Elateriformia (Coleoptera)

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J. F. Lawrence, A. M. Hastings, M. J. Dallwitz, T. A. Paine and E. J. Zurcher

Character List

Nomenclature and classification #1. <Synonymy>/

#2. <Classification (suborder, series, superfamily)>/

1. Archostemata/

2. Myxophaga/

3. Adephaga/

4. Polyphaga: Staphyliniformia/

5. Polyphaga: Scarabaeoidea/

6. Polyphaga: Elateriformia/

7. Polyphaga: Bostrichiformia/

8. Polyphaga: Lymexyloidea/

9. Polyphaga: Cleroidea/

10. Polyphaga: Cucujoidea/

11. Polyphaga: Tenebrionoidea/

12. Polyphaga: Chrysomeloidea/

13. Polyphaga: Curculionoidea/

14. Placement Uncertain/

The superfamilial classification used here follows Lawrence and Newton (1995 in General Bibliography), but the superfamilies of Staphyliniformia, Elateriformia and Bostrichiformia have been combined into their respective series.

Distribution #3. <Geographical distribution>/

#4. Biogeographic regions:/

1. Nearctic/

2. Palearctic/

3. Neotropical/

4. Afrotropical/

5. Oriental/

6. Australian/

The regions used here are based on the traditional biogeographic regions of Wallace (1876 and 1910) and Sclater and Sclater (1899), as discussed (or dismissed) by Mayr (1944), Whitmore (1981), Cracraft (1982), Nelson and Platnick (1984), Humphreys and Parenti (1986), Briggs (1987) and others. Although these regions are still useful for making generalizations about distribution patterns, they reflect ideas on biogeography prevalent during the late 19th and early 20th Centuries and not those of modern biogeography, based on plate tectonics. It is now well known that many of the insects found in southern temperate South America are more closely related to those in New Zealand and Australia than they are to other inhabitants of the Neotropical region. A more modern and more detailed system of biotic regions, based on plant distributions, is that of Takhtajan (1986). SEE General Bibliography.

The divisions between adjacent biogeographic regions were often been disputed (when such things were taken more seriously), so some latitude must be given when coding taxa from montane Mexico, Asia Minor or central China. The Australian region, as here delimited, includes the Lesser Sunda Islands (except Bali), the Moluccas, New Guinea, Australia, New Zealand, and the Pacific Islands, but excludes Sulawezi.

#5. <Occurrence in Australia>/

1. not occurring in Australia/

2. occurring in Australia/

A taxon is coded as state 2 if at least one species occurs as an established population in Australia, including Lord Howe Island and Tasmania, but not Norfolk Island, Christmas Island or the Cocos-Keeling Islands.

#6. <Occurrence in America north of Mexico>/

1. not occurring in America north of Mexico/

2. occurring in America north of Mexico/

A taxon is coded as state 2 if at least one species occurs as an established population on the North American continent north of the Mexican border (including Alaska, Canada, and the United States excluding Hawai'i).

#7. <Occurrence in Canada, Alaska or Greenland>/

1. not occurring in Canada, Alaska or Greenland/

2. occurring in Canada, Alaska or Greenland/

A taxon is coded as state 2 if at least one species occurs as an established population in Canada, Alaska, or Greenland.

#8. <Occurrence in New Zealand>/

1. not occurring in New Zealand/

2. occurring in New Zealand/

A taxon is coded as state 2 if at least one species occurs as an established population in New Zealand, including Campbell Island, the Auckland Islands and the Chatham Islands.

#9. <Occurrence in Europe>/

1. not occurring in Europe/

2. occurring in Europe/

A taxon is coded as state 2 if at least one species occurs as an established population in Europe, west of the Ural Mountains, Ural River and Caspian Sea.

#10. <Occurrence in Japan>/

1. not occurring in Japan/

2. occurring in Japan/

A taxon is coded as state 2 if at least one species occurs as an established population in Japan, including the Ryukyu Islands.

#11. <Occurrence in the Patagonian region>/

1. not occurring in the Patagonian region/

2. occurring in the Patagonian region/

A taxon is coded as state 2 if at least one species occurs as an established population in the Patagonian region of South America, south of a line joining the 36th Parallel on the coast of Chile and the 42nd Parallel on the coast of Argentina (SEE Morrone & Roig 1995 in General Bibliography). It is likely that some taxa recorded as state 2 (based on records from northern Argentina and-or central Chile) may not actually occur as far south as indicated above. It is also possible that a few more Neotropical groups extend into this region.

#12. <Occurrence in the Hawaiian Islands>/

1. not occurring in the Hawaiian Islands/

2. occurring in Hawaiian Islands/

A taxon is coded as state 2 if at least one species occurs as an established population in the Hawaiian Islands.

#13. <Occurrence in Costa Rica>/

1. not occurring in Costa Rica/

2. occurring in Costa Rica/

A taxon is coded as state 2 if at least one species occurs as an established population in Costa Rica.

Biology #14. <Biology>/

Notes #15. <Notes>/

#16. Data at least partly checked by/

References #17. <References>/

General appearance #18. Total length/

mm/

The total length is measured from above and extends from the anterior edge of the labrum (clypeus or frontoclypeus, if the labrum is absent or not visible from above) to either the abdominal apex (if this is visible from above), the conjoined elytral apices, or an imaginary transverse line joining the separated elytral apices. The mandibular length is not included.

#19. Ratio of body length to greatest body width/

The body length is measured from above and extends from the anterior edge of the pronotum (at mid point) to either the abdominal apex, if this is visible, or the conjoined elytral apices or an imaginary transverse line joining the separated elytral apices. The head length is not included. The body width is almost always synonymous with the elytral width; but it may correspond to the pronotal width in those taxa with a greatly enlarged prothorax (e.g. some Elateridae) or to the greatest width across the thorax or abdomen in adult beetles which lack elytra (e.g. female Phengodidae).

#20. Body <degree of convexity>/

1. strongly flattened/

2. slightly flattened to moderately convex/

3. strongly convex/

#21. Sides of body <whether evenly curved>/

1. not evenly curved/

2. evenly curved/

A taxon is coded as state 1 when the lateral outline is evenly curved and uninterrupted, as in regularly elliptical, ovoid or spherical beetles.

#22. Body <whether capable of conglobation>/

1. not capable of conglobation (rolling into a ball)/

2. capable of conglobation (rolling into a ball)/

Conglobation is a type of compaction mechanism in which the body is so constructed that the head and prothorax are capable of fitting against the underside of the shortened hind body to form a sphere. This is often referred to as "rolling into a ball". In many Anobiidae, the head and prothorax may be modified in a similar way, and fit against the hind body; however the abdomen is usually more or less exposed and these taxa are not considered to have conglobate (state 2) body.

#23. Upper surfaces of body <vestiture>/

1. glabrous or subglabrous/

2. clothed with distinct hairs, setae or scales/

A taxon is coded as state 1 when the upper surfaces have a small number of distinct setae (such as the tactile setae in Carabidae) or when there are large numbers of minute setae which are not or barely visible under lower magnifications (e.g. 10 X). A glabrous surface, as defined here, does not have to be smooth; taxa with tuberculate costate, or otherwise modified surfaces are still coded as state 1, if they lack obvious hairs, bristles or scales.

#24. Vestiture of upper surfaces <whether including stiff, erect bristles>/

1. not including stiff, erect, dark bristles/

2. including stiff, erect, dark bristles/

A taxon is coded as state 1 when the upper surfaces have erect, hyaline setae or light colored bristles. State 2 refers only to the stout, dark bristles characteristic of many Cleroidea.

#25. Vestiture of upper surfaces <whether including scales or scale-like setae>/

1. not including scales or scale-like setae/

2. including scales or scale-like setae/

A taxon is coded as state 1 when the upper surfaces have moderately stout, acute hairs, which are opaque and may form a pattern. State 2 refers to the presence of true, flattened scales or very stout, truncate setae, which are usually expanded apically.

#26. Upper surfaces of body <whether with deep foveae>/

1. without deep foveae/

2. with one to several deep foveae/

State 2 refers to relatively large, deep foveae, such as those found in pselaphine Staphylinidae. They may be located on the head, pronotum, elytra or exposed abdominal tergites.

#27. Prothorax, metathorax and-or abdomen <whether with extrusible glands>/

1. without extrusible glands/

2. with extrusible glands/

The prothoracic glands are anterolateral in location and often bifurcate or complex. Those at the base of the abdomen may also be divided. The others are usually relatively small, single eversible sacs (Evers 1968 in General Bibliography). Excluded are those glands associated only with the abdominal apex (segments 8 abd 9).

#28. Underside of body <whether with hydrofuge surface>/

1. without hydrofuge surface(s)/

2. with hydrofuge surface(s)/

An area of hydrofuge pubescence, often referred to as a plastron, is usually characterized by having a dull, silver, gray, yellow or brownish sheen. Under high magnifications (scanning electron microscope) these areas are often seen to be composed of dense, overlapping, flattened and often complex hairs.

Head #29. Ratio of head length to its greatest width (excluding eyes)/

1. greater than 1/

2. 1 or less/

Head length is measured from above and does not include the free labrum; head width is measured just behind eyes and does not include the eyes.

#30. Head width just behind eyes <whether distinctly greater than prothoracic width>/

1. not distinctly greater than prothoracic width/

2. distinctly greater than prothoracic width/

Head width is measured just behind eyes and does not include the eyes.

#31. Head <whether strongly declined>/

1. not or slightly declined <less than 45 degrees>/

2. moderately to strongly declined <45 or more>/

A taxon is coded as state 1 when the base of the head is horizontal or slightly inclined (angle of inclination less than 45 degrees). A head is described as having state 2 when its long axis forms an angle greater than 45 degrees with the horizontal plane of the prothorax. This character may be difficult to use, since the normal angle of the head at rest may be altered by the method of fixation or preparation. Carded specimens often have the head pulled out and glued horizontally, in order for the mouthparts to be visible. In a number of groups, the head base may be not be declined, but the frontoclypeal region is strongly and often abruptly declined, so that the mouth cavity becomes ventrally oriented. SEE "longitudinal axis of head (whether forming an angle)", "frontal region (whether vertical or inflexed)", and "mouth cavity (whether ventrally or posteroventrally oriented)".

#32. Head <whether concealed from above by pronotum>/

1. not entirely concealed from above by pronotum/

2. entirely concealed from above by pronotum/

A taxon is coded as state 2 when the head is completely concealed from above by the anterior edge of the prothorax. The head may or may not be stongly declined. Carded specimens sometimes have the head artificially pulled out, although normally it would not be visible from above in the resting position. If the base of the head is just visible beyond the edge of the pronotum, the specimen is coded as state 1.

#33. Head <whether with elongate rostrum>/

1. without elongate rostrum/

2. with elongate rostrum <longer than wide>/

The rostrum is an elongation of the frontoclypeal region, so that the mouthparts come to lie some distance in front of the eyes. Specimens are coded as state 2 only when the area between the anterior edges of the eyes and the mandibular bases (sometimes called the muzzle) is longer than wide. The clypeus or frontoclypeal region may be prolonged anteriorly over and in front of the mouthparts; the resulting projection is not considered to be a rostrum, since the mouthparts are not located at its apex.

#34. Head <whether abruptly constricted to form neck>/

1. not abruptly constricted posteriorly/

2. abruptly constricted posteriorly to form neck/

A taxon is coded as state 1 when the head is not or only gradually narrowed behind the eyes, or abruptly narrowed either immediately behind the eyes or immediately adjacent to the occpipital foramen at the posterior edge of the head. A taxon is coded as state 2 when the head behind the eyes is distinctly, more or less abruptly narrowed, and often slightly expanded again, forming a neck which is narrower than the area in front of the eyes, and a pair of temples or tempora, which are strips of cuticle lying in between the eyes and the neck constriction. Sometimes the neck region is short and may be concealed when the head is retracted into the prothorax or abuts the anterior edge of the pronotum. SEE "temples (whether closely adpressed to prothorax". If the head is gradually narrowed to a point just in front of the occipital foramen and then widened again to form a condyle-like neck; this condition is still coded as state 1.

#35. Temples <whether closely adpressed to prothorax>/

1. absent or not closely adpressed to prothorax/

2. closely adpressed to and abutting prothorax/

A taxon is coded as state 2 when the narrow neck is concealed from above, because the temples are closely adpressed to and abutting the anterior edge of the pronotum. In most cases the head also has a transverse occipital ridge which abuts the rim of the pronotum. If the temples are not visible because the head is inserted into the anterior opening of the pronotum, then the taxon is coded as state 1.

#36. Temples <presence and relative length>/

1. absent/

2. shorter than length of eye/

3. longer than length of eye/

The head is not considered to have temples, unless there is an abrupt constriction followed by a narrower neck. Eye length here is measured from above and does not refer to the longest eye diameter.

#37. Transverse occipital ridge or carina <whether present>/

1. absent/

2. present/

A taxon is coded as state 2 when there is a distinct transverse ridge or carina on the head. Those heads with a fine transverse line or groove are coded as state 1.

