Santocellus legrandi, Tauber, 2025

Tauber, Catherine A., 2025, Two new species in the green lacewing genus Santocellus (Neuroptera, Chrysopidae, Leucochrysini), ZooKeys 1226, pp. 195-215 : 195-215

Santocellus legrandi sp. nov.

Figs 1 , 2 , 3 a , 4 , 5 , 6 , 8 a, c, e , 10 d , 11 d , 12 d

Etymology of name.

The species carries the surname of Jean Legrand (see Dedication above). Jean had a special love for art (especially painting); it is thus fitting that his name is associated with a beautiful lacewing that has stunning wing markings. The species name is a singular noun in the genitive case following a masculine genus name.

Type specimens.

(sex and figures; verbatim label data [specimen location]) Holotype • Male (Figs 1 – 5 ). – (1) Nicaragua, Rio San Juan / Bartola , 15 m, 12–15.10. 09 / N 10 54 56 / W 84 17 57 / P. Duelli, U. & H. Aspoeck – (2) 11 [ CAS] . Allotype • Female (Fig. 6 ). – (1) Costa Rica, 22 km / N. San Ysidro , / San Jose Prov. / V- 24-85 – (2) J. T. Doyen, / P. A. Opler / Collectors [ SDMC] . Paratype • Female. – (1) Belize: Cayo / Chiquibul Forest. Las Cuevas / Nature Trail / 31 st July 2001 / G. C. McGavin Coll. (2) G. C. McGavin Coll. / Pres. G. C. McGavin / OUM-2002-006 [ OUMNH] . Paratype • Male. – (1) Honduras 1923 / Rio Paulaya: / Barraneo , iv- 17 / 303 T. H. Haldell (2) Phillip A. Adams / Collection / 1998 bequest to / Calif. Acad. Sci. [ CAS] . Paratype • Male. – (1) Chiriqui – (2) prob. Leucochrysa risi Esb-Pet , det. P. Adams ’ 80 [ NHMUK; collection site probably in Panama; abdomen in poor condition; pieces of exoskeleton in attached micro-vial] .

Description.

Adult. Medium-sized. Head (Fig. 1 ): Dorsum, including eyes 1.70 (F) – 1.85 (M) mm wide; ratio of head width to eye width = 1.71 (M) – 1.86 (F): 1; sutures not visible. Vertex raised, with small posterior fold. Antenna long (28 mm, n = 1), ~ 1.5 times length of forewing; scape large, with approximately equal length and width; length of flagellomeres (beyond # 5) ~ 1.6–1.8 times width, with setae in four rings; labrum emarginate. Torulus relatively large, extending slightly below mid-eye level. Frons with short to moderate length, pale setae, lower margin scalloped upward above clypeus (below toruli). Tentorial pits distinct, round, raised. Gena 0.24 mm long; ratio of genal length to distance between tentorial pits = 0.43: 1. Clypeus quadrate, except dorsal margin broadly curved upward below frons, distal margin relatively straight, pale mesally, dorsal surface rounded, slightly raised mesally, with sparse, short to medium length amber setae. Labrum tapering anteriorly, with bilobed frontal margin, distinct cleft between lobes moderately deep, lateral margins tapering, long setae basally, shorter setae distally. Labium with submentum slender, rounded ventrally, with rounded, slightly bulbous tip. Coloration: Markings on vertex, frons dark brown to black with tinge of deep red; broad, scalloped stripe across frons; vertex with large pentagonal mark, cream area within; clypeus with slender stripe across midsection, not reaching margins; labrum with dark reddish brown tinge mesally. Antenna with scapes pale frontally, dark brown dorsally, frontolaterally; pedicel cream with tinge of red distally; basal three flagellomeres dark brown to black frontally, medially; remaining flagellomeres pale. Maxilla with distal three palpomeres dark brown to black, basal two palpomeres pale with black markings laterally. Labium pale, with distal two palpomeres black, basal one pale.

