Simopone

Simopone seem to be rare, arboreal ants (with workers occasionally found foraging on ground or in rotten wood) and presumably are nocturnal (Bolton, 1973a; Brown, 1975; Kutter, 1977; Bolton & Fisher, 2012; Hita Garcia, Wiesel and Fischer, 2013). Knowledge about the natural history is very limited but from some species it is known that they are specialised predators of other ants (Brown, 1975; Bolton & Fisher, 2012).

Identification
Borowiec (2016) - Worker Workers of Simopone are unique among all dorylines in the combination of 11-segmented antennae, eyes and ocelli present, no spur on mid tibia, and the lack of metatibial gland. Simopone also possess a conspicuous groove on the interior surface of hind basitarsus. Other dorylines lacking mid tibial spur include certain species of Lioponera, all Tanipone, and Vicinopone. All these genera have 12-segmented antennae.

Male The males are easily identified by a combination of antennal sockets partially concealed by the torulo-posttorular complex in full-face view, 12-segmented antennae, presence of notauli, and lack of spurs on middle tibiae. The only other non-army ant doryline genus that lacks spurs on middle tibiae is Tanipone, although it is possible that Vicinopone males will turn out to lack them also, when discovered. All Tanipone males known thus far have fully exposed antennal sockets, 13-segmented antennae, and no notauli, in addition to characteristically long maxillary palps that reach the occipital foramen.

Species Groups
Simopone species groups

Distribution
Borowiec (2016) - Simopone is limited to the Old World. Most occur in Madagascar and in the Afrotropics but five rare species have been recorded from the Indomalayan Region (China, Thailand, Vietnam, Singapore, Philippines) and New Guinea.

Species by Region
Number of species within biogeographic regions, along with the total number of species for each region.

Biology
Borowiec (2016) - Simopone is limited to the Old World and currently contains 39 named species. Most occur in Madagascar and in the Afrotropics but five rare species have been recorded from the Indomalayan Region (China, Thailand, Vietnam, Singapore, Philippines) and New Guinea.

Borowiec (2016) - Simopone is a genus of arboreal predators of other ants, found in the Old World tropics. Despite relatively high species diversity very little is known about the biology of Simopone, although several species have been recorded nesting arboreally (Brown 1975, Bolton and Fisher 2012). One species, the Madagascan Simopone sicaria, was observed during a raid. The ants took the brood of arboreal Terataner alluaudi as prey (Bolton and Fisher 2012). Brood production appears not to be synchronized (author’s observations).

Castes
Bolton & Fisher (2012) described queens as 'worker-like', however this is misleading because they refer to total body size, not the thorax which is clearly that of a flying queen. A naive reader can conclude that Simopone queens are ergatoid, although this is not the case in Africa. In Madagascar however, no dealate queens have been found, and only ergatoid queens may exist.

Nomenclature

 *  SIMOPONE [Cerapachyinae: Cerapachyini]
 * Simopone Forel, 1891b: 139. Type-species: Simopone grandidieri, by monotypy.
 * Simopone subgenus of Cerapachys: Forel, 1892l: 243.
 * Simopone revived status as genus: Dalla Torre, 1893: 17.

Worker
Bolton and Fisher (2012) - Shared characters of the genera Simopone, Vicinopone and Tanipone - As well as the 20 features that are exhibited by all members of Cerapachyini these three genera share the following suite of seven characters in the worker caste. None of the characters is claimed as synapomorphic for the three, or for any two of the three.

1 Pretarsal claws with a single preapical tooth, at least on the metatarsus.

2 Mesotibial spurs absent (at most a setiform vestige may remain that cannot be distinguished by light microscopy from other setae at the tibial apex).

3 Metatibial spur single, pectinate.

4 Eyes present and conspicuous, always large (EL/HW 0.30–0.53).

5 Apical antennomere subcylindrical, not inflated and bulbous.

6 Ventrolateral margin of head without a continuous longitudinal carina that commences close to anterior margin below the mandible and extends the entire length of the head to the posterior margin (a carina is present posterolaterally that usually extends onto the ventral surface).

7 Frontal lobes widely separated by the relatively broad clypeus; frontal lobes not or only slightly elevated laterally on each side of the clypeus (never closely approximated and vertical).

Predominantly arboreal cerapachyine ants (workers occasionally found foraging on ground or in rotten wood). Workers of most (perhaps all) species exhibit considerable size variation. With the shared characters of Cerapachyini and also with the following combination of characters. Two undoubted apomorphies of the genus are in italics.

