Scaralina marmorata ( Spinola, 1839 ) Yanega & Goemans & Dam & Gómez-Marco & Hoddle, 2024

Yanega, Douglas, Goemans, Geert, Dam, Matthew Van, Gómez-Marco, Francesc & Hoddle, Mark, 2024, Description of a new genus of North and Central American planthoppers (Hemiptera: Fulgoridae) with fourteen new species, Zootaxa 5443 (1), pp. 1-53 : 18-24

Scaralina marmorata ( Spinola, 1839) , comb. nov.

( Figs 1 , 16 , 33 , 48 , 63 , 76 , 85 )

= Crepusia glauca Metcalf, 1923 (synonymy by Ball, 1933, here restored); for other combinations, see Metcalf, 1947.

Note on synonymy. The three related taxa that co-occur in Arizona ( metcalfi , cristata , and aethrinsula ) are readily distinguished from one another and from marmorata , including forms at the extremes of variability in color and maculation, by even a quick glance at the thorax and forewings. Given the large number of specimens of all three taxa in collections ( metcalfi is relatively rare, though the most similar to marmorata ), it is fairly surprising that no one had recognized that the variants of “ Alphina glauca ” fell into three or four consistent and discrete morphotypes, including (and especially) in areas where they co-occurred, and where one putative “variant” lacked a female supra-anal plate; nonetheless, this problem has persisted for most of a century, with the additional complication that Metcalf’s 1947 catalogue reversed Ball’s synonymy of Crepusia glauca Metcalf with Calyptoproctus marmoratus Spinola , which led to the almost complete abandonment of the latter name by taxonomists after 1947. Significantly, among specimens bearing Metcalf’s personal ID labels, we were able to find a number of specimens labeled as “ Crepusia glauca ”, all from Arizona (and all, thus far, of S. cristata ), as well as several specimens of a true Calyptoproctus , from Mexico and Guyana, that he had labeled as Ca. marmoratus ; that is, he mistakenly believed that Spinola’s species was a true Calyptoproctus , because he had evidently never seen the type.

In the absence of Metcalf’s holotype, and given that there are no specimens we have seen from any populations in habitat consistent with southern Texas that could be interpreted as belonging to a second, distinct species, and given that there is nothing in Metcalf’s description which differs from marmorata as we understand it, we will follow Ball’s synonymy (based, as it presumably was, on direct examination of Metcalf’s now lost specimen) as evidence that Metcalf’s holotype was indeed a specimen of Calyptoproctus marmoratus Spinola. We further note that Spinola’s type is a perfect match to the populations from southeastern Texas, even more so than populations of orientalis from Nuevo León and Tamaulipas, which are somewhat similar. We consider this the final compelling point for designating a Texan specimen as the neotype of Crepusia glauca Metcalf , in order to firmly establish its identity and synonymy, fulfilling all necessary requirements under the International Code of Zoological Nomenclature (Article 75.3; ICZN, 1999). Although Metcalf’s type locality was Brownsville (Cameron Co., TX), we have not seen any U.S. specimens from within 300 kilometers of Brownsville, though we have seen Mexican material (of orientalis ) that is marginally within this range (over 250 km). It does, in fact, seem improbable that specimens closer to the original type locality could be located: within Texas, this species seems restricted to the woodlands of east Texas, west to the wetter parts of the eastern Edwards Plateau, a habitat type which is not present near Brownsville. However, we consider it important to make Metcalf’s taxon an objective synonym of Spinola’s (by assigning it a neotype that clearly belongs to Spinola’s taxon) rather than to assign it a neotype belonging to a different taxon from northeastern Mexico simply to preserve Metcalf’s name, in large part because it would be very likely to cause people to assume that the former “ Alphina glauca ” and a reinstated “ Scaralina glauca ” are the same taxon, when the former name has been almost universally applied to multiple species from Arizona. We further feel that Spinola’s type is so readily and unmistakably associated with the southeastern Texas populations (e.g., Bastrop, Brazoria, Brazos, Leon, Polk, and Travis counties; compare Fig. 1 to Fig. 16 ) that it is unnecessary to replace it with a specimen whose provenance is more explicitly known; that is, a neotype designation for S. marmorata is not justifiable under the ICZN.