#38. Occiput <whether with stridulatory file or files>/

1. without stridulatory file/

2. with 1 median stridulatory file/

3. with 2 paramedian stridulatory files/

Under lower magnifications, a stridulatory file looks like a shiny area, which is usually narrow and elongate. Under higher magnification, a series of fine transverse ridges may be seen.

#39. Longitudinal axis of head (from occipital foramen to mouth cavity) <whether forming an angle>/

1. inclined at an angle of less than 45 degrees/

2. inclined at an angle of 45 to 90 degrees/

3. inclined at an angle of more than 90 degrees/

The longitudinal axis of the head is considered to be a line passing through the center of the occipital foramen and the center of the mouth cavity. If this is more or less straight, or inclined at an angle of less than 45 degrees, the character is coded as state 1. States 2 and 3 refer to heads in which the angle is between 45 and 90 degrees or more than 90 degrees, repsectively. Note that this character concerns the angle formed between the front and back of the head and has nothing to do with the angle which the base of the head forms with the prothorax. SEE "head (whether strongly declined)".

#40. Frontal region <whether vertical or inflexed>/

1. not to moderately, gradually declined/

2. strongly deflexed, vertical or inflexed at apex only/

3. strongly deflexed, vertical or inflexed from behind eyes/

The frontal region, for the purposes of this key, extends from just behind and between the eyes to the clypeus or clypeolabrum, from which it may be separated by a frontoclypeal suture. SEE "frontoclypeal suture (whether distinct)" and "frontoclypeal suture (whether strongly curved or angulate)". The posterior or upper part of this region is often called the vertex. In taxa having state 3, the frontal region is more or less vertical or sometimes posteriorly inflexed, due to a strong bend in the longitudinal axis of the head. SEE "longitudinal axis of head (whether forming an angle)". In taxa coded as state 2, only the apical part of this region is declined. State 3 is usually correlated with a ventrally or posterioventrally oriented mouth cavity, while state 2 is less likely to be. SEE "mouth cavity (whether ventrally or posteroventrally oriented)".

#41. Occipital region <whether with median groove or line>/

1. without median longitudinal groove or line (endocarina)/

2. with median longitudinal groove or line (endocarina)/

State 2 is present when a median longitudinal groove or line extends anteriorly from the upper edge of the occipital foramen. It may be of various lengths and is often accompanied by an internal endocarina to which muscles are attached. SEE "frontal region (whether with median groove or line)" for a more anteriorly placed groove or endocarina.

#42. Frontal region <whether with median groove or line>/

1. without median groove or line (endocarina)/

2. with median groove or line (endocarina)/

The median frontal groove or endocarina is sometimes relatively weak and may be short or may be continuous with the occipital groove. If there is a continuous line or groove from the posterior edge of the head well into the frontal region, then both this character and the previous one ("occipital region (whether with median groove or line)") are coded as 2.

#43. Head <whether with ocelli>/

1. without ocelli/

2. with 1 median ocellus/

3. with 2 ocelli/

Most ocelli or simple eyes consist of distinct circular lenses, which contrast with the surrounding cuticle. In a few cases, however, they may be reduced to a lightly pigmented areas on the cuticle. In some Derodontidae the ocelli are associated with a system of ridges and grooves and are somewhat more difficult to distinguish.

#44. Compound eyes <whether absent>/

1. present/

2. absent/

Specimens are coded as state 1 if they have compound eyes consisting of at least several, but usually many facets. A very few larviform females (e.g. Phengodidae, some Lycidae and Rhagophthalmidae) lack compound eyes but have a pair of single-lens larval stemmata; these are coded as state 2.

#45. Eyes <whether strongly protuberant>/

1. not or only slightly protuberant/

2. strongly protuberant/

#46. Vertical diameter of eye <whether at least 2 times as long as horizontal diameter>/

1. less than 2 times horizontal diameter/

2. at least 2 times horizontal diameter/

The horizontal diameter of the eye is more or less in line with the longitudinal axis of the head and is measured across the widest part of the eye (which is near one end when the eye is emarginate).

#47. Eyes <whether coarsely facetted>/

1. finely facetted/

2. coarsely facetted/

This character is difficult to code, since the two states are not rigorously defined. In coarsely facetted eyes the individual facets are usually strongly curved and somewhat projecting, while in finely facetted ones they tend to be flatter and not easily separable. Many eyes are not easily placed in one category or another and have been coded as 1/2.

#48. Eyes <whether with interfacetal setae>/

1. without interfacetal setae/

2. with interfacetal setae/

Some eyes coded as state 1 may have a few setae around the edge, but the great majority of facets will be bare. In some cases there may be very fine interfacetal setae which are not normally visible; these have been coded as state 1.

#49. Ommatidium <type>/

1. of the eucone type/

2. of the acone type/

3. of the exocone type/

This character is very difficult to observe without special preparation. A thin section of the eye is always necessary, and observation of the eucone condition requires special fixation. The exocone condition may be distinguished from either acone or eucone even in cleared specimens by the presence of long exocone lenses projecting in from the cornea; however some acone eyes may have relatively thick lenses and appear to be exocone. The coding of this character is usually based on published information from selected taxa within the group and not from original observations. SEE Caveney 1986 in General Bibliography.

#50. Eye <whether divided into upper and lower parts>/

1. entire/

2. divided into upper and lower parts by fine transverse line or narrow strip of cuticle/

3. divided into upper and lower parts by moderately to very wide strip of cuticle/

Eyes are considered to be entire (state 1) even when deeply emarginate (SEE "anterior or mesal edge of eye (whether emarginate)"). An eye is coded as state 2 when there is only a fine transverse line or very narrow strip of cuticle completely separating the upper and lower halves; the two portions of a divided eye may also differ in the size of the facets. In state 3 eyes, the two halves are more distant, and may be separated by a distance greater than the width of either half.

#51. Anterior or mesal edge of eye <whether emarginate>/

1. not or only barely emarginate/

2. shallowly emarginate or slightly divided by canthus/

3. deeply emarginate or strongly divided by canthus/

A canthus is a cuticular intrusion into the eye. In some cases the canthus projects out from the eye surface and appears to be superimposed over the eye, but in others it lies on the same plane as the eye or even below it. These two conditions are not distinguished here.

#52. Posterior edge of eye <whether emarginate>/

1. not or barely emarginate/

2. distinctly emarginate/

The posterior edge of the eye is often more or less truncate, but in state 2, there is a distinct emargination.

#53. Antennal insertions <whether concealed from above>/

1. exposed from above/

2. concealed from above/

The antennal insertion is an opening in the head capsule which receives the basal condyle of the scape or antennomere 1; in many cases this opening is countersunk forming a basal depression or antennal fossa. Specimens coded as state 1 must have the actual antennal insertions (as opposed to the fossae) at least partly visible from above. In some taxa, there is a projecting frontal ridge or paired supra-antennal ridges which conceal the insertions from above; in others there are no distinct ridges but the insertions are anteriorly or laterally directed and thus not visible from above (state 2).

#54. Antennal insertions <position relative to anterior edges of eyes>/

1. located anterior to imaginary line joining anterior edges of eyes/

2. located posterior to imaginary line joining anterior edges of eyes/

#55. Antennal insertions <whether approximate>/

1. moderately to widely separated <more than 0.5 X HW>/

2. narrowly separated <0.2–0.5 X HW>/

3. closely approximate or contiguous <less than 0.2 X HW>/

The antennal insertion is an opening in the head capsule which receives the basal condyle of the scape or antennomere 1; in many cases this opening is countersunk forming a basal depression or antennal fossa. For this character, it is the distance between the insertions themselves and not the fossae which is measured. SEE "antennal insertions (whether located in fossae)". A taxon is coded as state 1 when this distance is at least 0.5 times as great as the head width behind the eyes, state 2 when they are separated by a distance between 0.2 and 0.5 times as great as the head width, and state 3 when they are contiguous or separated by less than 0.2 times the head width.

#56. Antennae <whether borne on raised tubercles>/

1. not borne on raised tubercles/

2. borne on raised tubercles/

In Cerambycidae and a few other groups, the antennal insertions are raised up onto prominences, usually allowing the antennae to project directly backwards.

#57. Antennal insertions <whether located in fossae>/

1. flush with head capsule or raised, not in fossae/

2. countersunk so that they lie within saucer-like fossae/

#58. Subantennal groove or cavity on head <presence>/

1. absent or very weakly developed/

2. well developed/

The antennal groove or cavity is a shiny, concave area lying just mesad or mesoventrad of the eye, often between it and the mandibular articulation or the genal ridge. In many cases, it is short and houses the scape only, but sometimes it extends as a narrow groove below and behind the eye and houses several antennomeres. In Curculionoidea, this feature is called a scrobe, but that term is also used in Carabidae for a concavity on the outer edge of the mandible.

#59. Frontoclypeal suture <whether distinct>/

1. absent or incomplete/

2. indistinctly impressed/

3. distinctly impressed/

The frontoclypeal or epistomal suture is a transverse impression usually lying in front of the antennal insertions and behind the mandibular articulations and separating the frons posteriorly from the clypeus anteriorly. f It is often absent or vaguely indicated but may be seen through a cleared cuticle as a transverse internal ridge. This character refers only to the external indication of the suture. The frontoclypeal suture is an invagination, and it should not be confused with the line of demarcation between a basal, more heavily sclerotized region of the clypeus (postclypeus) and a more or less hyaline anterior region (anteclypeus) to which the labrum is attached.

#60. Frontoclypeal suture <whether strongly curved or angulate>/

1. straight, slightly curved or angulate/

2. strongly curved or angulate/

Sometimes the frontoclypeal suture is angulate and is joined at the angle with a short longitudinal suture called the median stem or midcranial suture.

#61. Clypeus <whether laterally emarginate>/

1. not laterally emarginate/

2. laterally emarginate to receive mandibular tubercles/

State 2 refers to a condition in which each side of an anteriorly projecting clypeus are concave, forming a slight notch into which a dorsal mandibular tubercle fits. SEE "dorsal part of mandible (whether with tubercle)".

#62. Anterior edge of clypeus or clypeolabrum <whether emarginate>/

1. straight to convex/

2. concave to shallowly emarginate/

3. deeply emarginate or excavate/

4. complexly lobed or dentate/

When there is no apparent labrum, the anterior edge of the head is considered to be the clypeus or clypeolabrum.

#63. Mouth cavity <whether ventrally or posteroventrally oriented>/

1. anteriorly or anteroventrally oriented/

2. ventrally or posteroventrally oriented/

The ventral orientation of the mouth cavity (state 2) is relative to the long axis of the head and does not refer to the ventral declination of the entire head. SEE "head (whether strongly declined)" and "longitudinal axis of head (whether forming an angle)".

#64. Pregular area <whether with laterally opening cavities>/

1. without laterally opening cavities/

2. with pair of laterally opening cavities/

These laterally opening cavities lie immediately behind the ventral mouthparts; they are relatively deep and filled with setae.

#65. Head ventrally <whether with paired subgenal ridges>/

1. without paired subgenal ridges/

2. with paired subgenal ridges/

The subgenal ridges are on the underside of the head, extending posteriorly from just beneath the eye; they are usually sharply defined and when the head is at rest they may lie against the large procoxae or, in highly compacted beetles, against the metaventrite (= metasternum).

#66. Head <whether with projecting genal processes>/

1. without anteriorly-projecting genal processes/

2. with anteriorly-projecting genal processes <visible from above>/

A taxon is coded as state 2 only if the genal processes project anteriorly and are visible from above the head, as in Prostomidae.

#67. Gular sutures <whether confluent>/

1. widely separated or absent/

2. narrowly separated/

3. at least partly confluent/

Gular sutures are coded as state 1 if they are widely separated or if they cannot be seen at all. State 3 gular sutures 3 may be confluent anteriorly or posteriorly, or may be entirely confluent, so that there is only a single line visible.

#68. Corporotentorium <width or absence>/

1. broad/

2. narrow/

3. incomplete or absent/

The corporotentorium, as defined here, is synonymous with the tentorial bridge, and is located in the posterior part of the head capsule. It is usually narrow and sometimes strongly arched. Newton and Thayer (1995, SEE reference under STAPHYLINIDAE (major part)) use the term corporotentorium for a more anterior fusion of the laminatentoria. General surveys of beetle tentoria include that of Stickney (1923 in General Bibliography).

#69. Corporotentorium <whether with median process>/

1. without median process/

2. with median process/

A narrow, anteriorly projecting process occurs in various taxa of Cucujoidea.

#70. Cervical sclerites <whether present>/

1. absent/

2. present/

The cervical sclerites are very small in some taxa and may be lost in dissection. Usually one or both remain attached to the head, but may also be found attached to the anterior edge of the prothorax. In some taxa these sclerites are subdivided.

Antennae #71. Number of antennomeres/

Counting the number of antennomeres is usually relatively simple, The first is often enlarged or widened and is called the scape, while the 2nd (pedicel) may be reduced in size and rarely enclosed within the apex of the scape. When there is an antennal club (SEE "antenna (whether distinctly clubbed)"), the antennomeres immediately preceding the club may be strongly transverse and not easy to distinguish from one another.

