::The Moths of Borneo::

INTRODUCTION

This part of the series covers the subfamily Geometrinae of the Geometridae (emerald moths and allies) and the groups traditionally (e.g. Prout, 1910) placed within the subfamily Oenochrominae. The Geometrinae represent a natural grouping that can be defined relatively clearly (Geometrinae, Geometrini), but it has been apparent in recent publications (Scoble, 1986; Scoble & Edwards, 1989) that the validity of the old concept of the Oenochrominae cannot be substantiated, and even Prout (1910, 1920) considered the subfamily heterogeneous but a grouping of convenience.

Therefore, before dealing with Bornean taxa currently in the Oenochrominae, it is necessary to investigate this higher classification problem.

The first step in this process is to review the family-group names available within the Geometrinae and Oenochrominae sensu lato.

Family-group names
Family-group names for Ennominae have already been listed in part 11 of this series (Holloway, 1993[4]), with two omissions, Dalimini and Phaseliini, discovered after the work had gone to press. These are mentioned briefly below, as well as all family-group names referred to Oenochrominae or Geometrinae by Prout (1910, 1912a, b, 1913), and Fletcher (1979), or published in those subfamilies since Prout's catalogues (e.g. Inoue, 1961; Ferguson, 1969; Viidalepp, 1981). The Orthostixini and two groups with genitalia features in common with the Ourapterygini (Diptychini, Oxydiini) may also be referable to the Ennominae.

The list is presented with the same caveat as that for the Ennominae, and provides a basis for the comments on higher classification presented in the next section. All are arbitrarily given tribal rank endings except where recognition as a subfamily-group can be substantiated.

ALETINI: Aletinae Hampson (1918). This name was originally proposed as a replacement for Oenochrominae, being based on the oldest generic name (Aletis Hübner) within the subfamily as then recognised. However, as indicated by Prout (1929: Gross-Schmett. Erde, 16: 7), this genus has male genitalia and eighth sternite and the signum of the bursa copulatrix in the female typical of the Scopulini. Aletini should therefore be treated as a synonym of Scopulini.

ALSOPHILINAE: Alsophilinae Herbulot (1962-3). This group has been subordinated to Oenochrominae by Inoue et al. (1982) and Seven (1991), but, given the polyphyletic nature of the group as currently recognised and the more restricted definition of Oenochrominae, this grouping of delicate Palaearctic taxa is best placed as a distinct subfamily. It does not appear to be related to the Desmobathrinae (for example, the pupal cremaster is bifid). Nakamura (1987) used pupal characters to suggest inclusion of the group in Ennominae.

AMETRIDINI: Ametridicae Prout (1910). See also Mecoceratini. This is a Neotropical group not considered in detail here. The female of Ametris nitocris Cramer has an elongate, pyriform bursa on a long, slender ductus. The signum is an elongate sclerotised band with short spines irregularly along its length set on transverse ridges. The status of the group requires further investigation.

ARCHAEOBALBINI: Archaeobalbini Viidalepp (1981). This name is available for the robust Geometrinae and represents Inoue's (1961) concept of Terpnini, but is probably a junior synonym of Pseudoterpnini.

ARACIMINI: Aracimini Inoue (1961). A group within the Geometrinae.

CHLOROCHROMINI: Chlorochromites Duponchel (1845); Seven (1991). Chlorochroma Duponchel is a junior synonym of Thalera Hübner: see also Thalerini for status in Geometrinae.

COMIBAENINI: Comibaenini Inoue (1961); Leraut (1980); Seven (1991). See Euchlorini and (Geometrini) for indications of relationship to Synchlorini within Geometrinae.

COMOSTOLINI: Comostolini Inoue (1961). (Geometrini) for discussion of status within Geometrinae.

DALIMINI: Daliminae Wehrli (1940; Gross-Schmett. Erde 4 (Suppl.): 349). The type genus was included in Boarmiini of the Ennominae by Holloway (1993 [4]).

DESMOBATHRINAE: Desmobathridae Meyrick (1886). Desmobathra Meyrick is a junior synonym of Ozola Walker. See below for definition of this group and justification of subfamily status.

