CERAMBYCIDAE Latreille, 1802
Ever popular with collectors, this group includes some of the most impressive species in the world, as well as in the UK.
POLYPHAGA Emery, 1886
CHRYSOMELOIDEA Latreille, 1802
Around the World
This is one of the largest families of the Coleoptera with about 35000 described species in 4000 genera and ten subfamilies, it has always been popular with collectors at all levels, hence the large number of described species, but the classification remains fluid because each subfamily varies widely and there are generally species in each group which defy the overall description, and this applies even at the family level (see below). In many parts of the world longhorns are collected commercially and sold to collectors at all levels, this remains a thriving business but recently many countries have tried to stop this by making it illegal without a licence. As to whether such commercial activity harms those desirable species that fetch high prices is a matter of debate but viewed against the environmental vandalism that continues in many tropical areas it can only be thought of as trivial. They are a very diverse group but in general, with due acknowledgment to the families outlined at the end of this page, they are recognizable. It is also a family where the life history of many species is understood, firstly because many are reared by collectors from samples of host material taken from the field and kept indoors, and secondly because many of the members are pest species of commercially important plant products, especially timber, and so research into their biology receives regular funding. Thus cerambycid faunas tend to be rather dynamic because of the worldwide movement of timber etc. and in many temperate areas the ‘occasionally imported’ list is lengthy compared with that of many other groups. The family is cosmopolitan, occurring from sea-level to about 4000m and in a very wide range of habitats although in temperate regions they are generally insects of wooded areas. They range in size from a few millimetres to some of the largest species of Coleoptera; the Titan beetle, Titanus giganteus (Linnaeus, 1771), a Neotropical Prionid, is generally considered the largest, if not the heaviest, at up to 16.7 cm. The world’s longest species in absolute terms i.e. including the antennae is the New Guinea species Batocerus kibleri Schwarzer, 1925 the antennae of which can measure 20cm. Most cerambycids are between 8 and 50 mm and vary in form from cylindrical to flat, and from elongate to short-oval and broad. Many species worldwide are dull-brown or black but bright, and often very spectacular. Cryptic or mimetic colouration is common and many are metallic, even in temperate regions. Many species are crepuscular or
Callidium violaceum larva
nocturnal and cryptically coloured while, probably the great majority, are diurnal and exhibit a very wide range of colours and patterns; part of the appeal for collectors is the extensive intraspecific variation, many have accumulated long lists of aberrations, varieties and subspecies, and some exhibit mimicry of ants, bees and wasps, in colour, morphology and behaviour. Ostensibly the defining feature of the group are the long antennae which are generally at least half as long as the body but usually much longer, sometimes ridiculously so, and in general they are longer in the male; though seemingly unwieldy, even the longest antennae are delicate and versatile organs, and this can only be appreciated by observing such species as the Timberman, Acanthocinus aedilis (Linnaeus, 1758) in the wild.
Cerambycids are mostly xylophagous and phytophagous and are associated with a very wide range of plants; in temperate regions mostly trees but more generally grasses, including bamboo, herbaceous plants, cacti and succulents. The larvae develop within all parts of the host; leaves, twigs and branches, seeds, trunks and roots, and they may continue to develop after timber has been processed, there are records of adult longhorns emerging from furniture and structural timber after remaining dormant for two decades. Some species are pests in forest plantations, orchards and among nursery stock, they generally attack damaged or weakened plants and then greatly accelerate the damage, and a large number of species develop in dead wood, fallen timber, decaying logs, fence-posts etc. and they are important decomposers in forest ecosystems. Adult longhorns are generally short-lived and have a brief season, in temperate regions from late spring to early summer, most fly well and may be observed around flowers in wooded areas, some are slow-moving and tend to remain around the host material while others can move very quickly and disperse long distances, they these can often be observed feeding upon bark, foliage, flower parts and pollen etc. Some are flightless and have the elytra fused, these are generally terrestrial, live on grasses or herbaceous plants and disperse only short distances, the populations are prone to fragmentation and so they tend to be speciose e.g. the Palaearctic genus Dorcadion Dalman, 1817. Most species take between one and three years to complete the life-cycle although in warmer climates this may be complete in two or three months. The life history of many species is typical; the females oviposit in damaged wood, bark-crevices or in the soil around the roots, in fallen timber or in herbaceous stems etc. and the larvae emerge after a week or two and immediately begin to feed upon the host material, either among the softer cambium or by boring into stems, some bore directly into roots or into the soft xylem of decaying logs etc., as they grow they bore deeper into the wood producing often characteristic tunnels, or up and down herbaceous stems, but in general they remain near the surface. Pupation occurs within the host or in an earthen cell among the roots, and adults eclose either in the autumn and remain in place until the following season or in the spring and emerge directly. Among those emerging in the autumn are species of Cerambyx Linnaeus, 1758, Molorchus Fabricius, 1792, Pogonocherus Dejean, 1821, Acanthocinus Dejean, 1821, Mesosa Latreille, 1829 and Phytoecia cylindrica (Linnaeus, 1758).
