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)), 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): 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), 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
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
[EPHYRINI]: Ephyridae Guenée (1857). The name is based on a
junior homonym (= Cyclophora) of a genus group name outside the
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) 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
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) 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) 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), 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
Two tribes are not represented in Borneo, but their affinities will be discussed
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
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
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