Insects of Britain and Ireland: the families of Lepidoptera

L. Watson and M.J. Dallwitz

Geometridae

Hydromenidae, Monocteniadae, Selidodesmidae, Sterrhidae

Adults diurnal, or crepuscular, or nocturnal; small to large; relatively short-bodied (rarely), or medium-bodied (mostly), or long-bodied (rarely); nearly always relatively slender-bodied (wingspan more than 15 times the thoracic width). The abdomen conspicuously patterned (rarely, e.g. Abraxas being yellow-bodied with dorsal black spots), or plain (mostly). Adults with fully developed wings (the outline always more or less butterfly-shaped), or wingless, or with vestigial wings (females, occasionally); wings in repose apposed vertically over the back (occasionally), or spread flat with their lower surfaces more or less appressed to the substrate (mostly), or packed with the forewings directed backwards to cover hindwings and abdomen (or approaching this).

Antennae very short to of medium length; extending to about 0.4–0.6 times the length of the forewing; not clubbed. Antennae of males bipectinate, or dentate, or simple; non-ciliate, or simply-ciliate, or fasciculate-ciliate. The antennal segments without rings of scales. Eyes hairy, or glabrous (?). Ocelli absent. Chaetosemata present. Maxillary palps much reduced; 1 segmented (usually), or 2 segmented. Labial palps usually ascending; 3 segmented. Proboscis fully developed; not scaly.

Wingspan (16–)20–60(–69) mm; 10.5–14–19 times the thoracic width, or 7.2–9.2 times the thoracic width (very rarely, e.g. Biston, Lycia). Forewings broad; 1.6–1.9(–2.2) times as long as wide (very rarely reaching 2.0). The outer and hind margins angled at (90–)95–100(–115) degrees (the wider-angled condition mainly represented in Biston and Eupithecia). The outer margin convexly curved (nearly always), or more or less straight, or angulated (notably in Ennomos s. str.), or sigmoid-curved (e.g., in Ourapteryx); forewings apically blunt (nearly always), or pointed. Hindwings broadly rounded (nearly always), or broadly angular (occasionally, or angulated); quite markedly narrower than the forewings to similar in breadth to the forewings; with a rounded apex; angulated to tailed (rarely), or not tailed; the upper surfaces conspicuously patterned above, or plain; with a discal spot and with transverse lines (usually, with at least the median, second and sub-terminal lines, but often with alternately light-and-dark transverse striae additional to the and obscuring them), or with neither discal spot nor transverse lines (when the markings are more or less ‘obsolete’); with a frenulum.

Neuration of forewings and hindwings dissimilar. Forewings 11 veined, or 12 veined (more often); with 1 anal vein (mostly, with one or both of the other two sometimes represented by vestiges), or with 2 anal veins (rarely). The anal veins of the forewings representing 1b only (usually, but this often accompanied by vestiges of 1a and 1c), or comprising 1a and 1b (cf. Meyrick's figure of Opisthograptis luteolata). Forewings lacking a tubular vein 1c (but this sometimes present as a vestige). Vein 1b of the forewings furcate proximally to simple. The transverse vein complete to incomplete. Discal cell of the forewings without a tubular media (M) vein. Hindwings 8 veined, or 9 veined; with 1 anal vein, or with 2 anal veins. The anal veins of the hindwings representing 1b only, or comprising 1a and 1b (or 1a only). Hindwings exhibiting vein 1a; lacking a tubular vein 1c. Vein 1b of the hindwings simple. The transverse vein complete to incomplete. The hindwing cell emitting more than six veins, or emitting no more than six veins. 6 veins arising from the hindwing cell, or 7 veins arising from the hindwing cell (usually, including the anastomosed vein 8, with vein 5 sometimes vestigial only). The cell-derived hindwing veins all arising independently of one another, or 3+4 proximally joined, or 6+7 proximally joined, or 3+4 proximally joined and 6+7 proximally joined. Vein 8 of the hindwings completely independent of the cell (e.g., Opisthograptis, Odezia), or anastomosing with the upper margin of the cell (mostly, from near the base to beyond the middle); not approximating to vein 7.

Fore-legs with a tibial epiphysis. Tibiae of middle legs 2-spurred. Posterior tibiae 2-spurred, or 4-spurred.

Tympanal organs present; abdominal.

Eggs, larvae and pupae. Eggs markedly flattened; ribbed and/or reticulate. Larval prolegs 4 (‘loopers’, lacking prolegs on segments 7 to 9). Larvae exposed feeders. On divers Dicot trees, shrubs and herbs, even a few on conifers and lichens; but curiously, very few indeed (or none?) on Monocots.

