Olivier, 1791) View in CoL —The sumac speckled lanternfly

( Figures 2–4A–C View FIGURE 2 View FIGURE 3 View FIGURE 4 , 5–7 View FIGURE 5 View FIGURE 6 View FIGURE 7 , 12A–B View FIGURE 12 , 18A View FIGURE 18 )

= Fulgora fuliginosa Olivier, 1791: 574 View in CoL (original combination). = Poblicia fuliginosa ( Olivier, 1791) View in CoL , comb. by Van Duzee (1917: 719). = Poblicia thanatophana Kirkaldy, 1907 View in CoL ; synonym by Ball (1933: 146) (error). = Crepusia fuliginosa ( Olivier, 1791) View in CoL , comb. by Nast (1951: 270) (error). Olivier 1791: 574 [described].— Metcalf 1947: 62 [cataloged, full synonymy list].— Nast 1951: 270 [erroneous combination].—

Wilson & McPherson 1980a: 16, 48 [key, illustrated].— Wilson & McPherson 1980b: 29 [listed].— Wilson & McPherson

1980c: 15 [listed].— Porion 1994: 21 [cataloged, illustrated].— Moran et al. 2005: 8803 [endosymbionts].— Urban & Cryan

2009 [phylogenetics].— Bartlett et al. 2011 [listed, key, illustrated].— Urban & Cryan 2012 [endosymbionts].— Bartlett et al. 2014: 31–32, 169 [illustrated, notes].— Broadley et al. 2023 [parasites].

Diagnosis. Body length between about 11.0– 15.5 mm, nearly black with minute pale spotting on the body, and larger bluish to white maculations on the forewing, area past nodal line bearing many pale dashes between longitudinal veins. Vertex very narrow (vs. P. misella ), in frontal view dorsum of head truncate. Fore- and middle tibiae with 2 pale bands. Hindwings smoky-transparent washed with fuscous in proximal half (enclosing two large irregular clear windows) and along wing apex. Rostrum reaching hind coxa. Abdominal dorsum red.

Amended description. Color. Overall coloration is nearly black ( Fig. 2 View FIGURE 2 , darker in vivo, Fig. 18A View FIGURE 18 , dry specimens notably browner in appearance) with faint brown to bluish wash; head with anterior margin distinctly paler; body with fine pale speckles everywhere, except forewings ( Fig. 3A View FIGURE 3 ) with larger pale maculations especially following costa (forewing spots pale blue proximally, whitish past nodal line), forewings uniformly darkened (differing in hue past nodal line, before nodal line bearing many fine, irregular contrasting pale reticulate lines in cells, mostly connecting veins; beyond nodal line more distinct white dashes between veins) usually with short whitish transverse bar at apex of clavus (often obscure in vivo, more evident in dry specimens, especially if washed with alcohol), and a matching marking from costa; hindwings ( Fig. 3B View FIGURE 3 ) mostly smoky-transparent, apex fuscous-gray, proximal portion near black with two large irregular pale spots (may fade to whitish or translucent in preserved specimens), veins dark except proximal portion of MA pale. Legs black with irregular white spots, front and middle tibiae ( Fig. 4C View FIGURE 4 ) with two white bands (proximal one close to tibial femoral joint, distal band before tibial apex). Abdomen ventrally near black, dorsally bright red (evident on lateral portion of abdomen in repose).