DICHORDOPHORINI: Dichordophorini Ferguson (1969, 1985). This small Geometrinae tribe appears to have no close affinities with the S.E. Asian fauna and will not be considered further here. (Geometrini).

DIPTYCHINI: Diptychini Janse (1932-5). Pinhey (1975) treated this African group as a subfamily rather than as a tribe of Oenochrominae. The signum of the female is ennomine, and the male genitalia have a valve and furca structure reminiscent of the Ourapterygini.

DYSPHANIINI: Dysphaniinae Warren (1895). See also Euschemini, Hazini. The status of this family-group is discussed below (Dysphaniini).

EUCHLORINI: Euchlorini Herbulot (1962-3). Euchloris Hübner is a junior homonym of Euchloris Billberg (Coleoptera). As applied by Herbulot, the concept includes Comibaena and Thetidia Boisduval, and is therefore referable to Comibaenini.

EUMELEINI: Eumeleinae Warren (1894). See below (Eumeleini) for discussion of the status of this monogeneric family-group.

EUSCHEMINI: Euschemidae Walker, (1862); Butler (1886). This name is based m Euschema Hübner, a junior synonym of Dysphania Hübner. See Dysphaniini.

GEOMETRINAE: Geometridae Stephens (1829).

HAZINI: Hazidae Guenée (1857). Hazis Boisduval is a junior objective synonym of Euschema Hübner. See Euschemini, Dysphaniini.

HEDYLIDAE: Hedylidae Guenée (1857). This family of Neotropical moths was included in the Oenochrominae by Prout (1910), but has been shown by Scoble (1986) not to be geometroid but to be related to the butterflies. A recent review of its status may be found in Scoble (1992: 302-4).

HELIOTHEINI: Heliothinae [sic] Exposito (1978). The name is based on Heliothea Boisduval, placed in the Oenochrominae by Fletcher (1979). The type species flies by day in Spain. The genus was indicated to have affinity with the Geometrinae by Prout (1912; Gross-Schmett. Erde 4: 6): the structure of the ovipositor lobes is somewhat geometrine, and the male genitalia have geometrine aedeagus and socii, with valve ornamentation similar to that of Spaniocentra Prout, associated with the group Rhomboristini. (Geometrini).

HEMISTOLINI: Hemistolini Inoue (1961). See (Geometrini) for discussion of status within the Geometrinae.

HEMITHEINI: Hemitheidae Bruand (1846). This name has been used as subfamily-group name for the Geometrinae as recognised here (e.g. Prout, 1912a) and more recently at tribal level (e.g. Ferguson, 1985). See (Geometrini) for discussion of its status.

JODINI: Jodiini Inoue (1961). See (Geometrini) for discussion of status within the Geometrinae.

LOPHOCHORISTINI: Lophochoristini Ferguson (1969, 1985). Male genitalic features suggest that N. American taxa in this group may be related to Spaniocentra Prout and the Rhomboristini: (Geometrini).

[LYRCEINI]: Lyrceini Meyrick (1883 [4]). This is based on Lyrcea Walker, a junior homonym of a name outside the Lepidoptera. Meyrick's concept refers to Xyridacma Prout, a New Zealand genus currently in the Oenochrominae.

MECOCERATINI: Mecoceridae Guenée (1857); Mecoceratinae Hulst (1896). Mecoceras Guenée is a junior objective synonym of Ametris Hübner: see Ametridini.

MONOCTENIINI: Monocteniadae Meyrick (1889). This name embraces robust Australian taxa that belong to the true Oenochrominae.

NEMORIINI: Nemorinae Gumppenberg (1887); Ferguson (1969, 1985); Pitkin (1993). This geometrine group is represented in the Old World by the Ochrognesiini (Geometrini and below): the two should be treated as synonymous, with Nemoriini having priority (Pitkin, in press).

NEOHIPPARCHINI: Neohipparchini Inoue (1961). See (Geometrini) for discussion of status within the Geometrinae.

OCHROGNESIINI: Ochrognesiini Inoue (1961). See Nemoriini (below) and Geometrini.

OENOCHROMINAE: Oenochromidae Guenée (1857). See Geometrini for discussion of a more restricted concept of this subfamily.