#72. Antennae when posteriorly extended <posterior extension>/

1. not reaching middle of prothorax/

2. reaching beyond middle of prothorax but not middle of elytra/

3. reaching beyond middle of elytra but not elytral apices/

4. reaching beyond elytral apices/

Since the antennae are rarely straight and cannot be moved in dried specimens, the best way to code this character is to make a rough estimate of antennal length and then compare this with the distance from the antennal insertion to the appropriate position on the body.

#73. Antennae <type>/

1. filiform/

2. moniliform/

3. serrate/

4. pectinate or bipectinate/

5. flabellate or biflabellate/

6. plumose or biplumose/

7. with articulated appendages/

8. incrassate or clavate/

9. capitate/

10. perfoliate/

11. of a unique configuration, none of above/

Filiform antennae (state 1) are more or less thread-like, with the antennomeres varying only slightly in width. These grade into the incrassate or clavate type (state 8), in which the antennomeres gradually increase in thickness and usually decrease in length towards the apex. Strongly incrassate antennae are often called clubbed, but we consider that term to be synonymous with capitate (state 9), in which one to several apical antennomeres are clearly set off from the preceding ones by their increased length and-or width. Moniliform antennae (state 2) are characterized by having bead-like or barrel-like segments. A gradation exists between state 1 or filiform antennae and the serrate type (state 3), in which the apical portion of each antennomere asymmetrically produced and somewhat acute, producing a saw-like effect. This, in turn, may grade into the pectinate and flabellate types (states 4 and 5). In the pectinate or comb-like type, most of the antennomeres are asymmetrically produced into a short, narrow, cylindrical or somewhat flattened process. In the flabellate antenna, each process is much longer and flattened. In the bipectinate or biflabellate types, each antennomere gives rise to a pair of processes. The plumose antenna (state 6) also has one or two processes on each antennomere, but these processes are very long and slender and often bear accessory processes or hairs. State 7 resembles the pectinate or flabellate type, but each process is articulated at the base. In all antennae representing states 4 to 7, characteristic enlargements or projections begin on the 3rd or or more apical antennomere and may not involve the apical one. Sometimes an incrassate antenna (state 8) may become wider apically but narrower again just before the apex. The perfoliate antenna is one in which most segments are flattened and expanded on either side of the middle.

#74. Antennomeres 3, 4 or 5 to 10 <whether biramose>/

1. without or with single rami (uniramose)/

2. with double rami (biramose)/

Paired rami are usually more or less equal, but in some groups of Lampyridae one is much shorter than the other (so that the antenna may appear uniramose).

#75. Antennae <whether partly pubescent>/

1. entirely subglabrous and without obvious modifications/

2. at least partly pubescent or with obvious modifications/

Although all antennomeres will have some setae on them, most antennae are more densely setose apically or have other types of sensory modifications on all or some of those antennomeres lying beyond the scape. These modifications may consist of serrations, rami or enlargements and include the apical club. Those taxa coded as having state 1 have simple, unmodified, more or less filiform antennae with no or few setae. State 1 is relatively rare, occurring mainly in aquatic beetles.

#76. Antennal modifications beginning on/

1. antennomere 3 (rarely 2)/

2. antennomere 4/

3. antennomere 5/

4. antennomere 5 or beyond/

Antennal modifications include serrations, various types of rami in pectinate, flabellate or plumose antennae, and club segments in capitate antennae.

#77. First antennomere (scape) <whether more than 3 times as long as 2nd>/

1. less than 3 times as long as 2nd (pedicel)/

2. more than 3 times as long as 2nd (pedicel)/

#78. Antenna <whether geniculate>/

1. not geniculate/

2. geniculate or elbowed/

In geniculate antennae, the scape (1st antennomere) has an irregular or asymmetrical articulation with the pedicel (antennomere 2), so that the latter can move in a single plane only. As a result, the antenna often looks "elbowed" with a distinct angle between the scape and remaining antennomeres. In most antennae the articulation between 1 and 2 is a ball and socket joint, and the antennae rarely appears "elbowed".

#79. Antenna <whether distinctly clubbed>/

1. without a distinct club/

2. with a distinct club/

A state 1 antenna may be gradually clavate (= incrassate).

#80. Antenna <whether with weak or strong apical club>/

1. without apical club/

2. with weak apical club/

3. with strong apical club/

#81. Antennal club <number of segments>/

1. 1-segmented/

2. 2-segmented/

3. 3-segmented/

4. 4-segmented/

5. 5-segmented/

6. with 6 or more segments/

Divisions between club segments must be complete if both of the segments are to be counted.

#82. Antennal club <whether 5-segmented with 2nd segment smaller than 1st or 3rd>/

1. not 5-segmented or with 2nd segment subequal to or larger than 1st/

2. 5-segmented with 2nd segment smaller than 1st or 3rd/

The state 2 type of antennal club usually occurs only in the family Leiodidae, but may also occur in a few other families, such as Corylophidae and Hobartiidae.

#83. Antennal club <whether compact>/

1. loose/

2. compact/

In some very compact antennal clubs, the divisions between the club segments are vaguely defined, and a 3-segmented club may appear to have a single segment; in this case the character "antennal club (number of segments)" may be coded as 1 or 3.

#84. Antennal club <whether lamellate>/

1. not lamellate/

2. lamellate/

A lamellate antennal club (state 2) is comprised of broad, flattened asymmetrical and more or less adpressed segments. The sensory structures are usually on the surfaces which are enclosed when the segments are pressed together.

#85. Antennal club <whether serrate-pectinate or flabellate>/

1. not or slightly serrate/

2. strongly serrate or pectinate/

3. flabellate/

State 2 is similar to the character "antennae (type)", state 4, except that the pectinate processes are restricted to the last few antennomeres. Similarly, state 3 is similar to the character "antennae (type)", state 5.

#86. Antennal club <whether cupuliform>/

1. not cupuliform/

2. cupuliform/

A cupuliform club (state 2) is one in which the apical club segments are partly enclosed within the cup-like basal club segment. This almost always refers to a lamellate club.

#87. Antennal club <whether strongly flattened>/

1. not or slightly flattened/

2. distinctly flattened/

#88. Antennal club <whether preceded by a cupule>/

1. not preceded by a cupule/

2. preceded by a cupule/

A cupule is a strongly transverse, glabrous, cup-like antennomere lying at the base of the antennal club and involved in respiration in most hydrophilid and hydraenid beetles. In some antennae, more than one antennomere may be cupule-like.

Mouthparts #89. Labrum <whether concealed by clypeus>/

1. at least partly visible/

2. concealed beneath clypeus or apparently absent/

A labrum is considered to be visible (state 1) if it can be seen from a dorsal or anterodorsal perspective. A state 2 labrum may be barely visible in anterior view when concealed by the clypeus. Sometimes a state 1 labrum may be represented by a narrow strip of cuticle only.

#90. Labrum <whether fused to clypeus or frontoclypeus>/

1. free, membranous or separated by suture/

2. partly or completely fused to clypeus or frontoclypeus, without or with incomplete suture/

A taxon is coded as state 1 when the labrum is separated from the clypeus or frontoclypeus by a complete suture or strip of membrane, and also when the entire labrum is membranous. It may be freely movable, but this is not always the case. A state 2 labrum is is always immobile and is either completely fused with the clypeus or frontoclypeus (apparently absent) or separated by a vague impression or incomplete suture.

#91. Major portion of labrum <shape>/

1. strongly transverse <less than 0.6 X as L as W>/

2. slightly transverse or subquadrate <0.6–1 X as L as W>/

3. distinctly longer than wide/

This usually refers to the visible or most heavily sclerotized portion of the labrum; however in some taxa in which the labrum is lightly sclerotized and concealed within the head capsule, it refers to the major portion, excluding basal membrane and tormae. A strongly transverse labrum (state 1) is less than 0.6 times as long as wide. A slightly transverse labrum (state 2) is greater than 0.6 times as long as wide.

#92. Apex of labrum <shape>/

1. subtruncate to slightly convex/

2. strongly convex, narrowly rounded or acute/

3. slightly concave or emarginate/

4. deeply emarginate or bilobed/

5. trilobed or with several lobes or teeth/

This character is not coded when the labrum is solidly fused to clypeus (without clypeolabral suture). SEE "labrum (whether fused to clypeus or frontoclypeus)".

#93. Labrum <whether mostly membranous>/

1. moderately to heavily sclerotized, except at base and-or apex/

2. mostly membranous or very lightly sclerotized/

Taxa are coded as having state 2 when the labrum is much more lightly sclerotized than the clypeus or frontoclypeus.

#94. Mouthparts <whether forming a piercing or sucking tube>/

1. not forming a piercing or sucking tube/

2. forming a piercing or sucking tube/

Tubular mouthparts are more or less similar wherever they occur, consisting of an elongate, acute labrum and stylet-like mandibles and maxillary lobes. Mouthparts which form a sucking or lapping structure composed of long and setose maxillary lobesor maxillary palps (Handschin 1929, Kaszab 1963 in General Bibliography) are coded as state 1.

#95. Mandibles <whether absent>/

1. present/

2. absent/

Mandibles are rarely absent in beetles. In some taxa they are very small, and they may appear to be absent when more or less enclosed within the mouth cavity. SEE "mandible (whether enclosed in mouth cavity)".

#96. Mandible <shape>/

1. short and broad <less than 2 X as L as W at base>/

2. moderately elongate <2–3 X as L as W at base>/

3. very narrow and elongate <more than 3 X as L as W at base>/

A taxon is coded as state 1, when the length of the mandible is less than 2 times as great as the basal width. State 2 mandibles are between 2 and 3 times as long as wide at base, and those coded as state 3 are more than 3 times as long as wide at base. The mandible length in slender, strongly curved mandibles is calculated by adding the lengths of 2 or 3 sections and not by measuring the shortest distance between the base and the apex.

#97. Mandibular apex <whether curved mesally>/

1. not or slightly and gradually curved mesally/

2. moderately to strongly, gradually curved mesally/

3. strongly and abruptly curved mesally/

Most mandibles are at least moderately curved towards the midline, so that the apices face one another or overlap (state 2). In a relatively small number of groups mandibles may not be mesally curved or may actually curve outwardly (state 1). In a state 3 mandible, there is a definite external angle where the outer edge of the mandible abruptly changes direction.

#98. Mandibular apex <number of teeth or lobes>/

1. unidentate, truncate or rounded/

2. bidentate or bilobed/

3. multidentate or multilobed/

State 1 mandibles also include those which do not have a distinct pointed apex but rather are truncate or rounded apically. SEE "mandibular apex (whether rounded)". When there is one apical tooth and a second smaller tooth located before the apex, there may be confusion as to whether to code this state 1 or 2. If the two teeth are relatively close to one another, they should be state 2. If the second tooth is well behind the apical one, then code this character as state 1. SEE "incisor edge of mandible (whether with one or more teeth)".

#99. Mandibular apex <whether rounded>/

1. subacute, bidentate or multidentate/

2. rounded or truncate/

State 2 is characteristic of those mandibles which are highly reduced and often membranous apically, but it also occurs in some heavy-bodied mandibles in which the apex is flattened and chisel-like.

#100. Dorsal part of mandible <whether with tubercle>/

1. without tubercle/

2. with tubercle fitting into cavity on clypeus/

The tubercle in a state 2 mandible is located on the dorsal surface and fits into a distinct cavity on the lateral portion of the clypeus. SEE "clypeus (whether laterally emarginate)".

#101. Dorsal part of mandible <whether with setose cavity>/

1. without setose cavity/

2. with dorsally-opening, setose cavity/

Mandibles with a setose cavity opening laterally are coded as state 1. In state 2 mandibles, there is a dorsally-opening, setose cavity, which is not visible unless the mandibles are open. This cavity is usually accompanied by a tubercle. SEE "dorsal part of mandible (whether with tubercle)".

#102. Mandible <whether lightly sclerotized or pigmented at apex>/

1. moderately to strongly, more or less evenly sclerotized or pigmented/

2. much more lightly sclerotized or pigmented at apex/

State 2 mandibles may be moderately well-developed basally, often with a well-developed mola, but the apical region is thin and more or less translucent or membranous.

#103. Mandible <whether enclosed in mouth cavity>/

1. visible in lateral view/

2. enclosed in mouth cavity or not visible in lateral view/

Taxa are coded as state 1 when at least the base of the mandible can be seen in lateral view. State 2 mandibles are highly reduced and-or more or less concealed within the mouth cavity.

#104. Incisor edge of mandible <whether with one or more teeth>/

1. simple/

2. with single tooth/

3. with 2 or more teeth/

Mandibles with a serrate cutting edge are coded as state 3.

#105. Mandible <whether with mola>/

1. with well developed mola/

2. with reduced mola/

3. without mola/

The mola is an enlarged area at the base of the mandible, which is provided with some type of armature (teeth, ridges etc.) and is in contact with the comparable structure on the opposing mandible. In some mandibles the armature may have been worn down.

