Entiminae | uk beetles

ENTIMINAE Schönherr, 1823

Broad-nosed Weevils

Characterised by their short rostrum, this group includes some of the largest British weevil genera. Many species are flightless, and most feed on a wide range of foodplants.

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POLYPHAGA Emery, 1886

CURCULIONOIDEA Latreille, 1802

CURCULIONIDAE Latreille, 1802

115

2.1-15mm

Introduction

This is a large group of generally distinct weevils; more than 12300 species of 1370 genera and 55 tribes are described and all regions have a diverse fauna, the greatest diversity is in tropical regions but they are known from most oceanic islands and many have a diverse endemic fauna, both northern and southern temperate regions are diverse in species and higher taxa. The group includes many large genera that are widely distributed while some are more restricted e.g. Otiorhynchus Germar, 1822 includes more than 1500 species in 105 subgenera and is cosmopolitan, Polydrusus Germar, 1817 includes about 225 species and is almost exclusive to Old World regions, Myllocerus Schöenherr, 1823 includes about 350 species and is widespread across Eurasia, Africa and Australia while Compsus Schöenherr, 1823, with more than 100 species, is a New World group almost entirely restricted to south America. On a world scale many of the tribes are restricted to a single biogeographical region or an area within a single region, about ten occur across two regions, generally Nearctic and Palaearctic or Palaearctic and Oriental, and only a few are generally distributed throughout the world. Among the tribes of more restricted distribution include Nothognathini Marshall, 1916 from India and Pakistan, Ophtalmorrhynchini Hoffmann, 1965 from Central Africa, Premnotrypini Kuschel, 1956 from the high Andes, Typhlorhinini Kuschel, 1954 from Madagascar and Laparocerini Lacordaire, 1863 from the Azores, Madeira and the Canary Islands, and several others are known from various oceanic island chains. Agraphini Horn, 1876, Hormorini Horn, 1876 and Ophryastini Lacordaire, 1863, along with twenty more widely distributed tribes are Nearctic while Phyllobiini Schönherr, 1826, Sciaphilini Sharp, 1891, Holcorhinini Desbrochers, 1898 and Otiorhynchini Schönherr, 1826 are mostly Palaearctic. Neotropical tribes include Anypotactini Champion, 1911, Entimini Schöenherr, 1823, Eudiagogini LeConte, 1874, Eustylini and Lordopini Schönherr, 1823 while Embrithini Marshall, 1942, Oosomini Lacordaire, 1863 and Tanyrhychini Schönherr, 1826 are Afrotropical. Mesostylini Reitter, 1913 and Nastini Reitter, 1913 are Oriental, Celeuthetini Lacordaire, 1863 and Ottistirini Heller, 1925 are Australasian and Anomophthalmini Morrone, 1998 is endemic to Patagonia. The Palaearctic fauna includes 28 tribes of which 14 extend to the UK, and the following discussion is an attempt to characterize this wider fauna in terms of our own. Tribes occurring in the UK are underlined.