The Western Palaearctic Fauna, with Special Reference to the UK
This will be no more than a brief look at the European fauna as much more detail will be given in the various pages. The European longhorn fauna includes about 700 species in about 6 subfamilies; the higher-order classification will be found to vary as some subfamilies will be listed as tribes e.g. the Necydylinae Latreille, 1825 is often listed as a tribe of the Lepturinae Latreille, 1802. Diversity is greatest in warmer regions and decreases from south to north along a temperature gradient. Turkey has about 600 species but is in some sense ‘well-placed’ between Asia, The Middle-East and Europe, while Italy has about 275, Croatia 220, Germany 195, France 250, Denmark about 75, Scandinavia more than a hundred and the U.K. about 70. These figures include the native or established species but all European countries have extensive lists of occasionally recorded species, both from surrounding countries and from those imported with timber etc. from farther afield; about 70 of these ‘occasional’ have been recorded in the U.K. or about the same number as the established species. These figures are in line with saproxylic groups generally where diversity is always greatest at southern and middle latitudes. Most northern European species, certainly those from the U.K. form parts of much wider European or Eurasian distributions and a few e.g. Rhagium inquisitor (Linnaeus, 1758), Callidium violaceum (Linnaeus, 1758) and Judolia sexmaculata (Linnaeus, 1758) are Holarctic.
As with many other saproxylic groups many xylophagous longhorns are in decline and have been for some decades, mostly due to the destruction of forested areas or their commercial management, and this continues today. Only two species are protected by law, Rosalia alpina (Linnaeus, 1758) and Cerambyx cerdo Linnaeus, 1758, neither of which occur in the U.K. Economic loss due to pest species is not so serious as in some other parts of the world but nonetheless a few species are noteworthy; Hylotrupes bajulus (Linnaeus, 1758), the House Longhorn, causes occasional and sometimes severe damage to property but this has declined over recent decades as new
Rosalia alpina (Linnaeus, 1758) - one of only two protected species of Cerambycidae in Europe
constructions materials are less prone to attack, species of Monochamus Dejean, 1821 and Tetropium Kirby, 1837 can cause serious damage in conifer plantations, and amenity trees and shrubs are attacked by a wide range of genera including Cerambyx Linnaeus,1758, Callidium Fabricius, 1775, Arhopalus Audinet-Serville, 1834 and Asemum Eschscholtz, 1830 but again such species are in general decline due to ‘improvements’ in environmental management e.g. the removal of fallen timber and damaged trees. A much greater threat is posed by invasive foreigners; in recent years two species in particular pose a serious threat, Anoplophora glabripennis (Motschulsky, 1853), the Asian Longhorn, and A. chinensis (Forster, 1771), the Citrus longhorn, both native to China and polyphagous on various hardwoods, have become established throughout Europe, at the moment (2017) only in scattered local populations but they are extremely invasive and damaging, and they have caused serious damage in various Nearctic regions. Another destructive invasive, Callidiellum villosulum (Fairmaire, 1900), the Brown Fur Longhorn which is another Chinese native, has recently been recorded established on Malta. Between 1990 and 2010 nineteen foreign species have become established in Europe and if this is a consequence of changing climate then perhaps we can look forward to many more.
Dorcadion circumcinctum Chevrolat, 1862
Dorcadion is a large genus in Europe, but is not found in the UK.