Thorax (Fig. 2 a ): Prothorax wider than long: 0.86–0.93 mm long, 1.11–1.27 mm wide, ratio of length to width = 0.69: 0.78; setae scattered, long to medium length, thin, golden. Legs robust, of moderate length, with dense, short, golden setae; tarsal claws without basal dilation, but with two prongs. Coloration: Pronotum cream mesally, dark brown laterally; anterior brown section broadly quadrate, extending about halfway across dorsal surface, with central section bearing cream spot; posterior section cream mesally, with brown stripe laterally. Mesonotum dark brown laterally, with patches of cream mesally on prescutum, scutum. Scutellum with large cream spot. Metanotum entirely dark brown dorsally. Thoracic pleuron, venter pale to light tan. Legs pale, mostly unmarked, with short to medium length, golden setae, few long, pale setae; terminal segment (base of tarsal claws) smooth, shiny, black dorsally, with pair of small (indistinct) swellings, two pairs of moderately long setae at tip (Fig. 7 e ). Tarsal claw moderately long, narrow distally, with single small spur; base slightly enlarged, without distinct dilation.

Wings (Fig. 3 a ): Measurements: male left wing (n = 1–2); female, right wing (n = 1). Forewing: 17.3–18.7 mm long, 6.2–6.9 mm wide (maximum). Costal area moderately tall; tallest costal cell (# 9, 10) 1.5–1.7 mm tall, 0.23–0.24 times maximum height of wing. First intramedian cell triangular, elongate, 1.5–1.6 mm wide, ~ 0.7–0.8 times width of third median cell. Origin of first radial crossvein ~ 0.5 mm distal to origin of Rs; radial area (between R and Rs) with single row of 13 or 14 closed cells; tallest radial cell 1.1–1.3 mm, 0.77–0.80 times taller than tallest costal crossvein. Three b cells, each full sized, slightly longer than tall; first b cell heavily marked on all sides. Two series of gradates; 8–10 inner gradates, 7 or 8 outer gradates; size of cells bounded by gradates fairly uniform, lengthening then shortening distally. Four to 5 b’ cells; last b’ cell wider than tall. Three intracubital cells, distal one open; first cubital crossvein located slightly basal to first mediocubital crossvein; icu 1, icu 2 each shorter than icu 3; icu 1 + icu 2 considerably longer than icu 3. Vein 1 A forked. Terminal margin with approximately 5 simple veinlets, 13 forked veinlets. Hindwing: 16.7–17.0. mm long, 5.0– 5.1 mm wide. Eleven to 14 radial cells (counted from origin of R, not false origin); t cell absent. Two series of gradates; 5–9 inner gradates, 7 outer gradates. Two to 4 b cells; 4 or 5 b’ cells beyond im 2. Terminal margin with 7 or 8 simple basal veinlets, 9–11 forked basal veinlets. Coloration: Wing surface hyaline, with bold, dark brown markings. Forewing: Stigma with moderately large, dark brown spot basally, second, smaller dark brown spot distally; veins pale except when surrounded by dark marks: e. g., distal part of R and Rs dark brown with surrounding area dark brown; slightly elongate dark brown mark at tip of wing; two dark spots in approximate center of wing; scattered dark brown marks along basal 2 / 3 of lower edge of wing; veins dark when within dark spots. Hindwing: stigma with one large dark brown mark; base of RP, basal b cell dark, within dark brown mark; distal section of cu (upper fork) dark; posterior margin of wing with dark brown section, including suffusion at intersections with marginal veinlets.

Male abdomen (Figs 4 , 5 ). Segments 1–3 not particularly long or slender; distal segments only slightly larger than basal segments (male and female). Tergites shallow, with slightly rounded margins; each tergite with distinctive brown marking having white center spot. Tergites, sclerites, pleural region with numerous, short to medium length setae, microsetae. Abdominal sternites moderately tall (S 6: ratio of length to width = 1.3: 1), with dorsal margins straight, slanting, spiracles not enlarged, without microtholi; S 9 with slightly long, robust setae distally, without microtholi. Callus cerci almost round (0.23– 0.19 mm diameter), with 33–35 long trichobothria; cupuliform bases of variable size [those within central section larger (~ 0.026 mm) than those on periphery (~ 0.014 mm)]. Dorsum of T 9 + ectoproct rounded distally, fused mesally, with very small to no cleft; no midline suture visible; with long setae, densest along distal margin; dorsal apodeme simple, straight, short, extending around proximal side of callus cerci. S 8 + 9 fused without suture, tapering to acute apex (lateral view); S 9 slightly darker than S 8; ventral apodeme extending across dorsal surface of S 8 + 9; distal margin of S 8 + 9 spatulate, with terminal membrane prominent; membrane bearing moderately heavy gonocristae in two broad (somewhat irregular) lateral patches, single, small mesal patch of several (~ 3) gonocristae. Coloration: Dorsum of each segment pale with large black central mark; marks on T 2 – T 8 with prominent white spot in center of black area; mark on T 1 without white spot; venter, pleuron white to cream. Setae golden, long; dorsal half of callus cerci light cream colored; ventral half black.

