Diagnostic morphological characters of geometrid subfamilies, based on the literature and on the present study. Currently, nine subfamilies are recognized, but the validity of Orthostixinae has not been tested in a phylogenetic context (see comments below). None of the listed characters is unambiguous at subfamily level and all of them show exceptions, but when considered together, they allow delimitation of subfamilies as monophyletic groups. A diurnal lifestyle, accompanied by atypical coloration compared to nocturnal relatives, has evolved independently in several lineages. This is one the reasons why wing colour and pattern are of limited value in diagnosing subfamilies. Structural characters show less variation and homologies are easier to interpret, but homoplasy is frequent in Geometridae (e.g. Sihvonen & Kaila, 2004) and Lepidoptera (Heikkilä et al., 2015). Diagnoses for the entire Geometridae are available in the literature (Minet & Scoble, 1999; Hausmann, 2001)
| Subfamily . | Diagnostic (synapomorphic) characters . | Remarks . |
|---|---|---|
| Sterrhinae | Most anterior discocellular vein (vein between the base of M1 and areole) in forewing long and oriented almost parallel with M veins (Forbes, 1948, our wording) (Fig. 16A); forewing fasciae single (e.g. Minet & Scoble, 1999); hindwing discal spot with pale marking in darker surround (Holloway, 1997); one or two areoles in forewings (absent in Mecoceratini); in forewing point of origin of vein M1 is either proximal or distal areole; absence of anterolateral extensions on male 2nd abdominal sternite (Sihvonen & Kaila, 2004; Sihvonen et al. 2020). | The first three characters were not included in a morphological phylogeny (Sihvonen & Kaila, 2004), because it was not possible to code the observed variation into discontinuous character states. Most Sterrhinae species have one or two forewing areoles, the character is absent in only a few species (Hausmann, 2004) and Mecoceratini (Sihvonen et al. 2020). Forewing fasciae are single in many Sterrhinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Sterrhinae and Larentiinae share a hammer-headed tip in tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1967, 1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Larentiinae | Hindwing veins Sc + R1 and Rs fused for long-distance (Meyrick, 1892, Hausmann & Viidalepp, 2012) (Fig. 16B); forewing fasciae are multiple (= composed of bands of closely juxtaposed lines) (e.g. Minet & Scoble, 1999); forewing radial veins Rs1–Rs4 or Rs2–Rs4 arising from the areole and are stalked or connate with M1, or, if separate, M1 proceeds in line with the anterior margin of the discal cell (Õunap et al., 2008); subcostal accessory cell (between veins Sc and R) missing; vein M2 thinner concerning M1 and M3 on both wings; socii absent (Hausmann & Viidalepp, 2012); gnathos generally strongly reduced (Schmidt, 2015). | Fusionof hindwing veins Sc + R1 and Rs is a typical feature of Larentiinae (in other families these veins may be parallel or connected via short cross-vein). These veins can be separate in Larentiinae also, for instance in monotypic Amygdaloptera testaria and Dyspteris (Hausmann & Viidalepp, 2012), the latter taxon being among the most basal Larentiinae (Murillo-Ramos et al. 2019). A long fused condition of the hindwing veins Sc + R is found in the Sterrhinae tribe Rhodometrini also (Õunap et al. 2008, Sihvonen et al. 2020). Forewing fasciae are multiple in many Larentiinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Hausmann (2004) lists further diagnostic characters, but none of them is without exceptions in Larentiinae. Contrary to the historical view, gnathos or remnants of gnathos are found in Larentiinae (Hausmann, 2004, Schmidt, 2015). Sterrhinae and Larentiinae share hammer-headed tip in the tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Archiearinae | Accessory tympanum lacking in metapostnotum and fenestra media narrow (Minet, 1983); larva with prolegs on segments A3–A6; hindwing M2 weak (Minet & Scoble, 1999); hindwing veins A2–A3 present and A1 present but not tubular (Hausmann, 2001); head with hair-like scales on the front of the head (Minet & Scoble, 1999). | Additional larval prolegs are also present in few Ennominae and Oenochrominae s.s. The Australian genera Dirce and Acalyphes, which were previously classified in Archiearinae (McQuillan & Edwards, 1996), are currently classified in Ennominae (Young, 2006; Murillo-Ramos et al. 2019). The systematic position of Mexican and Neotropical taxa, classified in Archiearinae, awaits further studies. |
