POLYPHAGA Emery, 1886
TENEBRIONOIDEA Latreille, 1802
These small beetles develop in rotting vegetation, and the adults can often be seen on flower heads. Most species are coastal.
This is a large cosmopolitan family of about 3500 described species in more than 100 genera and 8 subfamilies although some other groups are variously included; Ischaliidae Blair, 1920, the Broad-hipped Flower Beetles, is a distinct family formerly included as a subfamily, and Afreminae Levey, 1985 and Lagrioidinae Abdullah & Abdullah, 1968 are generally included but their placement is doubtful. The family has historically been used to accommodate many tenebrionoid genera or species not easily placed elsewhere and as there is no modern analysis of the family there may well be significant changes in the future, more especially as several hundred species have been added in recent years and the pace of discovery is continuing, but the subfamilies listed below are generally accepted as constituting the family. Anthicids occur on all continents but temperate faunas tend to be limited e.g. 230 species are listed from the Nearctic region and the European fauna includes 400 species but diversity decreases rapidly with latitude e.g. only 27 species are listed from Germany, and by far the greatest diversity is in tropical and subtropical regions; in some rainforest regions of Asia and some semi-arid Afrotropical regions they occur in huge numbers and are second only to ants in terms of biomass.
As the common name suggests these beetles superficially resemble ants and, with some few exceptions outlined above, this is generally true; there is a wide variation in general morphology but these are variations on a basic theme and most species are readily recognized as members of the family. In life they also mimic ants in behaviour; their movements are rapid and jerky and the way they run with the legs splayed laterally is strongly suggestive of ants. Most can be recognized by the over habitus; the head is usually angled inwards behind relatively long temples to a distinct and often narrow neck, the pronotum is either rounded laterally or widest in front of the middle and narrower at the base than the base of the elytra, the elongate elytra are narrow and parallel or only weakly rounded laterally and the appendages are long and slender. Confusion with some other families can be avoided by being aware of the 5-5-4 tarsal formula. Most species are rather drab coloured; many have red, yellow and brown
or black patterns to the pronotum and/or elytra; banding or maculae which are aposematic, but these tend to be subtle rather than striking. Dorsal pubescence is very variable; sparse and fine or almost glabrous to dense and relatively long but most species are to some extent distinctly pubescent. The following description will apply to the majority of species and certainly those of the Anthicinae which is most relevant to the UK fauna.
1.2-18.0mm, in UK species 2.5-4.5mm. Head prognathous or only slightly inclined with the labrum at least partly visible from above, vertex convex and finely punctured and/or microsculptured, sometimes longitudinally furrowed but otherwise generally without major structure, eyes round to transversely oval, convex and sometimes prominent but usually relatively small, often with short and fine interfacetal setae, temples distinct, often long and contracted towards a base that is often truncate or nearly so. Antennae 10-12 segmented but 11-segmented in the vast majority of species, short to long; about half the body length, and filiform or only weakly and gradually expanded towards the apex, insertions visible from above and placed laterally on the frons anterior to the eyes. Frontoclypeal suture usually distinct, sometimes only faintly impressed, labrum usually transverse and well-sclerotized, mandibles small and curved; with a well-developed mola and truncate to simple or bi- or multidentate apically. Labial palpi 3-segmented with the apical segment fusiform, maxillary palpi 4-segmented with the apical segment cylindrical, fusiform or truncate and expanded. Pronotum elongate and simply oval to trapeziform, often broadest anterior to the middle and with indistinct anterior angles or simply rounded to a narrow neck, often constricted before the base but without a sub-basal transverse impression, anterior and posterior margins simple or bordered, lateral margins without borders. In Notoxus modified anteriorly into a horn which extends over the head, in some with a longitudinal median furrow but otherwise usually without major structure e.g. in Omonadus floralis (Linnaeus, 1758) there are 2 small tubercles towards the anterior margin. Prosternum short anterior to posteriorly open and contiguous coxal cavities, if present the process is short, mesosternum broad and rounded or triangular apically, the coxal cavities contiguous or narrowly separated and open or closed laterally. Metacoxal cavities