1 Palp formula 6,4 or 5,3. Maxillary palp of moderate length: with mouthparts retracted the apex of the maxillary palp, when extended back on underside of head, does not reach beyond the level of the posterior margin of the eye.

2 Mandibles triangular, either edentate, bluntly denticulate, or with small, low, blunt crenulations at least on the basal half; mandibles without a basal groove and without a basal pit.

3 Antenna with 11 segments, gradually incrassate apically; apical antennomere large but subcylindrical, not swollen and bulbous, its maximum width generally no greater than that of the preapical segment.

4 Scape short (SI 33–56); when laid straight back in full-face view the scape apex merely reaches the level of the anterior margin of the eye or slightly beyond.

5 Eyes large (EL/HW 0.30–0.53), located from slightly in front of, to distinctly behind, the midlength of the head.

6 Ocelli present; usually distinct but may be small, inconspicuous and flush with the surface of the head.

7 Clypeus broad and more or less flat across; broadly inserted between frontal lobes.

8 Frontal lobes and carinae present; frontal lobes at most only weakly elevated and in full-face view conceal at least the inner margins of the antennal sockets; frontal carinae usually extend back to at least the level of the anterior margins of the eyes; frontal carinae never abruptly truncated posteriorly and never terminated immediately behind the frontal lobes.

9 Parafrontal ridges variably developed, sometimes absent.

10 Head capsule without a differentiated vertical posterior surface above the occipital foramen; instead the vertex slopes evenly down to the upper margin of the occipital foramen, which is visible in full-face view.

11 Head capsule, in ventral or ventrolateral view, with a carina that extends down the posterolateral margin and onto the ventral surface, where it terminates or fades out well before meeting the ventral midline; this carina is anterior to, and separate from, that which borders the occipital foramen.

12 Mesosoma dorsally with at least a transverse vestige of promesonotal suture; metanotal groove usually absent, although sometimes vestigially present to obvious.

13 Pronotum in dorsal view bluntly angulate to carinate anteriorly between anterior surface and dorsum; propodeum rounded to carinate between dorsum and declivity.

14 Mesosoma laterally usually with an impression or pit near the metapleural-propodeal junction, on approximately the same level as the propodeal spiracle.

15 Midpoint of metanotal groove (or midpoint of dorsum at that level if groove absent) with a pit or impression.

16 Mesotibia without spurs.

17 Metatibia with a single, pectinate spur.

18 Metatibial gland absent (at least no external orifice or indication is present).

19 Metabasitarsus ventrally with a longitudinal glandular groove that occupies at least the basal half of the tarsomere length.

20 Pretarsal claws of all legs with a single preapical tooth on the inner surface of each claw.

21 Propodeal lobe in profile broad-based, bluntly triangular or rounded apically.

22 AII (petiole) flattened or shallowly convex dorsally; angulate to marginate laterally in dorsal view; often with a longitudinal carina on the side above the level of the spiracle.

23 AIII broadly postpetiolate, more voluminous than AII (petiole) but usually smaller than AIV.

24 Prora of AIII a simple curved cuticular rim or carina that separates the anterior face of the poststernite from its lateral and ventral surfaces.

25 Pretergite of AIV in dorsal view strongly constricted with respect to posttergite of AIV.

26 Cinctus of AIV smooth, without cross-ribs.

27 Tergite of AIV without a pair of subovate glandular patches on the posterior half.

28 Pygidium variable in structure at its apex: either apical margin evenly curved and equipped with a row of 4–6 minute denticles; or apex with one or two pairs of somewhat enlarged teeth that are flanked by smaller teeth; or apex with a short but stout, bifid cuticular fork that is much more strongly developed than any other teeth on the margin.

Comments on worker characters (Numbers correspond to character numbers above)

1 Palp formula 6,4 was established by dissection in workers of conradti, dux, nonnihil, rex, sicaria, silens, trita and wilburi, and by in situ counts of everted mouthparts in grandis, latiscapa, marleyi, miniflava and persculpta. Palp formula 5,3 was established by dissection in elegans and grandidieri. In some other species the palp formula could be estimated even though the mouthparts were retracted (e.g. PF 6,4 in annettae, brunnea, dryas) but in some, and especially in specimens mounted flat on card, the mouthparts were obscured and PF could not be counted.

3 In most species all funicular segments except the apical are conspicuously broader than long, but in laevissima and some extralimital species of the grandidieri group segments 4–6 appear at least as long as broad.

