The Ennominae are the largest subfamily of the Geometridae. In the Bornean fauna just under half the species are ennomines. The traditionally diagnostic feature for the subfamily is loss (or reduction to a fold) of vein M2 on the hindwing. No other features have been located that are shared by all major tribal groupings within the Ennominae, so the monophyly of the group is only weakly supported.

The group as a whole has a wide ecological range, occurring in diversity at all except very high latitudes and, in the tropics, altitudes. In Borneo, Sulawesi and Seram, ennomines show their greatest diversity in the lower montane zone (Holloway, 1984a, 1993; Holloway, Robinson & Tuck, 1990).

Some ennomines show specificity to lowland forest types, such as heath forest, alluvial forest or mangrove. The subfamily also includes several species that appear to fly predominantly in the forest understorey. Ennominae are therefore highly suitable as an environmental indicator group except for poor representation in open habitats (Holloway, 1984a, 1985; Holloway & Barlow, 1992).

An attempt is made here to define tribal groupings and suggest a higher classification for the subfamily, though this is based primarily on groups represented in S.E. Asia. Host-plant specialism is also reviewed in relation to this classification.

Family-group names
The attempt to assess the higher classification of the Ennominae and to assign genera to tribal groupings necessitated an investigation of prior usage of family- group names. This in turn led to the accumulation of about 150 family-group names used for the Geometridae and some indication of when these names first appeared. For the subfamilies Oenochrominae, Geometrinae, and Sterrhinae the listings of Prout (1912, 1913, 1934-5) provide a foundation, though a number of family-group names have been erected subsequently, e.g. in the Geometrinae by Inoue (1961) and Ferguson (1969). Prout did not publish lists for the Larentiinae and Ennominae.

In the Code of Zoological Nomenclature, rules of priority for family-group names tend to apply, but somewhat less strictly than for genus-group and species-group names. After 1960, family-group names based on junior generic synonyms may not be replaced by a new name based on the senior generic synonym, but can be replaced by extant family group names based on senior generic synonyms or by senior family group names based on other genera within the concept of the reclassification. Before 1961, such replacements can stand if the replacement name has won general acceptance. Thus Bistonini should probably stand in relation to the older Amphidasini and Hyberniini (replaced by Erannini), though Seven (1991) uses Amphidasini (see list below for details). Reinstatement of the genus Macaria Curtis from synonymy with Semiothisa Hübner strengthens the case for preferring Macariini to Semiothisini (as does Seven), as Macariini is the senior name. 

It became apparent during this investigation that reclassification of the Sterrhinae, a subfamily name indicated by Fletcher (1979) to be preferred, will probably lead to even greater changes to family-group names. Indeed, Mikkola et al. (1985) and Seven (1991) have reverted to using Scopulinae for this subfamily based on the oldest name (Scopulites, Duponchel (1845)). The situation is therefore reviewed briefly here to exemplify the problems that can be encountered. Revision of the classification may show that the group is polyphyletic rather than monophyletic: even if monophyletic, it may contain seven groupings:

Scopulini (= Acidaliini Duponchel (1845) based on a junior homonym of a genus-group name outside the Lepidoptera). Scopula and allied genera;

Cyclophorini Moore (1887) (= Zonosomini White (1876, Scot. Nat. 3: 361), reintroduced probably on priority grounds by Seven (1991) but Cyclophora is a senior objective synonym of Zonosoma; Cosymbiini Prout (1911, Tr. City Lond. ent. soc. 20: 23)), which traces back to the Cyclophorae of Hübner. The name Ephyrini of Guenée (1857) is based on a junior homonym of a genus group name outside Lepidoptera but was applied to the concept of Cyclophorini. Another reason for preference of Zonosomini is the usage of the family-group name Cyclophoridae in Mollusca, based on Cyclophorus Montfort. The moth group includes genera with a distinctive girdle to the pupa (Holloway, Bradley & Carter, 1987);

Idaeini Butler (1881) (= Sterrhini Meyrick (1892), a seniority that must stand both on grounds of priority and because Sterrha Hübner is currently treated as a junior subjective synonym of Idaea Treitschke);

Timandridi Stephens (1850) (= Calothysanini Herbulot (1962-3)). This name is used by Hodges et al. (1983) and Seven (1991);

Rhodometrini Agenjo (1952);

Cyllopodini Warren (1895);

Rhodostrophini Prout (1934-5).

The family-group names (over 60) listed below have been located for possible usage within the Ennominae. Invalid names, based on junior homonyms, usually of genus-group names outside the Lepidoptera, are indicated by square brackets.