| Desmobathrinae | Tegumen with lateral loops, extending beyond the articulation with vinculum (Fig. 16C); male 3rd sternite with pair of longitudinally elongated patches; male 8th sternite often with a finely corrugate zone laterally; often with complex structures at the junction of male 3rd and 4th sternites; delicate moths with elongated appendices (Holloway, 1996); forewing with one areole or areole absent (our observation); hindwing veins Sc + R1 and Rs parallel for long-distance or separate and connected with a short cross-vein (our observation); male with apophyses arising from membrane between the 2nd and 3rd sternites (our observation); pupa cremaster with four hooked shaftlets (= setae) (Holloway, 1996). | The characters listed here for Desmobathrinae are based on Desmobathrini only (see the composition of this clade in Murillo-Ramos et al. (2019)). Male apophyses arising from membrane between 2nd and 3rd sternite (found widely in Desmobathrinae, our observation) have not been mentioned as a diagnostic character earlier (Fig. 16D). Many Ennominae and Sterrhinae have similar-looking sternotympanal processes, but those originate from the posterior margin of the 2nd sternite and they are outside the abdomen (see for instance Ekboarmia in Skou et al. (2017)). No diagnostic Desmobathrinae characters have been found in the female structures. The corpus bursae have longitudinal rows of spines in the type genus Ozola, but there is great variation between genera (Holloway, 1996). Hausmann (2001) reports that Desmobathrinae has one or two areoles present in the forewing. Areoles are absent in Ozola, type genus of Desmobathrinae (our observation). |
| Epidesmiinae | Male antennae unipectinate, labial palps (2nd segment) elongated; two signa (one stellate, one elongated); hindwing with one anal vein (A); fore wing with two areoles; cross-veins between M veins reduced in both wings; gnathos arms fused, granulate or dentate apically; vom Rath’s organ elliptical. | Epidesmiinae are diverse externally and structurally, but some characters are shared by genus-groups such as the shape of the ansa, a planiform or tectiform resting posture, and valvae with or without dorsally directed ampulla. |
| Oenochrominae s.s. | Robust-bodied; lacinia of tympanal organ circular, lying parallel to the tympanum; fultura superior fused with the transtilla, forming a rigid plate (Cook & Scoble, 1992); larva often with prolegs on A4–A6 (Scoble & Edwards, 1989); pupa cremaster with two hooked shaftlets (= setae) (Holloway, 1996). | Oenochrominae s.s. refers to the robust-bodied species that occur in Australasia and Indo-Malay realms, approximating Guenée’s original concept (Guenée, 1857). The female signum, when present, is a single, small, circular patch of sclerotization (Holloway, 1996). The cremaster hooks (or setae) in pupa are reduced to a single apical pair in the Australian genera (McFarland, 1988). We illustrate here two additional structures from the metathorax of Oenochroma vinaria, which are unique among the material examined, but more material is needed to confirm their diagnostic value. The anterior and basal laminas of the metafurca are fused (Fig. 16E) and dorsal tergite is triangular extending to the opening of the euphragma (Fig. 16F). |
| Geometrinae | The green pigment, (geoverdin) is predominant (Cook et al. 1994); the hindwing vein M2 arises closer to M1 than to M3 (Minet & Scoble, 1999); the ansa is narrow at the base, widening medially and tapered towards the apex (Cook & Scoble, 1992), the female pheromone glands are long (Bendib, 2001). | Although green pigment is found widely in Geometrinae, there are several non-green Geometrinae taxa, and green moths are in other subfamilies also. Other defining, widely present characters include: reduction of the frenulum; paired setal patches on the male 3rd sternite; socii well developed often with parallel reduction of uncus; aedeagus with sclerotization reduced to a ventral strip along the length; oblique, papillate ovipositor lobes; a bicornute signum (Holloway, 1996). Beljaev (2008) added further characters from the skeleto-muscular system of the male genitalia. The systematic position of Eumeleini has remained controversial, see discussion under Desmobathrinae and in Holloway (1996) and Murillo-Ramos et al. (2019). Beljaev (2008) classified Eumeleini in the Geometrinae. |
| Ennominae | Loss (or reduction to a fold) of hindwing vein M2; transverse setae on the 3rd male sternite; stellate signum, often with spiked concavity medially (e.g. Holloway, 1994, our wording). | Numerous molecular studies have provided support for a monophyletic Ennominae (e.g. Sihvonen et al. 2011, Murillo-Ramos et al. 2019), but the morphology-based definition of this species-rich subfamily is weak. For instance, hindwing vein M2 is present as tubular in numerous Ennominae lineages, e.g. Melanolophia (Boarmiini), Alsophila (Alsophilini), Epirranthis (Epirranthini), Zerenopsis (Diptychini) (Pitkin, 2002; Skou & Sihvonen, 2015; Sihvonen et al. 2015; Müller et al. 2019). Several lineages have additional prolegs on larva. Numerous other adult and larva characters define smaller groupings within Ennominae (Holloway, 1994; Young, 2008). |
| Orthostixinae | Putative characters: larva with prolegs on A5 (and occasionally on A4); hindwing veins Sc + R1 and Rs connected with short cross-vein; hindwing veins A2–A3 present; hindwing vein M2 tubular (Hausmann, 2001). | Subfamily status needs more research. Taxon shows similarities with Ennominae (Holloway, 1996) and Desmobathrinae (Hausmann, 1996b). It was noted as a ‘putative tribe of Ennominae’ (Hausmann & Sihvonen, 2019), based partly on the position of the tentative orthostixine genus Naxa (Sihvonen et al. 2011). Beljaev (2008) included Orthostixinae in Desmobathrinae based on skeleto-muscular structure. We find this position also to be possible (current study), because Orthostixis cribraria shares, with Desmobathrinae the following characters: male apophyses arising from membrane between 2nd and 3rd sternites; a tapering ansa, tegumen with weakly developed lateral loops; and presence of A2–A3 veins in the hindwing. Ecological features also support a relationship with Desmobathrinae (Hausmann, 2001). |
