SUBFAMILY LARENTIINAE

This subfamily tends to predominate over other groups of Geometridae with increasing latitude. In the tropics a similar increase in representation is seen with altitude (Holloway, 1986b). This renders it difficult to conduct a broad review of the higher classification of the subfamily as was attempted for other geometrid groups, as only a minority of the tribal level taxa, as listed in the next section, are represented in Borneo. The bulk of the Bornean fauna falls within the tribes Trichopterygini, predominantly tropical though mostly montane, and Eupitheciini, a highly diverse group that has been particularly successful amongst the Larentiinae in reaching remote oceanic islands and archipelagos such as those of Polynesia: the remarkable Hawaiian eupitheciine radiation with predatory caterpillars perhaps represents the pinnacle of this phenomenon (Montgomery, 1982).

Lowland tropical Larentiinae are few (See Introduction), but tend to be difficult to assign to temperate groups. They are often patterned with red and yellow rather than having the more cryptic facies of their temperate and montane relatives, though many of these have proved to belong to the Eupitheciini. Some Australasian groupings are also becoming evident: again these can be difficult to assign to higher categories based on north temperate genera.

Family-group names in the Larentiinae.
ASTHENINI: Astheninae Warren (1894). (See Eupitheciini), and the last few genera treated in Eupitheciini.

CATACLYSMINI: Cataclysmini Herbulot (1962-3). Treated as a valid tribe by Vives Moreno (1994). Not represented in Borneo.

CHESIADINI: Chesiadi Stephens (1850); Seven (1991). See also Eucestiini below. Treated as a valid tribe by Vives Moreno (1994). Not represented in Borneo.

CHLOROCLYSTINI: Chloroclystina Mironov (1990). Proposed as subtribe of Eupitheciini.

CIDARIINI: Cidarites Duponchel (1845); Seven (1991). (See Cidariini).

ENTEPHRINI: Entephrinae Pierce (1914). Subordinated to Larentiini by Vives Moreno (1994).

EPIRRHOINI: Epirrhoinae Pierce (1914). Subordinated to Larentiini by Vives Moreno (1994).

ERATEINIINI: Erateinidae Guenée (1857) This is a Neotropical group with hesperiid-like facies.

EUBOLIINI: Eubolites Duponchel (1845). Preferred to Larentiinae by Seven (1991) against predominant trend of usage this century (e.g. Fletcher(1979) and Hodges et al. (1983)). Subordinated to Xanthorhoini by Vives Moreno (1994).

EUCESTIINI: Eucestiinae Warren (1894). Eucestis Hübner in a junior objective synonym of Chesias Treitschke: see Chesiadini above.

EUCOSMIINI: Eucosminae Gumppenberg (1887). Eucosmia Stephens is a synonym of Hydria Hübner. Confusion with a family-group name in Tortricidae based on Eucosma Hübner probably explains why this name has been neglected!

EUDULINI: Eudulinae Warren (1897); Hodges et al. (1983). This is a New World group. Forbes (1948) noted members to have chaetosemata forming a wide band across the back of the head.

EUPHYIINI: Euphyiini Herbulot (1962-3); Seven (1991). Treated as a valid tribe by Vives Moreno (1994).

EUPITHECIINI: Eupitheciinae Tutt (1896); Hodges et al. (1983), Seven (1994). See Tephroclystiini below and Eupitheciini for discussion of this tribe, the most diverse in the Bornean fauna. Vives Moreno (1994) includes Chloroclystina Mironov (1990) as a subtribe.

HETERUSIINI: Heterusiinae Warren (1897). This is a New World group.

HYDRIOMENINI: Hydriomenidae Meyrick (1892); Hodges et al. (1983), Seven (1991). Treated as a valid tribe by Vives Moreno (1994) and Nielsen, Edwards & Rangsi (1994). None of the Australian genera included by the latter occur in Borneo.

LARENTIINAE: Larentites Duponchel (1845). The generally preferred name for the subfamily (see Euboliini above).

LOBOPHORINI: Lobophorinae Tutt (1896); Hodges et al. (1983), Seven (1991). (See Trichopterygini)

LYTHRIINI: Lythriini Herbulot (1962-3); Seven (1991). Treated as a valid tribe by Vives Moreno (1994).

MELANIPPINI: Melanippidae Bruand (1846). Melanippe Duponchel is a junior synonym of Rheumaptera Hübner (Fletcher, 1979). See Rheumapterini below.

MELANTHIINI: Melanthites Duponchel (1845); Seven (1991). Subordinated to Rheumapterini by Vives Moreno (1994). However, Patocka (1981) indicated the pupa of the type genus shared characters with that of the Hydromenini and was distinct from that of the Rheumapterini.

ODEZIINI: Odeziini Exposito (1978). Treated as a valid tribe by Vives Moreno (1994). Not represented in Borneo.

OPEROPHTERINI: Operophterinae Packard (1876); Hodges et al. (1983). This name should probably take precedence over Oporiniini, below, as in Vives Moreno (1994). It is essentially a group of high latitudes. The pupal cremaster is distinctly bifid. (See Eupitheciini)

OPORINIINI: Oporiniinae Pierce (1914); Seven (1991). See Operophterini above.

ORTHOLITHINI: Ortholithinae Tutt (1896). Not represented in Borneo.

