The application of this family-group name is based on usage rather than date priority. As can be seen from the following list of family-group names available within the Sterrhinae, there are several with equal claims, and some recent check-lists have preferred Scopulinae (reviewed briefly by Holloway (1993[4])), though the majority (e.g. Hodges et al. (1983), Vives Moreno (1994) and Nielsen, Edwards & Rangsi (1996)) have used Sterrhinae.

Before discussing the higher classification of the subfamily, a list is presented of family-group names potentially applicable within it. It is presented with the same caveat as that made for the list of Ennominae by Holloway (1993[4]): the search for names and the earliest reference thereto has been extensive but not exhaustive. All names are arbitrarily given tribal rank endings except for Sterrhinae.

Family-group names in the Sterrhinae
[ACIDALIINI]: Acidalites Duponchel (1845). The name is based on a junior homonym of a genus-group name outside the Lepidoptera.

ALETINI: Aletinae Hampson (1918). Holloway (1996) indicated this to be within the concept of the Scopulini.

CALOTHYSANINI: Calothysanini Herbulot (1962-3). (See Timandrini). This name is based on Calothysanis Hübner, a synonym of Scopula Schrank, and therefore cannot be applied (e.g. as by Hausmann (1993) and Vives Moreno (1994)) to include genera such as Timandra Duponchel.

COSYMBIINI: Cosymbiinae Prout (1911); Hodges et al. (1983). The name embraces the same concept as Cyclophorini and Zonosomini. See Holloway (1993[41) and Chrysocraspeda Swinhoe for discussion of the preferable name.

CYCLOPHORINI: Cyclophoridae Moore (1887); Herbulot (1962-3), Wiltshire (1990). As stated by Holloway (1993[4]), the family-group name Cyclophoridae is used extensively in Mollusca, based on Cyclophorus Montfort. It is recommended here that lepidopterists gracefully concede this on grounds of usage rather than attempt to establish priority. See Cosymbiini, Zonosomini and discussion of the most appropriate name for the group on Chrysocraspeda Swinhoe.

CYLLOPODINI: Cyllopodidae Kirby (1892); Warren (1895). This is a Neotropical group of highly coloured (yellow and black), day-flying genera (Covell, 1983). They bear some resemblance in facies to Bytharia Walker, but are probably related to the Rhodostrophiini. See Definition of subfamily and tribal groupings, below.

[EPHYRINI]: Ephyridae Guenée (1857). The name is based on a junior homonym (= Cyclophora) of a genus group name outside the Lepidoptera.

GONIACIDALIINI: Goniacidaliinae Packard (1876). Goniacidalia Packard is a synonym of Idaea Treitschke (Hodges et al. 1983), therefore this name becomes subordinate to Sterrhini. It does, however, have date priority over Sterrhini, as does Idaeini. See next entry.

IDAEINI: Idaeidae Butler (1881). This name refers to the same concept as Sterrhinae (see below), has priority, and is also based on the most senior of two generic synonyms. However, the extensive usage of Sterrhinae in preference to Idaeinae this century mitigates against revival of the latter.

[MICROPININI]: Micropinidae Kirby (1912). The name is based on an unnecessary replacement genus-group name, Micropinus Kirby, for Micropus Hübner, replaced earlier by Smicropus Warren. The genus is referable to the Cyllopodini (see above).

PROBLEPSINI: Problepsini Wiltshire (1990). This is within the concept of Scopulini as defined here (See Scopulini), but no earlier reference with a valid description has been located.

PTYCHOPODINI: Ptychopodinae Pierce (1914); Forbes (1948). Ptychopoda Curtis is a junior subjective synonym of Idaea Treitschke, therefore this name is subordinate to Sterrhinae.

RHODOMETRINI: Rhodometrinae Agenjo (1951). Herbulot (1962-3) treated this as a subfamily but it was placed as a tribe of the Sterrhinae by Seven (1991) and Vives Moreno (1994). The group does not occur in Borneo. See Definition of subfamily and tribal groupings.

RHODOSTROPHIINI: Rhodostrophiicae Prout (1935); Herbulot (1962-3); Seven (1991). This tribe is discussed on Rhodostrophini.