4 SI range includes all directly measured Afrotropical and Malagasy species. Among the extralimital species, chapmani has SI 33 (holotype measured), bakeri has SI 37 (MCZC gyne measured), oculata has SI ca 42 (estimated from Radchenko, 1993: 46, fig. 5) and gressitti has SI ca 41 (estimated from Taylor, 1965: 4, fig. 2).

5 In the schoutedeni group many conspicuous short setae arise between the ommatidia of the eye. Elsewhere in the genus such setae are much more sparse and usually indistinct, but in most species a few can be detected at high magnification.

7 In most members of the emeryi group the clypeus is very obviously strongly reflexed, so that the true anterior clypeal margin, at the clypeo-labral junction, is considerably below and behind the apparent anterior clypeal margin as seen in full-face view. The lateral portions of the clypeus, in front of the antennal sockets, tend to be flattened or evenly rounded in members of the grandidieri group, the schoutedeni group, and in grandis and persculpta of the emeryi group. In the remainder of the emeryi group the margin is produced anteriorly as a rounded lobe or short, blunt triangle in front of each socket, but these projections are reduced in trita and silens.

9 Parafrontal ridges (sensu Wilson, 1964, Borowiec, 2009; = genal carinae sensu Bolton, 2003) are usually distinctly present in the grandidieri group and the schoutedeni group, where they appear as fine carinae that extend back almost to the eye. In the emeryi group their development is more variable. They may be entirely absent or represented only by a slight tumulus immediately behind the clypeus (e.g. conradti, marleyi, rex, silens), or weakly present but short and inconspicuous (e.g. dignita, latiscapa, merita, persculpta, victrix). They are never extended backwards to near the anterior margin of the eye.

11 In direct contrast to the situation in Simopone, in both other genera treated here, Tanipone and Vicinopone, the carina extends to the ventral midline.

12 Promesonotal suture is sometimes vestigial or mostly absent from the dorsum (elegans, grandidieri), or it may be merely a weak impression or a fine, slightly incised line (consimilis, dignita, emeryi, marleyi); most often its track is indicated by a row of short cuticular ribs or linked punctures across the dorsum, which may be conspicuous or very faint (conradti, laevissima, latiscapa, rex and the schoutedeni group). The metanotal groove is sometimes absent, but often it is represented by a weak to vestigial transverse line (conradti, grandis, inculta, latiscapa, rex); only very rarely is it obviously, though shallowly, impressed (chapmani, laevissima).

13 Anterodorsally the pronotum may be merely angulate between its anterior and dorsal surfaces (rex, silens) but in most it is distinctly marginate to carinate. However, the pronotal dorsum never rounds broadly and evenly into its anterior surface.

14 This impression or pit perhaps represents the last visible trace of the metapleural spiracle on the surface of the mesosoma. It is quite conspicuous in the majority of species but is small and indistinct, or even absent, in the Afrotropical laevissima and the Malagasy species elegans, emeryi, fera, grandidieri, merita and nonnihil.

15 The median pit is usually distinct but may be small and inconspicuous in conradti and grandis, and may be difficult to distinguish from the predominant sculpture in those species of the schoutedeni group that have dense foveolate punctures on the dorsal mesosoma.

16 Apex of the mesotibia usually has one to several setae that are roughly aligned in the direction of the long axis of the shaft. It is probable that one of these represents the last vestige of a spur.

18 The metatibia, at the base of the pectinate spur, usually shows a small translucent patch of cuticle, or a small cuticular vesicle. This appears to be associated with the articulation of the spur and is not part of a metatibial gland structure. In large, strongly sclerotised species such as rex and silens, the small translucent patch is usually not apparent.

19 The ventral metabasitarsal groove is conspicuous in all species. It varies from a narrow slit to a broad trench that occupies well over half the width of the tarsomere and is often filled with white to yellowish flocculent material. Elsewhere in Formicidae a similar groove is exhibited only by Paraponera (Paraponerinae), Myrmecia and Nothomyrmecia (Myrmeciinae), and Tetraponera and Myrcidris (Pseudomyrmecinae) (summarised in Bolton, 2003).

20 The preapical tooth on each pretarsal claw is conspicuous on all legs in larger species, but may be small and more difficult to see in small species, especially on the forelegs, but is always present.