With the exception of a few iconic species such as Rosalia and Cerambyx the U.K. fauna, while only ten percent that of Europe, is a fair representation of the wider fauna. Many species are common along the northern coasts of Europe e.g. Spondylis buprestoides Linnaeus, 1758, an invasive and sometimes destructive species, and it is baffling why they have not become established here, and some that hold only a tenuous place on our U.K. list e.g. Dinoptera collaris (Linnaeus, 1758) are common and often abundant on the continent. On the other hand the widespread European Stictoleptura cordigera (Fussli, 1775) has recently become established in the south of England and so our fauna may be undergoing some modern changes. The destruction of woodland habitats across Europe since the twentieth century occurred much more thoroughly hundreds of years previous in the U.K. and ever since our countryside management has been appalling and so it is perhaps no wonder that our cerambycid fauna has become impoverished; many of those on our list are rare and local. One genus that occurs throughout central and southern Europe but is notably absent from the U.K. is Dorcadion Dalman, 1817 - a very speciose group of terrestrial and flightless longhorns, they are associated with herbaceous plants and some have been pests of cereal crops, they live in colonies in open situations which become easily separated and local populations develop in isolation leading to rapid differentiation and speciation, in Russia there are more than 150 species occurring in steppe habitats and so it might be imagined that parts of the U.K. would form ideal habitats. Some formerly common and widespread species have declined drastically over recent decades due to habitat loss and modification e.g. D. fulginator Linnaeus, 1758 is now generally endangered and extinct in some areas e.g. Luxembourg.
The majority of European longhorns are saproxylic; in Germany about 50% of species develop in softwood while about 26 develop in hardwoods, other species develop in a range of herbaceous plants although only two such species occur in the U.K., Phytoecia cylindrica (Linnaeus, 1758) and Agapanthia villosoviridescens (DeGeer, 1775, in each case members of much larger European genera. Most are active during May and June although some are later e.g. Prionus coriarius (Linnaeus, 1758), while others may occur earlier depending upon the season e.g. Clytus arietis (Linnaeus, 1758) and Acanthocinus aedilis (Linnaeus, 1758). A few occur year round and may be active during mild spells through the winter e.g. species of Pogonocherus Dejean, 1821. They may be collected by searching bark and logs etc. or flowers and sap in wooded areas, some have characteristic flight behaviour e.g. Pachytodes cerambyciformis (Schrank, 1781) hovers above flowers like a bee while Clytus darts quickly between flowers like a wasp. Keeping an eye along woodland margins or around solitary trees near wooded areas may reveal specimens in flight e.g. Leptura aurulenta Fabricius, 1792 or species of Tetropium Kirby, 1837. Some crepuscular species e.g. Prionus, Arhopalus Audinet-Serville, 1834 and Asemum Eschscholtz, 1830 may be seen in flight as the light fades, and many of these are attracted to light e.g. this is the easiest way to find Phymatodes testaceus (Linnaeus, 1758). Both phytophagous and saproxylic species are generally associated with a wide range of hosts, although our U.K. phytophages generally occur on umbels, and many genera occur on either hardwoods or softwoods but a few species within widely polyphagous genera may show a very restricted host range or even monophagy e.g. in the U.K. Semanotus russicus (Fabricius, 1776) is partial to hedging conifers, while Oberea oculata (Linnaeus, 1758) develop in species of Salix. On the other hand species of Monochamus Dejean, 1821, a Holarctic and African genus of more than 160 species, mostly attack conifers. Some may choose an unlikely range of hosts e.g. Pogonocherus hispidus (Linnaeus, 1758) usually develops in ivy but has also been recorded from Cornus (Dogwood), Corylus (Hazel), Ilex (Holly) and Euonymus (European spindle). In general the Cerambycinae and Lamiinae choose hardwoods while Spondylidinae choose conifers. Some groups choose both types of wood e.g. Rhagium Fabricius, 1775 and Oxymirus Mulsant, 1863. Timber in all stages of decay may host cerambycids, even healthy trees although they will seek out damaged areas of bark, generally low down on the trunk or among the roots, and fallen timber and logs in an advanced stage of decay will often be found to host them. It is the larvae that damage the wood as many species take two or three years to develop, tunnelling through the outer layers of xylem and going deeper as the surface is obliterated. Adults do feed but their effects are minimal; Lepturinae generally feed on flowers when they assemble to mate although some species e.g. Stenocorus meridianus (Linnaeus, 1758) and Stictoleptura rubra (Linnaeus, 1758) only occasionally visit flowers. Many Lamiinae as well as Cerambycinae visit flowers to feed and in particular may be found on umbels and blossom, among the more common of such
Longhorn Beetles - aptly named.