ORTHOSTIXINI: Orthostixidae Meyrick (1892); Seven (1991). This group has characteristics suggestive of relationship to the Ennominae: see p. (Orthostixini).

OXYDIINI: Oxydiidae Butler (1886). This Neotropical group is based on taxa with an ennomine signum in the female, and male genitalia with valve and furca characteristics reminiscent of Ourapterygini: the type genus was placed in Ourapterygini by Hodges et al. (1983). See also Diptychini above.

PHASELIINI: Phaseliinae Wehrli (1941; Gross-Schmett Erde 4 (Suppl.): 466). Wiltshire (1990) placed this group as a tribe within the Ennominae.

PINGASINI: Pingasini "Inoue" (1992), in Heppner & Inoue (1992). This appears to be the first usage of this name: see note in Holloway (1993[4]: 302) concerning the doubtful validity of some tribal nomenclature in this publication.

PSEUDOTERPNINI: Pseudoterpninae Warren (1893); Seven (1991). See Geometrini for discussion of status within the Geometrinae.

RHOMBORISTINI: Rhomboristini Inoue (1961). See Geometrini for discussion of status within the Geometrinae.

SYNCHLORINI: Synchlorini Ferguson (1969, 1985). See Geometrini for discussion of status within the Geometrinae and relationship to Comibaenini.

TERPNINI: Terpnini Inoue (1961). Terpne Hübner is a synonym of Geometra Linnaeus. Inoue's concept refers to Pachyodes Guenée (Terpna auctorum): see Pseudoterpnini and Geometrini .

THALASSODINI: Thalassodini Inoue (1961). See Geometrini for discussion of status within the Geometrinae.

THALERINI: Thalerini Leraut (1980), who included Hemithea Duponchel within the concept. See also Chlorochromini and Geometrini for discussion of status within the Geometrinae.

TIMANDROMORPHINI: Timandromorphini Inoue (1961). See Geometrini for discussion of status within the Geometrinae.

The higher classification of the Geometrinae and Oenochrominae sensu lato
Scoble & Edwards (1989) reviewed the past status and extent of the Oenochrominae and recommended a return to something approaching Guenée's original concept of the group, embracing only the type genus and its robust-bodied Australasian relatives. Their definition of the Oenochrominae sensu stricto is followed here, examined in more detail on Oenochrominae, in relation to what appears to be the only Oriental representative of the group, the genus Sarcinodes Guenée.

This poses the problem of the classificatory position of all the other Bornean genera traditionally placed in Oenochrominae. Inoue (1971) placed Eumelea Duncan in Ennominae and Derambila Walker and Ozola Walker in the Larentiinae, but has since reverted to the traditional concept of Oenochrominae (in Heppner & Inoue, 1992).

The genera Heteralex Warren and Naxa Walker appear to be better placed in the Ennominae, despite having a strong vein M2 in the hindwing. Naxa may be best associated with the Orthostixini family group. Both genera are discussed on Heteralex Warren et seq..

Holloway (1984b) recognised and defined a major pantropical grouping of slender-bodied taxa, a concept that he extended (Holloway, 1993 [4]: 10) to include also Celerena Walker. The addition of Ozola Walker and the Derambila Walker group of genera to this grouping (Desmobathrini) means that Desmobathridae of Meyrick, based on a synonym of Ozola, provides an available family-group name for the ensemble.

The genus Eumelea Duncan, though of similar build to the previous group and with a similar habit of frequenting the forest understorey, particularly where disturbance has occurred, and resting on the undersides of leaves, lacks the definitive features of the desmobathrines and exhibits several unique ones itself. The family-group name Eumeleinae of Warren is available for it.

A final consideration must be the position of the genus Dysphania Hübner (with Cusuma Moore) in relation to the rest of the Geometrinae. In a similar manner to Eumelea it lacks several features that appear to be in the ground plan of the rest of the Geometrinae, and shows several unique ones itself. The family- group name Dysphaniinae of Warren (= Hazidae of Guenée, Euschemidae of Walker, based on junior synonyms of Dysphania) is available.

There is the option of treating all these groups as full subfamilies. In the absence of any satisfactory synapomorphies (shared, uniquely derived characters) linking any of them together, this might be the preferred option, though it yields monogeneric subfamilies for Eumelea and Dysphania.