The literature references given are only of those considered to indicate both early usage and currency in modern literature. Whilst an attempt has been made to ensure the earliest reference has been located, this cannot be taken as categorical as the search was not exhaustive. The task is considerably complicated by both past and present reluctance of authors to indicate authority of the family-group names they use or to own up to new concepts.

However, the earliest dates located for names within Oenochrominae, Geometrinae and Sterrhinae up to the Catalogus Lepidopterorum publications of Prout (1912, 1913, 1934-5) were found to be the same as in Prout’s lists. Many of Duponchel's names are attributed to Guenée, but represent the first published occurrence.

ABRAXINI: Abraxinae Warren (1893); Swinhoe (1900), Hodges et al. (1983), Seven (1991). (See Abraxini.) Conceptually replaced the unavailable Zerenini, and see also Pantherini (q.v.).

AMPHIDASINI: Amphidasites Duponchel (1845); Bruand (1846), Guenée (1857). Amphidasis Treitschke is a junior objective synonym of Biston Leach (Fletcher, 1979). Used for concept of Bistonini by Seven (1991). (See Boarmiini.)

ANAGOGINI: Anagogini Forbes (1948); Hodges et al. (1983). Subordinated here to Hypochrosini.

ANGERONINI: Angeronini Forbes (1948); Hodges et al. (1983). Applicable to concept of Aspilatini (q.v.) as Aspitates and Angerona share a costal ampullate process to the valve and other features of the male genitalia (see McGuffin (1987)) and Higher Classification.

ASCOTINI: Ascotinae Warren (1893). Included in the strict concept of Boarmiini of Sato (1984a).

[ASPILATINI]: Aspilatites Duponchel (1845); Bruand (1846). Based on a misspelling of the genus-group name Aspitates Treitschke (see Fletcher (1979)). The concept is currently included in Angeronini (q.v.).

AZELININI: Azelinini Forbes (1948); Hodges et al. (1983). Larval characters suggest a relationship to Nacophorini (Heitzmann, 1985).

BAPTINI: Baptini Forbes (1948); Hodges et al. (1983) include the older name Palyadini (q.v.) within their concept, but see discussion of Baptini.

BISTONINI: Bistonidi Stephens (1850); Humphreys (1858-9), Hodges et al. (1983) but see Amphidasini and Boarmiini, Biston Leach.

BOARMIINI: Boarmites Duponchel (1845); Bruand (1846), Stephens (1991). (See Boarmiini.)

BRACCINI: Braccinae Warren (1894). (See Boarmiini.)

BUPALINI: Bupalini Herbulot (1962-3); Leraut (1980). Not investigated here, but the bifid cremaster indicates relationship to the Boarmiini group of tribes.

CABERINI: Caberites Duponchel (1845); Bruand (1846), Stephens (1850), Guenée (1857), Hodges et al. (1983), Seven (1991). See also Deiliniini (q.v.), and concept may also incorporate Erastriini and Catopyrrhini (q.v.). (See Tribe Caberini).

CAMPAEINI: Campaeini Forbes (1948); Hodges et al. (1983), Seven (1991). Metrocampini (q.v.) is an older name but based on a junior objective synonym of Campaea Lamarck (Fletcher, 1979). 

CASSYMINI: This volume. May embrace the N. American (Forbes, 1948) concept of Abraxini (See Tribe Cassymini).

CATOPYRRHINI: Catopyrrhinae Warren (1894). See Erastriini (q.v.) and Tribe Caberini.

CHEIMATOBIINI: Cheimatobiidi Tutt (1896). See Theriini (q.v.). Cheimatobia Stephens is a junior objective synonym of Theria Hübner.

CINGILIINI: Cingiliini Forbes (1948). Some usage in N. America but generally subordinated to Ourapterygini (q.v., Forbes et al. (1983)). The relationship between the two needs clarification; larval characters indicate a close relationship with the Ennomini (Heitzmann, 1985).

CLEORINI: Cleorites Duponchel (1845); Stephens (1850). Included in the strict concept of Boarmiini of Sato (1984a).

COLOTOINI: Colotoini Herbulot (1962-3); Seven (1991).

COMPSOPTERINI: Compsopterini Herbulot (1962-3); Leraut (1980). See Ligiini, Pachycnemiini and Prosopolophiini (q.v.). Status not investigated here. The type genus includes a number of robust Palaearctic species.

CROCALLINI: Crocallidi Tutt (1896). No current usage located.

DASYDIINI: Dasydites Duponchel (1845); Stephens (1850), Tutt (1896). Based on a junior objective synonym of Sciadia Hübner. Concept not investigated here.