Structure. Body size (including wings) males x̄ = 12.8 mm (11.0–13.5, n=5), females x̄ = 15.0 mm (14.0–15.5, n=5). Body in dorsal view ( Fig. 2A View FIGURE 2 ) broad and parallel-sided, in lateral view ( Fig. 2B View FIGURE 2 ), head and thorax in similar plane (not appearing hump-backed). Head (dorsal view, Fig. 2A View FIGURE 2 ) broad and strongly transverse, just narrower than prothorax, anteriorly weakly convex and posteriorly concave. Vertex much wider than long (about 4.5× wider than midlength), carinate on all margins (carina appearing thicker anteriorly), disc depressed, bearing weak median longitudinal carina, surface roughened; anterior margin of head (dorsal view) appearing to have second anterior carina. Head (lateral view, Fig. 2B View FIGURE 2 ) with face receding ventrally, inflection between vertex and frons sharp. Frons (frontal view, Fig. 2C View FIGURE 2 ) broad, roughly quadrate, much broader than high (midlength about 0.56× width at dorsal margin), dorsal margin truncate, lateral margins sinuate, laterally foliate, somewhat narrowing ventrad; face surface rugulose bearing faint transverse grooves, with median carina (not reaching fastigium) and diagonally oriented lateral (approximated at base, giving a V-shaped appearance, becoming obsolete ventrad). Frontoclypeal suture arched, clypeus triangular with rounding dorsal margin, median third longitudinally carinate. Antennae short, scape smooth, barely as tall as wide (hidden behind lateral foliations of frons from frontal view), pedicel somewhat bulbous, longer than wide bearing numerous sensory plaques, flagellum bristle-like with bulbous base. Eyes bulbous, laterally projecting, subcircular (in lateral view), with small callus behind ventrocaudal margin, lacking a vertical carina or spine in front of eye. Lateral ocelli conspicuous at anterior ventral margin of eye. Rostrum reaching hind coxae.

In dorsal view, pronotum broad, about 2× broader than vertex along midline (length at midline about 1/3 width), anterior margin arched and carinate, posterior margin nearly (roundly) truncate, surface irregularly rugulose bearing distinct median carina and obscure irregular tracings on disc. Mesonotum triangular, a little longer than pronotum + head at midline, bearing a distinct median carina, with lateral carinae obscure. Tegulae conspicuous. Front and middle legs ( Fig. 4C View FIGURE 4 ) with femora and tibiae expanded and flattened, hind tibia ( Fig. 4A View FIGURE 4 ) with 5–6 lateral teeth, apical ornamentation ( Fig. 11B View FIGURE 11 ) 7(2+5)-(8–9)-(6–9), with teeth at the apex of tibia large and those on the tarsi much smaller in uniform row (lateral teeth largest), interspersed with conspicuous setae. Forewings broad ( Fig. 3A View FIGURE 3 , length about 2.3× width), opaque and relatively short, costal and trailing margins subparallel (costal margin weakly convex, trailing margin inflected slightly at costal apex), apex (at apical branches of RP and MP) broadly rounding, claval apex just distad of midlength; venation dense with numerous secondary veinlets, veinlets in form of pale dashed in apical third, densely reticulate in proximal 2/3; apex of clavus near nodal line (at apical 1/3 of wing); ScP+R forked from MP at basal cell; PCu and A1 fused at about ¾ of clavus length. Hindwings ( Fig. 3B View FIGURE 3 ) broad, maximum breadth near base, apex broadly rounded, trailing margin indentated at claval fold; anal lobe well-developed.

Male terminalia. Pygofer in lateral view ( Fig. 5A View FIGURE 5 ) narrowly trapezoidal (narrower posteriorly), anterior margin concave, caudal margin convex, ventral margin diagonally linear, dorsal margin sinuate; in ventral view, posterior pygofer margin concave (medioventral lobe absent). Gonostyli spatulate ( Fig. 12B View FIGURE 12 ), proximal portion bearing a conspicuous laterally projected hook around basal third; distal portion roughly quadrate; dorsal surface bearing a large dentate process a little distad of midlength, distal dorsal margin truncate then angled to a second truncate portion before reaching rounded dorsoventral margin. Anal tube (lateral view) broad and elongately triangular, dorsal and ventral margins weakly sinuate, distally expanded in proximal ¾, then strongly narrowed into blunt median lobe (apex of anal tube below midpoint); in dorsal view ( Fig. 6E View FIGURE 6 ), apex deeply concave with lateral margins expanded to blunt lobes; epiproct hidden, paraproct elongated, exceeding caudal margin of anal tube.

Plant associations. Rhus copallinum L. ( Anacardiaceae , winged sumac). Also reported (specimen records) Ambrosia trifida L. ( Asteraceae ), and Acer rubrum L. ( Sapindaceae ).