BLOSYRINI Lacordaire, 1863 includes 5 genera and 50 species and occurs in the eastern Palaearctic and Oriental regions but is absent from Japan; the greatest diversity is China and the largest genus, Dactylotus Schönherr, 1847, with 35 species, is more or less exclusive to that country.
BRACHYDERINI Schönherr, 1826 is a large and widespread tribe of 21 genera and more than 200 species, most are confined to certain areas of Asia but the group is well-represented in Europe with 8 genera and 86 species. The UK fauna includes 8 species of 2 genera. Of our two species of Brachyderes Schönherr, 1826 one, B. incanus (Linnaeus, 1758) is widespread across central and western Europe while B. lusitanicus (Fabricius, 1781) occurs only in Spain and Portugal; their presence in the UK is from specimens introduced with horticultural stock and it is unlikely they will become established in the long term. The genus includes a further 6 species from the west of Europe and northwest Africa. Strophosoma Billberg, 1820 is the largest genus of the tribe with 75 species, the European fauna is diverse with about 50 species and the remainder occur in western regions; northwest Africa and Asia Minor. The UK fauna includes 6 species, all of which are widespread in Western Europe. The only other large genus, Pholicodes Schönherr, 1826, which includes 40 species, is primarily Asian although 15 species extend into eastern parts of Europe. The tribe is otherwise most diverse in western parts of Asia and North Africa.
BYRSOPAPAGINI Lacordaire, 1863 includes 15 genera and 113 Palaearctic species, it is a widespread group with several genera restricted to eastern regions, including Japan, but The European fauna is diverse with 43 species of 5 genera and many more occur in Asia Minor and North Africa. Graptus Schönherr, 1823, sometimes included in the Alophini LeConte, 1874 is a western genus with 29 species occurring in Europe of which the very widespread G. triguttatus (Fabricius, 1775) extends to the UK. Trophiphorus Schönherr, 1842, sometimes included in the Tropiphorini Marseul, 1863, includes 17 species, mostly of restricted distribution in southwest Europe and northwest Africa, but 3 are widespread and extend to the UK.
CELEUTHETINI Lacordaire, 1863 is a far-eastern group of 21 species in 8 genera, most occur in Japan while a few are endemic to Thailand.
CNEORHININI Lacordaire, 1863 includes about 100 species in 15 genera and is widespread across the region, several genera are restricted to eastern regions; China and Japan being diverse, but others are more western e.g. the 4 species of Polydius Dejean, are restricted to Europe, and north-west Africa is very diverse,28 species are listed from Europe. Of the 7 species of Philopedon Schönherr, 1826, all of which occur in the west of Europe or north-western Africa, the widespread and generally common P. plagiatum (Schaller, 1783) extends to the UK. Attactagenus Tournier, 1876 includes 18 species and is restricted to the western extremes of Europe and North-west Africa and only the most widespread species, A. plumbeus (Marsham, 1802), extends to the UK.
CRATOPODINI Hustache, 1919 includes only the monotypic genus Hemicratopus Voss, 1964 and is endemic to India.
CYPHICERINI Lacordaire, 1863 is a large group of about 470 species included in 85 genera and 5 subtribes, the greatest diversity is in Asian and oriental regions and only a 4 or 5 have been recorded from Europe, none of which have occurred in the UK.
EPISOMINI Lacordaire, 1863 includes 21 species of Episomus Schönherr, 1823 from southern Asia and the Oriental region and a single monotypic genus from India, none are known from Europe.
GEOMENINI Gistel, 1848 includes 31 species of 3 genera and a further 6 species of uncertain placement. They are almost entirely distributed in southern Europe and northwest Africa although 3 species of Barynotus Germar, 1817 are much more widespread and extend to the UK; the genus includes a further 22 species.
GONIPTERINI Lacordaire, 1863 includes 2 species of Gonipterus Schönherr, 1833 from southern Europe, neither of which has been recorded from the UK.
HOLCORHININI Desbrochers, 1989. This tribe is almost entirely confined to North Africa, from the extreme northwest east to Egypt, with only a very few occurring in the south of Europe; Spain, Italy and Greece. The Palaearctic fauna includes about 70 species in 23 genera.
LAPAROCERINI Lacordaire, 1863 is a very diverse group confined to the Atlantic island groups off North Africa. It includes a monotypic genus from the Canary Islands, and more than 200 species included in 20 subgenera of the genus Laparocerus Schönherr, 1834 which is widespread through Gran Canaria, Tenerife, Madeira and Lanzarote etc. So far as is known none are established in mainland Europe or Africa.