DEILINIINI: Deiliniinae Warren (1893). (See Tribe Caberini).

EMPLOCIINI: Emplocidae Guenée (1857). Based on a Neotropical genus; not investigated here.

ENNOMINAE: Ennomites Duponchel (1845); Bruand (1846), Guenée (1857), Hodges et al. (1982), Seven (1991). See also Odopterini (q.v.) and Cingiliini.

EPIONINI: Epionidae Bruand (1846); Stephens (1850), Humphreys (1858-9).  

EPIRRHANTHINI: Epirrhanthini Forbes (1948); Hodges et al. (1983). Not investigated. McGuffin (1981) notes some similarities with Lithinini (q.v.). A small N. American group.

ERANNINI: Eranniinae Tutt (1896); Dugdale (1961). See also Hyberniini (q.v.). Hybernia Berthold is a junior objective synonym of Erannis Hübner (Fletcher, 1979). Tutt included Eranniinae as a subfamily of Hyberniidae! Taxa usually included in modern concepts of Bistonini/Amphidasini.

ERASTRIINI: Erastriae Hübner (1825); Erastridae Herrich-Schäffer (1845); Stephens (1850). The original concept was applied to the acontiine Noctuidae for a considerable time. Classification of the generic type species (see Fletcher, 1979) leaves this family-group name applicable within the Ennominae. See Cryptopyrrhini and Tribe Caberini.

EUBYJINI: Eubyjinae Warren (1893). Eubyja Hübner is based on a species currently included in Biston Leach. See Amphidasini and Bistonini.

EUTOEINI: This volume; (See Eutoeini).

FERNALDELLINI: Fernaldellinae Hulst (1896) Based on a N. American genus currently placed within the concept (Hodges et al. 1983) of the Macariini/ Semiothisini (q.v. and see Macariini).

FIDONIINI: Fidonites Duponchel (1845); Bruand (1846), Stephens (1850), Guenée (1857). Has been used to include Macariini/Semiothisini subsequently (Packard, 1876) but is based on an unrelated S. European taxon. (See Macariini). The name has not been used by Herbulot (1962-3) or Leraut (1980).

GNOPHINI: Gnophites Duponchel (1845); Bruand (1846), Stephens (1850), Seven (1991). Not investigated in detail but probably subsumed by the concept of Boarmiini in this volume.

GONODONTINI: Gonodontini Forbes (1948); Hodges et al. (1983). Applied in the sense of Odontoperini (q.v.). Redefined in this volume (See Gonodontini).

HYBERNIINI: Hibernites Duponchel (1845); Bruand (1846), Stephens (1850), Guenée (1857). See Erannini. Falls within current concepts of Amphidasini/ Bistonini.

HYPOCHROSINI: Hypochrosidae Guenée (1857); Kirby (1897). See Anagogini and (See Hypochrosini).

LACARIINI: Lacarini Orfila & Schajovskoy (1959). Associated with Lithinini by these authors and Rindge (1986). (See Lithinini).

[LIGIINI]: Ligidae Guenée (1857). Based on a junior homonym of a genus group name outside Lepidoptera. See Compsopterini, Pachycnemiini and Prosopolophini (q.v.).

LITHININI: Lithinini Forbes (1948); Hodges et al. (1983). (See Lithinini).

MACARIINI: Macaridae Guenée (1857); Moore (1887), Swinhoe (1900), Seven (1991). See Semiothisini (q.v.), Fernaldellini and Macariini.

MELANCHROIINI: Melanchroiinae Hulst (1896); Kirby (1897). Applicable to a small New World group; not investigated here.

MELANOLOPHINI: Melanolophini Forbes (1948); Hodges et al. (1983). A New World group that probably falls within the broad concept of Boarmiini applied here.

METROCAMPINI: Metrocampidae Tutt (1896). See Campaeini.

MILIONIINI: Milioniini Inoue (1992) is the only reference so far located. In the absence of a published definition, or mention before 1930, the name is not available. The type genus falls within the concept of Boarmiini here (See Boarmiini).

NACOPHORINI: Nacophorini Forbes (1948); Hodges et al. (1983). A New World group outlined by Rindge (1961). The tribe is possibly related to the Azelinini (see above).

NEPHODIINI: Nephodiinae Warren (1894). A Neotropical group not studied here.

OBEIDIINI: Obeidiini Inoue (1992) is the only reference so far located. In the absence of a published definition, or mention before 1930, the name is not available. The type genus falls within the concept of Boarmiini here (See Boarmiini ).

ODONTOPERINI: Odontoperinae Tutt (1896). Fulfils the N. American concept of Gonodontini (see above and (See Gonodontini).