Biology. Poblicia fuliginosa is a univoltine species found on Rhus copallinum . They have five nymphal instars with adults starting to appear in July (in North Carolina). Females lay eggs in rows and cover them with a waxy covering ( Fig. 7A View FIGURE 7 ). Egg masses range in number of eggs (6 to 38) with an average of 25 eggs per mass (n= 40 egg masses; Hagerty 2024). Nymphs ( Fig. 7B View FIGURE 7 ) and adults are messy feeders and often leave seeping wounds on trees, which turn into nodules in subsequent years. Favored tree behavior has been observed with this species, where certain trees will have several individuals while none will be on other nearby trees. Nymphs appear to prefer hiding in foliage in the lower portions of the host plant. This species has been reared has been reared in association with spotted lanternfly biocontrol non-target host suitability studies and we intend to describe details of its biology in a separate publication.

Distribution. USA: AL, AR, GA, IL, KS, LA, MD, MO, MS, NC, OK, SC, TN, TX (except upper Rio Grande region), VA; Mexico: Coahuila ( Fig. 24 View FIGURE 24 ).

Remarks. We have found that the eastern and western populations of Poblicia differ morphologically, biologically, and molecularly and therefore deserve status as separate species. Poblicia thanatophana was described from Nogales, Arizona (Kirkaldy 1907, Fig. 8 View FIGURE 8 ) and is an available name for the western species, so we restore P. thanatophana out of synonymy with P. fuliginosa . The diagnostic differences between P. fuliginosa and P. thanatophana are that the former species is slightly larger, with the abdominal terga red (orange in P. thanatophana , at least laterally) and that P. fuliginosa usually bears two white bands on the front and middle legs ( Fig. 4C View FIGURE 4 , one band in P. thanatophana , Fig. 4F View FIGURE 4 ). The proximal forewing reticulations are more strongly contrasting in P. fuliginosa ( Fig. 3A View FIGURE 3 ) than in P. thanatophana ( Fig. 9A). The terminalia of the two species are quite similar, but can be distinguished by several subtle differences in the shape of the gonostyli ( Fig. 12 View FIGURE 12 ), and that the anal tube in P. fuliginosa ( Fig. 6E View FIGURE 6 ) appears more deeply bilobed apically than P. thanatophana ( Fig. 11E View FIGURE 11 ). Poblicia fuliginosa primarily utilizes winged sumac ( Rhus copallinum L.) as a host species, whereas P. thanatophana is mainly on Asteraceae , in particular desertbroom ( Baccharis sarothroides A. Gray ).

Nast (1951: 270), in his remarks concerning the genus Poiocera De Laporte, 1832 , synonymized Poiocera venosa Walker, 1851 under Crepusia fuliginosa ( Olivier, 1791) (which was a new combination for fuliginosa ). This combination was missed in Bartlett et al. (2014) and is currently followed by Bourgoin (2025) because the issue had not been formally addressed in prior work. Nast (1951) described his reasons for excluding a series of taxa from Poiocera , but not his basis for placing fuliginosa under the South American genus Crepusia Stål, 1866 ( type species Lystra miniacea Germar, 1830 from Brazil; Costa Lima 1935, Metcalf 1947). While early authors seemed uncertain about the diagnosis of Crepusia (e.g., Metcalf 1923, 1938; Ball 1933), the genus ( Fig. 23 View FIGURE 23 ) appears to be robust forms with the apex of the forewing transparent and the base of the hindwings usually colored red or orange (e.g., Porion 1994, figs 36, 42). An underlying issue is that the limits of many of the genera in the Poiocerini deserve reassessment, and many species are doubtfully placed to genus. We feel the placement of fuliginosa in the South American genus Crepusia by Nast (1951) was likely a result of a misunderstanding of the latter genus, and we here reaffirm the traditional placement of fuliginosa under Poblicia as it has been considered by all authorities since Nast (1951).

The type specimen of Poblicia fuliginosa could not be located. The collection of G.A. Olivier is reported to be primarily in Paris, with a portion in Scotland ( Horn & Kahle 1937, Osborn 1952). The location of the type specimen of this species was not reported in Porion (1994).

Records on iNaturalist from Mexico (e.g., observation 16700436 from Coahuila State) appear quite similar to P. fuliginosa and may represent valid records of this species, but P. fuliginosa and P. thanatophana can not always be definitively diagnosed based on the views presented.