MESOSTYLINI Reitter, 1913 includes 12 species in 5 genera, all have quite restricted distributions in southern and central Asia and none extend to Europe.
MYORHININI Marseul, 1863 is a ‘western’ tribe occurring in Asia Minor and Europe, it includes 7 species in 5 genera of which Aspis Germar, 1820 (2 spp.) and the monotypic Haptomerus Faust, 1899 and Subhaptomerus Hoffmann, 1960 are local in southern Europe but none have been recorded from the UK.
NASTINI Reitter, 1913 includes about 40 species of the single genus Nastus Schönherr, 1842, all have a central or western distribution centred around Kazakhstan, Uzbekistan and southern Russia although a few extend into Asia Minor and the southeast of Europe. None have been recorded from the UK.
NAUPACTINI Gistel, 1848 includes 3 species of Naupactus Dejean, 1821, one is widespread in southern Europe and two are endemic to the Azores, all have become widely established in North and South America as well as variously in Africa and Australasia but none have been recorded from the UK.
NOTHOGNATHINI Marshall, 1916 includes two genera; Lathrotiorrhynchus Voss, 1964 with 6 species from India, and Nothognathus Marshall, 1916 with 2 species endemic to Pakistan.
OMIINI Shuckard, 1840 is a widespread tribe which is most diverse in Europe, Asia Minor and North Africa and relatively poorly represented in eastern regions although Asphalmus Sharp, 1896, with about 45 species, is endemic to Japan. The tribe includes about 230 Palaearctic species in 21 genera, of which about 110 species of 14 genera occur in Europe, and some are almost exclusively European e.g. Amicromias Reitter, 1913 with 17 species mostly from south-eastern regions, and the widespread Omias Germar, 1817 which includes almost 50 species. The tribe includes many endemic groups as well as many widespread European species e.g. the genus Omiamima Silferberg, 1977 includes 7 species restricted to relatively small areas in Asia Minor, Southern Europe and North Africa but a further species, O. mollina (Boheman, 1834) is widespread across Europe and extends into the UK, it is the only member of the tribe to do so.
OPHRYASTINI Lacordaire, 1863 includes 28 species of the genus Deracanthus Schönherr, 1823 distributed throughout Asia from Kazakhstan to China.
OTIORHYNCHINI Schönherr, 1826 is the largest tribe with more than 1300 Palaearctic species in 18 genera, by far the greatest diversity is in Europe where more than 1050 species of 12 genera are listed and beyond this many more occur in adjacent regions of North Africa and Asia Minor. Much of this diversity is within the single genus Otiorhynchus Germar, 1822; of the 1200 or so Palaearctic species, included in numerous subgenera, about 950 occur in Europe and there are many regional subspecies. Otiorhynchus is the only genus to occur in the UK where it is represented by 17 subgenera and 25 species.
PACHYRHYNCHINI Schönherr, 1826 includes 9 species in 2 genera and occurs in Japan and on various islands off Taiwan.
PERITELINI Lacordaire, 1863 is a large tribe of about 320 species in 30 genera, by far the greatest diversity is in Mediterranean regions; there are many genera endemic to Morocco, Tunisia, Yemen and the Atlantic islands, and many more are confined to southern areas of Spain, France and Italy. Two genera are represented by single species in the UK. Simo Dejean, 1821 includes 4 species, one endemic to Italy (Sicily) and another endemic to Algeria, two are widespread and of these S. hirticollis (Herbst, 1795) has recently been added to our list. Peritelus Germar, 1824 includes two species from northwest Africa and two that are widespread in Europe, of which one, P. sphaeroides Germar, 1824 occurs in the UK.
PHYLLOBIINI Schönherr, 1826 includes about 200 species and occurs throughout the Palaearctic region but is most diverse in Europe and surrounding areas, especially around the Mediterranean. More than half are included in the large genus Phyllobius Germar, 1823 which is classified into 19 subgenera and includes many species from central Asia and Europe. Among the more common and widespread species 10 extend to the UK, representing 7 subgenera, and they are the only members of the tribe to occur here.