#106. Mandible <whether with prostheca>/

1. with well developed prostheca/

2. with reduced prostheca/

3. without prostheca/

The prostheca usually consists of a membrane, often accompanied by a fringe of hairs, distal to the mola or at the base of the inner edge of the mandible in the absence of a mola; however it may consist solely of a tuft or fringe of hairs. Occasionally, an articulated process may be involved. SEE "prostheca (whether with articulated sclerotized process)". A prostheca may exist in the absence of a mola. unlike the situation in beetle larvae where structures unaccompanied by a mola have been given other names (Lawrence et al. 1993 in General Bibliography). . For a brush of hairs or pubescent process at the inner basal angle of the mandible, SEE character "inner basal angle of mandible (whether with brush or pubescent process)".

#107. Prostheca <whether with articulated sclerotized process>/

1. absent or without articulated, sclerotized process/

2. including articulated, sclerotized process/

A taxon is coded as state 2 when a tooth-like or hook-like process forms part of the prostheca. It may be subdivided and is always separated from the incisor edge of the mandible by membrane (and so appears articulated). It should not be confused with solid teeth or denticles which may form part of the incisor edge.

#108. Inner basal angle of mandible <whether with pubescent process>/

1. without pubescent process/

2. with pubescent process/

This character is usually coded only in those mandibles without a basal mola. In mandibles with a mola, molar asperities occasionally grade into hair-like microtrichia basally; these are not considered to form a basal brush. The pubescent process (state 3) is often elongate and extends posteriorly into the pharynx; this structure occurs only in Curculionoidea and has been called the pharyngeal process by Ting (1936 in General Bibliography).

#109. Maxilla <apical lobes>/

1. with distinct galea and lacinia/

2. with single apical lobe/

3. without apical lobes/

The galea or outer lobe is always laterad of the lacinia or inner lobe. Either the galea or lacinia or both may be reduced in state 1 maxillae, and the lacinia may no more than a slight expansion of the stipes. In some doubtful cases, the maxilla has been coded as 1/2. In state 2 maxillae, the single lobe may represent the galea, lacinia or a fusion of the two lobes. A state 3 maxilla may appear to consist of a palp only.

#110. Maxillary lobe(s) <whether stylet-like>/

1. not stylet-like/

2. stylet-like/

Stylet-like maxillary lobes are very long and slender, often with fine hooks at the end.

#111. Apex of galea or maxillary lobe <vestiture>/

1. densely setose or spinose/

2. without or with few setae or spines/

#112. Apex of galea or maxillary lobe <whether with teeth or hooks>/

1. without heavily sclerotized teeth or hooks/

2. with heavily sclerotized teeth or hooks/

The galea (or maxillary lobe when galea and lacinia are fused) is usually relatively lightly sclerotized and setose, but in some scarabaeoid taxa it may be well sclerotized apically with relatively stout teeth or spines (state 2).

#113. Lacinia <whether with apical hooks or spines>/

1. without hook(s) or spine(s)/

2. with apical or subapical hook(s) or spine(s)/

The hooks or curved spines on the lacinia may be single and simple or apically divided, and more than one may be present. Most of these structures are located at or very near the apex, but in some cases there may be a hook-like process on the inner edge of the lacinia well before the apex.

#114. Apical maxillary palpomere <shape>/

1. cylindrical to fusiform/

2. slightly expanded and truncate to subtriangular/

3. securiform to cultriform/

4. aciculate/

5. of a unique shape/

State 1 includes apical palpomeres which are either cylindrical, wider at the middle and gradually narrowed basally and apically, or widest at the base and slightly narrowed apically (subconical). An extreme condition, where the sides are strongly rounded to produce a more or less globular palpomere, could be coded as either state 1 or state 2, depending on whether the widest point is towards the base or apex, respectively. State 2 includes those apical palpomeres which are slightly widened apically with a truncate apex which may be somewhat oblique, to those which form a more or less inveted equilateral triangle with the widest point at the apex. State 3 includes both securiform and cultriform palpomeres; the former, often defined as triangular, is here restricted to an asymmetrical triangle with sides of different lengths, while the latter has the form of a pairing knife and represents an extreme asymmetrical elongation of the securiform type. The aciculate or awl-like apical palpomere (state 4) is one which is either entirely narrow and needle-like or sopmetimes broad at the base and abruptly narrowed before the middle, so that it looks like an awl. The maxillary palp is often called aciculate when the apical palpomere is much narrower than the preapical one. SEE "preapical maxillary palpomere (whether much larger than apical one)".

#115. Apical maxillary palpomere <whether shorter and narrower than preapical one>/

1. at least as wide as or longer than preapical one/

2. distinctly shorter and narrower than apical one/

The apical maxillary palpomere is coded as state 2 when it is both shorter and narrower than the preapical one. One which is slightly narrower but distinctly longer than the preapical one is coded as state 1.

#116. Maxillary palp <whether with palp organ>/

1. without complex palp organ/

2. with complex palp organ/

A taxon is coded as state 2 when the penultimate maxillary palpomere bears a complex, multilobed sensory organ. This occurs only in the family Lymexylidae.

#117. Apical labial palpomere <shape>/

1. cylindrical to fusiform/

2. slightly to strongly expanded apically/

#118. Ligula <whether emarginate, bilobed or absent>/

1. undivided or finely cleft/

2. shallowly to moderately emarginate/

3. deeply emarginate or bilobed/

4. indistinct or absent/

Some bilobed ligulae (state 4) are further subdivided, so that 4 lobes are formed.

Prothorax #119. Ratio of pronotal length to greatest pronotal width/

The length of the pronotum is measured from above and extends from the anterior edge at midpoint and the posterior edge at midpoint. Pronotal width is the greatest width. This is the same as the prothoracic length and width, since pleural and sternal elements are never visible from above.

#120. Prothorax widest/

1. anteriorly/

2. at middle/

3. posteriorly/

#121. Sides of prothorax <shape>/

1. more or less straight/

2. moderately to strongly curved/

3. straight posteriorly, curved anteriorly/

4. sinuate/

5. variously lobed/

A laterally sinuate prothorax (state 4) is usually outwardly curved anteriorly and inwardly curved posteriorly; however this may be reversed, or two or more s-curves may be present. State 5 includes prothoraces with distinct lateral tubercles or teeth as in many Cerambycidae.

#122. Prothorax <whether laterally compressed forming cavities for legs>/

1. not laterally compressed to form cavities for legs/

2. laterally compressed forming cavities for legs/

A bilateral compression and excavation of the posterior portion of the prothorax occurs in a number of beetles which live in tunnels and have undergone modifications allowing the movement of prothoracic legs in a confined space.

#123. Sides of prothorax <whether explanate>/

1. not or slightly explanate/

2. moderately to strongly, horizontally explanate/

3. moderately to strongly, obliquely or vertically explanate/

A taxon is coded as state 3 when the explanate pronotal margins extend ventrally and are either oblique or almost vertical. This condition is often accompanied by oblique, ventrally projecting elytral epipleura.

#124. Base of prothorax <whether much narrower than elytral bases>/

1. not or slightly narrower than elytral bases/

2. distinctly narrower than elytral bases/

The width of the combined elytral bases is not measured at the very base of the elytra, but rather just behind this, where the humeri are usually located.

#125. Greatest prothoracic width <whether much narrower than greatest elytral width>/

1. not or slightly narrower than greatest elytral width/

2. distinctly narrower than greatest elytral width/

#126. Lateral pronotal carinae <whether incomplete or absent>/

1. complete/

2. incomplete/

3. absent/

The lateral pronotal carina is the sharp lateral edge of the prothorax, which separates the disc from the hypomeron on each side. It is sometimes called the lateral margin, but that term here is restricted to a lateral carina accompanied by a raised bead. In some groups, the carina does not extend to the anterior edge of the prothorax, and in a few it may not reach the posterior edge; in either case the taxon is coded as state 2. If the carina is completely absent, so that the disc is not separated from the hypomeron, then the taxon is coded as state 3.

#127. Lateral pronotal carinae <whether crenulate, serrate or dentate>/

1. simple/

2. finely crenulate or denticulate/

3. coarsely dentate/

The lateral pronotal carina is the sharp lateral edge of the prothorax, which separates the disc from the hypomeron on each side. State 2 refers to fine teeth or crenulations, whereas state 3 includes larger and more easily visible teeth or serrations.

#128. Lateral pronotal carinae <whether visible for their entire lengths from above>/

1. visible for their entire lengths from above/

2. not visible for their entire length from above/

When the pronotal disc is strongly convex, the lateral carina may not be visible from above. A taxon is coded as state 2 when it is impossible to see the entire lengths of both carinae from directly above.

#129. Lateral pronotal carinae <whether margined>/

1. without a raised margin/

2. with a raised margin or narrow bead/

State 2 includes not only the normal narrow margin or bead but also the broader margin defined by a more or less submarginal groove. The latter might be confused with the submarginal carina.SEE "pronotal disc (whether with lateral, submarginal lines, ridges or carinae)."

#130. Lateral portion of prothorax <whether with deep pit>/

1. without deep pit/

2. with deep pit near procoxal cavity/

A taxon is coded as state 2 when there is a deep pit near the lateral edge of the procoxal cavity, which represents an internal apodeme and is often accompanied by a groove.

#131. Pronotum <whether with anterolateral callosities>/

1. without anterolateral callosities/

2. with anterolateral callosities/

Anterolateral callosities (state 2) are laterally projecting enlargements which have a more or less flattened, polished surface containing a gland opening.

#132. Anterior angles of pronotum <whether produced forward>/

1. absent or not produced forward/

2. distinctly produced forward/

The anterior pronotal angles are coded as state 2 when they are produced forward in front of the remainder of the anterior edge.

#133. Anterior angles of pronotum <whether produced>/

1. absent, right or rounded, not produced/

2. produced and broadly rounded or obtusely angulate/

3. produced and narrowly rounded or acute/

#134. Posterior angles of pronotum <whether absent, obtuse or acute>/

1. absent or broadly rounded/

2. obtuse or right/

3. moderately to strongly acute/

#135. Posterior angles of pronotum <whether produced and acute>/

1. not produced and acute/

2. strongly produced and narrowly acute/

Specimens coded as state 2 have the posterior angles narrowly acute and projecting well behind the posterior edge of the prothorax, as in most Elateridae.

#136. Posterior edge of pronotum <whether sinuate or lobed>/

1. more or less straight or evenly rounded/

2. moderately to strongly produced forming mesal lobe/

3. distinctly sinuate or variously lobed/

#137. Posterior edge of pronotum <whether crenulate>/

1. simple/

2. distinctly crenulate/

Crenulations are occasionally difficult to see when the prothorax is fitted tightly to the elytral bases; however they also tend to be lost in taxa usually characterized by their presence.

#138. Posterior edge of pronotum <whether margined>/

1. not or vaguely margined/

2. with narrow raised margin or bead/

3. with broad margin and submarginal groove/

#139. Discal carinae of pronotum <presence and type>/

1. absent/

2. located on posterior angles only/

3. basal but mesad of posterior angles/

4. extending beyond middle of disc/

Submarginal lines or carinae usually lie well in from the lateral edge but may be confused with the edges of broad lateral margins. SEE "lateral pronotal carinae (whether margined)". Basolateral carinae more or less restricted to the produced posterior angles, as in various Elateridae (state 2), are distinguished from shorter (state 3) or longer (state 4) sublateral or paramedian carinae. In some taxa long, sublateral carinae may be accompanied by a second pair of paramedian carinae. If the only longitudinal carinae present are paramedian ones (lying closer to the midline than to the lateral edge, then these are considered to be submarginal lines and coded as such (state 3 or 4).

#140. Pronotal disc <whether with paired basal pits or impressions>/

1. without paired basal impressions/

2. with paired basal impressions/

State 2 includes well marked pits, as well as larger, shallow impressions. Sometimes the pits or impressions may be accompanied by short longitudinal carinae. SEE "lateral submarginal carinae (presence and type)". They may also be joined by a transverse groove or impression. SEE "posterior edge of pronotum (whether margined)".

#141. Pronotum <whether with median groove or line>/

1. without median longitudinal groove or line/

2. with median longitudinal groove or line/

Taxa coded as state 2 for this character may have a distinct median longitudinal groove or a fine line, usually associated with an internal endocarina.

#142. Anterior edge of pronotum <whether margined or with collar>/

1. simple, without margin/

2. with narrow margin or bead/

3. with broad collar or flange/

Taxa coded as state 2 have a very narrow bead-like margin at the anterior edge of the pronotum, similar to those often on the lateral carinae. State 3 refers to those prothoraces with a broader flange or collar, often continued ventrally onto the prosternum. SEE "anterior edge of prosternum (whether margined)".

#143. Hypomeron <whether with large, flat-bottomed pit>/

1. without pit/

2. with large, flat-bottomed pit/

The shallow, flat-bottomed pits in state 2 are located just in front the procoxal cavities. When examined closely, they are seen to be deeper pits each of which is partly blocked by a flattened cuticular flap. Their function is unknown.

#144. Anterior portion of prosternum at midline <length relative to that of prosternal process>/

1. shorter than prosternal process/

2. as long as prosternal process/

3. longer than prosternal process/

The anterior portion of the prosternum is not clearly delimited but can be defined as the midline distance between the anterior edge and an imaginary line drawn across the base of the prosternal process.