Male genitalia. Tignum, gonapsis, parameres, quadrate hood, pseudopenis, spinellae absent; S 8 + 9 fused, elongate, not extending beyond T 9 + ectoproct. Gonarcus broad, with very small gonocornua on frontolateral margin of gonarcal bridge, with relatively large, rounded gonarcal arms expanded forward slightly; mediuncus in two sections; basal section extending from below gonarcal bridge, with pair of internal apodemes dorsomesally, with dorsal surface membranous, broad basally, tapering distally; distal section of mediuncus extending perpendicularly from tip of basal section, distally with flange-like lateral arms partially surrounding mesal beak; gonosaccus beneath mediuncus, stiff, flat, laterally with a few gonosetae on chalazae.

Female abdomen (Fig. 6 ; n = 2, both slightly teneral). Callus cerci round, 0.14 mm in diameter, with 36–37 trichobothria of variable length; cupuliform bases also of variable diameter (0.010 –0.029 mm). Dorsum of T 9 + ectoproct rounded posteriorly, tall (lateral view), extending approximately to lateral margin of gonapophysis lateralis. Sternite 7 with dorsal margin relatively straight; distal margin blunt, not particularly heavy, with setae mostly missing. Gonapophyses laterales slender, occupying approximately half of abdominal posterior margin; interior membranous area not greatly expanded or inflatable. T 9 + ectoproct truncate posteriorly, extending slightly below gonapophyses laterales (lateral view). Gonapophyses laterales occupying ~ 60–70 % of posterior margin of abdomen.

Female genitalia. Entire genital structure compact; spermathecal complex, bursal duct, base of subgenitale all contained below bursa copulatrix. Subgenitale broadly based, with distinct quadrate knob distally; distal section of knob bilobed, with broad, flat trough between lobes; lower frontal region of subgenitale with elongate, downward-projecting, snout-like protuberance (knob). Spermatheca round, doughnut shaped, with broad invagination, distal end with relatively large tapering, sclerotized, coiled and curved tubule (velum), with slit along most or full length, apparently opening to bursa; velum ~ 0.9 mm long, ~ 0.14 mm wide at widest point, 0.04 mm wide near middle. Spermatheca doughnut shaped, 0.38 mm in diameter, 0.16 mm deep, with invagination in center, 0.19 mm in diameter, 0.11 mm deep. Spermathecal duct very faint. Bursa copulatrix folded, pleated, small, barely covering dorsum of spermatheca, with distal end tapered, attached directly to sclerotized spermathecal velum, without bursal duct; bursal glands not found. Colleterial complex, transverse sclerite not discerned.

Variation.

Unknown.

Known distribution.

Central America. Belize (Cayo), Costa Rica (San Jose), Honduras (Colón, prob.), Nicaragua (Rio San Juan), Panama (Chiriquí).

Figures

Figure 1. Head of Santocellus legrandi, sp. nov. Holotype, male, Nicaragua, Río San Juan, Bartola (CAS) a frontal b head, prothorax, dorsal c head, frontolateral d ventral. Width across head (dorsal, including eyes), 1.85 mm.

Figure 2. External features of Santocellus legrandi, sp. nov. Holotype, male, Nicaragua, Río San Juan, Bartola (CAS) a head, thorax, dorsal b body, lateral c abdomen, dorsal d terminus, lateral e protarsal claw, lateral. Width across head (including eyes), 1.85 mm; diameter of callus cerci, ~ 0.2 mm.

Figure 3. Wings of two new Santocellus species a S. legrandi sp. nov., holotype, male, Nicaragua, Río San Juan, Bartola (CAS); forewing length, 18.7 mm b S. helene, sp. nov., holotype, male, Bolivia, Cochabamba (CAS); forewing length, 19.4 mm. Arrows highlight interspecific differences in markings.