| Subfamily . | Diagnostic (synapomorphic) characters . | Remarks . |
|---|---|---|
| Sterrhinae | Most anterior discocellular vein (vein between the base of M1 and areole) in forewing long and oriented almost parallel with M veins (Forbes, 1948, our wording) (Fig. 16A); forewing fasciae single (e.g. Minet & Scoble, 1999); hindwing discal spot with pale marking in darker surround (Holloway, 1997); one or two areoles in forewings (absent in Mecoceratini); in forewing point of origin of vein M1 is either proximal or distal areole; absence of anterolateral extensions on male 2nd abdominal sternite (Sihvonen & Kaila, 2004; Sihvonen et al. 2020). | The first three characters were not included in a morphological phylogeny (Sihvonen & Kaila, 2004), because it was not possible to code the observed variation into discontinuous character states. Most Sterrhinae species have one or two forewing areoles, the character is absent in only a few species (Hausmann, 2004) and Mecoceratini (Sihvonen et al. 2020). Forewing fasciae are single in many Sterrhinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Sterrhinae and Larentiinae share a hammer-headed tip in tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1967, 1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Larentiinae | Hindwing veins Sc + R1 and Rs fused for long-distance (Meyrick, 1892, Hausmann & Viidalepp, 2012) (Fig. 16B); forewing fasciae are multiple (= composed of bands of closely juxtaposed lines) (e.g. Minet & Scoble, 1999); forewing radial veins Rs1–Rs4 or Rs2–Rs4 arising from the areole and are stalked or connate with M1, or, if separate, M1 proceeds in line with the anterior margin of the discal cell (Õunap et al., 2008); subcostal accessory cell (between veins Sc and R) missing; vein M2 thinner concerning M1 and M3 on both wings; socii absent (Hausmann & Viidalepp, 2012); gnathos generally strongly reduced (Schmidt, 2015). | Fusionof hindwing veins Sc + R1 and Rs is a typical feature of Larentiinae (in other families these veins may be parallel or connected via short cross-vein). These veins can be separate in Larentiinae also, for instance in monotypic Amygdaloptera testaria and Dyspteris (Hausmann & Viidalepp, 2012), the latter taxon being among the most basal Larentiinae (Murillo-Ramos et al. 2019). A long fused condition of the hindwing veins Sc + R is found in the Sterrhinae tribe Rhodometrini also (Õunap et al. 2008, Sihvonen et al. 2020). Forewing fasciae are multiple in many Larentiinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Hausmann (2004) lists further diagnostic characters, but none of them is without exceptions in Larentiinae. Contrary to the historical view, gnathos or remnants of gnathos are found in Larentiinae (Hausmann, 2004, Schmidt, 2015). Sterrhinae and Larentiinae share hammer-headed tip in the tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Archiearinae | Accessory tympanum lacking in metapostnotum and fenestra media narrow (Minet, 1983); larva with prolegs on segments A3–A6; hindwing M2 weak (Minet & Scoble, 1999); hindwing veins A2–A3 present and A1 present but not tubular (Hausmann, 2001); head with hair-like scales on the front of the head (Minet & Scoble, 1999). | Additional larval prolegs are also present in few Ennominae and Oenochrominae s.s. The Australian genera Dirce and Acalyphes, which were previously classified in Archiearinae (McQuillan & Edwards, 1996), are currently classified in Ennominae (Young, 2006; Murillo-Ramos et al. 2019). The systematic position of Mexican and Neotropical taxa, classified in Archiearinae, awaits further studies. |
| Desmobathrinae | Tegumen with lateral loops, extending beyond the articulation with vinculum (Fig. 16C); male 3rd sternite with pair of longitudinally elongated patches; male 8th sternite often with a finely corrugate zone laterally; often with complex structures at the junction of male 3rd and 4th sternites; delicate moths with elongated appendices (Holloway, 1996); forewing with one areole or areole absent (our observation); hindwing veins Sc + R1 and Rs parallel for long-distance or separate and connected with a short cross-vein (our observation); male with apophyses arising from membrane between the 2nd and 3rd sternites (our observation); pupa cremaster with four hooked shaftlets (= setae) (Holloway, 1996). | The characters listed here for Desmobathrinae are based on Desmobathrini only (see the composition of this clade in Murillo-Ramos et al. (2019)). Male apophyses arising from membrane between 2nd and 3rd sternite (found widely in Desmobathrinae, our observation) have not been mentioned as a diagnostic character earlier (Fig. 16D). Many Ennominae and Sterrhinae have similar-looking sternotympanal processes, but those originate from the posterior margin of the 2nd sternite and they are outside the abdomen (see for instance Ekboarmia in Skou et al. (2017)). No diagnostic Desmobathrinae characters have been found in the female structures. The corpus bursae have longitudinal rows of spines in the type genus Ozola, but there is great variation between genera (Holloway, 1996). Hausmann (2001) reports that Desmobathrinae has one or two areoles present in the forewing. Areoles are absent in Ozola, type genus of Desmobathrinae (our observation). |