22 Tergite of abdominal segment AII (petiole) shows a wide range of structures. Transverse carinae may be present anterodorsally, posterodorsally, or both, where the dorsum meets the anterior and posterior surfaces. The lateral surface of AII often has a curved longitudinal carina that extends its length. This carina is located below the dorsolateral margination but above the level of the spiracle. In the grandidieri group it is sometimes short and confined to the area in front of and above the spiracle (elegans, grandidieri) but it is entirely absent in bakeri, chapmani and laevissima. In the emeryi group it is usually present but is weak in conradti and marleyi, and absent in dignita, dux, sicaria and victrix. In the schoutedeni group it is universally present and generally strongly developed. The posterior corners of AII are frequently extended into lobes or teeth that project posteriorly or laterally.

24 In a few species, for instance grandidieri, laevissima and some members of the schoutedeni group, the prora is only feebly developed ventrally.

25 In dorsal view the maximum width of the pretergite of AIV is 0.52–0.69 × the maximum width of the posttergite of the segment. The ratio in Vicinopone falls within the same range (0.62–0.64) but the pretergites of Tanipone species are relatively much wider, 0.76–0.89. This ratio has not yet been investigated in other cerapachyine genera.

28 Pygidial armament varies among the three species groups recognised within the genus. In the grandidieri group the posterior pygidial margin is lined with an arc of small, usually peg-like denticles that are all approximately the same size, without an enlarged pair apically. In the schoutedeni group the posteromedian pair or two pairs of denticles on the arc are somewhat enlarged, immediately above the sting. In the emeryi group the margin of the pygidium is produced posteriorly into a short but stout cuticular fork that is very obviously larger and more strongly developed than any marginal denticle.

Borowiec (2016) - Head: Antennae with 11 segments. Apical antennal segment not enlarged, not broader and longer than two preceding segments combined. Clypeus without cuticular apron. Lateroclypeal teeth absent. Parafrontal ridges absent or reduced. Torulo-posttorular complex horizontal. Antennal scrobes absent or present. Labrum with median notch or concavity. Proximal face of stipes projecting beyond inner margin of sclerite, concealing prementum when mouthparts fully closed. Maxillary palps 6- or 5-segmented. Labial palps 4- or 3-segmented. Mandibles triangular, edentate. Eyes present, composed of more than 20 ommatidia. Ocelli present. Head capsule without differentiated vertical posterior surface above occipital foramen. Ventrolateral margins of head without lamella or ridge extending towards mandibles and beyond carina surrounding occipital foramen. Posterior head corners dorsolaterally immarginate. Carina surrounding occipital foramen ventrally absent. Mesosoma: Pronotal flange often separated from collar by distinct ridge, occasionally ridge absent. Promesonotal connection with suture present, weakly differentiated, immobile. Pronotomesopleural suture visible, unfused up to notal surface. Mesometapleural groove weakly impressed or not impressed. Transverse groove dividing mesopleuron absent or present. Pleural endophragmal pit concavity present. Mesosoma dorsolaterally immarginate. Metanotal groove on mesosoma absent or shallowly impressed but well-defined line. Propodeal spiracle situated low on sclerite. Propodeal declivity often without distinct dorsal edge or margin but occasionally marginate, rectangular in posterior view. Metapleural gland without bulla visible through cuticle. Metapleural gland with bulla visible through cuticle. Propodeal lobes present, well developed. Metasoma: Petiole anterodorsally immarginate or marginate, dorsolaterally immarginate, and laterally above spiracle marginate. Helcium in relation to tergosternal suture placed at posttergite and axial. Prora simple, not delimited by carina, a U-shaped margin, or U-shaped margin with median ridge. Spiracle openings of abdominal segments IV–VI circular. Abdominal segment III anterodorsally immarginate and dorsolaterally immarginate. Abdominal segment III more than half size of succeeding segment IV, which is weakly constricted at presegmental portion (uninodal waist). Girdling constriction of segment IV present, i.e. pre- and postsclerites distinct. Cinctus of abdominal segment IV gutter-like, not sculptured. Abdominal segment IV not conspicuously largest segment. Abdominal tergite IV not folding over sternite, and anterior portions of sternite and tergite equally well visible in lateral view. Girdling constriction between pre- and posttergites of abdominal segments V and VI absent. Girdling constriction between pre- and poststernites of abdominal segments V and VI absent. Pygidium large, with impressed medial field, armed with modified setae, and in some species deeply notched at apex. Hypopygium unarmed. Legs: Mid tibia without spurs. Hind tibia with single pectinate spur. Hind basitarsus not widening distally, circular in cross-section. Posterior flange of hind coxa not produced as raised lamella. Metatibial gland absent. Metabasitarsal gland present. Hind pretarsal claws each armed with a tooth. Polymorphism: Monomorphic.