Diagnosis. S. marmorata is the most widespread species in the genus, and it shows considerable, though highly localized, variation in color. Within any given population, individuals are fairly uniform in appearance, but over fairly short geographic distances the color patterns can change considerably between populations that are structurally indistinguishable. We consider the structural features to be taxonomically reliable, especially the very short notal pubescence, and male genitalia (though these too show slight population-level variation). Molecular analysis may ultimately reveal that this is a complex of species, each inhabiting relatively discrete “islands” of habitat and/or host plants; nonetheless, the structural similarities are too great to confidently exclude most of these populations from the circumscription of the species as a whole. The male gonostyli do not appear to vary significantly within the SE U.S. populations, nor does the short notal pubescence, and these are perhaps the most reliable diagnostic features for recognizing marmorata . The remaining taxa appear to be quite distinct from marmorata , and while there is ample tradition and precedent to designate subspecies for geographically isolated populations, as most of these taxa appear to be, we refrain from doing so here, and treat them all as full species. All of the other taxa described here have long notal pubescence, especially the other members of the marmorata species group ( Fig. 3 ). The most similar taxa are the geographically proximate orientalis and rileyi , which share an impressed frons, and very similar male gonostyli; the primary difference is the long versus short notal setae, and this may be at best a subspecific feature, though the male genitalia also differ slightly. It is probably not a coincidence that marmorata is the only species routinely found at low elevations; all but three of the hundreds of confirmed records for species other than marmorata are over 1200m. We suspect that the reduction of notal pubescence is unique (within Scaralina ) to marmorata , and associated with living at low elevations. The other species in the marmorata species group all possess long setae on the dorsal thorax, slightly longer than most members of the cristata species group.

Description. Head ( Figs 33 , 48 ). Rim of vertex dark anteriorly and laterally, with three prominent pale marks anteriorly; supra-ocular lobes pale apically; posterior rim mostly pale, except darkened adjacent to supra-ocular lobes; dark spots of vertex posterior in position; black markings very well-defined and restricted (generally only lateral bands and anterior spots), no tiny spots as in preceding species. Frons with a broad pale upper reflexed margin, very dark just below this (pale markings at corner of frontal crease near ocellus generally well-defined as in aethrinsula ), and the ground color below a little less dark, except for darker spots surrounding setal bases; lower frons with surface very finely wrinkled, and with two low but distinct converging ridges forming a “V”, surface somewhat impressed; sometimes a trace of a third, medial, ridge just below reflexed margin. Lateral margin of frons nearly straight, lower lobes of frons not distinctly expanded. Clypeus quite variable, typically palest in center and apically. Rostrum typical for genus, or slightly darker.

Thorax ( Fig. 48 ). Pronotal spots well-defined, including an extra set of small spots behind the usual post-ocular spots (sometimes confluent); at least anterior portion of dorsal pronotal face pale, becoming darker towards posterior and center, medial carina entirely pale; may be more or less extensively dark over surface in different populations. Mesonotum center typically mostly pale (mesonotum also variably dark-marked); medial carina prominent; lateral carinae angled inward anteriorly and very weakly sinuate posteriorly; inner posterior carinae weak, granular dark spots large; posterior mesonotal lobe somewhat acutely rounded, and usually pale medially and dark laterally. Setae of dorsal pronotum shorter than interspaces between them, and slightly recumbent, but still visible in oblique view; Legs as in generic diagnosis, but pro- and metatarsi with light dorsal mark on third tarsomere sometimes nearly absent.

Wings ( Figs 16 , 63 ). Forewings with dark markings on basal half of 1st claval vein distinctly interrupted; other markings quite variable, but most commonly with a well-developed, continuous dark transverse patch from costa to clavus (with pale crossveins), at the level of the first branch of M, and translucent to opaque pale olivaceous, ochre, or orange basal markings (the most variable wing color feature among populations), often with extremely tiny red flecks along the venules; a few nearly opaque dark spots near the end of the claval vein; postcostal cell extensively black, with pale spots and at least 3 distinct, wide pale bands; remainder of wing mostly clear with variable, irregular dark but translucent spotting. Costal crossveins fairly numerous, mostly straight or slightly arcuate but oblique, often obscure basally, or crossveins not reaching costa; M with 4–5 main branches, crossveins between them mostly straight, well-spaced, making square to rectangular cells; at first branch of M, M is closer to CuA than to Sc; CuA with 3–4 branches, rarely more than 1 of these coming directly off of CuA, and the first branch is typically near or slightly beyond the claval vein juncture; greatest distance between Pcu and A 1 veins slightly exceeds greatest distance between A 1 and wing margin; fused vein posterior to juncture relatively short. Hind wing hyaline except base, which is orange-infused.