Figure 4. Male abdomen of Santocellus legrandi, sp. nov., holotype, Nicaragua, Río San Juan, Bartola (CAS) a dorsum of abdominal segments A 2 - terminus b lateral view of abdominal segments A 6 - terminus c dorsum of abdominal segments A 7 to terminus with gonarcus everted d lateral view of abdominal terminus. Length of T 8 along dorsal margin, 0.68 mm. Abbreviations: c. c. callus cerci; d. ap. dorsal apodeme of T 9 + ectoproct; gcr field of gonocristae on terminal membrane of S 9; gs gonarcus; S 8 + 9 fused eighth and ninth sternites; T 3, T 7, T 8 third, seventh, and eighth tergites; T 9 + e fused ninth tergite and ectoproct; v. ap. ventral apodeme of S 8 + 9.

Figure 5. Male genitalia of Santocellus legrandi sp. nov., holotype, Nicaragua, Río San Juan, Bartola (CAS) a gonarcal complex, dorsal b same, lateral c same, frontal d same, posteroventral. Abbreviations: gcr, field of gonocristae on terminal membrane of ninth sternite; gsac gonosaccus; gst gonosetae; g. ap. gonarcal apodeme; mu-d mediuncus, dorsal view; mu-f mediuncus.

Figure 6. Cleared abdominal terminus of female Santocellus legrandi sp. nov., allotype, Costa Rica, San José (SDMC) a segments A 7 to terminus, lateral b genitalia, lateral c subgenitale, distal view d spermathecal complex below bursa copulatrix, lateral view. Abbreviations: b. c. bursa copulatrix; c. c. callus cerci; g. l. gonapophysis lateralis; k-sg distal knob of subgenitale; sg-b base of subgenitale; sp spermatheca; sp. d. spermathecal duct; sp. i. spermathecal invagination; S 7 seventh sternite; T 8 eighth tergite; vel, spermathecal velum.

Figure 7. Head and thoracic region of Santocellus helene sp. nov., holotype, male, Bolivia, Cochabamba, Alto Palmar (CAS) a upper body (head, thorax) dorsal b head, prothorax, lateral c head, frontal d head, ventral e tarsus (dorsum) and tarsal claw (lateral). Width across head (including eyes) 1.82 mm.

Figure 8. Side-by-side comparisons of male external abdominal features a, c, e Santocellus legrandi sp. nov., holotype, Nicaragua, Río San Juan, Bartola (CAS) b, d, f Santocellus helene sp. nov., holotype, Bolivia, Cochabamba, Alto Palmar (CAS) a, b abdominal segments A 6 - terminus, lateral c, d terminus of S 8 + 9, with terminal membrane bearing distinctive patches of gonocristae e, f callus cerci.

Figure 10. Side-by-side comparison of head and body (dorsum) of currently known Santocellus species a S. atlanticis, male; Brasil, Rio Grande do Sul b S. riodoce, female; Brasil, Espírito Santo c S. risi, male; Madre de Dios, Peru d S. legrandi sp. nov., male; Nicaragua, Río San Juan, Bartola e S. helene sp. nov., male; Bolivia, Cochabamba f sketch of thoracic sclerites (from Séméria 1976).

Figure 11. Side-by-side comparison of the head (frontal) of currently known Santocellus species a S. atlanticis, male; Brasil, Rio Grande do Sul b S. riodoce, female; Brasil, Espírito Santo c S. risi, male; Peru, Madre de Dios d S. legrandi sp. nov., male; Nicaragua, Río San Juan, Bartola e S. helene sp. nov., male; Bolivia, Cochabamba.

Figure 12. Side-by-side comparison of the wings of currently known Santocellus species a S. atlanticis, male; Brasil, Rio Grande do Sul b S. riodoce female; Brasil, Espírito Santo c S. risi, male; Peru, Madre de Dios d S. legrandi sp. nov., male; Nicaragua, Río San Juan, Bartola e Santocellus helene sp. nov., male; Bolivia, Cochabamba. The numbers on the right of each image indicate the maximum length of the forewing recorded for that species.