Queen
Bolton and Fisher (2012) - Known queens are entirely worker-like except that the mesosoma has a full complement of flight sclerites. Therefore all the worker characters listed above, except for numbers 12 and 15, are duplicated here. Queens are known for the Afrotropical species annettae, conradti, latiscapa, marleyi, matthiasi, miniflava, persculpta, wilburi, and for the extralimital species bakeri; most specimens are dealate. No queen recognisable by external morphology has been seen in any Malagasy species, the queens of which are suspected to be remarkably ergatoid, or perhaps even replaced by gamergates, but proof of this must await dissection of reproductive systems to assess the presence of spermathecae and enlarged ovaries. For venation, see discussion of character 15 under males.

Borowiec (2016) - Alate or extremely ergatoid/ replaced by fertile workers. Alate and dealated queen specimens are known in all three Simopone species-groups recognized by Bolton and Fisher (2012), e.g. in Simopone annettae of the schoutedeni group, Simopone latiscapa of the emeryi group, and Simopone bakeri of the grandidieri group. Members of all three species-groups occur also on Madagascar. However, no morphologically distinct gynes have ever been collected among the 16 species occurring on the island, despite multiple nest samples available. It is possible that queens have been replaced there by reproductively active workers, the so-called gamergates (Bolton and Fisher 2012).

Male
Bolton and Fisher (2012) - Of the Malagasy species males are known for dux, mayri, nonnihil, rex, silens and seven unassociated forms of the emeryi group, and also for grandidieri of the grandidieri group. The males of two species are known from the Afrotropical region, marleyi and an unassociated male of the schoutedeni group. No males have been recorded for any of the extralimital taxa.

1 Palp formula 6,4 or 5,3.

2 Mandibles triangular and edentate, masticatory margins straight to shallowly concave, meeting at full closure.

3 Antenna 12-segmented, filiform; funicular segments 3 to apex longer than broad; apical segment the longest, tapering apically and not swollen, longer than funicular segment 11 but no broader.

4 Eyes large and very conspicuous.

5 Ocelli present.

6 Frontal lobes and frontal carinae present and strongly developed; lobes somewhat elevated on each side of the relatively broad clypeus but concealing at least the inner margins of the antennal sockets in full-face view.

7 Head capsule, in ventral or ventrolateral view, with a carina that extends down the posterolateral margin and onto the ventral surface, where it terminates or fades out well before meeting the ventral midline.

8 Pronotum slightly to very conspicuously visible in dorsal view.

9 Notauli present.

10 Parapsidal grooves present.

11 Propodeal lobes present and conspicuous.

12 Mesotibia without spurs.

13 Metatibia with a single, pectinate spur.

14 Pretarsal claws each with a preapical small tooth.

15 Venation as discussed below.

16 Prora present as a transverse cuticular flange.

17 AIII postpetiolate, with AIV > AIII > AII.

18 Pygidium with posterior margin not denticulate.

Comments on male characters

Too few males are known to give as detailed a diagnosis as the workers, but the listed characters should serve to successfully isolate Simopone males from other cerapachyines. Numbers below correspond to character numbers above.

1 PF 6,4 has been confirmed by dissection or in situ count in dux, marleyi, rex, silens and three unassociated Malagasy species; PF 5,3 was confirmed by in situ count of grandidieri.

3 In all males except the one from the schoutedeni group the length of funicular segment 4 is equal to or > 3 > 2 > 1; in the schoutedeni group male funiculus segment 4 is about equal to 3, and 2 is about equal to 1, with 1 and 2 shorter than 3 and 4. The scape in all known Malagasy species of the emeryi group has a flange-like or broadly tooth-like prominence apically, in front of the insertion of the first funicular segment. This is not developed in marleyi, grandidieri, or the schoutedeni group male.

4 In the schoutedeni group male the short, bristly setae that arise between the ommatidia, which are so obvious in workers and queens, are absent.

8 Two quite distinct degrees of pronotal exposure are exhibited. In all the known Malagasy males the pronotum, in dorsal view, merely forms a narrow collar in front of the mesoscutum, as is also the case in Tanipone. But in the Afrotropical marleyi and in the unassociated male of the schoutedeni group the visible pronotum forms an extensive, transverse sclerite in front of the mesosocutum. The midline length of the pronotum (its narrowest point) is 0.25 (marleyi)–0.35 × the midline length of the mesoscutum. By contrast, in Malagasy males the pronotum is only 0.10–0.15 × the mesoscutum midline length.