POLYDRUSINI Schönherr, 1823 includes about 260 species in 12 genera, the majority of species occur in Europe and the surrounding areas of North Africa and Asia Minor are also very diverse, most of the remainder occur in western parts of Asia and very few extend to the far east. Our UK fauna includes 14 species of 3 genera. Of the 10 species of Liophloeus Germar, 1817, 9 have a rather restricted and mostly southern European distribution while one, L. tessulatus (Muller, O. F., 1776) occurs throughout the region, extending north to Fennoscandia and the UK. Pachyrhinus Schönherr, 1823 includes 29 species in 3 subgenera and is most diverse in North Africa and southern Europe, a few extend into central and eastern Asia and 2 are endemic to Japan. Most are endemic to Morocco, Tunisia, Algeria and various northern Mediterranean countries and only two are widespread in Europe, extending north into the UK. The tribe is dominated by the western, and mostly European, genus Polydrusus Germar, 1817, which includes almost 200 species, most of which have a restricted distribution but of the relatively few widely-distributed species 11 extend north to the UK.
PASLLIDIINI Lacordaire, 1863 is a mostly Asian group with many species of restricted distribution in central Asia, Asia Minor and Greece. The tribe includes 73 species in 3 genera, 2 of the genera are widespread while Sphingorrhinotus Voss, 1957, with 2 species, is endemic to Pakistan.
SCIAPHILINI Sharp, 1891 is a large tribe of more than 150 species in 33 genera, most of the smaller genera are restricted to areas of Europe, North Africa or central Asia while the larger genera are generally widespread, the exception is the largest genus, Brachysomus Schönherr, 1823 which is generally European with a few species in North Africa and Asia Minor; it includes 60 species but of these only 5 are at all widespread and of these 2 extend to the UK. Exomias Bedel, 1883 includes 33 species and is, with a single exception from Mongolia, entirely European. Most species are of very restricted and generally southern distribution but among the most widespread are E. araneiformis (Schrank, 1781) and E. pellucidus (Boheman, 1834) which occur throughout and extend to the UK. Our UK list also includes 2 other species; E. curvimanus (Jacquilin du Val, 1854), otherwise known only from France, and E. pyrenaeus (Seidlitz, 1868) which occurs also in Spain, France and Switzerland. Sciaphilus Schönherr, 1823 includes 5 species, 2 occur in the west of Europe, one is endemic to Algeria, one from Asia and the very widespread European S. asperatus (Bonsdorff, 1785) which extends to the UK.
SITONINI Gistel, 1848 includes about 140 species in 7 genera, it ranges east to Mongolia and Central Asia but by far the greatest diversity is in Europe and North Africa. Nine species of 2 genera are restricted to Asia, 4 small genera are European and Sitona Germar, 1817, with about 110 species occurs throughout the region but is mostly European and North African, about 30 are widespread in Europe and of these 15 extend to the UK. The monotypic Andrion Velázquez de Castro, 2007 occurs throughout Europe and extends to northwest Africa and the UK. Charagmus Schönherr, 1826 includes 6 widespread European species, 2 of which occur in the UK. Coelositona González, 1971 includes 10 species, all of which occur in or adjacent to Europe; 5 are widespread and of these 2 occur in the UK.
TANYMECINI Lacordaire, 1863. Among the largest of the tribes, this group includes more than 950 Palaearctic species in 53 genera and 3 subtribes, it is represented in Asia Minor and North Africa by only a few species and a very few occur in southeast Europe but the majority occur in far eastern regions, and more especially in China. Piazomiina Reitter, 1913 is the largest subtribe with about 600 species and is entirely Asian and Oriental while Tainophthalmina Desbrochers, 1873, with 45 species, and tanymecina Lacordaire, 1863, with about 320 species, extend further west and include a few European species. Tanymecus Germar, 1817 is one of the few ‘western’ genera, it includes 33 species which are mostly of very restricted distribution in Asia Minor, North Africa and Mediterranean Europe but one, T. palliates (Fabricius, 1787), occurs throughout Europe and western Asia and extends north to the UK.
TRACHYPHLOEINI Gistel, 1848 imcludes about 340 species in 13 genera and diverse throughout the region; Europe and North Africa are the most speciose areas but to the east Japan is notable for having two endemic genera, one is monotypic while Trachyphilus Faust, 1887 includes about 90 species. The UK fauna includes 16 species of 4 genera. Caenopsis Bach, 1854 includes 17 species, most are local to areas in the south and west of the region but two are more widespread and extend to the UK. Cathormiocerus Schönherr, 1842 is widespread in Europe and North Africa, it includes about 95 species of which 6 have been recorded in the UK. Ronualdius Borovec, 2009, with 11 species, and Trachyphloeus Germar, 1817, with about 80, occur mostly in the extreme southwest of Europe and in North West Africa, and most species are of very restricted distribution but both include widespread species that occur in the UK.