| Epidesmiinae | Male antennae unipectinate, labial palps (2nd segment) elongated; two signa (one stellate, one elongated); hindwing with one anal vein (A); fore wing with two areoles; cross-veins between M veins reduced in both wings; gnathos arms fused, granulate or dentate apically; vom Rath’s organ elliptical. | Epidesmiinae are diverse externally and structurally, but some characters are shared by genus-groups such as the shape of the ansa, a planiform or tectiform resting posture, and valvae with or without dorsally directed ampulla. |
| Oenochrominae s.s. | Robust-bodied; lacinia of tympanal organ circular, lying parallel to the tympanum; fultura superior fused with the transtilla, forming a rigid plate (Cook & Scoble, 1992); larva often with prolegs on A4–A6 (Scoble & Edwards, 1989); pupa cremaster with two hooked shaftlets (= setae) (Holloway, 1996). | Oenochrominae s.s. refers to the robust-bodied species that occur in Australasia and Indo-Malay realms, approximating Guenée’s original concept (Guenée, 1857). The female signum, when present, is a single, small, circular patch of sclerotization (Holloway, 1996). The cremaster hooks (or setae) in pupa are reduced to a single apical pair in the Australian genera (McFarland, 1988). We illustrate here two additional structures from the metathorax of Oenochroma vinaria, which are unique among the material examined, but more material is needed to confirm their diagnostic value. The anterior and basal laminas of the metafurca are fused (Fig. 16E) and dorsal tergite is triangular extending to the opening of the euphragma (Fig. 16F). |
| Geometrinae | The green pigment, (geoverdin) is predominant (Cook et al. 1994); the hindwing vein M2 arises closer to M1 than to M3 (Minet & Scoble, 1999); the ansa is narrow at the base, widening medially and tapered towards the apex (Cook & Scoble, 1992), the female pheromone glands are long (Bendib, 2001). | Although green pigment is found widely in Geometrinae, there are several non-green Geometrinae taxa, and green moths are in other subfamilies also. Other defining, widely present characters include: reduction of the frenulum; paired setal patches on the male 3rd sternite; socii well developed often with parallel reduction of uncus; aedeagus with sclerotization reduced to a ventral strip along the length; oblique, papillate ovipositor lobes; a bicornute signum (Holloway, 1996). Beljaev (2008) added further characters from the skeleto-muscular system of the male genitalia. The systematic position of Eumeleini has remained controversial, see discussion under Desmobathrinae and in Holloway (1996) and Murillo-Ramos et al. (2019). Beljaev (2008) classified Eumeleini in the Geometrinae. |
| Ennominae | Loss (or reduction to a fold) of hindwing vein M2; transverse setae on the 3rd male sternite; stellate signum, often with spiked concavity medially (e.g. Holloway, 1994, our wording). | Numerous molecular studies have provided support for a monophyletic Ennominae (e.g. Sihvonen et al. 2011, Murillo-Ramos et al. 2019), but the morphology-based definition of this species-rich subfamily is weak. For instance, hindwing vein M2 is present as tubular in numerous Ennominae lineages, e.g. Melanolophia (Boarmiini), Alsophila (Alsophilini), Epirranthis (Epirranthini), Zerenopsis (Diptychini) (Pitkin, 2002; Skou & Sihvonen, 2015; Sihvonen et al. 2015; Müller et al. 2019). Several lineages have additional prolegs on larva. Numerous other adult and larva characters define smaller groupings within Ennominae (Holloway, 1994; Young, 2008). |
| Orthostixinae | Putative characters: larva with prolegs on A5 (and occasionally on A4); hindwing veins Sc + R1 and Rs connected with short cross-vein; hindwing veins A2–A3 present; hindwing vein M2 tubular (Hausmann, 2001). | Subfamily status needs more research. Taxon shows similarities with Ennominae (Holloway, 1996) and Desmobathrinae (Hausmann, 1996b). It was noted as a ‘putative tribe of Ennominae’ (Hausmann & Sihvonen, 2019), based partly on the position of the tentative orthostixine genus Naxa (Sihvonen et al. 2011). Beljaev (2008) included Orthostixinae in Desmobathrinae based on skeleto-muscular structure. We find this position also to be possible (current study), because Orthostixis cribraria shares, with Desmobathrinae the following characters: male apophyses arising from membrane between 2nd and 3rd sternites; a tapering ansa, tegumen with weakly developed lateral loops; and presence of A2–A3 veins in the hindwing. Ecological features also support a relationship with Desmobathrinae (Hausmann, 2001). |