[ODOPTERINI]: Odopteridi Stephens (1850), Humphreys (1858-9). Odoptera Sodoffsky is an unnecessary objective replacement name of Ennomos Treitschke. See Ennominae.

ONYCHORINI: Onychorini Herbulot (1962-3); Leraut (1980). Based on W. Palaearctic genus not studied here.

OURAPTERYGINI: Urapteridae Bruand (1846); Guenée (1857), Warren (1893), Hampson (1918; preferred to Ennominae for subfamily, as based on the oldest genus-group name), Forbes (1948; ?first spelling as Our- rather than Ur-), Hodges, et al. (1983). See Cingiliini and Ourapterygini.

PACHYCNEMIINI: Pachycnemiidae Kirby (1903); as replacement name for Ligidae. See above and also Compsopterini and Prosopolophini.

PALYADINI: Palyadae Guenée (1857); Moore (1887), Warren (1894), Hulst (1896), Kirby (1897). Subordinated to Baptini in Hodges et al. (1983) but see Baptini. Guenée's concept of this group is valid today if Eumelea Duncan is excluded (M.J. Scoble, pers. comm.).

[PANTHERINI]: Pantheridae Moore (1887), as replacement for Zerenini (q.v.) but Panthera Hübner is a junior homonym of a genus-group name outside the Lepidoptera (Fletcher, 1979). (See Abraxini.)

PLUTODINI: Plutodinae Warren (1894); Swinhoe (1900). (See Plutodini).

PROSOPOLOPHINI: Prosopolophinae Warren (1894); Swinhoe (1900). Proposed to replace Ligidae (see above, and also Compsopterini and Pachycnemiini).

RUMIINI: Rumiinae Tutt (1896). Based on Rumia Duponchel, a junior objective synonym of Opisthograptis Hübner. No current usage located.

SCARDAMIINI: Scardamiinae Warren (1894). (See Scardamiini).

SELENIINI: Seleniidi Tutt (1896). Probably best subsumed by Ennomini; (See Introduction).

SELIDOSEMINI: Selidosemidae Meyrick (1892); Warren (1893). Based on a European and N. African genus not studied here. The concept of earlier authors probably coincides with Boarmiini as treated here.

SEMIOTHISINI: Semiothisinae Warren (1894); Hodges et al. (1983). See Macariini and Fernaldellini.

SIONINI: Sionites Duponchel (1845); Bruand (1946), Stephens (1850), Guenée (1857). Based on a European species included in Gnophini by Herbulot (1962-3).

SPHACELODINI: Sphacelodini Forbes (1948); Hodges et al. (1983). A New World group not investigated here.

THERIINI: Theriini Herbulot (1962-3); Leraut (1980). See Cheimatobiini. This is based on a Palaearctic genus that, from pupal cremastral characters, probably falls within the broad concept of Boarmiini applied here.

THINOPTERYGINI: This volume. (See Thinopterygini )

[ZERENINI]: Zerenites Duponchel (1845); Bruand (1846), Stephens (1850), Guenée (1857). Based on a junior homonym of a genus-group name outside the Lepidoptera. See Abraxini and Pantherini.

The higher classification of the Ennominae

An attempt has been made in the systematic section to group Bornean genera of ennomines into tribes. In some instances (e.g. Caberini, Baptini) it was possible to recognise a major generic grouping defined by several characters, but association of this grouping with the type genus of the tribe was more tenuous.

The absence as a tubular vein or weakening of vein M2 in the hindwing currently defines the subfamily. In this section potential groupings of tribes within it will be assessed.

Forbes (1948) divided N. American tribes on characters of the pupal cremaster: whether it was apically bifid only (Group D of Mosher (1916)) or with more than two spines (usually eight: four pairs). McGuffin (1981) suggested that the bifid condition was the more primitive, reversing an earlier opinion which, on surveying cremaster characters in Geometroidea generally, was probably correct: the bifid condition is derived.

Data on geometroid pupae from Forbes, McGuffin, Mosher (1916), Sato (1984a), Carter (1984), Khotko (1977) and the manuscript notes on the Indian fauna by T.R.D. Bell indicate that possession of eight cremastral hooked shafts is the commonest and more generally distributed condition seen in the Cyclophorini, Naxa Walker amongst the Oenochrominae (?Orthostixini), the majority of Geometrinae and Larentiinae, and also in the Drepanidae, Uraniidae and Epiplemidae.