Vine weevil larva (Otiorhynchus sulcatus)

http://data.nhm.ac.uk/dataset/collection-specimens

Ecology

A feature of the group is larvae that develop freely in soil, feeding externally on roots and various other parts of plants. Only adults are found above the soil; they generally oviposit in the soil and, following larvae development, they will emerge from subterranean pupal cells before making their way to the surface. Most species oviposit in the spring or summer and winter is passed in the larval stage, pupation occurs early in the year and adults feed nocturnally on leaves and shoots, and to the experienced eye many species may be detected by their characteristic notched feeding patterns. In colder northern latitudes the life cycle may be extended over several years but this is exceptional. Adults of many species are wingless and so populations may be very localized, nonetheless some species have become very widespread through being transported in soil along with nursery stock and some have become serious pests, the most notorious being the Vine Weevil, a native European species that has become widespread in temperate regions throughout the world.

Adults of many species are long-lived, in the case of the Vine Weevil up to three years, and this contributes to spread of pest species and their ability to generate large local populations, and under the ideal conditions often found in commercial nurseries they may become continuously brooded. Unlike those of many weevil groups females do not use the rostrum to prepare or assist with oviposition; instead in the majority of cases eggs are deposited directly into the soil, usually singly or in small batches close to or upon host material, some species will deposit them on the soil beneath decaying vegetation or on the host stem at ground level, some simply drop them onto the surface and other strategies are sometimes used e.g. some Pachyrhynchini oviposit in bark. Compared with other weevils entiminids are very fecund; females may deposit batches of between 20 and 80 eggs which they coat with a gelatinous secretion for protection, and oviposition may continue over long periods. Eggs generally hatch within two or three weeks, depending upon temperature and humidity, and larvae begin searching for host material and feeding immediately. Larvae of most are polyphagous, and most can develop on plants of a wide range of angiosperm families, they feed on soft roots and may move through the soil in search of roots when a host is exhausted, here they can switch host species and continue developing normally. The number of instars varies and up to 11 have been recorded. Soil type and condition is sometimes important e.g. in the UK some species are prefer sandy or chalky districts but many seem to tolerate a wide range of soils. A few groups have developed specialized feeding habits; larvae of some Ectemnorhinini-a small group restricted to various islands in the southern Indian Ocean-live on the soil surface beneath deep leaf-litter and feed on algae, while larvae of some Pachyrhynchini feed on aerial parts of trees and shrubs, tunnelling in branches and pupating in a subcortical cell. Some tropical groups, especially in humid rainforest habitats, are known to feed on various lichens, algae and mosses both as larvae and adults, and here some adults have formed very special relationships with cryptogams which grow on various body surfaces and so provide protection through camouflage, other tropical species feed on algae in coastal situations and have very high salt tolerences. Monocotyledons of all kinds host entenimids, among dicotyledons the most popular group seems to be legumes but various families are popular including Fabaceae, Fagaceae, Malvaceae, Rosaceae, Rutaceae and Solanaceae and families include hosts. Most species are polyphagous and many very widely so; there are many pest species which have become widely established outside their native ranges and these tend to be polyphagous both as adults and larvae, some have lists of hundreds of hosts and this is another factor contributing to their success, adults tend to do little damage but the long-lived larvae can be very destructive. Some do have host preferences and some are fairly restricted, e.g. Phyllobius oblongus which can develop on a range of trees and shrubs but is more generally associated with elms, limes and hawthorne, and in the UK Barynotus moerens is usually associated with dog’s mercury. Larvae and adults can have different hosts or, most commonly, both stages are polyphagous.