Diagnostic morphological characters of geometrid subfamilies, based on the literature and on the present study. Currently, nine subfamilies are recognized, but the validity of Orthostixinae has not been tested in a phylogenetic context (see comments below). None of the listed characters is unambiguous at subfamily level and all of them show exceptions, but when considered together, they allow delimitation of subfamilies as monophyletic groups. A diurnal lifestyle, accompanied by atypical coloration compared to nocturnal relatives, has evolved independently in several lineages. This is one the reasons why wing colour and pattern are of limited value in diagnosing subfamilies. Structural characters show less variation and homologies are easier to interpret, but homoplasy is frequent in Geometridae (e.g. Sihvonen & Kaila, 2004) and Lepidoptera (Heikkilä et al., 2015). Diagnoses for the entire Geometridae are available in the literature (Minet & Scoble, 1999; Hausmann, 2001)
| Subfamily . | Diagnostic (synapomorphic) characters . | Remarks . |
|---|---|---|
| Sterrhinae | Most anterior discocellular vein (vein between the base of M1 and areole) in forewing long and oriented almost parallel with M veins (Forbes, 1948, our wording) (Fig. 16A); forewing fasciae single (e.g. Minet & Scoble, 1999); hindwing discal spot with pale marking in darker surround (Holloway, 1997); one or two areoles in forewings (absent in Mecoceratini); in forewing point of origin of vein M1 is either proximal or distal areole; absence of anterolateral extensions on male 2nd abdominal sternite (Sihvonen & Kaila, 2004; Sihvonen et al. 2020). | The first three characters were not included in a morphological phylogeny (Sihvonen & Kaila, 2004), because it was not possible to code the observed variation into discontinuous character states. Most Sterrhinae species have one or two forewing areoles, the character is absent in only a few species (Hausmann, 2004) and Mecoceratini (Sihvonen et al. 2020). Forewing fasciae are single in many Sterrhinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Sterrhinae and Larentiinae share a hammer-headed tip in tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1967, 1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Larentiinae | Hindwing veins Sc + R1 and Rs fused for long-distance (Meyrick, 1892, Hausmann & Viidalepp, 2012) (Fig. 16B); forewing fasciae are multiple (= composed of bands of closely juxtaposed lines) (e.g. Minet & Scoble, 1999); forewing radial veins Rs1–Rs4 or Rs2–Rs4 arising from the areole and are stalked or connate with M1, or, if separate, M1 proceeds in line with the anterior margin of the discal cell (Õunap et al., 2008); subcostal accessory cell (between veins Sc and R) missing; vein M2 thinner concerning M1 and M3 on both wings; socii absent (Hausmann & Viidalepp, 2012); gnathos generally strongly reduced (Schmidt, 2015). | Fusionof hindwing veins Sc + R1 and Rs is a typical feature of Larentiinae (in other families these veins may be parallel or connected via short cross-vein). These veins can be separate in Larentiinae also, for instance in monotypic Amygdaloptera testaria and Dyspteris (Hausmann & Viidalepp, 2012), the latter taxon being among the most basal Larentiinae (Murillo-Ramos et al. 2019). A long fused condition of the hindwing veins Sc + R is found in the Sterrhinae tribe Rhodometrini also (Õunap et al. 2008, Sihvonen et al. 2020). Forewing fasciae are multiple in many Larentiinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Hausmann (2004) lists further diagnostic characters, but none of them is without exceptions in Larentiinae. Contrary to the historical view, gnathos or remnants of gnathos are found in Larentiinae (Hausmann, 2004, Schmidt, 2015). Sterrhinae and Larentiinae share hammer-headed tip in the tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Archiearinae | Accessory tympanum lacking in metapostnotum and fenestra media narrow (Minet, 1983); larva with prolegs on segments A3–A6; hindwing M2 weak (Minet & Scoble, 1999); hindwing veins A2–A3 present and A1 present but not tubular (Hausmann, 2001); head with hair-like scales on the front of the head (Minet & Scoble, 1999). | Additional larval prolegs are also present in few Ennominae and Oenochrominae s.s. The Australian genera Dirce and Acalyphes, which were previously classified in Archiearinae (McQuillan & Edwards, 1996), are currently classified in Ennominae (Young, 2006; Murillo-Ramos et al. 2019). The systematic position of Mexican and Neotropical taxa, classified in Archiearinae, awaits further studies. |