Abdomen ( Fig. 16 ) Posterolateral pale markings varying in size, grayish-yellowish, sometimes fairly small, commonly with pale markings medially on terga 2 and 3, and a narrow pale midline. Female supra-anal plate large, typically dark medially. Pleurites either entirely black or with pale posterior margin. Sternites typically pale tan except laterally and/or basally black.

Male terminalia ( Figs 76 , 85 ). Dorsal setose bulge weakly convex, setae generally reddish, short, and fairly dense. Dorsal margin curves inward at bulge, forming rounded right angle, and there is usually a low oblique ridge running from the setose bulge to the apex. Lateral hooks small, acute, tapering.

Type material.

Crepusia glauca Metcalf, 1923 .

Neotype, male (here designated): “ TEXAS: Brazos Co. ”, College Station , Lick Ck. Pk., VI-5-1996, E.G. Riley-353, UV” “NEOTYPE Crepusia glauca Metcalf , det. D. Yanega 2022” ( TAMU).

Calyptoproctus marmoratus Spinola, 1839

Holotype, male (photo only): “ United States ” ( NMW) ( Fig. 1 ).

Other material examined. UNITED STATES: ARKANSAS: Bradley Co.: Moro Bay State Park , 5.vi.1988, D.C. Hawks ( UCRC ENT 54885 ) ; FLORIDA: Clay Co.: Mike Roess Gold Head State Park Campground, 27 mi NE Gainesville, 29°50’N, 81°57’W, 18.vii.1995, J.D. Oswald & L.A. Stange, “ex. Turkey oak scrub,” 1M ( TAMU) ; Levy Co.: Manatee Springs State Park , 21.vii.1973, 1F ( AMNH) ; GEORGIA: Charlton Co.: Folkston, Trader Hills Rec. Area , N 30 46.74’, W 82 01.61’, 22.v.2004, N.H. Nazdrowicz, Hg vapor lamp, 1M; same but 15.vi.2004, 1M ( UDCC) ; Whitfield Co.: Dalton , 31.viii.1990, J.K. Adams, 1F ( UCRC ENT 104993 ) ; LOUISIANA: Rapides Par.: Kisatchie National Forest , 1.viii.1970, C.W. Griffin, UV light, 1F ( TAMU) ; St. Tammany Par.: 4.2 mi NE Abita Springs , 30°30.986’N, 89°57.276’W, T6S R12E sec. 24, V.A. Brou, 1M ( TAMU) ; NORTH CAROLINA: Brunswick Co.: Bald Head Island, 47 Fort Holmes Tr. , 2.vi.2005, N.H. Nazdrowicz, Hg vapor lamp, 1M ( UDCC) ; SOUTH CAROLINA: Oconee Co.: Oconee State Park , 10.vi.1991, E.G. Riley, 1M, 1F ( TAMU) ; TENNESSEE: Cooke Co.: Cosby Ranger Station, Great Smoky Mountains National Park , 1800 ft, 35°46’21”N, 83°12’49”W, 5–8.vii.2004, E.G. Riley, UV, 2F ( TAMU) ; TEXAS: Angelina Co. : 17.vi.1993, T, Bishop, 1F ( UCRC ENT 156920 ) ; Bastrop Co.: “ Stengl Ranch ,” 17.vi.1993, J. Gillaspy, 1M ( UTIC) ; “ Camp Swift. Nat. Gd. ,” 8.6 km N Bastrop, 25.v–28.vii.2009, J.C. Abbott, 1M ( UTIC) ; Bexar Co.: Friedrich City Park, San Antonio , 11.v.1986, S. Hanselmann, 1M ( TAMU) ; Judson Nature Trail, San Antonio , 23.vi.1987, J. & S. Hanselmann, “black lght,” 1M ( TAMU) ; Brazoria Co.: 4 mi S West Columbia , 11–15.vii.1976, 1F ( UTIC) ; Brazos Co.: Bryan , 5.vi.1988, E.G. Riley, “mercury vapor & blacklight,” 1F; same but 8.viii.1988, 1F ( TAMU) ; College Station, Lick Creek Park, 24.viii.1997, E.G. Riley, UV, 2F; same but 13.vii.1997, 1M; same but 27.iv.1996, 1M ( TAMU) ; 9 km SSE College Station, 15889 Woodlake Drive , 30°32”N, 96°17”W, 26–27.vii.1997, J. Oswald, UV, 1M; same but 3.viii.1997, 1F; same but 20.viii.1997, 1F; same but 24.viii.1997, 1F ( TAMU) ; Houston Co.: Ratliff Lake Recreational Area , 11.vii.1996, W.F. Chamberlain, “at light,” 1M, 1F ( TAMU) ; Kerr Co.: Kerrville , 31.vii.1983, W.F. Chamberlain, “at light,” 1F ( TAMU) ; Leon Co ,: 5 mi N Flynn, 24.v.1994, E. Riley, UV light, 1M ( TAMU) ; Nacogdoches Co.: Nacogdoches, Park St. , 8.viii.1990, W. Godwin, black light, 1F ( TAMU) ; Polk Co.: Big Sandy Creek at FM 1276, 3 mi E Segno, 14.vii.1994, J.C. Abbott #204, J.W. Chirhart, M. Passanante, UV light, 1M ( UTIC) ; Travis Co.: Austin , “Brackenridge Field Lab,” 170m, 21.vi.1992, J.E. Gillaspy, 1M; same, but 30.vii.1992, 1M ( UTIC) ; Tyler Co.: Kirby State Forest , 30°34’30”N, 94°25’03”W, 19.v–8.vi.2003, E. Riley, Lindgren funnel trap, 1M; same but 20.vii–24.vii.2003, 2F ( TAMU) Wharton Co.: Mackay , 23.v.1984, Marlin Rice, black light, 1M ( UDCC) . There are also numerous images of this species online, such as iNaturalist, showing specimens from as far north as Kentucky and Virginia at the eastern end of the range, and Oklahoma at the western end.