Parthenogenesis is relatively frequent in the subfamily although the vast majority of species reproduce sexually; this is a rarely seen in beetles generally, it is known from only 3 subfamilies of weevils, and of the 75 or so species known to reproduce in this way more than 50 belong in the present subfamily although in some cases they may be parthenogenetic only in certain parts of their range. Certain genera e.g. Otiorhynchus, are known to include several parthenogenetic lineages, these are most diverse in the Palaearctic region but other genera are known from across the world, and, curiously such species are almost always flightless. The UK fauna includes several examples although in some cases males have been found in Europe, these include such familiar species as Barynotus moerens, Strophosoma melanogrammum, and various Otiorhynchus among others. In such species females develop from unfertilized eggs and because there is no meiotic stage the generations are genetically identical unless there is a mutation at the egg stage. Apomictic weevils-those that bypass the fertilization and meiotic phases-are frequently polyploidy and typically triploid, and compared with the more usual diploid condition they often have a larger body size. Apometic weevils are more common among the better-studied faunas but it may be just as common across the world as it has arisen repeatedly in the subfamily, it may generate more potential for rapid adaptation and diversification than in diploid species and contribute to understanding the great diversity of some genera.

Description

Broad-nosed weevils soon become obvious, especially among the limited UK fauna, they are appropriately named as almost all have a characteristic short and broad rostrum, but there are many exceptions; among the UK fauna species of Tropiphorus Schönherr, 1842 have a relatively long rostrum, and there are other weevils with a short rostrum, most obviously scolytids but also e.g. Rhinocyllus conicus (Frölich, 1792). Another feature seen in most groups, although notably absent in e.g. Sitonini Gistel, 1848 and some genera of Alophini LeConte, 1874, is an oval or round scar on the mandibles which is produced soon after the adult leaves the pupa by the shedding of a ‘deciduous mandibular process’, a sickle-shaped appendage used by the adult free itself of the pupal skin and surrounding soil. In most species, and all of those in the UK, the scar is present on both mandibles, they are usually quite large and easily seen but in many species of Trachyphloeini Gistel, 1848 they may be small and inconspicuous. A more general characteristic of the group is the large mentum which covers the apical excision of the rostrum, and the absence of a submentum, but these features are only visible from underneath and are often difficult to appreciate. Most species are medium to large; among our fauna from about 2mm in Brachysomus Schönherr, 1823 to 15mm in Otiorhynchus Germar, 1822 but most are between 4 and 10mm, they are mostly elongate, discontinuous in outline and very convex. In temperate regions most are drab brown, grey or black, often patterned or tessellated and with widely varying vestiture, they may be glabrous, entirely and densely, patchily or sparsely scaled, finely and densely pubescent or have long and sparse setae on the dorsal surface; only a few groups e.g. Phyllobius Germar, 1824 and Polydrusus Germar, 1817 are brightly coloured. The morphology of both the setae and the scales varies widely and several distinct types of each may be present on a single species. The head is clearly visible from above; the vertex and frons are usually wide and may be simply convex, longitudinally impressed or have various fovea or raised areas, especially around the eyes, the temples are generally short and may be variously enlarged, the eyes are usually relatively large and visible from above, they may be simply round and convex, flattened, reniform or asymmetrical. The rostrum may be parallel-sided or expanded towards the apex and is often longitudinally impressed, sometimes simply so but very often with series of parallel or oblique channels, the apex is variously notched and the mandibles are usually obvious when viewed from in front. The scrobes are very variable; when placed laterally they tend to be parallel-sided and quite short, they may be straight and terminate in front of the eyes or curved down in front of the eyes, almost onto the ventral surface, in many genera they are placed on the dorsal surface and here they are often expanded and rounded anteriorly and fully-visible from above, this is best appreciated in some species of Otiorhynchus. The antennae always consist of three well-developed and sharply-defined parts; the scape is usually long, often