species are Clytus arietis, species of Tetrops Stephens, 1829 and Phytoecia Dejean, 1835. Most Lamiinae are bark and stem feeders; adults of Acanthocinus aedilis and Saperda populnea (Linnaeus, 1758) etc. may be observed chewing into bark, often prior to oviposition, while Cerambyx species will feed upon sap as well as fallen fruit. Foliage feeders include Lamia textor, and species of Leiopus Audinet-Serville, 1835 and Phytoecia Dejean, 1835. Some species of Leiopus as well as Stictoleptura rubra have been observed feeding upon fungi fruiting among bark crevices.
Although very variable longhorns are generally recognizable as the elongate and cylindrical or flattened form is distinctive, certain Tenebrionoidea groups include superficially similar species but here the form of the tarsi and the tarsal formula are different. The basic form of the Lamiinae, Cerambycidae and Lepturinae will soon become familiar. Most species are finely pubescent but this often matches the colour of the underlying cuticle and so it is often not obvious. The head is always visible from above; in some groups it is partially recessed into the prothorax so that the temples are not, or only narrowly, visible e.g. Prioninae and Lamiinae, while in others, typified by the Lepturinae, it is freely articulated beyond the anterior pronotal margin. Among the European fauna the Lamiinae are distinctive with the head vertically inclined (hypognathous), in the Lepturinae it lies about in line with the plane of the body (orthognathous) while in the Cerambycinae it is oblique, these orientations soon become familiar and most species can be placed in the field. The eyes are well-developed, especially in nocturnal species where they are larger and more coarsely faceted, they usually follow the contour of the head or are only weakly convex and they are most prominent from above in Lepturinae. The shape varies from oval to reniform or narrow and curved around the antennal insertions; in extreme cases e.g. Tetropium, they may be completely divided. The mandibles are robust and heavily sclerotized, generally broad and triangular in form, sharp internally and often bifurcate apically. The maxillary palpi are four-segmented with the terminal segment varying from securiform to cylindrical and truncate; the labial palpi are three-segmented and generally at least to some extent expanded to securiform. The vertex and frons are flat to weakly concave or convex and there is often a median longitudinal furrow or carinae between the eyes, the clypeus is well-sclerotized and separated from the frons by a distinct suture and the labrum is freely articulated and generally covered by the clypeus; it is often only visible by the fringe of long setae lying along the anterior margin. The antennae are in some sense the defining character of the group, they are typically long or very long and thin, often reaching to the elytral apex or beyond although in some groups they are short and rather broad e.g. Clytus, and in Spondylis they are more-or-less moniliform. In most groups they are 11-segmented although in the male of Prionus and in species of Agapanthia they have twelve. The basal segment is long, broad and often curved or expanded towards the apex, and sometimes elongate-pyriform, the second is short and quadrate to transverse, often globular and sometimes simply cylindrical, the remainder; 3-11 are usually elongate, sometimes extremely so, and cylindrical or expanded towards the apex but the morphology varies widely from strongly serrate in Prionus to very elongate in Acanthocinus, in most species they are pubescent from the second or third segment and in some tropical species they have elaborate arrangements of setae towards the apex of various segments, a condition approached in Rosalia. The pronotum varies from simply cylindrical or barrel-shaped to campanulate with produced posterior angles, in many it is flattened dorsally and there may be surface structure as e.g. in Hylotrupes. With the exception of the Prioninae the lateral margins are not bordered but there may be tubercles, spines or small teeth. The surface is usually punctured, often strongly so, and microsculptured. The prosternum is transverse and deep with a distinct and very variable process that extends beyond the coxae which are round to transverse-oval in shape; the prosternal epimera and episterna vary in shape but are generally proportionally small. The mesosternum is small, variable and produced posteriorly into a rounded or truncate process which is often incised to receive the produced anterior margin of the metasternum. In most groups the elytra cover the abdomen but there are many exceptions e.g. Necydalinae Latreille, 1825, Molorchus Fabricius, 1792 or Glaphyra Newman, 1840, they are either broadest at the shoulders and narrow towards the apex, or are parallel or nearly so, only rarely do the broaden posterior to the middle e.g. in Stenostola Dejean, 1835. The apex may be rounded, oblique or truncate and in some groups e.g. Pogonocherus Dejean, 1821 it has teeth or spines at the angles. They vary from cylindrical to flattened; the surface is randomly punctured and usually smooth i.e. without distinct striae although in Spondylis there are longitudinal ridges. In most species the hind wings are well-