PERIZOMINI: Perizomini Herbulot (1962-3). Treated as a valid tribe by Vives Moreno (1994). (See Eupitheciini)

PHASIANINI: Phasianinae Gumppenberg (1887). The name is based on Phasiane Duponchel. The family-group name is also used in Aves (Phasianus Linnaeus), and would undoubtedly take priority there.

PHILEREMINI: Philereminae Pierce (1914). Subordinated to Rheumapterini by Vives Moreno (1994), supported by pupal characters according to Patocka (1981).

PSYCHOPHORINI: Psychophorinae Hampson (1918). This was proposed as a substitute for Larentiinae, being based on the oldest generic name in the group. Subordinated to Larentiini by Vives Moreno (1994)

RHEUMAPTERINI: Rheumapterini Herbulot (1962-3). The name Melanippini (see above) refers to the same concept, is older, but is based on a junior synonym of Rheumaptera. Treated as a valid tribe by Vives Moreno (1994).

SCOTOPTERYGINI: Scotopteryginae Warren (1895). Not represented in Borneo.

SOLITANEINI: Solitaneini Leraut (1980). Not represented in Borneo.

STAMNODINI: Forbes (1948); Hodges et al. (1983). Not represented in Borneo.

TEPHROCLYSTIINI: Tephroclystiinae Warren (1895). The name is based on a genus for which the type species is currently placed in Eupithecia Curtis: see Eupitheciini above.

THERINI: Therinae Pierce (1914). This concept has not found favour; confusion with Theriini in Ennominae is inevitable. Therini is the older name.

TRICHOPTERYGINI: Trichopteryginae Warren (1894); Dugdale (1980). See Trichopterygini. The group is diverse in Borneo.

TRIPHOSINI: Triphosidi Tutt (1896). Not represented in Borneo.

XANTHORHOINI: Xanthorhoinae Pierce (1914); Hodges et al. (1983). (See Xanthorhoini).

Definition of subfamily and tribal groupings
The Larentiinae were defined by Minet (1983) on possession of a hammer-headed ansa, but this occurs also amongst the sterrhines (Holloway, Bradley & Carter, 1987; Cook & Scoble, 1992), and in the geometrine genus Dysphania Hübner (Cook & Scoble, 1992; Holloway, 1996a).

Holloway, Bradley & Carter (1987) noted more reliable facies characters for distinguishing the group: each fascia of the forewing tends to be multiple rather than single and to meet the dorsum at right-angles rather than obliquely. A high proportion of the species rest with the forewing mostly or wholly covering the hindwing (Common, 1990), the latter tending to be exposed only where the pattern is as strong as that of the forewing (see Tweedie & Emmet, 1991).

Males usually lack a hair-pencil in the hind-tibia (Phthonoloba Warren in the Trichopterygini is an exception ), and secondary sexual characteristics in that sex tend to be found on the hindwing or in the form of coremata towards the distal end of the abdomen. However, the Trichopterygini are characterised also by a structure on the second sternite (See Trichopterygini).

The eggs of Larentiinae (Salkeld, 1983) are generally sculptured with numerous polygonal cells, but these are often weak to the point of smoothness. Some Cidariini and Hydriomenini have strong pitting within the polygons. No pitting is seen in Xanthorhoini but aeropyles along the polygon walls can be numerous. In Asthenini the polygons can be elevated, swollen, separated by sharp grooves.

Singh (1953) characterised the larentiine larva in his key as having the subventral group of setae on segments A1-5 bisetose (occasionally unisetose on A1) and by having the subanal plate only weakly produced to a blunt point rather than strongly so, though this is also the case in some Sterrhinae. Though Singh gave much detail of chaetotaxy, it is difficult to interpret this in terms of apomorphy.

Patocka (1995) described the larentiine pupa as rather sturdy, compact, expanded centrally, conical at the posterior end and only moderately attenuated, rounded anteriorly. There is a distinct coronal suture separating the frons and vertex, and the metanotum has rounded or obtuse frontolateral processes: there can also be a distinctive sculpturing of the first abdominal segment. These features enable larentiine pupae to be distinguished from those of the Ennominae and Sterrhinae.

The pupal cremaster of a number of Larentiinae has been illustrated or described by Khotko (1977), Patocka (1980, 1981, 1995), Hashimoto (1982, 1985) and Bell (MS). Unlike in other geometrid groups there seem to have been several instances where the terminal pair of hooklets has become predominant, with reduction or loss of the other three pairs. This has definitely occurred in the Trichopterygini and Eupitheciini, even within the genus Eupithecia Curtis (Patocka, 1980), and also in Xanthorhoini and Cidariini. Patocka did not initially (1980) reach any clear conclusion about classification with the tribal groupings recognised by Herbulot (1962-3). In subsequent papers (Patocka, 1981, 1995), he noted a number of groupings supported by pupal morphology. Some of these are referred to in the comments on family-group names above. Others are relevant to the tribes occurring in Borneo and are discussed in the accounts of those tribes.

The Trichopterygini are rather set apart from the rest of the tribes and may well bear a sister-relationship to them. The forewing fasciae tend to be rather evenly spaced across the wing rather than bunched together in distinct fasciae, though this is far from being a hard and fast distinction, particularly with regard to Palaearctic trichopterygine taxa. There tends to be more frequent angling of the forewing postmedial in non-trichopterygine groups, though again this is inconsistent. Fusion or not of veins Sc and Rs in the hindwing (Forbes, 1948) is also not fully reliable as lack of fusion is seen in some Trichopterygini but also in the Hydriomenini.

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