Distribution: This species is found from southeast and central Texas (Bexar and Kerr counties) up to Arkansas and across to Fauquier county in Virginia, and all areas south and east of this, in areas where oaks are found.

Figures

FIGURE 1. Holotype male of Calyptoproctus marmoratus Spinola (courtesy Dr. Harald Schillhammer, NMW).

FIGURE 3. Mesonotum of Scaralina rileyi showing long dorsal pubescence.

FIGURES 13–20. Habitus photos of Scaralina species. (13) S. aethrinsula (male); (14) S. chapina (male); (15) S. cristata (male); (16) S. marmorata (female); (17) S. durango (type female); (18) S. gigantea (type female); (19) S. hawksi (female, crushed somewhat); (20) S. metcalfi (male).

FIGURES 27–35. Faces of Scaralina species. (27) S. aethrinsula; (28) S. chapina; (29) S. cristata; (30) S. durango; (31) S. gigantea; (32) S. hawksi; (33) S. marmorata; (34) S. metcalfi; (35) S. monzoni.

FIGURES 42–50. Heads and nota of Scaralina species. (42) S. aethrinsula; (43) S. chapina; (44) S. cristata; (45) S. durango; (46) S. gigantea; (47) S. hawksi; (48) S. marmorata; (49) S. metcalfi; (50) S. monzoni.

FIGURES 63–68. Forewings of Scaralina species. (63) S. marmorata; (64) S. metcalfi; (65) S. monzoni; (66) S. obfusca (male); (67) S. obrienae; (68) S. orientalis.

FIGURES 73–81. Male genitalia of Scaralina species, dorsal view, anal tube removed. (73) S. aethrinsula; (74) S. chapina; (75) S. cristata; (76) S. marmorata; (77) S. metcalfi; (78) S. monzoni; (79) S. obrienae; (80) S. orientalis; (81) S. rileyi. NOTE: obfusca, sullivani not included.

FIGURES 82–90. Male genitalia of Scaralina species, lateral view. (82) S. aethrinsula; (83) S. chapina; (84) S. cristata; (85) S. marmorata; (86) S. metcalfi; (87) S. monzoni; (88) S. obrienae; (89) S. orientalis; (90) S. rileyi.