SCOPULINI DUPONCHEL, 1845

Sihvonen, Pasi, 2005, Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae), Zoological Journal of the Linnean Society 143 (4), pp. 473-530 : 499-501

publication ID	https://doi.org/ 10.1111/j.1096-3642.2005.00153.x
persistent identifier	https://treatment.plazi.org/id/03E78792-E13D-2F69-D511-8C8DFC0AFF77
treatment provided by	Diego
scientific name	SCOPULINI DUPONCHEL, 1845
status

Treatment

Definition of the tribe is based on a cladistic analysis of the Sterrhinae by Sihvonen & Kaila (2004), where two synapomorphies were found: the signum of the female genitalia being ovoid, granulate, with spines pointing away from the centre ( Fig. 130 View Figures 118–132 ), and the absence of a medial ridge on the epinotum of the male metathorax (character state ‘present’ is figured in Sihvonen & Kaila, 2004). In that analysis neither of these characters was considered to be unique to the tribe, see Results. In the present study it was found that they were unique. In addition, 20 other characters, 18 of which are unique, are proposed as possible tribal synapomorphies ( Table 3). The male 8th tergite with the posterior margin concave with two round lateral lobes (88: 1, Figs 118, 122 View Figures 118–132 ), may be a true synapomorphic character. Also characteristic are the absence of uncus and gnathos, and the presence of socii.

Wings

Wingspan c. 11–70 mm, usually 25-40 mm. Shape usually triangular, forewing apex elongated or termen occasionally rounded; hindwing margin often concave between vein ends or elongated at M 3. Facies very variable; many are straw-coloured, with numerous wavy transverse lines running from forewing to hindwing, and with small, dark discal spots (e.g. Figs 8, 21 View Figures 1–25 ); ocellate discal markings and various other fasciae on white or grey background are also common; iridescent scales present in many species; forewings with fine striae and wide, dark medial band are not unusual; underside often fuscous, above patterns weakly repeated underside; underside ground colour uniform ochreous in some; transverse lines variably developed, usually all present, occasionally absent; sub- and adterminal shaded lines usually present, occasionally pronounced; terminal line continuous, discontinuous or absent; fringes unicolorous or chequered. Forewings with one or two areoles; R 5 from proximal or distal areole, either free or stalked with R 2 -R 4 or stalked with M 1. Hindwing Sc + R 1 and Rs fused shortly or Sc and Rs separate but connected via R 1.

Head

Labial palpus upturned, 3-segmented, organ of vom Rath distinct. Pilifer present, rarely absent. Proboscis fully developed; variably shaped sensilla styloconica at distal end, absent in many species. Frons convex. Two openings or extensions on anterior part of dorsal sclerite between antennae either present or absent. Compound eye round to elliptic; proximal pigment layer circular or semicircular. Scales between antennae flat or flat and hair-like. Male antenna usually fasciculate, occasionally bipectinate to two-thirds; if bipectinate, proximal rami sometimes elongated; sensilla in single or multiple rows. Female antenna usually fasciculate, simple, occasionally sensilla arranged in single or even multiple rows. Antennal segments dorsally uni- or bicolorous. Chaetosemata usually from naked surface, occasionally from scaled surface.

Thorax

Foreleg with well-developed epiphysis; tibial spurs absent; thick spines on tarsus usually absent. Midleg with two tibial spurs, symmetrical or asymmetrical; tarsus normally developed, 5-segmented, thick spines usually absent, sometimes present. Male hindleg tibia often with outer and/or inner hair pencil proximally ( Fig. 113 View Figures 109–117 ), or absent; distal spurs two, one or absent; reduction in tarsus length common; all or part of tarsus segments may be fused; yet usually fully developed, 5-segmented; thick spines usually absent; two apical spines present in most species; usually two claws, symmetrical or asymmetrical, occasionally one or absent; arolium and pulvillus usually present. Female hindleg tibia usually with 2 + 2 spurs, occasionally with one medial spur or absent altogether; hindleg tarsus normally developed. Anterior and posterior ventral lamina of male metafurca usually present, occasionally absent; margins of metafurca basally opened or straight ( Fig. 110 View Figures 109–117 ); pouch on posterior margin of metathorax coxa usually present ( Fig. 108 View Figures 100–108 ), semiround, occasionally pronounced; medial ridge absent on epinotum of male metathorax dorsal sclerite; lateral processes of male coxa elongated in many species, variable in size and shape, rarely absent; absent in females.

Abdomen

Male 2nd sternite usually with distinct, round pouch ( Fig. 116 View Figures 109–117 ) between tympanal organs, posterior margin often invaginated, occasionally with two chambers, surface smooth, with fine ornaments or stout setae; anterolateral extensions present or absent. Male 6th segment occasionally with setae. Membranous sac between male 7th and 8th sternite usually present ( Fig. 66 View Figures 55–79 ). Male 8th sternite very variable; anterior margin convex, concave, tri- or bifurcate; cerata and mappa present in many species ( Figs 62, 76 View Figures 55–79 ); posterior margin membranous or sclerotized; posterolateral appendices occasionally present. Male 8th tergite variable; posterior margin usually concave, with two round lateral lobes ( Figs 118, 122 View Figures 118–132 ). Female abdomen simple, without modifications.