Pupae exposed, with no coccoon, or concealed; suspended from the tail (cremaster), with no median silk girdle (mostly), or not suspended, but attached at the tail and secured by a median girdle of silk (attached to a leaf, and sometimes having a silken girdle in Cyclophora); subterranean, or on the surface of the ground, or above the ground. Empty pupae not becoming protruded.

British representation. Genera 130 (2 adventive); about 330 species (19 adventive, 1 extinct). See the separate Geometridae Intkey package for descriptions of all genera, and illustrations of nearly all the species.

Classification. Macromoths. Suborder Ditrysia. Superfamily Geometroidea.

Comments. This family exhibits numerous examples of melanism, including a few spectacular examples of near exclusive industrial melanism (notably in Biston betularia, in which the effects on habitats of industrial pollution boosted melanism from the status of a very rare mutation to almost 100% in some regions), as well as some examples of exclusively non-industrial rural melanism, with relatively large incidence of melanics for which “natural” selective factors have been postulated; and there is clear evidence in some species of drift to industrial melanism superimposed on the latter. Examples are depicted in illustrations provided with this family description, and under Agriopis, Alcis, Angerona, Apocheima (Phigalia), Biston, Ectropis, Epirrita, Erannis, Odontopera, Operophtera and Peribatoides in the accompanying ‘Genera of Geometridae’ package.