The bifid condition is not unique to the Ennominae but is much less frequent elsewhere. It occurs in some, but not all, Scopulini where the cremaster consists of two parallel shafts rather than divergent ones. The Alsophilini have the cremaster bifid, almost T-shaped as does Operophtera (but not Epirrita, where the 'T' is associated with two pairs of reduced hooklets) of the larentiine tribe Operophterini (Packard, 1876; = Oporiniini of Seven (1991)) (Carter, 1984). In the Eupitheciini all conditions occur from eight shaftlets, through two strong ones and six weaker ones, to two shaftlets; the Epiplemidae show a similar range of conditions. The bifid condition is seen in some other larentiine genera such as Aplocera Stephens and Lithostege Hübner. In the Archiearinae the cremaster resembles that of Alsophila (Mosher, 1916; Nakamura, 1987).

Hence it is probable that, within the Ennominae, the divergently bifid cremaster represents a derived condition. It brings together the Boarmiini sensu lato (See Boarmiini) with the Macariini, Cassymini and Eutoeini. The inclusion of Abraxini by Forbes rests on his examination of N. American genera he assigned that tribe but which are probably better placed in Cassymini (See Cassymini n.).

All the tribes with a bifid cremaster have the male forewing foveate in a high proportion of genera, although the fovea is structurally different in each (Fig 1). Occurrence of a forewing fovea is rare elsewhere in the Ennominae, e.g. in Corymica Walker of the Hypochrosini and Borbacha Moore, tentatively assigned here to the Baptini. Debauche (1937) defined a subgeneric taxon in the Palyadini on presence of a fovea, somewhat as in Borbacha, but with a spur from vein CuA as in the Macariini. Outside the Ennominae a fovea occurs in Noreia venusta Warren, uniquely in a pantropical complex of genera currently in the Oenochrominae that was identified by Holloway (1984b: 139). This complex must now be expanded to include the Indo-Australian genus Celerena Walker as it shares the definitive features of the male abdomen. The geometrine genus Dysphania Hübner also has a strong fovea, but again structurally different from ennomine types.

Nevertheless, the frequency of the foveate condition supports the relationship of these ennomine tribes suggested by the pupal character. A further character that may serve to unite them is the presence of a cucullate anterior apical portion of the valve that supports a concentration of the valve setae; these setae extend more sparsely down the costal zone of the valve. In the Macariini and Eutoeini this zone is separated from the sacculus as the dorsal arm of the valve by a c!eavage that extends almost to the base of the valve. In the Cassymini the valve is similarly cleft but the dorsal arm is reduced to a slender process that usually only bears a few apical setae. In other tribes the setae tend to be more generally and sparsely distributed over the anterior of the valve lamina or, as in the Palyadini and a generic grouping in the Baptini, concentrated elsewhere on the valve. However, not all Boarmiini exhibit this cucullus.

The division of the valve may provide a synapomorphy to group the Cassymini, Eutoeini and Macariini, but it is not clear whether the mode of cleavage is homologous or homoplasious: it appears somewhat different in each case.

Another feature common to all four tribes is the presence in many genera of a transverse comb of posteriorly directed setae on the male third abdominal segment (e.g. Figs 232, 251, 296-8, 349, 369, 494). This appears often to be correlated with the presence of a hair pencil sheathed within the hind tibia (e.g. Rindge, 1983) and may serve as a means of distributing scent from that pencil. But this feature is also seen in other tribes, e.g. for the Bornean fauna: Abraxini, Gonodontini, Lithinini, Ourapterygini.

Setal ornamentation is seen elsewhere in the Geometridae but not in the form of a transverse comb. In the Noreia/Celerena group of the Oenochrominae mentioned earlier the setae occur in two robust patches arranged symmetrically on either side of the mid-line, separated by a group of finer setae forming together a roughly circular area (Celerena) or in two sparser, more widely separated patches with no setae in between (Noreia Walker). In the Geometrinae there is usually a diffuse patch on each side linked by a region with sparser setae in between. These are more easily detached than in Ennominae, except in Pingasa Moore and allied genera where they are more robust.

In the Noreia group and Ozola Walker (Desmobathrini) in the Oenochrominae and in the Geometrinae, occurrence of a tibial hair pencil is frequent but the transverse comb arrangement of the sternal setae appears to be unique to the Ennominae. The occurrence of a hair pencil/setal connection in groups outside the subfamily makes it unclear whether the comb is a feature of the Ennominae ground plan, and therefore its consistent absence in a tribe is a derived feature, or whether it can be treated as a synapomorphy for the tribes that possess it, i.e. grouping the Abraxini, Gonodontini, Lithinini and Ourapterygini with the bifid cremaster complex of tribes.