#145. Lateral portion of prosternum in front of coxae <length relative to mid length of procoxal cavity>/

1. shorter than mid length of procoxal cavity/

2. as long as mid length of procoxal cavity/

3. longer than mid length of procoxal cavity/

The length of the lateral portion of the prosternum is measured at a point about midway between the mesal and lateral edges of the procoxal cavity (excluding iits lateral extension, if present). This length is compared with the length (or shortest diameter) of the procoxal cavity at the same point.

#146. Anterior edge of prosternum <whether forming chin piece>/

1. not produced anteriorly/

2. distinctly produced forming chin piece/

A taxon is coded as state 2 when the anterior edge of the prosternum is rather abruptly and distinctly produced forward to produce a chin piece beneath the head.

#147. Prosternum in front of coxae <whether concave, convex or carinate>/

1. concave or biconcave/

2. flat to moderately convex/

3. strongly convex without carina/

4. strongly convex with median carina/

A biconcave prosternum (state 1) may also have a median carina which separates the concavities, but this type of prosternum is not strongly convex (as seen from below). A state 3 prosternum may have a pair of carinae enclosing a flattened or slightly concave area. SEE "prosternum in front of coxae (whether with paired lines or carinae)".

#148. Prosternum in front of coxae <whether with paired lines or carinae>/

1. without paired lines or carinae/

2. with paired lines or carinae/

The paired lines or carinae (state 2) extend from the base of the prosternal process or the anterior edges of the procoxal cavities.

#149. Anterior edge of prosternum <whether margined>/

1. without margin/

2. with narrow margin or bead/

3. with broad margin/

#150. Anterior edge of prosternum <whether with mesal excavation>/

1. without mesal excavation/

2. with mesal excavation/

State 2 involves a sharply delimited mesal excavation into which the head fits.

#151. Anterolateral or ventrolateral portions of prothorax <whether with cavities or grooves>/

1. without cavities or grooves/

2. with broad cavities/

3. with narrow grooves/

A taxon is coded as state 1 if the hypomera are broadly concave, of if there are tibial or tarsal grooves present in their posterior portions. Discrete broad cavities (state 2) usually house the antennal clubs only, but sometimes it may be the eyes or the entire antenna, which is curled to fit the space. Antennal grooves are narrower and usually house unclubbed antennae or portions of the flagellum. In rare cases, anterolateral grooves house the protarsi.

#152. Prothoracic cavities <presence and type>/

1. absent/

2. external, open anteriorly, visible from above/

3. external, open anteriorly, not visible from above/

4. external, not open anteriorly/

5. internal/

The cavities coded as states 2 and 3 usually house antennal clubs. State 2 cavities may open anteriorly or dorsally. State 3 cavities are ventral but are continuous with the anterior edge. State 4 cavities do not reach the anterior edge and may house antennae or legs or both. State 5 cavities are entirely internal and house the entire curled antenna; since they often have a relatively narrow opening, they may be confused with antennal grooves.

#153. Prothoracic grooves <presence and type>/

1. absent/

2. longitudinal, below lateral edges of prothorax/

3. longitudinal or oblique, elsewhere on hypomera or prosternum/

Anterolateral prothoracic grooves usually house the antennae, but some of those coded as state 3 are tarsal grooves. State 3 grooves may be on the prosternum or hypomeron or may be oblique and cross both sclerites.

#154. Prosternal process <whether incomplete or absent>/

1. complete/

2. incomplete/

3. absent/

4. complete, but interrupted/

An interrupted prosternal process (state 4) ends before the middle of the procoxae, which are thus contiguous; however an isolated remnant remains behind the coxae and may even be contiguous with postcoxal projections.

#155. Prosternal process <whether narrowed or variously expanded apically>/

1. narrowed apically/

2. parallel-sided/

3. gradually expanded or narrowed and then expanded/

4. gradually expanded and then narrowed/

5. strongly and abruptly expanded at apex/

6. solidly fused to postcoxal projections/

Taxa coded as state 1 typically have the prosternal process broad at base and gradually narrowing to an acute apex; however some taxa have a very short process which rapidly narrows to a broadly rounded or angulate apex. Those taxa with state 2 have the process more or less parallel for most of its length, sometimes slightly narrowing to form the apex. A state 3 process may gradually and evenly widen posteriorly or may become narrower and then widen at apex. If the process is expanded at the middle and narrowed apically, it is coded as state 4. State 5 includes those prosternal processes which abruptly expand at the apex, and there is often some gradation between this and state 3. In state 6, the process is fused with the postcoxal projections, and the dividing lines are often obscured, so that the shape of the prosternal apex cannot be determined. SEE "postcoxal projection".

#156. Prosternal process <whether curved or elevated apically>/

1. flat, concave, or only slightly elevated or curved behind coxae/

2. strongly elevated and curved dorsally behind coxae/

3. slightly to strongly elevated but not curved behind/

The state 4 type of prosternal process normally extends well onto the mesoventrite (= mesosternum) and may even join directly to the metaventrite (= metasternum). SEE "prosternal process (whether produced posteriorly)". In some cases, however, the process may terminate abruptly at or even before the posterior edges of the coxae.

#157. Prosternal process <whether produced posteriorly>/

1. not extending to mesoventrite/

2. slightly overlapping mesoventrite/

3. moderately to strongly overlapping mesoventrite/

4. concealing most or all of mesoventrite/

The state 3 type of prosternal process may conceal most of the mesoventrite (= mesosternum), so that only a narrow strip of cuticle is visible. The state 4 type extends posteriorly to the edge of the metaventrite (= metasternum).

#158. Apex of prosternal process <shape>/

1. acute or narrowly rounded/

2. broadly rounded, angulate or truncate/

3. singly or multiply cleft or emarginate/

#159. Prosternal process <whether with transverse groove>/

1. without transverse groove/

2. with transverse groove/

#160. Accessory (mesal) procoxal articulation <type>/

1. absent/

2. formed by prosternal tongue and coxal groove/

3. formed by coxal knob and prosternal cavity/

A accessory procoxal articulation may be formed when each side of the prosternal process is slightly widened to form a tongue which fits into a groove or cavity in the procoxa (state 2). In Adephaga,however, a small knob on the procoxa fits into a cavity on the prosternal process (state 3). In both cases, this secondary articulation restricts movement of the procoxa, while increasing the structural integrity of the coxal joint.

#161. Ventral portion of prothorax on each side <sutures>/

1. with notopleural and pleurosternal sutures/

2. with notopleural suture only/

3. with notosternal suture only/

4. without sutures or with incomplete notosternal suture/

The notopleural suture lies just beneath the lateral pronotal carina and separates the pronotal hypomeron from the propleuron. The pleurosternal suture separates the pleuron from the sternum. Both are present in state 1. In the state 2 prothorax the notosternal suture is distinct, but the pleuron is fused to the sternum. The notopleural suture is absent in state 3, but this is due, not to the fusion of pleuron and hypomeron, but to the internalization of the entire pleuron, which can be seen in cleared specimens. The single suture remaining is the notosternal suture, separating the hypomeron from the sternum. state 4 applies to those taxa in which all sutures have disappeared. In some taxa, like Micromalthidae, there is no internal pleuron, and an external pleuron has fused with both notum and sternum. Most other taxa are polyphagans with an internal pleuron, which is visible externally only as the so-called trochantin (pleurotrochantin). SEE "prothoracic trochantin or pleurotrochantin (whether concealed)".

#162. Propleuron <whether extending to anterior edge of prothorax>/

1. extending to anterior edge of prothorax/

2. not extending to anterior edge of prothorax/

In taxa coded as state 1 the propleuron extends to the anterior edge of the prothorax, thus separating the anterior edge of the notum from that of the sternum. In state 2 taxa the pleuron never reaches the anterior edge.

#163. Propleuron or pleurotrochantin <whether extending behind coxa>/

1. extending behind coxa/

2. not extending behind coxa/

In state 1 taxa, the external pleuron extends behind the procoxa and may form a postcoxal projection. This is not homologous to the postcoxal projection in Polyphaga, which is formed by the pronotal hypomeron. SEE "postcoxal projection".

#164. Procoxae <whether projecting below prosternum>/

1. not or slightly projecting below prosternum/

2. projecting well below prosternum/

#165. Procoxa <whether with long concealed, lateral extension>/

1. without or with short concealed lateral extension/

2. with long concealed, lateral extension <more than 0.7 X exposed portion of coxa>/

A concealed procoxal extension is considered to be long (state 2), if it is more than 0.7 times as long as the exposed portion of the coxa.

#166. Procoxal cavities <whether absent>/

1. present, procoxae countersunk/

2. absent, procoxae attached externally/

In almost all groups of Coleoptera, the procoxae are countersunk into the prothorax forming the coxal cavities. Each cavity is bordered anteriorly and often mesally by the prosternum and laterally and often posteriorly by the pleuron (Adephaga, Myxophaga and Archostemata) or pronotal hypomeron (Polyphaga) and is at least partly lined internally by a thin cuticular cowling formed by the prosternum. In a number of derived groups, such as the families Cantharidae, Lampyridae and Lycidae, these cavities undergo extreme reduction. The prosternal process disappears, the internal cowling is reduced, and what remains of it is no longer countersunk, so that it lies at the same level as the rest of the prosternum. The coxa is attached externally by membrane to an opening, which corresponds to the internal opening of the original coxal cavity. SEE "procoxal cavities internally (whether closed)". This condition is coded as state 2.

#167. Procoxal cavity <shape>/

1. strongly transverse/

2. slightly transverse/

3. circular or longer than wide/

A taxon is considered to have state 1 when the transverse diameter of a coxal cavity is at least twice the longitudinal diameter. State 2 cavities have the transverse diameter less than twice the longitudinal one, and state 3 cavities are about as long as wide.

#168. Procoxal cavities at middle <separation>/

1. contiguous/

2. narrowly separated/

3. moderately to widely separated <more than 0.4 X shortest diameter of coxal cavity>/

Widely separated procoxal cavities (state 2) are separated by more than 0.4 times the shortest diameter of the coxal cavity.

#169. Procoxal cavities externally <whether closed>/

1. open/

2. closed/

State 2 coxal cavities are closed behind by the meeting of the prosternal process and postcoxal projections or occasionally by the meeting of the two postcoxal projections behind the prosternal process.

#170. Procoxal cavities externally <degree of closure>/

1. broadly open/

2. narrowly open/

3. narrowly closed/

4. broadly closed/

#171. Postcoxal projection <presence and extent>/

1. absent or very short/

2. moderately long but not meeting prosternal process/

3. meeting prosternal process/

4. meeting opposing postcoxal projection/

5. solidly fused to opposing postcoxal projection, and sometimes to prosternal process/

A very short postcoxal projection (state 1) does not extend beyond the lateral 3rd of the coxal cavity. A moderately long postcoxal projection (state 2) extends beyond the lateral 3rd but does not meet the prosternal process. A state 3 projection meets the prosternal process (usually fitting into a notch at its apex), and a state 4 projection meets the opposite projection at midline, behind, below or above the prosternal process. A state 5 condition is when the two notal processes are completely fused, so that there is no suture between them; they also may be fused to the apex of the prosternal process. SEE state 6 of the character "prosternal process (whether narrowed or variously expanded apically)".

#172. Procoxal cavities <whether with lateral extensions>/

1. without narrow lateral extensions/

2. with narrow lateral extensions/

State 1 procoxal cavities may be broadly expanded laterally, but do not have the narrow, slit-like extension. The presence of this extension, which is due to the incomplete meeting of the prosternum and hypomeron at the coxal end of the notosternal suture, is sometimes described as having an "open procoxal fissure" (Herman 1970, pp. 349–353) or "open procoxal cavity" (Zimmerman 1994, p. 250–251); however the condition is totally unrelated to the posterior closure of the coxal cavity.

#173. Procoxal cavities internally <whether closed>/

1. open/

2. closed by slender bar/

3. broadly closed/

A procoxal cavity is considered to be internally open when there is a opening occupying much of the cavity, the cowling located mesally and anteromesally. Sometimes there is a narrow strip of cuticle or bar extending from the cowling to the inside of the postcoxal or notal process; such a coxa is considered to be internally "narrowly closed" and is coded as state 2. When the internal cowling is more extensive, occupying most of the cavity, except for a relatively small opening at the anterolateral end, to which the coxa is attached, then the cavity is said to be "broadly closed" and is coded as state 3.

#174. Prothoracic trochantin or pleurotrochantin <whether concealed>/

1. at least partly exposed/

2. completely concealed or absent/

In Archostemata and Adephaga, the trochantin is a distinct sclerite, separated from both pleuron and coxa; it is exposed in Archostemata and usually concealed in Adephaga. In Myxophaga and Polyphaga, the trochantin fuses with the pleuron forming a pleurotrochantin. In many Polyphaga, the pleurotrochantin is entirely concealed beneath the walls of the notum and sternum, which join to form the ventrolateral walls of the prothorax, but in others it may be partly or entirely exposed and is usually referred to as the trochantin.