A - Lepturinae
B - Cerambycinae
C - Lamiinae
developed and flight is strong. In most species the legs are long and slender although in many Spondylidinae and Lamiinae they are short and robust. The coxae project above the surrounding cuticle and vary from conical or globular to transverse-cylindrical with the meta-coxae reaching the elytral epipleura, and the cavities may be open or closed posteriorly. The femora are generally long and sub-parallel or distinctly thickened beyond the middle and in many cases strongly clavate. The tibiae are usually simple, although variously grooved in Lamiinae, and straight with two small spurs on the inner apical angle, in some Spondylidinae they are toothed along the external margin. The tarsi are 5 segmented although the fourth segment is often small or rudimentary or hidden within the bilobed third segment; this Chrysomelid arrangement of simple basal segments and a deeply bilobed third is a good guide to the group when compared to superficially similar Tenebrionoid families. In many species the basal segment, especially on the meta-tarsi is very long compared to the others. Sexual dimorphism is common among the family and often manifest in general habitus, antennal length, body proportions, and many females have a prominent and heavily sclerotized ovipositor.
Most Cerambycid larvae are pale and soft-bodied, never darkened or heavily-sclerotized, they are elongate and usually cylindrical or nearly so but flattened forms do occur, the mandibles are short and robust, generally triangular or curved, the legs short or much reduced, and abdominal ampullae are always present although sometimes very small, and they lack segmented urogomphi. Many larvae develop over several years and grow disproportionally large compared with their adult forms.
PRIONINAE Latreille, 1802
This large and almost cosmopolitan subfamily includes more than 1000 species in about 300 genera and 20 tribes, by far the greatest diversity is in tropical regions and very few extend to high temperate latitudes e.g. there are more than 350 species in 100 genera and 9 tribes in the New World but only 37 species of 13 genera and 5 tribes occur in the Nearctic. Only 3 species of 3 tribes occur in Northern Europe of which one, Prionus coriarius (Linnaeus, 1758), among the largest of European beetles, extends to the U.K. All species are saproxylic, developing in living or decaying wood, especially near the soil into which the larvae burrow to pupate, and many species have gregarious larvae. Most adults are nocturnal and may be found among wood and foliage; they generally fly strongly and are attracted to light. The group was formerly much larger as it included the Parandrinae Blanchard, 1845 or ‘False Stag Beetles’, a pan-tropical subfamily with more than 1000 species. Members of the Prioninae are among the largest insects in the world; they are very robust and mostly dark coloured, characterized by the distinct lateral border to the pronotum and the lack of a mesonotal stridulatory area. The head is usually large and robust with reniform eyes that may be very large and curved around the antennal insertions, distinct temples and generally a longitudinal impression on the frons, the mandibles are large and often produced forward, especially in some tropical species, the palps are to some extent truncate and the antennae 11- or 12-segmented and robust or massively developed with long spines or extensions to individual segments, sometimes giving a feathered effect. Many species are dimorphic; the male having exaggerated mandibles, no doubt for fighting other males, and more developed antennae. The pronotum is often short and transverse, sometimes appearing disproportionately so, with the lateral margins smooth to elaborately developed with teeth or spines. The elytra are entire and cover the abdomen, except in females where the ovipositor is visible, usually smooth i.e. without striae but in many there are longitudinal costae and in many they are narrowly explanate.