Rindge (1983) treated presence of setal comb and hair pencil as apomorphic states within his concept of the Nacophorini, a New World tribe with some representation in Australia (McQuillan, 1981) that has variable pupal cremastral ornamentation from two to eight shaftlets, and was assigned by Forbes (1948) to his non-bifid group of tribes.

McQuillan (1985) revised the Australian ennomine genus Mnesampela Guest, where the larvae have a full set of prolegs albeit with those on A3 somewhat reduced; it was suggested by McQuillan to be one of a number of rather generalised Australian taxa. The pupa has four cremastral hooks and the male genitalia have socii and a furca (see below). There is a setal comb on the male third sternite and a hind-tibial hair pencil. This may indicate that the hypothesis of presence a setal comb being a relatively basal feature in the ennomine phylogeny and subsequent loss being the derived state is correct.

Consistent absence of a setal comb throughout a tribe, perhaps associated with loss of a tibial hair pencil, might serve to unite a number of robust-bodied tribes around the Ennomini, including here the Hypochrosini and Scardamiini. These share with the Epionini extension and looping of the vinculum to support lateral coremata. Other tribes recognised in the Palaearctic and N. American faunas may associate with such a group, such as the Angeronini, Colotoini, Crocallini and Seleniini.

Many of these groups have a strong furca arising between the bases of the valves in the male genitalia, but a similar feature is seen in Ourapterygini where a setal comb is present, and some Boarmiini. Anellar processes, or processes from the juxta, probably not homologous with the furca in these tribes, occur in the Lithinini and Nacophorini.

Presence of socii associated with the uncus is probably a primitive feature as they also occur in Geometrinae. In the Ennominae they are much reduced or absent in the 'bifid cremaster' group of tribes and also the Abraxini, Thinopterygini and Gonodontini.

In the female genitalia a single, often mushroom-shaped, usually spined or dentate signum is more or less general to the subfamily (Fig 2). It is modified into a ridged structure in Lithinini. More general scobination with no signum is seen in the Plutodini and the Campaeini (or Metrocampini).

Study of the larvae of Ennominae has probably not yet been comprehensive enough to provide any clear pointers to higher classification. Sato (1984a) suggested that presence of four external setae on the A6 proleg was apomorphic for his Hypomecis group, but this characteristic was regarded as diagnostic for the subfamily relative to Geometrinae by Singh (1953), though not exclusive to it, and is a feature of the putative primitive ennomine genus Mnesampela (McQuillan,1985). In Geometrinae the proleg has three external setae.

Dugdale (1961) compared New Zealand ennomine larvae with published accounts from the Nearctic (earlier papers by McGuffin) and India (Singh, 1953). He identified four characteristics common to ennomines treated in these works and possibly diagnostic: (1) anal shield with seta SD1 anterior to seta D1; (2) hypoproct produced to a fine point, strongly developed; (3) four to eight annulets (transverse sclerotised bands that are muscle attachments) on A1 -5; (4) SV group tri- (to 6) setose on A2-5 (cf. bisetose).

Heitzman (1985) performed cladistic analyses on 70 larval characters for genera of Nearctic Ennomini and related tribes. He found no apomorphies for the Ennominae as a whole but defined the Ennomini (including the Cingiliini and the N. American concept of Ourapterygini) on possession of the V2 primary seta. He noted more than four external SV setae on the A6 proleg in Angeronini, Epirranthini and Seleniini as well as in the groups referred to below.

Heitzman (1985) and Poole (1987) indicated that the Azelinini (Pero Herrich-Schäffer) and Nacophorini shared a unique arrangement of multiple setae on the proleg on A6 (he illustrated 16 setae). Singh recorded up to 16 setae on the proleg of Odontopera similaria Moore (as Gonodontis) so the Odontoperini may be related to the other two tribes. The Lithinini appear to be characterised by the presence of six external setae (Heitzman, 1985; J. Weintraub, pers. comm.)

The Scardamiini and Hypochrosini studied by Singh had five external setae on the proleg, support for the relationship of these groups suggested above from consideration of the vinculum modification. Five external setae are found also in the larva of Traminda Warren in the Sterrhinae (Singh). The group of genera including Anagoga indicated by Heitzman to be closely related are referable to Hypochrosini. His other anagogine genera may be misplaced.

Salkeld (1983) surveyed the eggs of Canadian Geometridae. Amongst the 'bifid cremaster' grouping, particularly the Boarmiini sensu lato, there is a tendency for the walls of the polygonal cells ornamenting the surface to be heavy and for the cells to be aligned in longitudinal rows. In the putative Cassymini ('Abraxini') the wavy polygonal cell walls are overlain by a prominent reticulum, reminiscent of some Larentiinae. The Macariini appear to be a homogeneous group, with eggs of a rugged appearance, the polygonal cells with broad and elevated walls, sometimes enhanced by large, domed aeropyles at the wall junction and a roughly pitted texture to the chorion.