#175. Promesothoracic clicking mechanism <whether present>/

1. absent/

2. present/

The promesothoracic clicking mechanism (state 2) is composed of a relatively large and deep mesocoxal cavity and a long prosternal process extending well behind the procoxae. In addition, the dorsal (concealed) surface of the apex of the prosternal process bears a notch or step-like process and the anterior part of the mesoventral cavity (= mesosternal cavity) is raised slightly forming a lip. SEE character "mesoventral cavity". For information on the nature of the jumping mechanism, SEE Evans 1972 and 1973 in General Bibliography.

Elytra #176. Elytra <whether absent>/

1. present/

2. absent (anelytrous or larviform)/

Those taxa coded as state 2 include neotenic, larviform adults (almost always females), which not only lack elytra and hind wings, but also have larviform antennae, eyes, and-or legs; in other cases, however, the absence of elytra and hind wings is the only obvious difference between these forms and normal adults.

#177. Ratio of elytral length to greatest elytral width/

The length of the elytra is measured from above at midline and extends from the base of the scutellum (or posterior edge of the pronotum if the scutellum is concealed) to the conjoined elytral apices or an imaginary line joining the separated elytral apices. The elytral width usually is the greatest combined width; however, when the elytra are widely separated at the base, the width used is not actually with combined widths of the two, but the length of the longest line joining their two outside edges.

#178. Ratio of elytral length to pronotal length/

The length of the prothorax is measured from above and extends from the anterior edge at midpoint and the posterior edge at midpoint. The elytral length is measured from above at midline and extends from the base of the scutellum (or posterior edge of the pronotum if the scutellum is concealed) to the conjoined elytral apices or an imaginary line joining the separated elytral apices.

#179. Elytra <whether with distinct puncture rows or striae>/

1. apunctate, irregularly punctate, or with 5 or fewer distinct puncture rows or striae/

2. with more than 5 distinct puncture rows/

3. with more than 5 distinct impressed striae/

A taxon is coded as state 3 when there are at least 6 distinct puncture rows or striae which can be counted. If the most of the elytral punctures are irregular or confused and a few form striae, especially laterally or along the suture, the taxon is also coded as state 1.

#180. Elytra <whether with scutellary striole>/

1. without scutellary striole/

2. with scutellary striole/

The scutellary striole (state 2) is an abbreviated stria or puncture row lying near the scutellum and mesad of the first complete stria or puncture row.

#181. Number elytral puncture rows or striae/

1. 12 or more/

2. 11/

3. 10/

4. 9/

5. 8 or fewer/

The number of striae or puncture rows does not include the scutellary striole.

#182. Sutural stria <whether deeply impressed>/

1. absent or not deeply impressed near apex/

2. deeply impressed near apex/

A taxon is coded as state 2 only when there is a deep impression near the apex of the sutural stria.

#183. Abdominal tergites exposed by elytra <number>/

1. none or apex of 1/

2. most of one/

3. at least one but less than 2/

4. at least 2 but less than 3/

5. 3 or more/

The number of exposed tergites does not include the genital capsule or pregenital segments (segments 9 and 10), which sometimes project beyond the apex of segment 8. If one tergite is exposed (states 2 or 3), it is tergite 8. If 2 are exposed (state 4), they are tergites 7 and 8. If 3 are exposed (state 5) they are 6 to 8, etc. Some Pselaphidae coded as state 5 apparently have only 2 visible tergites; however the first of these is very long and represents the fusion of several tergites.

#184. Exposed abdominal segments <whether rigid>/

1. more or less flexible/

2. rigid and compact/

When a taxon is coded as state 2, the elytra are truncate, exposing several abdominal tergites which are not at all flexible. Occasionally basal tergites may be solidly fused (in which case the apparent basal tergite is very large). This condition separates pselaphine Staphylinidae from other members of the family.

#185. Elytral apices <whether independently rounded>/

1. meeting or almost meeting at the suture/

2. independently rounded or acute and separated by broad gap/

If a taxon is coded as state 1, the elytral apices form a continuous curve or straight line (in truncate elytra), without or with a relatively narrow gap between them.

#186. Elytral suture <whether deflected at or near apex>/

1. not deflected near apex/

2. deflected at or near apex/

State 2 of this character is not the same as state 2 of the character "elytral apices (whether independently rounded)". It occurs in elytra which meet at the apex (state 1 of "elytral apices") and involves a deflection of the suture, so that a portion of the elytral flange is visible when the elytral apices are in the closed position.

#187. Elytral apex <whether with internal interlocking tongue>/

1. without internal interlocking tongue/

2. with internal interlocking tongue/

The interlocking tongue is relatively broad-based apodeme located close to the elytral apex, projecting laterally and fitting under the inflexed laterosternite on ventrite 5 (sternite 7).

#188. Epipleuron <whether complete to apex>/

1. absent or incomplete/

2. complete/

In order for the elytral epipleuron to be coded as complete, it must extend to the apex at the suture. Thus truncate elytra with epipleura extending to the apex laterally are coded as state 1.

#189. Epipleuron <whether abruptly narrowed>/

1. not or gradually narrowed/

2. abruptly narrowed or excavated/

A taxon is coded as state 2 when the epipleura is abruptly narrowed at about the level of the metacoxae.

#190. Lateral edge of elytron <whether deeply emarginate>/

1. straight or weakly sinuate/

2. strongly sinuate or emarginate, exposing part of body from above/

A taxon is coded as having state 2 when a portion of the metathorax and-or abdomen on each side can be seen from above, due to a distinct emargination or sinuation in the lateral edge of each elytron.

Pterothorax #191. Scutellum <whether absent or not visible>/

1. well developed/

2. highly reduced/

3. absent or not visible/

#192. Scutellum <whether abruptly elevated>/

1. not abruptly elevated/

2. abruptly elevated/

A taxon is coded as state 2 when it is abruptly and more or less sharply elevated above the level of the mesoscutum as seen from the side. This is not visible externally when the pronotum and elytra are closely adpressed. A state 1 scutellum may be clearly separated from the mesoscutum.

#193. Scutellum anteriorly <whether notched or crenulate>/

1. simple/

2. notched/

3. crenulate/

A taxon is coded as state 1 when the anterior edge of the scutellum is neither notched nor crenulate and sometimes is not clearly separated from the mesoscutum.

#194. Scutellum posteriorly <shape>/

1. narrowly rounded or acute/

2. broadly rounded or obtusely angulate/

3. truncate/

4. emarginate/

#195. Mesoscutum <whether with stridulatory file>/

1. without stridulatory file/

2. with undivided stridulatory file/

3. with divided stridulatory file/

Under lower magnifications, a stridulatory file looks like a shiny area, which is usually narrow and elongate. Under higher magnification, a series of fine transverse ridges may be seen. A taxon is coded as state 3 when there is an external, longitudinal division of the file, and not when there is an internal endocarina beneath it (which may give the impression that the file is divided).

#196. Mesoventrite <whether with paired procoxal rests>/

1. without paired procoxal rests/

2. with paired procoxal rests/

Paired procoxal rests are slightly to strongly concave, shiny areas located at the anterior edge of the mesoventrite (= mesosternum); they form housings for the procoxae when the latter overlap the mesoventrite at rest. Sometimes these rests extend laterally onto the mesepisterna, but this character refers only to those portions formed from the mesoventrite. When the entire mesoventrite is concave or biconcave the taxon is coded as state 1.

#197. Paired mesoventral procoxal rests <presence and angle of inclination>/

1. absent, horizontal or slightly oblique/

2. moderately to strongly oblique/

3. vertical/

Procoxal rests on the anterior edge of the mesoventrite are paired concave housings into which the posterior portions of the procoxae fit. They are usually present only when the procoxal cavities are externally open. This character is coded as state 1 when the procoxal rests are absent or when they are only slightly concave and are more or less horizontally oriented and on the same plane as the remainder of the mesoventrite. When the rests are more obliquely oriented and usually more deeply concave, they are coded as state 2. State 3 procoxal rests are almost vertical in orientation, so that the procoxae fit into them anteriorly.

#198. Anterior edge of mesoventrite <whether with prosternal rest>/

1. without prosternal rest/

2. with median prosternal rest/

3. with broadly transverse prothoracic rest/

A prosternal rest (state 2) is usually a median flattened area on which the prosternal process rests. Sometimes it may be slightly concave or have a median carina. A taxon is coded as state 3 when the entire anterior portion of the mesoventrite (= mesosternum), and usually the associated mesepisterna, are smooth and rounded, forming a pedicel over which the entire prothorax fits like a ball and socket joint. State 3 is always correlated with broadly closed procoxal cavities.

#199. Mesoventrite <whether longitudinally divided>/

1. not divided by longitudinal groove or discrimen/

2. at least partly divided by longitudinal groove or discrimen/

A taxon is coded as state 2 when the mesoventrite (= mesosternum) is partly or completely divided by a distinct longitudinal groove or discrimen (Ferris 1940 in Genberal Bibliography). This state sometimes occurs in conjunction with a mesoventral cavity which obliterates much of the groove.

#200. Anterior edge of mesoventrite at midline <whether on different plane than metaventrite>/

1. on same plane as metaventrite/

2. on different plane than metaventrite/

A taxon is coded as state 2 when the anterior portion of the mesoventrite (= mesosternum) is distinctly and more or less abruptly elevated above the level of the metaventrite (= metasternum) as seen from a side view. The mesoventrite thus lies in two planes with a more or less vertical shelf in between them.

#201. Mesoventral cavity <presence, size and depth>/

1. absent/

2. small and shallow/

3. moderately large and shallow/

4. moderately to very large and deep/

The mesoventral cavity (= mesosternal cavity) is a small to very large, shallow to deep cavity in the mesoventrite (= mesosternum) which receives the apex of the prosternal process.

#202. Mesocoxa <shape>/

1. not conical and projecting/

2. conical and projecting/

#203. Mesocoxal cavities <whether absent>/

1. present, mesocoxae countersunk/

2. absent, mesocoxae attached externally/

In almost all groups of Coleoptera, the mesocoxae are countersunk into the mesothorax forming the coxal cavities. Each cavity is bordered anteriorly and often mesally by the mesoventrite (= mesosternum), laterally by the mesoventrite and mesepimeron, sometimes the mesepisternum, and rarely the metepisternum, and posteriorly and sometimes mesally by the metaventrite (= metasternum); internally the cavity is lined by a cowling composed of elements of the mesoventrite and metaventrite. In a number of derived groups, such as the families Cantharidae, Lampyridae and Lycidae, these cavities undergo extreme reduction. The median processes of the mesoventrite and metaventrite are obliterated, and the cowlings loose their concavity and come to lie in the same plane as the major portions of the meso- and metasterna. The coxae are externally attached by membrane to the opening which was originally within the original cavity.

#204. Mesocoxal cavities at middle <separation>/

1. contiguous/

2. narrowly separated/

3. moderately to widely separated <more than 0.4 X shortest diameter of coxal cavity>/

A taxon is coded as state 2 when the coxal cavities are separated by less than 0.4 times and state 3 when they are separated by more than 0.4 times the shortest diameter of a coxal cavity.

#205. Mesocoxae separated by/

1. less than shortest diameter of coxal cavity/

2. more than shortest diameter of coxal cavity/

#206. Mesocoxal cavities <whether strongly transverse>/

1. circular to slightly transverse/

2. strongly transverse/

#207. Mesocoxal cavities <whether strongly oblique>/

1. not or slightly oblique/

2. moderately to strongly oblique/

#208. Mesoventrite <whether fused on each side to mesepisternum>/

1. separated by complete sutures from mesepisterna/

2. partly or completely fused to mesepisterna/

#209. Mesepisterna <whether meeting at midline>/

1. distinctly separated at midline/

2. meeting or very narrowly separated at midline/

3. broadly joined at midline/

A taxon is coded as state 1 when the distance between the mesepisterna is more than and state 2 when this distance is less than 0.25 times the shortest diameter of a mesocoxal cavity. State 3 mesepisterna are broadly joined at the midline, so that the mesoventrite (= mesosternum) is separated from the anterior edge of the mesothorax. This character is not coded (considered as inapplicable) when the mesepisterna are not or incompletely separated from the mesoventrite (character "mesoventrite (whether fused on each side to metspisterna)", state 2).

#210. Mesepisterna <whether with deep pockets>/

1. without deep pockets/

2. with deep pockets/

The mesepisternal pockets usually project mesally, so that their openings face laterally. They are thought to be glandular.

#211. Mesepimeron <whether visible from above>/

1. not visible from above/

2. visible from above between pronotum and elytral base on each side/

#212. Mesocoxal cavities <lateral closure>/

1. open laterally/

2. closed laterally/

Mesocoxal cavities are considered to be laterally closed (state 2) when they are closed by the meeting of the mesoventrite (= mesosternum) and metaventrite (= metasternum). Laterally open mesocoxal cavities (state 1) are closed by the mesepimera, sometimes in conjunction with the mesepisterna and-or (rarely) the metspisterna, SEE "mesocoxal cavities (whether partly closed by metepisterna)".