LEPTURINAE Latreille, 1802
Of all the subfamilies the Lepturinae is that best appreciated from a study of the U.K. fauna, it is a large and cosmopolitan group of about 150 genera in 9 tribes but is mostly Holarctic in distribution e.g. of the 360 species of 80 genera occurring in the New World about 200 species of 60 genera are Nearctic, proportionally much less ‘tropical’ than the other large subfamilies, and they do not display the exaggerated morphological development seen in e.g. Lamiinae or Cerambycinae. About 70 species occur in central Europe and 25 species of 18 genera in 2 tribes in the U.K. Beyond our U.K. fauna much of the superficial variation lies in colour and pattern, and there are some impressively coloured species, the basic body form being rather stable; long and slender with the head narrower than the thorax and the thorax narrower than the elytra which are broadest across the shoulders and tapering towards the apex. The eyes are prominent and generally oval, unlike many groups in which they are reniform and curved around the antennal insertions, and the temples are generally long and obvious. They resemble the Cerambycinae in having the terminal segment of the palps blunt or truncate at the apex but differ from them in having conical coxae. The appendages are long and slender and in many the antennae are dimorphic. Unlike most longhorn groups they often display sexual dichromatism. Most species are saproxylic with adults being very active on flowers in wooded areas.
SPONDYLIDINAE Audinet-Serville, 1832
This small subfamily includes about 100 species in 5 tribes; 4 of these were formerly included in the Aseminae Thomson, J., 1860 while the Spondylidinae Audinet-Serveille, 1832 included the single genus Spondylis Fabricius, 1775, a group of 4 species, one of which is widespread in Europe but does not extend to the U.K. This isolation was due to Spondylis being considered to be related to the Prioninae Latreille, 1802 as they are superficially similar on several points of morphology as well as general habitus. Spondylis was initially included incorporated in the Aseminae on features of the wing venation but more recent studies of the larval morphology strongly suggest that all 5 tribes are part of the same lineage, and as the Spondylidinae was the first group to be named it has priority over the others and the subfamily becomes the Spondylidinae. All species are elongate and rather parallel-sided with rounded pronotal margins and elytral apices, with the exception of Spondylis they are flattened dorsally and all have an obliquely inclined head and short appendages. Males and females are closely similar. They are distinct from the Cerambycinae Latreille, 1802 in that the stridulatory area on the mesonotum (beneath the elytra) is either divided into two parts by a smooth central area or it is missing altogether; in the Cerambycines it is entire. The larvae are similar to those of Lepturinae Latreille, 1802, but differ widely from those of Cerambycinae, with a rounded head and large labrum, and they possess a pair of small and closely approximated spines on the terminal abdominal segment. They are mostly insects of boreal coniferous forests at higher temperate latitudes although a few occur in tropical and sub-tropical conifer forests of the New World and Africa, including Madagascar. With the exception of a few members of the Saphanini Gistel, 1856 and Anisarthrini Mamaev & Danilevsky, 1973, the larvae develop in conifer wood. Only members of the Holarctic genus Tetropium Kirby, 1837, which tend to have much finer faceted eyes, are diurnal, the others being crepuscular or nocturnal, often attracted to light and spending the day under bark or among logs and fallen timber etc. The subfamily is represented in the U.K. by 6 species in 3 genera of the Asemini.