A columnar arrangement of polygonal cells, leading to the walls separating adjacent rows forming longitudinal ridges is seen in Lithinini and Caberini. In the former the eggs are not attached to the substrate.

A thick rolled collar encircling the anterior pole is recorded from several genera in the Ennomini, Hypochrosini (as Anagogini) and Ourapterygini (including Cingiliini).

The features of adults and early stages just discussed suggest phylogenetic relationships amongst ennomine tribes somewhat different to those suggested by McGuffin (1987). His basal groups can be brought together if the bifid cremaster is considered derived, supported by fovea and valve cucullus features, as well as loss of socii. This would include a broad concept of Boarmiini (embracing Bistonini and probably Melanolophiini) together with Macariini, Cassymini (his Abraxini) and Eutoeini (an Old World group defined here). True Abraxini and Gonodontini may be related to that group (absence of socii), and Thinopterygini can also be tentatively placed here, though the setal comb has been lost.

Loss of setal comb, presence of a strong furca and occurrence of collared eggs could bring together Hypochrosini (Anagogini), Scardamiini, Epionini and perhaps Ennomini, Angeronini, other small Holarctic tribes, with Ourapterygini (+ Cingiliini) also associated though a setal comb is present. The first two tribes group on larval characters, and the others also share possession of additional SV setae on the A6 proleg, a feature developed to an extreme in the next group.

The Nacophorini, Azelinini and Odontoperini appear to form a group on the basis of larval characters. None is represented in Borneo.

The Lithinini and Caberini may group on features of the egg. The Baptini and Plutodini appear relatively isolated, though a grouping in the former shows parallels with the Neotropical Palyadini.

The tribes are therefore treated in the systematic section in the order: Hypochrosini, Scardamiini, Ourapterygini, Baptini, Plutodini, Lithinini, Caberini, Thinopterygini, Gonodontini, Abraxini, Cassymini, Eutoeini, Macariini, Boarmiini (sensu lato). The assignation of genera to these tribes is based as far as possible on an appreciation of shared, derived features (synapomorphies): it therefore differs in many respects from the more traditional arrangement of Inoue (1992).

Host-plant specialisation

The majority of ennomines are arboreal feeders as larvae. Host-plant specialisation ranges from extreme polyphagy in some tribes to restriction to a particular plant family in others.

The Hypochrosini are broadly polyphagous but with some specialisation at a generic level, e.g. Heterolocha Lederer on Caprifoliaceae and the sister-genera Fascellina Walker and Corymica Walker on Lauraceae and Illiciaceae. The few records for Achrosis Guenée are from the Rubiaceae.

The Scardamiini show some association with Flacourtiaceae.

The Ourapterygini are polyphagous.

In the Baptini, the typical genus Lomographa Hübner is a Rosaceae feeder, but the complex of genera making up the majority of the Bornean fauna in the tribe is associated with the Aquifoliaceae and, in one instance, the Flacourtiaceae. There are some records from Piper (Piperaceae).

No records for the Plutodini have been located.

The Lithinini are fern specialists. Indeed, Entomopteryx Guenée was associated tentatively with the tribe on the basis of adult morphology in an early draft of the text, the placement has been supported by biological information discovered between then and going to press (See Lithinini et seq.).

The Caberini are strongly associated with the Rhamnaceae, apart from the type genus which feeds on Betulaceae and Salicaceae.

The only records for the Thinopterygini are from Vitaceae.

The Gonodontini and Abraxini are polyphagous.

Some Cassymini genera show preference for Leguminosae, but there is one record from Guttiferae. Possible Palaearctic representatives are more polyphagous.

The few records for the Eutoeini suggest a preference for Melastomataceae.

The Macariini as a whole are polyphagous, with some temperate groups specialist on conifers, but the tropical representatives show a strong preference for Leguminosae.

The Boarmiini include some highly polyphagous genera but also a number that are much more narrowly specialised. Milionia Walker is only recorded from conifers, particularly southern hemisphere genera such as Araucaria, Dacrydium and Podocarpus. Several genera show some preference for the Lauraceae, such as Racotis Moore, Xandrames Moore, Amblychia Guenée and the related pair, Krananda Moore and Zanclopera Warren. Racotis has been recorded from the Annonaceae and Amblychia from the Illiciaceae (also Pogonopygia Warren), families distantly related to the Lauraceae within the Magnoliidae. Two genera, Amraica Moore and Microcalicha Sato, are associated with the Celastraceae.