curved and dilated distally, the funiculus consists of 4-8 segments which vary greatly in morphology even within a genus, and a club which usually consists of three closely-united segments, the vestiture of various parts, especially the scape can be diagnostic and may consist of scales, setae or pubescence. The prothorax is usually near-quadrate and when distinctly elongate or transverse then only weakly so, the surface is usually distinctly and often strongly and densely punctured, in many there are tubercles and these may be strongly raised and cover the entire dorsal surface and in some there is a longitudinal keel or impression, the margins, especially the lateral margins, usually lack distinct borders. The basal margin is usually closely fitting against the base of the elytra and the anterior margin may be concave, straight or produced, in many the lateral margin is produced behind the eyes although in Tanymecus this is replaced by a series of stiff setae, the so called ‘postocular bristles’. The prosternum is often densely-scaled, it may be long or short in front of the coxae which are usually round, convex and often projecting, and placed close together without a process or (rather obviously) a rostral channel. The elytra vary from globular to elongate and parallel-sided, the shoulders are usually well-developed and rounded or sloping, in fully-winged species they tend to be well-developed, the lateral margins generally reflexed ventrally and closely associated with the abdomen, each has up to ten regularly punctured striae, the tenth being abbreviated, but this is very variable and even in a single genus species may lack striae altogether or have very strongly punctured and complete striae. Similarly the interstices are very variable, from smooth to very roughly rugose or tuberculate, from flat to very convex and in many species alternate interstices may be raised and flat, in many flightless forms the suture is fused and in the majority of species the elytra completely cover the abdomen. Abdomen with two basal ventrites connate but with distinct sutures, ventrites 3-5 freely articulated. The mesosternum is short and wide, the anterior margin often produced between the coxae and the posterior margin simply closing the meta-coxae, in most it s weakly convex and devoid of structure. The meso-coxae are round or a little transverse and closely approximated, in most they are convex and projecting although not so strongly as the pro-coxae. Metasternum deep, wide and usually roundly-produced between the meta-coxae to meet the meso-sternum. Meta-coxae variously transverse and rather flat, usually widely separated, sometimes they are virtually circular in which case they are widely separated and displaced towards the lateral margins. Trochanters proportionately small, sometimes globular and often difficult to see among dense scaling. Femora robust, often very much so, and clavate, often excavate ventrally for the reception of the tibiae, they may be smooth but are often toothed, the form of the femoral tooth varies widely in shape and size and there may be two or three or a single tooth may be bifid or trifid. The tibiae are long and robust, the inner margin may be smooth or toothed and the inner apical angle may bear a small tooth or ‘mucro’ (where this is present they are termed mucronate), and the outer apical angle may be modified into a tooth-like process called an uncus (in which case the tibiae are termed ‘uncinate’). The terminal margin may be delimited by an oval area fringed with bristles, a corbel, in which case the tarsi may be attached inside or outside the corbel, in the former case the tarsi appear at the apex and the tibiae are termed –open’, in the latter case they appear to originate on the lateral margin and the tibiae are termed ‘closed’. The tarsi are typical of the family; 5-segmented with the third widely bilobed and enclosing a small fourth segment, a long and gradually expanded terminal segment and free or connate claws. This description is appropriate for many species in both southern and northern temperate regions but, as would be expected with such a large and cosmopolitan group, species from warmer regions are very diverse and not immediately obvious as members of the subfamily.

Some tropical species exceed 30mm in length and have exaggerated tubercles or other structures to the pronotum and/or elytra, those from arid regions tend to be drab, dark brown to black, lack scales but have variously developed setae and/or pubescence, in a few cases they are covered in a brightly-coloured waxy secretion over a dense covering of small adpressed scales. Those from tropical forests tend to be large and covered with dense scales that form striking, often aposematic, colour patterns.