transverse and separate. Scutellum small and usually triangular. Elytra entire and continuously rounded, only rarely abbreviated in some Anthicini, smooth and finely to strongly punctured, rarely striate, epipleura absent or narrow and variously continued to the apex. Surface without major structure. Wings variable; when present with a distinct anal lobe. Abdomen with 2 basal ventrites free and movable, remainder fused and immovable, segment 8 apparently without spiracles. Legs long and slender, femora often claviform, femora and tibiae sometimes with tooth-like expansions, all tibiae with apical spurs and tarsi always 5-5-4 with some, generally at least the penultimate, segment lobed ventrally. Claws simple or with a basal lobe, serrate, pectinate or bifid, sometimes with an empodium. Larvae are small <3mm when fully grown (UK species), elongate and more or less parallel-sided or slightly widened apically, cylindrical and poorly sclerotized but for the head, mouthparts and, where present, urogomphi. Head prognathous; without a distinct frontoclypeal suture but with a complete suture between the head capsule and the labrum, with a rounded labium and asymmetric bidentate mandibles. Antennae 3-segmented, labial palpi 2-segmented. Prothorax longer than the meso- and metathorax and a little narrower than the abdomen, all segments with 5-segmented legs, mesothorax with annular spiracles. All legs with a single moveable claw. Abdomen with 10 visible segments and much longer than the thorax; basal segments with annular spiracles and the terminal segment with paired cerci or fixed and unsegmented urogomphi.
In the UK both adults and larvae occur in a wide variety of habitats but many species occur in areas of sandy or loose soil with patchy vegetation and a supply of decaying organic matter. Larvae generally live among accumulated organic debris, within the soil or among decaying wood and under bark and, like the adults are mostly saprophagous or detritivores. Adults will occur through general sweeping of vegetation etc. and may sometimes be found in numbers on umbels and other flowers, in reed beds or among decaying plant matter, and sieving compost or accumulated litter over a sheet is a good way of finding them. Similar methods should be employed at the coast where diversity is generally higher, but searching under debris in salt marsh and estuary situations can be the most rewarding method, more especially as they tend to occur in large numbers in such situations. Notoxus may be found by sieving dry and loose sand in sparsely vegetated areas on sand dunes and slacks.
ANTHICIDAE Latreille, 1819
Ant Flower Beetles
The UK fauna includes Notoxus monoceros (Linnaeus, 1760), a widespread Palaearctic, African and Oriental species which is widespread and locally common on sandy soils around the coasts of England and Wales with occasional inland records, especially from the southeast, East Anglia and Wales. A further species, N. numidicus (Lucas, 1843) has been recorded on a single occasion from the North Somerset coast but is thought to be adventive and is not included in our checklist. Our remaining genera are all members of the Anthicini. Anthicus Paykull, 1798 is the largest genus of the family with about 600 species and is cosmopolitan in distribution; 5 species extend to the UK and most occur in sandy or salt marsh environments on the coast. A. angustatus Curtis, 1838 is very local and rare with only a few southern records. A. antherinus (Linnaeus, 1761) is widespread throughout central and southeast England, occurring in open dry habitats inland as well as coastal. A. bimaculatus (Illiger, 1801) is rare and sporadic around the coasts of Wales and Southern England. A. flavipes (Panzer, 1797) is a very local species from coastal North Wales, Northwest England and Southwest Scotland. A. tristis Schmidt, 1842 is a very local species of the Hampshire coast. Cordicollis Marsuel, 1879 includes about 80 species from the Palaearctic, African and Oriental regions. A single species, C. instabilis (Schmidt, 1842) occurs in sandy and salt marsh habitats around the southeast and southern coasts of England. Cyclodinus Mulsant & Rey, 1866 is a cosmopolitan genus of about 90 species, 2 of which; C. salinus (Crotch, 1867) and C. constrictus (Curtis, 1838) are local around the southeast coast of England. The cosmopolitan genus Omonadus Mulsant & Rey, 1866 includes about 40 species of which 3 occur in the UK; O. bifasciatus (Rossi, 1792) is a local insect of south and southeast England while O. floralis (Linnaeus, 1758) occurs throughout the mainland and O. formicarius (Goeze, 1777) is widespread in the south, all are associated with decaying vegetation and adults occur on flowers, none are particularly coastal. The cosmopolitan genus Stricticollis Marseul, 1879) includes about 50 species, of which one, the cosmopolitan C. tobias (Marseul, 1879), is of local occurrence in southeast and central England.