Material examined. LOUISIANA: Natchitoches parish, Kisatchie National Forest, 22–23 Sep 1995, M.J. Travitz ( 1 male, UDCC) ; same, Sep 2002, R. M. Krause ( 1 female, UDCC) ; same, 14 Sep 2002, W. Burnside ( 1 male, UDCC) ; Kisatchie Ranger District , 09 Oct 2002, M Kleinhenz ( 1 female, UDCC) ; Tangipahoa parish, Arcola, Sandy Hollow Wildlife Management Area, 7 Sep 2002, S. T. Dash & M. Seymour, Rhus copallina ( 1 male, 1 female, UDCC) ; Same , 7 Sep 2002, P. L. Vasseur ( 1 male, UDCC) . MARYLAND: Dorchester Co.: Linkwood Wildlife Management Area , 31 Aug 1994, M. J. Rothschild, Ambrosia trifida L. ( 1 male, UDCC) . N ORTH CAROLINA: Bladen Co., 2.5km N of Jones Lake Co. Rd. 1509, 17 Sep 1994, G. V. Fodor ( 1 male, NCSU) ; Bladen Lakes St Forest Co. Rd 1509, 6km NE Hwy 242, 7 Sep 1990, H. Zhong ( 1 male, 1 female, NCSU) ; same, 07 Sep 1990, M. A. Wolff ( 1 male, NCSU) ; same, 7 Sep 1990, RJ Kopanic Jr. ( 1 male, 2 female, NCSU) ; Bladen Lakes State Forest , 22–27 Aug 2020, T. J. Hagerty, Rhus copallinum ( 5 males, UDCC; 3 male, 2 female, RBINS) ; Bladen Lakes State Forest , 5.5 km S of Ammon, SR-1508 [Lulu Long Rd] at Turnbull Creek, 8 Sep 2001, M. A. Bertone ( 1 male, NCSU) ; near Bladen Lakes State Forest , 17 Sep 1994, C. R. Bartlett, Acer rubrum L. ( 2 female, NCSU) ; same, 18 Sep 1994, C. R. Bartlett, Rhus copallina ( 1 male, 1 female, NCSU, 1 female UDCC) ; Cumberland Co., Fayetteville , 6 Sep 1945, D. L. Wray ( 1 female, NCSU) ; Durham Co., 20 Km N. of Durham , on Hill Forest, 3 Sep 1999, J. R. Carlson ( 1 female, NCSU) ; same, 3 Sep 1999, C. M. Newman ( 1 female, ( NCSU) ; Durham , 21 Sep 1995, H. A. King, Rhus copallinum ( 1 male, UDCC) ; Moore Co., Southern Pines , 6 Oct 1908 ( 2 female, NCSU) ; same, 7 Oct 1908, A. H. Manee ( 1 female, NCSU) ; same, 26 Jun 1919 ( 4 male, NCSU) ; Southern Pines , Sep 1906, R. Woglum (2 sex undetermined, UCRC) ; same, 27 Sep 1906, R. Woglum (2 sex undetermined, UCRC) ; same, 7 Sep 1908, A. H. Manee (1 sex undetermined, UCRC) ; Pasquotank Co.: Elizabeth City , Late Aug 1919, F. Sherman ( 1 female, NCSU) ; Richmond Co., Hoffman , 26 Sep 1981, R. Webster (1 sex undetermined, UCRC) , Wake Co. , ( 13 specimens, NCSU) . OKLAHOMA: Payne Co., Stillwater , 7 Oct 1993, M. Gates (2 sex undetermined, UCRC) . SOUTH CAROLINA: Oconee Co., Seneca , 3 Sep 1989, Debbie Oliver ( 1 male, UDCC) . TEXAS: Tyler Co.: Tyler County , 7 Oct 1989, C. Bordelon ( 1 female, UDCC) . VIRGINIA: Northampton Co.: Kiptopeke State Park , 3.7km SW Towsend, 4 Oct 2014, S. T. Dash, Rhus copallina ( 1 male, UDCC) .

iNaturalist State Records: ALABAMA: Chilton Co. (137657115), Colbert Co. (97167950). ARKANSAS: Pulaski Co. (59383343), Faulkner Co. (125299876); Cleburne Co. (136038966) .