#213. Mesocoxal cavities <whether partly closed by metepisterna>/

1. not partly closed by metepisterna/

2. partly closed by metepisterna/

#214. Mesoventral process <presence and extent>/

1. absent or not extending to middle of mesocoxal cavity/

2. extending at least to middle of mesocoxal cavity/

#215. Mesometaventral junction <presence and shape>/

1. absent or a point/

2. a straight line/

3. a posteriorly curved, angulate or acute line/

4. an anteriorly curved, angulate or acute line/

5. a complex fitting/

6. absent due to fusion/

7. concealed by prosternal process/

8. concealed by metaventral process/

The meso-metaventral junction is the meeting of the intercoxal processes of the mesoventrite and metaventrite. A taxon is coded as state 1 when the mesoventral process (= mesosternal intercoxal process) is absent, does not meet the metaventrite (= metasternum), or meets it at a point. In the first two cases, the mesocoxal cavities are contiguous (state 1 of the character "mesocoxal cavities at midline (separation)"). A state 3 junction involves a convex, overlapping mesoventral process (as seen from below), and a state 4 junction consists of a convex, overlapping metaventral process. State 5 includes a variety of complex fittings, such as ball and socket or tongue and groove. In a state 6 junction, the line between the mesoventrite and metaventrite is vaguely indicated or absent and the two plates are immovably fused. States 7 and 8 are special conditions in which the junction is completely concealed by either the prosternal or the metaventral process, which may be joined directly to the metaventrite or prosternum, respectively.

#216. Mesoventral and metaventral processes at midline <whether fused>/

1. separated by gap, groove, or suture/

2. solidly fused or separated at most by weakly impressed line/

In most beetles, the external portions of the mesoventrite (= mesosternum) and metaventrite (= metasternum) at midline, if contiguous, are separated by a distinct groove or suture (state 1); internally (within the coxal cowling) the two are almost always solidly fused together. A taxon is considered to have state 2 when the mesoventrite and metaventrite are solidly fused externally, separated at most by a weakly impressed line.

#217. Accessory (mesal) mesocoxal articulation <type>/

1. absent/

2. formed by mesoventral tongue and coxal groove/

3. formed by metaventral tongue and coxal groove/

4. formed by coxal knob and mesoventral cavity/

The accessory mesal or internal mesocoxal articulation may be of three different types. In some groups of Cerambycidae and Chrysomelidae, the mesoventral process has a pair of lateral projections near the apex, each of which fits into a cavity or groove on the mesocoxa. In some Dermestidae, the articulation is formed by a pair of lateral processes on the metaventral process. In Haliplidae, a knob on each mesocoxa fits into a cavity on the underside of the mesoventral process. These structures limit the movement of the mesocoxae but increase the structural integrity of this region.

#218. Metaventral discrimen or median line <presence and extent>/

1. moderately to very long/

2. short <less than 0.25 X body of metaventrite>/

3. absent/

The metaventral discrimen is a line or groove representing the invagination of the true sternum and metendosternite; it is often called a median line or median metasternal suture (Ferris 1940 in General Bibliography). A state 1 discrimen is more than and a state 2 discrimen less than 0.25 times as long as the body of the metaventrite (excluding the anterior metaventral process).

#219. Paired postcoxal lines of metaventrite <whether present>/

1. absent/

2. present/

Postcoxal lines are fine, elevated ridges on the metaventrite (= metasternum), which extend posteriorly or posterolaterally from the hind edges of the mesocoxal cavities. They sometimes mark the edge of a concavity for the reception of the mid femora, and are sometimes called femoral lines.

#220. Postcoxal lines of metaventrite <type>/

1. absent/

2. arched and strongly recurved/

3. arched but not or slightly recurved/

4. sinuous or complexly curved/

5. straight or slightly, mesally curved/

There is usually a single pair of postcoxal lines, but rarely (state 6) two pairs are present. State 2 postcoxal lines curve posterolaterally and then curve anteriorly again to meet the lateral edges of the coxal cavities. State 3 lines also extend posterolaterally, but they are not recurved and often reach the metepisternum, forming a triangular area between them and the coxal cavity. Sometimes they follow the lateral edge of the coxal cavity and diverge only near the metspisternum. State 4 lines are more complex and sinuate, extending posteriorly then laterally and then posteriorly again, meeting the metepisternum near its posterior end. State 5 lines are more or less straight or only slightly curved mesally. State 6 is used when there are 2 pairs of lines which differ in form.

#221. Metaventrite <whether shorter than first abdominal ventrite>/

1. longer than first abdominal ventrite/

2. not longer than first abdominal ventrite/

A taxon is coded as state 2 when the body of the metaventrite (= Metasternum) is as long as or shorter than the first abdominal ventrite. This state is correlated with wing reduction and flightlessness.

#222. Postcoxal pits of metaventrite <whether present>/

1. absent/

2. present/

Postcoxal pits are relatively small, posteriorly opening cavities lining the posterior edge of the mesocoxal cavity.

#223. Metaventrite <degree of convexity>/

1. flat to slightly convex/

2. moderately to strongly convex/

#224. Transverse groove of metaventrite <whether present>/

1. absent/

2. present/

The transverse metaventral groove (= transverse metasternal suture) crosses the discrimen near the posterior end of the ventrite, but it rarely extends to the lateral edges. In some taxa coded as state 2, it does not extend far from the midline on either side.

#225. Anterior edge of metaventrite <whether with transverse carina>/

1. without transverse carina between mesocoxal cavities/

2. with transverse ridge or carina joining posterior edges of mesocoxal cavities/

A taxon is considered to have state 2 when there is a transverse, curved or straight line on the anterior portion of the metaventrite (= metasternum) connecting the posterior edges of the two mesocoxal cavities; this is often assocated with an abrupt change in the elevation of the meso-metaventral area. SEE "anterior edge of mesoventrite at midline (whether on different plane than metaventrite)".

#226. Exposed portion of metepisternum <shape>/

1. short and broad <less than 2.5 X as L as W>/

2. moderately elongate <2.5–4 X as L as W>/

3. very long and narrow or absent <more than 4 X as L as W>/

A taxon is coded as state 1 when the exposed portion of the metepisternum is less than 2.5 times as long as wide, state 2 when it is between 2.5 and 4 times as long as wide, and state 3 when it is more than 4 times as long as wide or sometimes absent (concealed by the edge of the elytron).

#227. Metacoxae <whether widely separated>/

1. contiguous or narrowly separated/

2. widely separated <at least 0.5 X longest coxal diameter>/

Metacoxae are considered to be widely separate (state 2) when they are separated by a distance which is at least 0.5 times as great as the transverse (longest) diameter of a metacoxa.

#228. Metacoxae <whether separated by 1 or 2 times longest coxal diameter>/

1. separated by less than longest coxal diameter/

2. separated by more than 1 but less than 2 times longest coxal diameter/

3. separated by more than 2 times longest coxal diameter/

#229. Metacoxae <whether extending laterally to meet elytra or sides of body>/

1. not extending laterally to meet elytra or sides of body/

2. extending laterally to meet elytra or sides of body/

A state 1 metacoxa does not extend laterally to meet the elytra at rest or the sides of the body (when the elytra are reduced or dehiscent), so that the junction between the metathorax and venrtite 1 is visible.

#230. Metacoxae <whether partly or completely fused to metaventrite>/

1. completely separated from metaventrite by suture/

2. at least partly fused to metaventrite, suture incomplete or absent/

Although most Adephaga have relatively immobile metacoxae, the complete fusion of the coxae to the metaventrite (= metasternum) with no separating line or suture is a rare condition.

#231. Metacoxal plates <degree of development>/

1. well developed, more or less uniform/

2. well developed mesally, weak laterally/

3. weakly developed/

4. absent/

A metacoxal plate is formed when the coxa is posteriorly excavate to receive the hind femur; it represents the ventral wall of the excavation. Sometimes it is posteriorly expanded to conceal a portion of the leg or part of the abdomen. SEE "metacoxal plates (whether concealing basal ventrites)".

#232. Metacoxal plates <whether concealing basal abdominal ventrites>/

1. not concealing most of basal abdominal ventrite/

2. concealing most of first abdominal ventrite/

3. concealing most of first three abdominal ventrites/

#233. Metacoxae <whether greatly enlarged>/

1. not greatly enlarged/

2. greatly enlarged/

A taxon is coded as state 2 when the metacoxa is longer at middle than the adjacent portion of the metaventrite (= metasternum). These enlarged metacoxae are often more or less fused to the metaventrite. In some cases (e.g. Haliplidae), the coxae are relatively small, but the attached coxal plates are very large and conceal the hind legs.

#234. Metacoxae <whether obliquely oriented>/

1. horizontally oriented/

2. obliquely oriented/

A taxon is coded as state 2 when the lateral end of the metacoxa lies well in front of the mesal end (horizontally oblique); thus it refers only to their orientation in ventral view. If the ventral portion of the metathorax is strongly convex, the metacoxae may be vertically oblique, but in this case they are coded as state 1.

#235. Lateral arms of metendosternite <whether moderately to very long>/

1. short or absent/

2. moderately to very long/

The metendosternite is usually a fork-like internal apodeme representing the invaginations resulting in the formation of the discrimen and transverse metaventral groove (= transverse metasternal suture). It is only visible in a cleared or completely dissected specimen. It usually consists of a median stalk and lateral arms, but may also have an anterior process, ventrolateral processes, laminae (on the arms), and a pair of anterior tendons. The median stalk varies in length and may be absent, especially in wingless forms and those with widely separated metacoxae.

#236. Metendosternal laminae <whether reduced or absent>/

1. well developed/

2. reduced/

3. absent/

#237. Ventrolateral processes of metendosternite <whether strongly developed>/

1. absent or weakly developed/

2. strongly developed/

The ventrolateral processes may be confused with laminae, but they are located on the stalk and project basally.

#238. Anterior process of metendosternite <whether short or absent>/

1. moderately long/

2. short or absent/

The anterior process may be deeply divided into two parts.

#239. Anterior tendons of metendosternite <whether widely separated or on arms>/

1. moderately or very close together/

2. widely separated but not on lateral arms/

3. on lateral arms or not apparent/

When there are no obvious anterior tendons, the taxon is coded as state 3.

#240. Apical portion of metendosternite <whether deeply emarginate>/

1. not or only slightly emarginate/

2. broadly, deeply and abruptly emarginate/

The state 2 condition is one in which the anterior edge of the metendosternite is abruptly, deeply and moderately broadly emarginate, so that the anterior process is divided into two halves, each one bearing a tendon. In some cases, the emargination extends basally beyond the origins of the short lateral arms; the resulting structure appears to have a pair of broad, bifurcate arms.

Hind wing #241. Hind wing <whether highly reduced or absent>/

1. well developed/

2. highly reduced or absent/

A reduced wing may be elongate and narrow with one or more longitudinal veins and sometimes an expanded apex bearing a remnant of the medial fleck (as in the figure) or it may consist of a minute flap without or with only remnants of venation. When wings are reduced or absent, the elytral humeri (shoulders) are often rounded and the metasternum is usually shortened (SEE "Correlated characters" image).

#242. Hind wing <whether lacking transverse folds>/

1. with normal transverse folds/

2. lacking transverse folds/

Almost all hind wings have at least one transverse or oblique fold, which allows the wing to decrease in length when at rest beneath the elytra. The most obvious exception (state 2) is the genus Atractocerus, in which the wings are never folded beneath the highly reduced elytra (SEE image)..

#243. Radial cell of hind wing <whether highly reduced or absent>/

1. well developed/

2. highly reduced or absent/

The term radial cell is used here in a special sense and refers to a type of cell found only in the wings of polyphagan beetles and absent (coded as inappplicable) in beetles belonging to the other three suborders. Where the cell occurs in Polyphaga, it is formed when vein RA forks at about the middle of the anterior edge of the wing and the two forks meet again enclosing an eyelet-like space between them. This type of cell, which is shown on screens A and B, is always adjacent to the anterior edge of the wing, and may be transversely elongate with an abrupt base. Radial cells in Adephaga, Myxophaga and Archostemata are usually formed by cross-veins joining the two forks of RA, and they are usually further modified by a series of transverse or oblique folds which occur in this region of the wing. These "false" radial cells, which are shown on screen C, are separated from the anterior edge and often vertical, irregular and-or subdivided.

#244. Radial cell of hind wing <shape>/

1. elongate/

2. shorter and broader/

3. incomplete or absent/

An elongate radial cell (state 1) is at least 2 times as long as wide. An incomplete radial cell (state 3) lacks a proximal end (the base of RA3+4).

#245. Inner posterior angle formed at base of radial cell <whether acute>/

1. right or obtuse/

2. acute/

The angle referred to is not the basal angle where RA3+4 forks with RA1+2, but rather the inner posterior angle formed where RA3+4 meets a radial cross-vein.