SCOPULINI: Scopulites Duponchel (1845). See Holloway (1993[4]) for arguments concerning subfamily priority for this name. As treated here it also embraces the concepts of Aletini and Problepsini.

STERRHINAE: Sterrhidae Meyrick (1892); Hodges et al. (1983). This name has priority of usage over Goniacidaliini and Idaeini (See Sterrhinae).

TIMANDRINI: Timandridi Stephens (1850); Hodges et al. (1983), Seven (1991). An older concept of Calothysanini (above) that has been preferred in recent literature. For discussion of current status see Timandrini.

ZONOSOMINI: Zonosomatidi White (1876, Scot. Nat. 3: 361); Seven (1991). See Holloway (1993[4]) and Chrysocraspeda Swinhoe for discussion of validity of this name relative to Cosymbiini. See also Cyclophorini above.

Definition of subfamily and tribal groupings
Recent general accounts of the Lepidoptera, such as Holloway, Bradley & Carter (1987), Scoble (1992) and Common (1990), give no clear definition of the Sterrhinae in terms of apomorphic characteristics. Definition of the tribes comprising the subfamily is much clearer, but features that pull them all together are elusive.

Forbes (1948) noted the length and longitudinal orientation of the most anterior discocellular vein (between the base of M1 and the areole) of the forewing. This is very short or transverse in other subfamilies except for some groups of the Ennominae where the sterrhine condition is approached. It is also seen in the Desmobathrinae (compare figures 24 and 27 with 25, 26 and 28 in Holloway (1986a)).

The only other adult feature that appears to be general to all tribes except Cyllopodini and Rhodometrini, occurring in at least some component taxa, is a hindwing discal spot with pale markings in a darker surround, the pale colour distinct from that of the general ground. Covell (1987) mentioned the presence of black discal spots on both wings, and indeed they are of more general occurrence than in other subfamilies, but pale centering is unusual. It occurs in all the tribes found in Borneo except for the Sterrhini, but these can be linked with other tribes on separate grounds.

The larvae and pupae may offer further characters.

Singh (1953) defined larvae of Sterrhinae as having the crotchets on the prolegs interrupted centrally (though this can also occur in some Ennominae and is general to Geometrinae) in combination with seta CD2 on the anal proleg being below the level of seta SV2. Within the Sterrhinae he recognised two major divisions on differences in chaetotaxy. In Division A the SV group is unisetose on A1 and bisetose on A2, whereas there is an extra seta in each in Division B. There are differences also in the position of CD2 on the anal proleg, SD1 and A1, and D1 on the anal shield.

Division A included Traminda mundissima Walker and species of Chrysocraspeda Hampson, i.e. the Timandrini and Cosymbiini as recognised here. Division B was subdivided on the shape of the subanal plate and other features of chaetotaxy, separating the Scopulini, with the subanal plate produced into a point posteriorly, from the Sterrhini where it is entire. Idaea larvae have clubbed setae.

This division is supported also by the analysis of pupal characters by Nakamura (1994), where Timandrini and Cosymbiini were grouped together, and Sterrhini were sister to Scopulini + Rhodostrophiini in the other lineage. This division is seen particularly strongly in the character of the cremaster (Khotko, 1977; Bell, MS) where the eight hooklets characteristic of the geometrid ground plan (Holloway, 1993[4]) are more or less equal in the first lineage and tend to be reduced sometimes to just the apical pair (usually enlarged) in the second lineage, with the other pairs small or absent, though this feature is less emphatic in the Sterrhini.

Nakamura noted some general features of sterrhinae pupae: the first and ninth abdominal segments lack setae (Timandra is an exception); the mesothoracic leg is never tapering at its extremity; the proximal end of the prothoracic leg appears from further caudad than in other geometrids.