| Desmobathrinae | Tegumen with lateral loops, extending beyond the articulation with vinculum (Fig. 16C); male 3rd sternite with pair of longitudinally elongated patches; male 8th sternite often with a finely corrugate zone laterally; often with complex structures at the junction of male 3rd and 4th sternites; delicate moths with elongated appendices (Holloway, 1996); forewing with one areole or areole absent (our observation); hindwing veins Sc + R1 and Rs parallel for long-distance or separate and connected with a short cross-vein (our observation); male with apophyses arising from membrane between the 2nd and 3rd sternites (our observation); pupa cremaster with four hooked shaftlets (= setae) (Holloway, 1996). | The characters listed here for Desmobathrinae are based on Desmobathrini only (see the composition of this clade in Murillo-Ramos et al. (2019)). Male apophyses arising from membrane between 2nd and 3rd sternite (found widely in Desmobathrinae, our observation) have not been mentioned as a diagnostic character earlier (Fig. 16D). Many Ennominae and Sterrhinae have similar-looking sternotympanal processes, but those originate from the posterior margin of the 2nd sternite and they are outside the abdomen (see for instance Ekboarmia in Skou et al. (2017)). No diagnostic Desmobathrinae characters have been found in the female structures. The corpus bursae have longitudinal rows of spines in the type genus Ozola, but there is great variation between genera (Holloway, 1996). Hausmann (2001) reports that Desmobathrinae has one or two areoles present in the forewing. Areoles are absent in Ozola, type genus of Desmobathrinae (our observation). |
| Epidesmiinae | Male antennae unipectinate, labial palps (2nd segment) elongated; two signa (one stellate, one elongated); hindwing with one anal vein (A); fore wing with two areoles; cross-veins between M veins reduced in both wings; gnathos arms fused, granulate or dentate apically; vom Rath’s organ elliptical. | Epidesmiinae are diverse externally and structurally, but some characters are shared by genus-groups such as the shape of the ansa, a planiform or tectiform resting posture, and valvae with or without dorsally directed ampulla. |
| Oenochrominae s.s. | Robust-bodied; lacinia of tympanal organ circular, lying parallel to the tympanum; fultura superior fused with the transtilla, forming a rigid plate (Cook & Scoble, 1992); larva often with prolegs on A4–A6 (Scoble & Edwards, 1989); pupa cremaster with two hooked shaftlets (= setae) (Holloway, 1996). | Oenochrominae s.s. refers to the robust-bodied species that occur in Australasia and Indo-Malay realms, approximating Guenée’s original concept (Guenée, 1857). The female signum, when present, is a single, small, circular patch of sclerotization (Holloway, 1996). The cremaster hooks (or setae) in pupa are reduced to a single apical pair in the Australian genera (McFarland, 1988). We illustrate here two additional structures from the metathorax of Oenochroma vinaria, which are unique among the material examined, but more material is needed to confirm their diagnostic value. The anterior and basal laminas of the metafurca are fused (Fig. 16E) and dorsal tergite is triangular extending to the opening of the euphragma (Fig. 16F). |
| Geometrinae | The green pigment, (geoverdin) is predominant (Cook et al. 1994); the hindwing vein M2 arises closer to M1 than to M3 (Minet & Scoble, 1999); the ansa is narrow at the base, widening medially and tapered towards the apex (Cook & Scoble, 1992), the female pheromone glands are long (Bendib, 2001). | Although green pigment is found widely in Geometrinae, there are several non-green Geometrinae taxa, and green moths are in other subfamilies also. Other defining, widely present characters include: reduction of the frenulum; paired setal patches on the male 3rd sternite; socii well developed often with parallel reduction of uncus; aedeagus with sclerotization reduced to a ventral strip along the length; oblique, papillate ovipositor lobes; a bicornute signum (Holloway, 1996). Beljaev (2008) added further characters from the skeleto-muscular system of the male genitalia. The systematic position of Eumeleini has remained controversial, see discussion under Desmobathrinae and in Holloway (1996) and Murillo-Ramos et al. (2019). Beljaev (2008) classified Eumeleini in the Geometrinae. |
| Ennominae | Loss (or reduction to a fold) of hindwing vein M2; transverse setae on the 3rd male sternite; stellate signum, often with spiked concavity medially (e.g. Holloway, 1994, our wording). | Numerous molecular studies have provided support for a monophyletic Ennominae (e.g. Sihvonen et al. 2011, Murillo-Ramos et al. 2019), but the morphology-based definition of this species-rich subfamily is weak. For instance, hindwing vein M2 is present as tubular in numerous Ennominae lineages, e.g. Melanolophia (Boarmiini), Alsophila (Alsophilini), Epirranthis (Epirranthini), Zerenopsis (Diptychini) (Pitkin, 2002; Skou & Sihvonen, 2015; Sihvonen et al. 2015; Müller et al. 2019). Several lineages have additional prolegs on larva. Numerous other adult and larva characters define smaller groupings within Ennominae (Holloway, 1994; Young, 2008). |
| Orthostixinae | Putative characters: larva with prolegs on A5 (and occasionally on A4); hindwing veins Sc + R1 and Rs connected with short cross-vein; hindwing veins A2–A3 present; hindwing vein M2 tubular (Hausmann, 2001). | Subfamily status needs more research. Taxon shows similarities with Ennominae (Holloway, 1996) and Desmobathrinae (Hausmann, 1996b). It was noted as a ‘putative tribe of Ennominae’ (Hausmann & Sihvonen, 2019), based partly on the position of the tentative orthostixine genus Naxa (Sihvonen et al. 2011). Beljaev (2008) included Orthostixinae in Desmobathrinae based on skeleto-muscular structure. We find this position also to be possible (current study), because Orthostixis cribraria shares, with Desmobathrinae the following characters: male apophyses arising from membrane between 2nd and 3rd sternites; a tapering ansa, tegumen with weakly developed lateral loops; and presence of A2–A3 veins in the hindwing. Ecological features also support a relationship with Desmobathrinae (Hausmann, 2001). |