CERAMBYCINAE Latreille, 1802
This large subfamily includes about 4500 described species in more than 800 genera and 70 tribes; it is cosmopolitan with by far the greatest diversity in tropical and subtropical regions, more especially the Neotropics. About 80 species occur in central Europe of which 18 extend to the U.K. By comparison about 440 species in 160 genera and 37 tribes occur in the Nearctic. Diversity of form is very extensive but the Holarctic temperate fauna and much else is represented by our U.K. species, beyond this some pages are given to illustrate the wider diversity of the group. They are characterized by the truncate terminal segment of the maxillary palpi and the oblique orientation of the head; most are rather parallel-sided or only weakly narrowing posteriorly and many are vividly coloured. The head is often partially retracted into the prothorax but the long, parallel-sided or tapering temples are usually visible, the eyes are large and reniform and the mandibles robust, sharp and lacking a molar tooth. The antennae are inserted between the eyes and are usually long, often extending well beyond the elytral apices and often sexually dimorphic, being longer in the male, the morphology varies widely but in some groups e.g. Cerambyx Linnaeus, 1758 they may be very robust, and in some e.g. Elaphidiini Thomson, 1864, a large, mostly Neotropical tribe of 92 genera, the segments have prominent spines. The pronotum is very variable; smoothly rounded laterally or toothed etc. but not margined, and the pro-coxal cavities may be open or closed. The mesonotum has a large undivided stridulatory area. The elytra are usually entire, covering the abdomen, generally parallel-sided or nearly so and rounded or truncate apically, in some groups e.g. Gracilia Audinet-Serville, 1834 they are dilated towards the apex and in some e.g. Molorchini Gistel, 1848 and Psebiini Lacordaire, 1868 they are abbreviated, exposing parts of the abdomen. The legs are long and slender, the femora straight and parallel to thickened or clavate, the tibiae only weakly expanded and with two small apical spurs. The tarsi are 5-segmented and pseudotetramerous.
LAMIINAE Latreille, 1825
This is the largest and most diverse of all the longhorn subfamilies with about 20000 described species in 3000 genera worldwide, the group is cosmopolitan with by far the widest diversity in tropical and subtropical regions; about 80 species occur in central Europe, 40 in Scandinavia and 15 in the U.K. The Nearctic fauna includes about 250 species in 50 genera and 20 tribes, while the total New World fauna includes about 5000 species in more than 800 genera and 40 tribes. The group is extremely diverse and so any description will be lengthy and incomplete, we will try to give some idea of this diversity when describing individual tribes etc. but even among our small U.K. fauna some idea can be gained by comparing e.g. Agapanthia villosoviridescens (DeGeer, 1775), Saperda Fabricius, 1775 and Tetrops Stephens, 1829. The species are characterized by the large and flat head which is partially retracted into the prothorax; the vertex and frons are perpendicular or form an acute angle, this head structure makes most species instantly obvious, even in the field, the antennae are 11- or 12-segmented and mounted high up on the frons, between the eyes, the basal segment is large, the second small and the remainder generally elongate; in this subfamily we see species with extremely long antennae e.g. the British Acanthocinus aedilis (Linnaeus, 1758), but in many groups they are much shorter e.g. in Tetrops. The eyes are deeply emarginate and generally curve around the antennal insertions, in some groups e.g. Tetropiini they may be completely divided. The terminal segment of the maxillary palpi is slender and pointed, another good guide to the group, the labrum is freely articulated and the mandibles acute or bifid and lack a molar tooth. The pronotum is very variable, often with lateral tubercles, teeth or spines but never margined, the mesonotum has a single, undivided stridulatory area and the metasternum lacks secretory glands. The anterior coxae are globose or sub-conical, the cavities usually angled externally and closed. The legs are short and generally robust with thickened or clavate femora and slender tibiae with the usual 2 apical spines, the pro-tibiae have an oblique inner groove and the meta-tibiae are variously grooved. The tarsi are typical of the family, 5-segmented and pseudotetramerous; the third segment bilobed and the fourth reduced.
NECYDALINAE Latreille, 1825
Originally included as a tribe of the Lepturinae Latreille, 1802, this is a small group of about 70 species in 12 genera, they are most diverse in Neotropical regions although 2 genera extend into the United States and some members of the genus Necydalis Linnaeus, 1758 occur in the Old World, of which 2 extend into Europe. The group is very distinctive with a long, narrow and cylindrical body, long and slender appendages and short elytra which do not extend beyond posteriorly beyond the metanotum. The eyes are large and reniform and the antennae extend back along the abdomen which is much narrower than the elytral apices. The long hind wings are folded over the abdomen and so are obvious. In life they mimic bees and wasps both in appearance and behaviour, and may also be mistaken for rove beetles as they move quickly among foliage. The European species, N. ulmi Chevrolat, 1838 and N. major Linnaeus, 1758 are large insects at 18-35mm, and very similar in appearance, both are widespread but generally rare, and both are widely polyphagous on various hardwood trees. Adults occur in July and August and frequent flowers.