Male genitalia

Genitalia capsule appearing ovoid when viewed ventrally, characterized by enlarged vinculum and usually short, separate valvula and sacculus of valva. Uncus and gnathos absent. Socii usually setose, separate; occasionally fused at apex, lying ventrally or dorsally of tuba analis. Ventral margin of tegumen straight or with various processes, sometimes with distinct teeth. Valvula and sacculus of valva usually separate (e.g. Fig. 41 View Figures 30–54 ), fused in a few cases; sacculus lying ventrally of valvula; both processes often asymmetrical; valvula curved ventrally, weakly sclerotized, setose, sometimes strongly sclerotized, acute; sacculus wide at base, tapering towards apex, blunt or acute; lateral and ventral extensions in many species. Coremata present ( Fig. 123 View Figures 118–132 ) at lateral end of transtilla, often pronounced or even bent ventrally; lost easily, but surface of genitalia distinctly granulate. Dorsal part of juxta tunnel-shaped; ventral part often with wing-like processes ( Fig. 52 View Figures 30–54 ), often asymmetrical. Transtilla usually present, occasionally anterior margins of tegumen fused replacing it. Vinculum enlarged , often with posteriorly directed lobe on top ( Fig. 49 View Figures 30–54 ). Aedeagus shape and size variable, often with dorsal lobes or carina at apex; caecum small or moderate; outer membrane of aedeagus expanded to semiround sac covering proximal end of ductus ejaculatorius; vesica very variable; simple sac with moderate diverticula or very complex with numerous coiled, twisted diverticula; plate-shaped sclerotization at proximal end of ductus ejaculatorius usually present ( Fig. 126 View Figures 118–132 ); otherwise variably sclerotized, occasionally with distinct cornuti. Spermatophore elongated sac, size and shape variable; occasionally frenum distinct, curved structure.

Note: special care is needed to interpret homologies of the male genital structures. Socius, and especially valvula, sacculus and juxta often replace each other functionally. For example, the left arm of the juxta in Somatina anthophilata Guenée has functionally replaced the left sacculus of the valva, the latter being diminished in size.

Female genitalia

Papillae anales longitudinally grooved, setose, sometimes bilobed into rounded dorsal and shorter, narrower ventral part. Apophyses posteriores usually longer than apophyses anteriores. Lamella postvaginalis pronounced in some species, usually weakly developed or absent. Ostium bursae cup-shaped, a complex of structures, occasionally simple or virtually absent. Lamella antevaginalis usually flap-like ( Figs 99 View Figures 80–99 , 132 View Figures 118–132 ), flexible or undeveloped in those species that have a well-developed lamella postvaginalis. Ductus bursae membranous or variably sclerotized; ductus seminalis arises from ductus bursae, corpus bursae neck or corpus bursae. Corpus bursae sac-like, occasionally heavily folded; signum an ovoid field of separate spines ( Fig. 130 View Figures 118–132 ), pointing away from centre, either fused or absent along medial axis in some species; spines on inner surface of corpus bursae common.

Distribution and species diversity

Virtually cosmopolitan with 912 described species and many more undescribed.

Immature stages

Due to the shortage of material, it was not possible to include immature stages in the cladistic analysis. Patocka (2003: 270), who worked with Central European fauna only, was able to find only quantitative delimiting pupal features for Scopulini . He diagnosed the pupa as follows: the ‘front leg verges on antennae as far as the compound eye extends’. In the Rhodostrophini , by contrast, the ‘front leg verges on antennae beyond the compound eye’. In addition, cremaster setae D2 of Scopulini are closer to each other than in Rhodostrophiini .

The Glossotrophia species group of Scopula View in CoL has the unique condition of a free and loped elongated proboscis extending beyond the distal part of the pupa ( Hausmann, 2001; Patocka, 2003). Chaetotaxy and pupal characters are described under generic concepts, where data are available. The narrow larvae of many species rest in a stick-like posture at an angle of 45 ∞. Some specialization in larval host-plants can be seen: Oleaceae View in CoL in Problepsis and View in CoL Rubiaceae View in CoL in some Somatina View in CoL . However, frequent polyphagy in Scopula View in CoL and the worldwide distribution of the tribe make generalizations difficult.

Biology

Species can be found in forested and open habitats. A few Scopula species are known to be of minor pest status (see Introduction).

Comments on systematics

In total, 22 characters were found to support the monophyly of Scopulini ( Fig. 134 View Figure 134 , Table 3). This is partially artificial since only two taxa from the putative sister group, Sterrhini , were used as outgroup. It is known from the cladistic analysis of the tribes of Sterrhinae ( Sihvonen & Kaila, 2004) that many of the characters that appear here as synapomorphies supporting the monophyly of Scopulini would not do so if more outgroup taxa were present. Scopulini also includes Aletini ( Hampson, 1918) and Problepsini ( Wiltshire 1990) as indicated by Holloway (1996, 1997). Both of these tribes share with Scopulini their characteristic structural features, including those of male 8th sternite, male and female genitalia and wing venation.