Illustrations. • Sterrhinae and Ennominae: Kirby. STERRHINAE. 1, Scopula marginepunctata (Mullein Wave); 2, Scopula nigropunctata (Sub-angled Wave); 3, Scopula ornata (Lace Border); 4, Scopula decorata; 5, Problepsis ocellata; 6, Cyclophora pendularia (Birch Mocha); 7, Cyclophora annularia (Mocha); 8, Cyclophora porata (False Mocha); 9, Cyclophora punctaria (Maiden's Blush); 10, Cyclophora linearia (Clay Triple-lines); 11, Timandra comae (Blood-vein); 12, Rhodostrophia vibicaria; 13, Rhodostrophia calabraria; 14, Arichanna melanaria; 15, Abraxas grossulariata (Magpie Moth); 16, Abraxas pantaria (Panther); 17, Abraxas sylvata (Scarce or Clouded Magpie); 18, Ligdia adustata (Scorched Carpet); 19, Lomaspilis marginata (Clouded Border); 20, Lomographa bimaculata (White-pinion Spotted); 21, Lomographa temerata (Clouded Silver); 22, Cabera pusaria (Common White Wave); 23, Cabera exanthemata (Common Wave); 24, Plagodis pulveraria (Barred Umber); 25, Pungeleria capreolaria; 26, Hylaea fasciaria (Barred Red); 27, Campaea margaritata (Light Emerald). From Kirby 45, with updated names. See the separate GENERA OF GEOMETRIDAE data set for comprehensive illustrations of this family. • Larentiinae (‘Carpets’): Kirby. LARENTIINAE. 1. Anticlea badiata (Shoulder Stripe); 2, Eustroma reticulatum (Netted Carpet); 3, Eulithis prunata (Phoenix); 4, Eulithis populata (Northern Spinach); 5, Cidaria fulvata (Barred Yellow); 6, Cosmorhoe ocellata (Purple Bar); 7, Plemyria rubiginata (Blue-bordered Carpet); 8, Thera britannica (Spruce Carpet); 9, Thera juniperata (Juniper Carpet); 10 and 11, Chloroclysta siterata (Red-green and Autumn-green Carpets); 12, Chloroclysta truncata (Common Marbled Carpet); 13, Colostygia olivata (Beech Green Carpet); 14, Colostygia pectinataria (Green Carpet); 15, cf. Colostygia turbata; 16, Xanthorhoe fluctuata (Garden Carpet); 17, Xanthorhoe montanata (Silver-ground Carpet); 18, Xanthorhoe quadrifasiata (Large Twin-spot Carpet); 19, Xanthorhoe ferrugata (Dark-barred Twin-spot Carpet); 20, Lampropteryx suffumata (Water Carpet); 21, Xanthorhoe biriviata (Balsam Carpet); 22, Xanthorhoe designata (Flame Carpet); 23, Orthonama vittata (Oblique Carpet); 24, Epirrita dilutata (November Moth); 25, Entephria caesiata (Grey Mountain Carpet); 26, Coenotephria tophaceata; 27, Catarhoe cuculata (Royal Mantle); 28, Epirrhoe galiata (Galium Carpet); 29, Epirrhoe alternata alternata (Common Carpet); 30, Euphya unangulata (Sharp-angled Carpet); 31, Melanthia procellata (Pretty Chalk Carpet). From Kirby 50, with updated names. • Larentiinae (‘Carpets’, ‘Pugs’, etc.): Kirby 51. LARENTIINAE. 1, Mesoleuca albicillata (Beautiful Carpet); 2 and 5, Spargania luctuata (White-banded Carpet); 3, Rheumaptera hastata (Argent and Sable); 4, Epirrhoe tristata (Small Argent and Sable); 5, Spargania luctuata (White-banded Carpet); 6, Epirrhoe molluginata; 7, Perizoma alchemillata (Small Rivulet); 8, Perizoma blandiata (Pretty Pinion); 9, Perizoma albulata (Grass Rivulet); 10, Asthena albulata (Small White Wave); 11, Hydrelia sylvata (Waved Carpet); 12, Hydrelia flammeolaria (Small Yellow Wave); 13, Camptogramma bilineata (Yellow Shell); 14, Hydriomena furcata; 15, Hydriomena impluviata (May High-flier); 16, "Larentia capitata", cf. Ecliptopera capitata; 17, Electrophaes corylata (Broken-barred Carpet); 18, Pareulype berberata (Barberry Carpet); 19, Anticlea derivata (The Streamer); 20, Catarhoe rubidata (The Flame); 21, Perizoma sagittata (Marsh Carpet); 22, Pelurga comitata (Dark Spinach); 23, Horisme vitalbata (Small Waved Umber); 24, Anticollix sparsata (Dentated Pug); 25, Eupithecia centaureata (Lime-speck Pug); 26, Eupithecia venosata (Netted Pug); 27, Eupithecia linariata (Toadflax Pug); 28, Eupithecia abietaria(?); 29, Pasiphila rectangulata (Green Pug); 30, Eupithecia nanata (Narrow-winged Pug); 31, Eupithecia innonata (Angle-barred Pug); 32, Eupithecia satyrata (Beech Pug); 33, Eupithecia lanceata. From Kirby 51, with updated names. • Crocallis elinguaria (Scalloped Oak): photo, Giles Watson. Crocallis elinguaria (Scalloped Oak, male). Godstow Priory, Wolvercote, Oxford, July 2008. Giles Watson. • Ennominae (‘Thorns’, etc.): Kirby. ENNOMINAE. 1, Ennomos quercinaria (August Thorn); 2, Ennomos autumnaria (Large Thorn); 3, Ennomos alniaria (Canary-shouldered Thorn); 4, Ennomos erosaria (September Thorn); 5, Selenia dentaria (Early Thorn); 6, Selenia lunularia (Lunar Thorn); 7, Selenia tetralunaria (Purple Thorn); 8, Apeira syringaria (Lilac Beauty); 9, Artiora evonymaria; 10, Odontopera bidentata (Scalloped Hazel); 11, Colotois pennaria (Feathered Thorn); 12, Crocallis elinguaria (Scalloped Oak); 13, Plagodis dolabraria (Scorch-wing); 14, Angerona prunaria (Orange Moth); 15, Ourapteryx sambucaria (Swallow-tailed Moth); 16, Opisthograptis luteolata (Brimstone Moth). From Kirby 36, with updated names. • Assorted Ennominae (‘Thorns’, with larvae, from Humphreys. ‘THORNS’. 1 & 2, Colotois pennaria (Feathered Thorn); 3 & 4, Crocallis elinguaria (Scalloped Oak); 5 & 6, Odontopera bidentata (Scalloped Hazel); 7, Ennomos autumnaria (Large Thorn); 8 & 9, Ennomos alniaria (Canary-shouldered Thorn); 10 & 11, Ennomos erosaria (September Thorn); 12 & 13, Ennomos quercinaria (August Thorn); 14, Selenia tetralunaria (Purple Thorn). From Humphreys (1860, Plate XXXIII). • Ennominae: Kirby. ENNOMINAE. 1, Cleora cinctaria (Ringed Carpet); 2, Peribatodes rhomboidaria (Willow Beauty); 3, Peribatodes secundaria (Feathered Beauty); 4, Deileptenia ribeata (Satin Carpet); 5, Alcis repandata (Mottled Beauty); 6, Boarmia roboraria (Great Oak Beauty); 7, Serraca punctinalis (Pale Oak Beauty); 8, Fagivorina arenaria (Speckled Beauty); 9, Cleorodes lichenaria (Brussels Lace); 10, Ectropis crepuscularia (Small Engrailed); 11, Ectropis (Paradarisa) consonaria (Square-spot); 12, Ectropis similaria (Brindled White-spot); 13, Aethalura punctulata (Grey Birch Moth); 14, Gnophos furvata; 15, Gnophos obscuratus (Dark Annulet); 16, cf. Charissa (Rhopalognophos) glaucinaria; 17, Gnophos (Charissa) obscuratus (Annulet); 18, Gnophos obfuscatus (Scotch Annulet); 19, Psodos quadrifaria; 20, Macaria fusca; 21, "Fidonia fasciolaria", = Isturgia?; 22, Istugia famula; 23, Isturgia roraria; 24, "Fidonia plumistraria", = ?; 25, Ematurga atomaria (Common Heath); 26, Bupalus piniaria (Bordered White); 27, Selidosema brunnearia (Bordered Grey); 28, Itame vincularia; 29, Semiothisa (Macaria) wauaria (V-Moth); 30, Semiothisa (Itame) brunneata (Rannoch Looper); 31, Macaria artesiaria; 32, Petrophora chlorosata (Brown Silver Line); 33, Semiothisa (Macaria) clathrata (Latticed Heath). From Kirby 48, with updated names. • Biston betularia (Peppered Moth, typical and melanic forms), cf. Biston strataria (photos). Figs. 1–6, Peppered Moth (Biston betularia; Figs. 7–8, Oak Beauty (Biston strataria).