Ecology and geography

The Ennominae are probably the most diverse and generally distributed of the geometrid subfamilies. The Geometrinae have a strong tropical focus, whereas the Larentiinae are more diverse in temperate latitudes and at high altitudes in the tropics (Holloway, 1986, 1993). In Borneo the greatest diversity of ennomines is in the lower montane zone at around 1000m where an association of species specific to that zone overlaps with the upper reaches of the lowland forest fauna and the lower reaches of an upper montane zone fauna. The Ennominae are particularly strongly represented within this lower montane association.

The subfamily also has a higher incidence of specificity to particular types of lowland forest, such as kerangas (heath) forest types or that on limestone. Examples of the former include Achrosis longifurca sp. n., Achrosis kerangatis sp. n., Synegia botydaria Guenée, Synegia eumeleata Walker, Bulonga schistacearia Walker, Peratophyga sobrina Prout (also montane forests), Sundagrapha lepidata Prout and Diplurodes kerangatis sp. n.

Limestone specialism is exemplified by Achrosis calcicola sp. n., and Achrosis alienata Walker.

There are two definite mangrove specialists: Gonodontis clelia Cramer and Cleora injectaria Walker.

Whilst there are no purely open habitat species, there are a number that respond positively to forest disturbance and are probably associated with plant taxa characteristic of regenerating forest. Hypochrosis binexata Walker, Hypochrosis pyrrhophaeata Walker, Bracca georgiata Walker and Craspedosis arycandata Guenée were identified as such by Holloway, Kirk-Spriggs & Chey (1992), and some Cleora Curtis, Hyposidra Guenée and Godonela Boisduval species may fall into this category too.

Just over 40% of the species are characteristic of the lowlands, and just under 40% are montane, with the remainder being recorded over a wide altitude range. About a quarter of the lowland species are endemic whereas this proportion increases to a half for montane species, the overall proportion of endemics being 30%. A third of all ennomines consists of species restricted to land on the Sunda Shelf with a further 6% extending from Sundaland to the Philippines or Sulawesi. About a quarter are more widely distributed in the Oriental tropics. Only about 5% range more widely through the Indo-Australian tropics (over 30% in the noctuid groups studied in the series to date).

There is some variation in proportion of various ecological and geographic categories amongst the tribes, and data on this have been published by Holloway & Barlow (1992). The Cassymini show a higher degree of endemism and restriction to lowland forest than the other major tribes, with endemism lowest in the Boarmiini. The Boarmiini and Baptini are more strongly represented in montane habitats.

Genera endemic to Borneo are few. Two are monobasic: Sundasclelia Gen. n. and Lampadopteryx Warren. Bornealcis Gen. n. has three species. Some genera such as Peratophyga Warren have a majority of species in Borneo.

Further collecting in Sumatra and Peninsular Malaysia is likely to reduce the proportion of species apparently endemic to Borneo. The faunas of the three areas are typically very similar, and the collecting activities of the Heterocera Sumatrana Society have reduced the number of Bornean ‘endemics’ considerably.

Localisation of species within Borneo is uncommon. Two of the Bornealcis species are restricted to particular mountain ranges, and further examples are found in Milionia and Dasyboarmia. Abraxas subhyalinata Röber is only known from Pulo Laut in southern Borneo and in the Lesser Sundas, a distribution type already noted for some agaristine noctuids (Part 12). Ruttellerona pulverulenta Warren is known only from Taganak, a small island off the north coast of Borneo, but has affinities with taxa in Sulawesi and further east. This is a pattern seen in some butterflies such as Eurema alitha and Cupha arias that just reach the northern part of Borneo or occur on small offshore islands (e.g. Pulo Mangalum), but have ranges extending through the Philippines and further east. Pseudeuchromia maculifera Felder & Rogenhofer is another ennomine of this geographic type.

Most genera represented in Borneo exhibit their greatest species richness in Southeast Asia or the eastern Himalayan region, and attenuate in richness eastwards into the Australasian tropics. A few, however, are more diverse in the Australasian tropics and attenuate westwards. Bracca Hübner and Craspedosis Butler reach their most westerly extremity in Sundaland, but Milionia Walker extends further onto the Asian mainland, reaching Japan.

One species is adventive. Macaria abydata Guenée is a Neotropical species that has established itself widely round the western Pacific margin during the past decade and a half, probably with Hawaii as a staging post. Its spread parallels to some extent that of the Leucaena psyllid (See Macaria abydata Guenée).

Copyright © Southdene Sdn. Bhd. All rights reserved.