Around the World
Tomoderinae Bonadona, 1961. This large group of more than 350 species in 6 genera is almost exclusively tropical although a few species of the pantropical genus Tomoderus LeFerté-Sénectére, 1849 extend into temperate regions; 3 occur in the United States and several have been recorded from southern Europe. They are generally distinct within the family in having the pronotum strongly constricted near the middle or towards the base so that it appears to be constructed of two distinct lobes, they are otherwise very diverse morphologically although overall they are distinctively ant-like and typical of the family. Most species are omnivores inhabiting rainforest litter, decaying timber, rot-holes etc. or living in the canopy and most species fly well, are widespread and occur in large numbers although a few are apterous and confined to the forest floor, among litter or living within the soil. Some south-east Asian species are associated with ants or termites and some are strictly riparian. Includes both diurnal and nocturnal species.
Steropinae Jacquelin du Val, 1863. Two genera and nine species are described. The monotypic genus Anisotria Young, 1984, originally described as a pyrochroid, occurs in North America while members of Steropes Stevens, 1806 are mostly Asian; several extend to eastern Europe and a single species occurs in Vietnam. Members are diurnal and mostly associated with arid regions where they are active on vegetation or on the ground although the quaintly-named S. hercules Telnov, 2007 is found in rainforests in Vietnam. They are rather more elongate and flattened than most anthicids and, with the exception of Anisotria, densely pubescent and so might be mistaken for several tenebrionoid families; Pyrochroidae, Melandryidae or Tenebrionidae. Males are distinct in having the three terminal antennomeres greatly elongated.
Macratriinae Leconte, 1862 includes two New World genera, the monotypic Thambopasta Werner, 1974 from the United States, the monotypic Salimuzzamania Abdullah, 1968 from Guatemala, and Macratria Newman, 1838 which includes about 200 species and occurs mostly in warmer areas throughout the world. Most species occur in rainforests and may be very abundant in riparian situations, many are nocturnal and come to light in huge numbers, and many are polyphagous on a range of monocotyledon and dicotyledons plants. In appearance they are typical of the family but, at least in Macratria, the elytra have distinct longitudinal lateral sulci; in many the elytra are striate and the pronotum is transversely depressed before the base, often giving the impression of a false basal margin.
Lemodinae Lawrence & Britton, 1991. This small subfamily includes 6 genera and about 40 species and, with the exception of 2 species of the Chilean genus Protoanthicus Moore & Vidal, 1991, is Australasian, extending to New Guinea and the Solomon Islands. The monotypic Zealanthicus Werner & Chandler, 1995 is a New Zealand endemic and the monotypic Lemodinus Blair, 1913 is Australian, while the others are more widespread; Trichananca Blackburn, 1891 and Lemodes Boheman, 1858 throughout the region, and Cotes Sharp, 1877 in Polynesia including New Zealand. Species occur from dry lowland savannahs to mountain altitudes but most are associated with rainforest habitats; on foliage of a wide range of plants or among litter or wood and some Trichananca seem to be epigean. So far none have been recorded associated with other insects. Many will be recognized as rather typical of the family; elongate and with a broad head which is produced anterior to convex and prominent eyes, many have deep longitudinal pronotal sulcus and strongly punctured elytra, either randomly or striate. Some Lemodes and Trichananca are strikingly coloured or patterned with red, yellow and black; Lemodes coccinea (Boheman, 1858) is a fine example.