| Subfamily . | Diagnostic (synapomorphic) characters . | Remarks . |
|---|---|---|
| Sterrhinae | Most anterior discocellular vein (vein between the base of M1 and areole) in forewing long and oriented almost parallel with M veins (Forbes, 1948, our wording) (Fig. 16A); forewing fasciae single (e.g. Minet & Scoble, 1999); hindwing discal spot with pale marking in darker surround (Holloway, 1997); one or two areoles in forewings (absent in Mecoceratini); in forewing point of origin of vein M1 is either proximal or distal areole; absence of anterolateral extensions on male 2nd abdominal sternite (Sihvonen & Kaila, 2004; Sihvonen et al. 2020). | The first three characters were not included in a morphological phylogeny (Sihvonen & Kaila, 2004), because it was not possible to code the observed variation into discontinuous character states. Most Sterrhinae species have one or two forewing areoles, the character is absent in only a few species (Hausmann, 2004) and Mecoceratini (Sihvonen et al. 2020). Forewing fasciae are single in many Sterrhinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Sterrhinae and Larentiinae share a hammer-headed tip in tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1967, 1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Larentiinae | Hindwing veins Sc + R1 and Rs fused for long-distance (Meyrick, 1892, Hausmann & Viidalepp, 2012) (Fig. 16B); forewing fasciae are multiple (= composed of bands of closely juxtaposed lines) (e.g. Minet & Scoble, 1999); forewing radial veins Rs1–Rs4 or Rs2–Rs4 arising from the areole and are stalked or connate with M1, or, if separate, M1 proceeds in line with the anterior margin of the discal cell (Õunap et al., 2008); subcostal accessory cell (between veins Sc and R) missing; vein M2 thinner concerning M1 and M3 on both wings; socii absent (Hausmann & Viidalepp, 2012); gnathos generally strongly reduced (Schmidt, 2015). | Fusionof hindwing veins Sc + R1 and Rs is a typical feature of Larentiinae (in other families these veins may be parallel or connected via short cross-vein). These veins can be separate in Larentiinae also, for instance in monotypic Amygdaloptera testaria and Dyspteris (Hausmann & Viidalepp, 2012), the latter taxon being among the most basal Larentiinae (Murillo-Ramos et al. 2019). A long fused condition of the hindwing veins Sc + R is found in the Sterrhinae tribe Rhodometrini also (Õunap et al. 2008, Sihvonen et al. 2020). Forewing fasciae are multiple in many Larentiinae (e.g. Minet & Scoble, 1999), but the wing pattern characters are variable. Hausmann (2004) lists further diagnostic characters, but none of them is without exceptions in Larentiinae. Contrary to the historical view, gnathos or remnants of gnathos are found in Larentiinae (Hausmann, 2004, Schmidt, 2015). Sterrhinae and Larentiinae share hammer-headed tip in the tympanal organ’s ansa (Cook & Scoble, 1992, see also Viidalepp, 2011) and a few other characters (summarized in Hausmann, 2004). McGuffin (1987) provided diagnostic larva and pupa characters, but those were based on limited Canadian material. |
| Archiearinae | Accessory tympanum lacking in metapostnotum and fenestra media narrow (Minet, 1983); larva with prolegs on segments A3–A6; hindwing M2 weak (Minet & Scoble, 1999); hindwing veins A2–A3 present and A1 present but not tubular (Hausmann, 2001); head with hair-like scales on the front of the head (Minet & Scoble, 1999). | Additional larval prolegs are also present in few Ennominae and Oenochrominae s.s. The Australian genera Dirce and Acalyphes, which were previously classified in Archiearinae (McQuillan & Edwards, 1996), are currently classified in Ennominae (Young, 2006; Murillo-Ramos et al. 2019). The systematic position of Mexican and Neotropical taxa, classified in Archiearinae, awaits further studies. |
| Desmobathrinae | Tegumen with lateral loops, extending beyond the articulation with vinculum (Fig. 16C); male 3rd sternite with pair of longitudinally elongated patches; male 8th sternite often with a finely corrugate zone laterally; often with complex structures at the junction of male 3rd and 4th sternites; delicate moths with elongated appendices (Holloway, 1996); forewing with one areole or areole absent (our observation); hindwing veins Sc + R1 and Rs parallel for long-distance or separate and connected with a short cross-vein (our observation); male with apophyses arising from membrane between the 2nd and 3rd sternites (our observation); pupa cremaster with four hooked shaftlets (= setae) (Holloway, 1996). | The characters listed here for Desmobathrinae are based on Desmobathrini only (see the composition of this clade in Murillo-Ramos et al. (2019)). Male apophyses arising from membrane between 2nd and 3rd sternite (found widely in Desmobathrinae, our observation) have not been mentioned as a diagnostic character earlier (Fig. 16D). Many Ennominae and Sterrhinae have similar-looking sternotympanal processes, but those originate from the posterior margin of the 2nd sternite and they are outside the abdomen (see for instance Ekboarmia in Skou et al. (2017)). No diagnostic Desmobathrinae characters have been found in the female structures. The corpus bursae have longitudinal rows of spines in the type genus Ozola, but there is great variation between genera (Holloway, 1996). Hausmann (2001) reports that Desmobathrinae has one or two areoles present in the forewing. Areoles are absent in Ozola, type genus of Desmobathrinae (our observation). |