A survey of morphological features, particularly of the abdomen, has revealed a number of potential synapomorphies, though none is entirely convincing. There is also one chemical one, the presence of unusually high concentrations of the pigment geoverdin, the basis of the green colour of the Geometrinae, in Dysphania and Celerena as well as the bulk of the geometrines (Cook et al., 1994). Such a concentration was not recorded from Eumelea by these authors in a sample of 30 lepidopteran taxa of which 24 were Geometroidea sensu lato.

Cook & Scoble (1992) surveyed tympanal structure in a wide sample of geometrid taxa. They noted great heterogeneity of morphology in the broader concept of the Oenochrominae, and found one feature to support the inclusion of Sarcinodes in a more strict definition. They suggested that the Geometrinae could be defined by a characteristically shaped ansa (Dysphaniini), a definition that would exclude Dysphania where it is hammer-headed, and lacking a central expansion.

The ansa in most Oenochrominae sensu lato is tapering from base to apex, a condition suggested to be primitive by Cook & Scoble, but that in Eumelea is long, slender, apically hammer-headed, with a small circular central expansion. The desmobathrine group have the ansa generally tapering, occasionally with some central expansion (Derambila, Alex) and with a slight hammer-head in Alex.

The ornamentation of male abdominal sternite 3 consists of a pair of setal patches in both the geometrines sensu stricto and the desmobathrines, though in a few genera of each (e.g. some Derambila, Berta Walker) there is a single central patch, though always with setae dispersed rather than in a tight transverse comb as in many tribes of Ennominae. Eumelea lacks setae (as do a number of geometrine genera) but Dysphania has a central cluster, more concentrated than in those desmobathrines and geometrines with a broad central patch, though not as much as in Ennominae.

The presence of a forewing fovea in Dysphania sets it apart from the geometrines. This feature is present in diverse forms in several tribes of Ennominae and in one desmobathrine genus. Only in Dysphania is it present in both sexes.

Both Eumelea and the desmobathrines have the tegumen of the male genitalia diagnostically modified, though in different ways: the apparent lack of homology means that treatment of such modifications as a synapomorphy is difficult to justify.

One feature of Dysphania that may indicate a relationship with the Geometrinae is the somewhat cruciform structure of the vinculum. (see Dysphaniini, Geometrini)

However, these adult features taken together do not yield any clearly unambiguous sister-relationships amongst the groups involved.

Evidence from early stages is sparse, and, as has happened elsewhere in the Lepidoptera, larval characters may hold the key to resolving these higher classificatory problems. Fresh, spirit-preserved material of larvae will be necessary for a study of chaetotaxy, and this is not currently available (nor is dried material) for desmobathrines and eumeleines. Descriptions available in the literature and in manuscript (e.g. of T.R.D. Bell) are not specific enough to facilitate setal comparisons.

There is some information on the pupal cremaster in the various groups, providing an indication of relationship between the eumeleines and desmobathrines. In both Ozola (see Ozola Walker) and Eumelea (see Eumelea Duncan) the number of hooked shaftlets is reduced to four from the geometrid ground plan number of eight (Holloway, 1993[4]). In Dysphania and the majority of geometrines there are eight, and the Oenochrominae sensu stricto have the number reduced to two. The cremaster is disc-like in Ozola, semicircular in Eumelea.

Thus there are a number of rather unsatisfactory pointers to a sister-relationship between the desmobathrines and Eumelea: the cremaster; modification of the tegumen; slender build; habitat and behaviour. It would be a conservative policy also to retain the association between the bulk of the geometrines and Dysphania. These two pairings may also be sister groups, potential synapomorphies being a high concentration of geoverdin pigment in at least some members of each group and the presence of a pair of setal patches on the male third sternite in many genera in each.

It is therefore suggested here that two subfamilies can be recognised, Desmobathrinae and Geometrinae, each consisting of two tribes: Desmobathrini + Eumeleini; Geometrini + Dysphaniini. Authors such as Inoue (1961), Ferguson (1969, 1985) and Viidalepp (1981) have proposed a number of tribal groupings for the Geometrinae that would have to be placed at subtribe level under this system. This will be discussed further in the introductory section to the Geometrini (see Geometrini).