Copobaeninae Abdullah, 1969. Four species of the single genus Copobaenus Fairmaire & Germain, 1863 are known from mountain forests in Chile and Argentina. They are convex and very elongate species with strongly and densely punctured elytra; among the family they are unusual in having metallic green or blue elytra and a pale forebody. Adults are diurnal but very little is known of their biology.
Eurygeniinae LeConte, 1862 is a large subfamily of about 30 genera in 3 tribes. Species occur worldwide but the group is most diverse in the New World and Australia. Mitraelabrini Abdullah, 1969 includes only the monotypic genus Mitraelabrus Solier, 1969 from Chile. Ictistygnini Borchmann, 1937 is a small group of 5 genera and about 20 species; the monotypic genera Diacallina Champion, 1916 and Ictistygnina Champion, 1916 occur in West Africa and Brazil respectively while the remainder are Australian. Eurygeniini LeConte, 1862 is pantropical, extending into North America, where the fauna is diverse with 50 species in 11 genera, Japan and southern and eastern Asia but absent from the western Palaearctic. Species are often difficult to distinguish from those of other subfamilies or some other tenebrionoid families e.g. Oedemeridae, but the group includes some of the largest members of the family, reaching 18mm. They are typically elongate and slender with large, anteriorly emarginate eyes, finely or moderately strongly and randomly punctured elytra and dorsally pubescent.
Notoxinae Stephens, 1829. Commonly known as monoceros beetles due to the anterior part of the pronotum being produced over the head in the fashion of a horn, and so among the most readily recognizable of beetle groups. About 400 species are known from 7 genera and, with the exception of the Neotropical region, occur worldwide with the greatest diversity in Africa. Species occur in a wide range of habitats from lowlands to subalpine zones and they are often prolific in rainforests, many are associated with light or sandy soils where the pronotal horn is used for burrowing. Adults are generally omnivorous, although a few have been reported to be predatory, feeding on plant parts or remains and pollen while the larvae develop among roots. In temperate regions mating and oviposition occurs in the spring; eggs are laid in groups in a subterranean tunnel excavated by the female for the purpose. Development is generally rapid with larvae emerging within two weeks and becoming fully-grown within two months, larvae burrow into the soil to pupate within a puparium prepared from vegetable debris and sand and adults eclose about two weeks after pupation. Many species of the worldwide genus Notoxus are attracted to the poisonous chemical cantharidin and may be found alongside oil beetles, males of some have specialized structures at the elytral apices which concentrate the chemical, this is passed to females as they feed upon male secretions and so it is passed to the eggs and larvae, it is thus used as a deterrent against predators and parasites.
Anthicinae Latreille, 1819. This is by far the largest subfamily with about 2500 species of 41 genera in 4 tribes; the group is cosmopolitan with the greatest diversity in tropical and subtropical regions. Temperate faunas tend to be well-represented by the Anthicini Latreille, 1819; 120 species occur in North America and almost 400 in Europe but most are from warmer southern regions and the central European fauna includes about 25 species. The tribe otherwise includes 32 genera of which many are restricted to Old World or New World tropical regions. Endomiini Bonadona, 1958 includes the single genus Endomia Laporte, 1840 which is widespread throughout the Old World and includes a single central European species. Formicomini Bucciarelli, 1980 is a mostly tropical group; it is absent from The Nearctic area although well-represented in the eastern Palaearctic and Oriental regions, European genera include Stenidius LeFerté-Sénectére, 1849, Anthepiscopus Becker, 1891 and the very large genus Anthelephila Hope, 1833 which extends into central Europe but not north to the UK. Microhorini Bucciarelli, 1980 is a widespread Old World group of 5 genera, it is well represented in all regions including Europe but only 2 species of Microhoria Chevrolat, 1877 extend into central Europe.