There is insufficient information about the eggs to provide strong pointers to the higher classification. What there is (e.g. Salkeld, 1983) shows consistency with the lineages noted above. Eggs in the cosymbiine lineage are sculptured with the irregular polygonal cells with aeropyles within the polygon walls that appears to be the geometrid ground plan. In Scopula and some Idaea the eggs have longitudinal ridges, within which are the aeropyles, and rugosity, pitting and sometimes weaker transverse ridges in between. Longitudinal ridges are also seen in the ennomine tribes Lithinini and Caberini (Salkeld, 1983; Holloway, 1993[4]), but these are more numerous with aeropyles more robust and of greater frequency along the ridges. Prout (1913, Gross-Schmett. Erde 4: 39) referred to longitudinal ridging in the eggs of Rhodostrophia Hübner.

Male secondary sexual characters also support the monophyly of a group containing Rhodostrophiini, Cyllopodini, Scopulini and Sterrhini. All have long scent pencils on the hind-tibia. This appears to be associated with a zone of setae scales or rugosity (Cyllopodini, Rhodostrophiini) or a pouched structure (Scopulini, some Sterrhini, situated between the tympanic bullae on the second sternite. In other geometrid subfamilies, such as Ennominae, Geometrinae and Desmobathrinae, occurrence of a tibial scent pencil is usually correlated with patches or a comb of setae on the third abdominal sternite. The hind-leg itself is often shortened, with tarsi reduced and the apex of the leg upturned, presumably to manipulate the pouch or its contents on the second sternite.

In the other lineage scent pencils do not occur, but in Perixera Meyrick of the Cosymbiini the tibia is shortened and often invested with thick tufts of scales. The abdominal correlate in this instance appears to be the presence of a pair of corematous structures on the fourth sternite (See Perixera Meyrick Gen.rev.).

Two tribes are not represented in Borneo, but their affinities will be discussed briefly here.

The Cyllopodini are Neotropical, the species relatively large, brightly patterned in yellow with black bands and borders. The type species, Cyllopoda claudicula Dallman, has characters, some already mentioned, that indicate an affinity with the Rhodostrophiini. The male hind-leg is very much reduced, halved in size, apically upturned; there is a semicircular setose plate on the second sternite. The uncus of the male genitalia is elongate, setose, apically broadened and bilobed somewhat as in Erythrolophus Swinhoe (See Erythrolophus Swinhoe). The gnathus is strong, apically acute. In the female the ovipositor lobes are longitudinally grooved, and there are two densely spined longitudinal bands of sclerotisation in the bursa copulatrix.

The Rhodometrini consist principally of the genus Rhodometra, distributed over semi-arid regions of Africa, Asia and in the Mediterranean area. The facies is consistent with this, the forewings pale yellow with an oblique reddish postmedial band. The type species, R. sacraria Linnaeus, has an enlarged central flap on the tympanic ansa, though not as extreme as in Traminda Saalmüller: the apex of the ansa is tapering, not hammer-headed. The male genital capsule is ovate, somewhat reminiscent of that in Scopula Schrank. The pupa has a projecting head case (Buckler, 1899) but this does not have the two protruberances seen in Timandra Duponchel . The pupa was stated by Buckler to be enclosed in an open network of silk between the stems of the host-plant or on other surfaces. This, and some indication of a host association with Polygonaceae (Buckler; Allan, 1949), suggest affinity with Timandrini, though the cremaster, with all eight hooklets set together on the distal edge of a trapezium, is similar to that in Cosymbiini.

Another feature indicating association with the Timandrini/Cosymbiini lineage is the signum of the female genitalia, a longitudinal, ridge-like invagination. Hausmann (1993) proposed this character as a synapomorphy bringing the three tribes together, suggesting its absence in Traminda and most Cosymbiini to be a secondary loss. The two genera he studied, Pseudosterrha Warren and Chlorerythra Warren, both have this feature of the female genitalia and were suggested to be in a somewhat basal position (plesiomorphic characteristics) within the Timandrini/ Cosymbiini lineage. The uncus of some Pseudosterrha species he illustrated resembles that of Chrysocraspeda Warren and Cyclophora Hübner. Hausmann also drew attention to "striate" valve ornamentation in the complex, but it is debatable whether the strong fields of basally directed setae in Cyclophora, the carinae of Traminda, and the rather weak setation in other Timandrini and Cosymbiini represent a strong homology.

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