| Epidesmiinae | Male antennae unipectinate, labial palps (2nd segment) elongated; two signa (one stellate, one elongated); hindwing with one anal vein (A); fore wing with two areoles; cross-veins between M veins reduced in both wings; gnathos arms fused, granulate or dentate apically; vom Rath’s organ elliptical. | Epidesmiinae are diverse externally and structurally, but some characters are shared by genus-groups such as the shape of the ansa, a planiform or tectiform resting posture, and valvae with or without dorsally directed ampulla. |
| Oenochrominae s.s. | Robust-bodied; lacinia of tympanal organ circular, lying parallel to the tympanum; fultura superior fused with the transtilla, forming a rigid plate (Cook & Scoble, 1992); larva often with prolegs on A4–A6 (Scoble & Edwards, 1989); pupa cremaster with two hooked shaftlets (= setae) (Holloway, 1996). | Oenochrominae s.s. refers to the robust-bodied species that occur in Australasia and Indo-Malay realms, approximating Guenée’s original concept (Guenée, 1857). The female signum, when present, is a single, small, circular patch of sclerotization (Holloway, 1996). The cremaster hooks (or setae) in pupa are reduced to a single apical pair in the Australian genera (McFarland, 1988). We illustrate here two additional structures from the metathorax of Oenochroma vinaria, which are unique among the material examined, but more material is needed to confirm their diagnostic value. The anterior and basal laminas of the metafurca are fused (Fig. 16E) and dorsal tergite is triangular extending to the opening of the euphragma (Fig. 16F). |
| Geometrinae | The green pigment, (geoverdin) is predominant (Cook et al. 1994); the hindwing vein M2 arises closer to M1 than to M3 (Minet & Scoble, 1999); the ansa is narrow at the base, widening medially and tapered towards the apex (Cook & Scoble, 1992), the female pheromone glands are long (Bendib, 2001). | Although green pigment is found widely in Geometrinae, there are several non-green Geometrinae taxa, and green moths are in other subfamilies also. Other defining, widely present characters include: reduction of the frenulum; paired setal patches on the male 3rd sternite; socii well developed often with parallel reduction of uncus; aedeagus with sclerotization reduced to a ventral strip along the length; oblique, papillate ovipositor lobes; a bicornute signum (Holloway, 1996). Beljaev (2008) added further characters from the skeleto-muscular system of the male genitalia. The systematic position of Eumeleini has remained controversial, see discussion under Desmobathrinae and in Holloway (1996) and Murillo-Ramos et al. (2019). Beljaev (2008) classified Eumeleini in the Geometrinae. |
| Ennominae | Loss (or reduction to a fold) of hindwing vein M2; transverse setae on the 3rd male sternite; stellate signum, often with spiked concavity medially (e.g. Holloway, 1994, our wording). | Numerous molecular studies have provided support for a monophyletic Ennominae (e.g. Sihvonen et al. 2011, Murillo-Ramos et al. 2019), but the morphology-based definition of this species-rich subfamily is weak. For instance, hindwing vein M2 is present as tubular in numerous Ennominae lineages, e.g. Melanolophia (Boarmiini), Alsophila (Alsophilini), Epirranthis (Epirranthini), Zerenopsis (Diptychini) (Pitkin, 2002; Skou & Sihvonen, 2015; Sihvonen et al. 2015; Müller et al. 2019). Several lineages have additional prolegs on larva. Numerous other adult and larva characters define smaller groupings within Ennominae (Holloway, 1994; Young, 2008). |
| Orthostixinae | Putative characters: larva with prolegs on A5 (and occasionally on A4); hindwing veins Sc + R1 and Rs connected with short cross-vein; hindwing veins A2–A3 present; hindwing vein M2 tubular (Hausmann, 2001). | Subfamily status needs more research. Taxon shows similarities with Ennominae (Holloway, 1996) and Desmobathrinae (Hausmann, 1996b). It was noted as a ‘putative tribe of Ennominae’ (Hausmann & Sihvonen, 2019), based partly on the position of the tentative orthostixine genus Naxa (Sihvonen et al. 2011). Beljaev (2008) included Orthostixinae in Desmobathrinae based on skeleto-muscular structure. We find this position also to be possible (current study), because Orthostixis cribraria shares, with Desmobathrinae the following characters: male apophyses arising from membrane between 2nd and 3rd sternites; a tapering ansa, tegumen with weakly developed lateral loops; and presence of A2–A3 veins in the hindwing. Ecological features also support a relationship with Desmobathrinae (Hausmann, 2001). |
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