Figs 1 and 4–6 depict the Peppered Moth as represented in mixed deciduous woodland around Leek, N. Staffs, during the decade 1948–1958 before implementation of a "Clean air Act" in Britain. Localities "rural", but about 10 and 30 miles miles from the heavily industrial regions of Stoke-on-Trent and Manchester, respectively. Melanic individuals commonly came to light in the summer. They are exemplified by Fig. 6, seemingly representing the full melanic carbonaria form, and Figs. 4 and 5 perhaps representing the insularia complex. Fig. 1 portrays the only “normal” (typica) specimen encountered there in the decade, with Figs. 2 and 3 representing typica specimens from Holme Fen and Cirencester (Glos.) respectively.

The Peppered Moth affords a spectacular example of industrial melanism, and has been intensively studied in that context by population geneticists. For a brief summary of the phenomenon, and of the status of the Peppered Moth in that connection, select ‘Melanism in British Moths’ in the ‘Remaining taxa’ window; and for detailed discussion, see Majerus (1998, 2002).

Imagines of the Oak Beauty (B. strataria, Figs 7 and 8) were to be found resting by day on the trunks of oak trees in Spring. Unlike B. betularia, this species has never exhibited industrial melanism in Britain, although it does so in mainland Europe.

Specimens: 1, Leek, Staffs., July 1955; 2, Holme Fen, August 1956; 3, Cirencester, Glos., 1951 (S.G. Smith); 4–6, Leek, Staffs., June 1954; 7–8, Leek, Staffs., April 1952. From Watson’s collection. • Ectropis crepuscularia (Small Engrailed: typical and melanic forms, photos). Ectropis crepuscularia (Small Engrailed). The melanic form of this common species of deciduous woodland exemplifies spectacular drift from relatively infrequent “rural” melanism in unpolluted habitats to “industrial melanism”. In the decade 1948–1958, melanic individuals (below) predominated in the Leek district, in ostensibly rural habitats located 10 and 30 miles from industrial Stoke-on-Trent and Manchester, respectively. The upper photo portrays one of only two specimens of the light (“normal”) form that were encountered there during that time.

From Watson's collection. • Apterous Larentiinae: Phigalia (Pale Brindled Beauty), Erannis (Mottled Umber, Scarce Umber, Spring Usher, Dotted Border: Newman). • Agriopis leucophaearia (Spring Usher), A. marginaria (Dotted Border), A. aurantiaria (Scarce Umber): polymorphism and melanism (photos). Specimens of all three species were commonly found resting on tree trunks by day, in mixed deciduous woodland around Leek, N. Staffs, from 1948–1958. Localities ostensibly "rural", but about 10 and 30 miles miles from the heavily industrial regions of Stoke-on-Trent and Manchester, respectively.

Figs. 1–6, Agriopis leucophaearia (Spring Usher): exemplifying male colour variants of this common, polymorphic species, of which the lighter forms were numerically predominant. Figs. 7–9, A. aurantiaria (Scarce Umber): 2 typical males and (9) a female. By contrast with the other two representatives of this genus (and with Erannis defoliaria, q.v.), the Scarce Umber is not noticeably polymorphic, and melanism is either absent or very rare. Figs. 10–16, A. marginaria (Dotted Border): 3 light (“normal”) males, 3 melanics, and an apparently melanic female. Light and melanic forms were equally common, without obvious intermediates. Fig. 14 is a 1911 specimen from Ilfracombe (Devon, R.A. Nichols coll.), for comparison with Figs. 11 & 12.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, and distributions of character states within any set of taxa. See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 2003 onwards. Insects of Britain and Ireland: the families of Lepidoptera. Version: 14th April 2022. delta-intkey.com’.

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