15 Wing venation is best represented in larger males, such as those of mayri, rex and silens. Diagnostically, on the forewing the pterostigma is large and pigmented, usually strongly so; C is absent; R1•f3 is a short stub or absent distal of the pterostigma; 2rs-m is absent; Rs•f2-3 is faint, sometimes spectral, detached basally from Rs+M so that its base floats freely in the membrane; 2r-rs and Rs•f4-5 are usually weakly developed and sometimes very faint indeed, and the latter does not distinctly meet the wing margin; 2r-rs may occasionally be absent; 1m-cu is usually present but may be very faint; cu-a arises from M+Cu (before it splits into M•f1 and Cu•f1); M fades out distally. In smaller males and in the only known alate queen (annettae) the forewing venation shows reduction from this pattern until the wing is veinless from a point well proximal of the pterostigma to the margin except for Sc+R1, linked to the pterostigma. On the hindwing rs-m is always absent.

Borowiec (2016) - Head: Antennae with 12 segments. Clypeus without cuticular apron. Parafrontal ridges present. Torulo-posttorular complex horizontal. Maxillary palps 6- or 5-segmented. Labial palps 4- or 3-segmented. Mandibles triangular, edentate. Ventrolateral margins of head without lamella or ridge extending towards mandibles and beyond carina surrounding occipital foramen. Carina surrounding occipital foramen ventrally absent. Mesosoma: Pronotal flange separated from collar by distinct ridge. Notauli present. Transverse groove dividing mesopleuron absent. Propodeal declivity with distinct dorsal edge or margin. Metapleural gland opening absent. Propodeal lobes present. Metasoma: Petiole anterodorsally marginate, dorsolaterally immarginate, and laterally above spiracle marginate. Helcium in relation to tergosternal suture placed at posttergite and axial. Prora forming a U-shaped protrusion. Spiracle openings of abdominal segments IV–VI circular. Abdominal segment III more than half size of succeeding segment IV; latter weakly constricted at presegmental portion (uninodal waist). Girdling constriction of segment IV present, i.e. pre- and postsclerites distinct. Cinctus of abdominal segment IV gutter-like and cross-ribbed. Girdling constriction between pre- and postsclerites of abdominal segments V and VI absent. Abdominal segment IV not conspicuously largest segment. Abdominal sternite VII simple. Abdominal sternite IX distally armed with two spines, with lateral apodemes about as long as medial apodeme, directed anteriorly (towards head). Genitalia: Cupula long relative to rest of genital capsule and shorter ventrally than dorsally. Basimere broadly fused to telomere, with no sulcus trace at junction, and ventrally with left and right arms separated. Telomere expanded at apex. Volsella variable. Penisvalva laterally compressed, rounded at apex. Legs: Mid tibia without spurs. Hind tibia with single pectinate spur. Posterior flange of hind coxa not produced as raised lamella. Metatibial gland absent. Metabasitarsal glands absent. Hind pretarsal claws each armed with a tooth. Wings: Tegula present, broad, demiovate in shape. Vein C in fore wing absent. Pterostigma broad. Abscissa R·f3 absent. Abscissae Rs·f2–3 present, connecting with Rs+M&M·f2. Cross-vein 2r-rs present, differentiated from Rs·f4 by presence of Rs·f2–3 or absent. Abscissae Rs·f4–5 present, fused in absence of 2rs-m. Abscissa M·f2 in fore wing present, separated from Rs+M by Rs·f2. Abscissa M·f4 in fore wing present, reaching wing margin. Cross-vein 1m-cu in fore wing present or absent. Cross-vein cu-a in fore wing present, arising from M+Cu and proximal to M·f1. Vein Cu in fore wing present, with both branches Cu1 and Cu2. Vein A in fore wing with abscissae A·f1 and A·f2 present. Vein C in hind wing absent. Vein R in hind wing absent. Vein Sc+R in hind wing present. Abscissa Rs·f1 in hind wing unknown. Abscissa Rs·f2 in hind wing unknown. Cross-vein 1rs-m in hind wing present, about as long as M·f1, never tubular. Vein M+Cu in hind wing present. Abscissa M·f1 in hind wing present. Abscissa M·f2 in hind wing present. Cross-vein cu-a in hind wing present. Vein Cu in hind wing present. Vein A in hind wing with abscissae A·f1 and A·f2 present.

Larva
Cocoons absent.