#246. Radial cell <whether forming equilateral triangle>/

1. not forming equilateral triangle/

2. forming equilateral triangle/

#247. Ratio of length of apical area to total wing length/

1. less than 0.2/

2. 0.20–0.35/

3. 0.36–0.5/

4. greater than 0.5/

The apical area is usually measured from the end of the radial bar (the thickened, sclerotized portion of the anterior edge of the wing) to the wing apex, and the total length is measured from the base of the humeral plate. However, in some wings, the anterior sclerotization extends out onto the apical area, in which case there is either an abrupt hinge or a spring (a region bissected by fine transverse lines) at some point. The apical area is considered to be that portion of the wing which is folded beneath the elytra, and is measured from the hinge or from the center of the spring.

#248. Apical area of hind wing <whether with veins or sclerotizations>/

1. without veins or sclerotizations/

2. with one or more vague sclerotizations or pigment patches/

3. with one or more distinct veins (branches of RA and-or RP)/

Sclerotizations in the apical region of the wing are usually remnants of RA or RP, the former more anteriorly placed and the latter more posteriorly placed and curving posteriorly. In taxa coded as state 3, these remnants are clearly marked and vein-like, but in state 2 wings, they are not as obviously veins and in some cases may not be homologues of RA or RP. Commonly in Elateriformia, three of these sclerotizations or wing remnants form a trident-like structure. In some Staphyliniformia, RA4 and RP1 converge and fuse apically.

#249. Medial bar of hind wing <whether crossed by fold>/

1. not crossed by fold/

2. crossed by fold/

A taxon is coded as state 2 when the medial bar (MP1+2) is crossed by a transverse or oblique fold. In Adephaga and Myxophaga there is a sharp hinge at this point, but in Archostemata there may be a spring. This crossing is located near the apex of the bar but basad of the oblongum cell, when this is present. In Polyphaga, any transverse folds occur beyond the apex of the medial bar or at least do not cross it.

#250. Free veins in medial area of hind wing <number>/

1. 5 or 6/

2. 4/

3. 3 or fewer/

The medial area is that region of wing membrane lying in between the medial bar (MP1+2) and the anal fold. The veins enclosed by these two features include MP3, MP4 and the branches of Cu and AA. Free veins are those with a free apex (not joined apically to another vein). In the terminology usually used for beetle wing venation (Crowson 1955 in General Bibliography), these veins are referred to as "anal veins in the main group".

#251. Oblongum cell of hind wing <whether absent>/

1. present/

2. absent/

The oblongum cell is formed by cross-veins joining RP and MP1+2. It is usually vertical (with its long axis perpendicular to the long axis of the wing) or more or less triangular, and sometimes it is incomplete.

#252. Medial fleck of hind wing <whether present>/

1. absent/

2. present/

The medial fleck is a more or less granulate binding patch which allows the two hind wings to remain attached to one another and-or to another binding patch on the elytra. This structure has also been called the subcubital fleck or binding patch.

#253. Medial fleck of hind wing <whether partly bisected by a vein>/

1. absent or not partly bisected by a vein/

2. partly bisected by a vein/

The medial fleck often appears to be divided into two parts, but a taxon is considered to have state 2 only when one of the branches of MP actually extends into the fleck, more or less bisecting it.

#254. Wedge cell of hind wing <whether reduced or absent>/

1. well developed/

2. reduced/

3. absent/

The wedge cell is formed by the joining of portions of the cubital and anterior anal veins. It is also called the anal cell or the 2nd cubito-anal cell, and it is the only completely closed cell in the medial area of the wing (the 1st cubito-anal cell extends to the wing base).

#255. Apex of wedge cell of hind wing <shape>/

1. squarely truncate/

2. obliquely truncate/

3. acute/

In a wing coded as state 1, the apical end of the wedge cell is squarely cut off by a cross-vein that is perpendicular to the long axis of the cell. In a state 2 wing, this end of the cell is more or less obliquely truncate, whereas state 3 refers to a condition in which the two veins making up the sides of the cell gradually approach one another and form an acute apical angle.

#256. Anal lobe of hind wing <whether present>/

1. absent/

2. present/

An anal lobe is formed when a distinct incision in the wing edge at the apex of the anal fold isolates the anal area from the remainder of the wing.

#257. Posterior edge of hind wing <whether with fringe of long hairs>/

1. without fringe of long hairs/

2. with fringe of long hairs/

A number of wings of small beetles have a marginal fringe, but a taxon coded as state 2 must have a relatively broad fringe consisting of long setae.

Legs #258. Femoral attachment of mid trochanter <whether transverse or oblique>/

1. transverse or slightly oblique/

2. strongly oblique with base of femur separate from coxa/

3. strongly oblique with base of femur abutting coxa/

The state 3 condition of the trochanterofemoral joint has been called the heteromeroid type, since it characterizes many members of the superfamily Tenebrionoidea. However, the condition occurs throughout the order.

#259. Mesotrochanter <whether reduced and concealed or absent>/

1. not reduced or concealed from below/

2. highly reduced, strongly oblique and concealed from below/

3. absent/

#260. Metafemur <whether much wide than mesofemur>/

1. not much wider than mesofemur/

2. much wider than mesofemur/

#261. Mesotibia <whether strongly widened>/

1. not strongly widened/

2. strongly widened, widest at or near apex/

3. strongly widened, widest well before apex/

#262. Outer edge of mesotibia <whether spinose>/

1. simple, crenulate or denticulate/

2. with distinct teeth or long spines/

A taxon is coded as state 1 if the outer edge of the tibia forms a simple or crenulate ridge, or when it is covered with small denticles or stiff setae. A state 2 mesotibia may have distinct lobes, teeth, or long and-or stout spines along the outer edge.

#263. Outer subapical edge of mesotibia <whether with antenna cleaner>/

1. without antenna cleaner/

2. with antenna cleaner/

The mesotibial antenna cleaner is located along the outer egde just before the apex and may consist of a distinct concavity lined with hairs, an oblique setose groove, or a densely setose area. SEE "inner subapical edge of protibia (whether with antenna cleaner)".

#264. Preapical surfaces of mesotibia <whether with ridges or combs>/

1. without ridges or combs/

2. with transverse or oblique ridges or combs/

A taxon coded as state 2 has a series of short, transverse or oblique ridges, often crenulate or comb-like and darkly pigmented, but sometimes bearing spines.

#265. Outer apical angle of mesotibia <armature>/

1. simple or slightly produced, without lobe, teeth or spines/

2. with one or more straight or outwardly facing teeth or spines/

3. with rounded lobe or process, sometimes bearing spines/

4. with inwardly curved or hooked spine (uncus)/

When a mesotibia is coded as state 4, it may have a curved or hooked spine projecting apically and inwardly curved only at apex, or it may have a spine which projects inwardly and crosses over the apex of the tibia. This differs from a mucro, which originates at the inner apical angle and projects inwardly. SEE "inner apical angle of mesotibia (whether forming tooth)".

#266. Inner apical angle of mesotibia <whether forming tooth>/

1. not or slightly produced, without tooth/

2. produced forming tooth (mucro)/

The mucro is a short, stout tooth or spine arising from the internal apical angle of the tibia and facing inwards; it may be confused with an uncus which sometimes arises at the outer angle but crosses the apex to face inwardly. SEE state 4 of the character "outer apical angle of mesotibia (whether forming tooth)".

#267. Mesotibial spurs <whether pubescent, serrate or pectinate>/

1. glabrous or absent/

2. pubescent/

3. serrate or pectinate/

A state 2 tibial spur is more or less evenly clothed with short hair-like processes. State 3 spurs may be distinctly serrate or pectinate, but sometimes they have a single longitudinal row of short, hair-like appendages.

#268. Mesotibial spurs <whether single or absent>/

1. double/

2. single/

3. absent/

Mesotibial spurs are usually double, while those on the protibia or metatibia may be reduced or absent. SEE characters "protibial spurs (whether single or absent)" and "metatibial spurs (whether single or absent)". In smaller beetles or those with distinct setal fringes at the tibial apex, the spurs are difficult to see; in general, when the spurs cannot be clearly distinguished from fringe setae, the specimen should be coded as state 3. When an apical setal fringe is viewed from an angle, several fringe setae are seen superimposed and may appear to be a broader, more heavily sclerotized spur.

#269. Mesotarsus <number of distinct and reduced tarsomeres>/

1. with 5 distinct tarsomeres (pentamerous)/

2. with 4 distinct tarsomeres and reduced penultimate one (pseudotetramerous)/

3. with 4 distinct tarsomeres (tetramerous)/

4. with 3 distinct tarsomeres and reduced penultimate one (pseudotrimerous)/

5. with 3 distinct tarsomeres (trimerous)/

6. with 2 or fewer tarsomeres/

The pseudotetramerous condition (state 2) is that in which the tarsi appear to be 4-segmented because the penultimate tarsomere is highly reduced and concealed within the lobe of the preceding tarsomere; this is also called cryptopentamerous. The pseudotrimerous condition (state 4) is similar, but it is the 3rd segment which is reduced and concealed within the apex of the 2nd, making a 4-segmented tarsus appear 3-segmented.

#270. Tarsomeres on hind leg <whether 1 fewer than on mid leg>/

1. at least as many as on mid leg/

2. 1 fewer than on mid leg/

#271. Tarsomeres on fore leg <whether 1 fewer than on mid leg>/

1. at least as many as on mid leg/

2. 1 fewer than on mid leg/

#272. Mesotarsomere 1 <whether reduced>/

1. well developed and visible/

2. reduced and partly concealed/

#273. Preapical mesotarsomeres together <whether shorter than apical one>/

1. longer than apical one/

2. shorter than apical one/

#274. Penultimate mesotarsomere <whether shorter than antepenultimate>/

1. not distinctly shorter than antepenultimate/

2. distinctly shorter than antepenultimate/

#275. Ventral mesotarsal lobes <presence, location and number>/

1. absent/

2. on penultimate tarsomere only/

3. on antepenultimate tarsomere only/

4. on basal tarsomere only/

5. on more than one tarsomere/

In the pseudotetramerous or pseudotrimerous type of tarsus, the lobe is on the antepenultimate tarsomere, but it appears to be on the penultimate, since the true penultimate tarsomere is highly reduced and fused to the following one.

#276. Mesotarsal claws <whether single or absent>/

1. paired/

2. single/

3. absent/

#277. Mesotarsal claws <whether differing in length, form or angle of inclination>/

1. subequal in length and similar in form and angle of inclination/

2. differing in length/

3. differing in form/

4. differing in angle of inclination/

Tarsal claws of different thickness are coded as state 3. State 4 is really just an artifact produced in dried specimens when the claws are independently movable along a single plane. When the beetle is killed, the two claws usually dry in two different positions.

#278. Mesotarsal claws <armature>/

1. simple/

2. toothed or bifid/

3. serrate, denticulate or pectinate/

A taxon is coded as state 2 if each tarsal claw is either split at the apex (bifid) or bears a single tooth, usually near the base. A state 3 tarsal claw may have 2 to many small teeth, serrations, or comb-like processes (pectinate).

#279. Appendage on each tarsal claw <type>/

1. absent/

2. membranous or lightly sclerotized, usually pubescent/

3. heavily sclerotized, blade-like/

4. heavily sclerotized, spine-like or bristle-like/

Tarsal claw appendages arise from the base of each claw and are usually nearly or quite as long as the claw itself.

#280. Mesotarsal claws <whether with setae near base>/

1. without setae near base/

2. with one or more setae near base/

The claw setae are usually located near the base of the outer surface of each claw; there is usually only a single seta on each claw, but there may be more.

#281. Mesotarsal empodium <whether absent>/

1. present and exposed/

2. absent or concealed/

If the empodium is reduced and does not extend beyond the bases of the tarsal claws, then the taxon is coded as state 2.

#282. Mesotarsal empodium <whether with 3 or more setae>/

1. absent or with 2 or fewer setae/

2. with 3 or more setae/

Most empodia are bisetose, but some may be unisetose or asetose. A state 2 empodium is relatively large and has more than 2 setae at the apex.

#283. Outer edge of protibia <whether with distinct lobes or teeth>/

1. simple and rounded to carinate but without lobes or teeth, except at apex/

2. with one or more distinct lobes or teeth/

The armature of the anterior edge of the protibia may consist of blunt, rounded processes or sharp teeth, and the former often represent sharp teeth or spines which have been worn down. If there is a single process or one or two small teeth only at the outer apical angle, then the taxon is coded as state 1. SEE "outer apical angle of protibia (armature)".

#284. Outer apical angle of protibia <armature>/

1. simple or slightly produced, without lobe, teeth or spines/

2. with one or more straight or outwardly facing teeth or spines/

3. with rounded lobe or process, sometimes bearing spines/

4. with inwardly curved or hooked spine (uncus)/

When a protibia is coded as state 4, it may have a curved or hooked spine projecting apically and inwardly curved only at apex, or it may have a spine which projects inwardly and crosses over the apex of the tibia; either type is called an uncus (particularly by those specializing in Curculionoidea). The uncus differs from a mucro, which originates at the inner apical angle and projects inwardly. SEE "inner apical angle of mesotibia (whether forming tooth)".

#285. Inner subapical edge of protibia <whether with antenna cleaner>/

1. without antenna cleaner/

2. with antenna clearner/

The protibial antenna cleaner is located on the inner edge just before the apex and may consist of a deep concavity or excavation lined with hairs or spines (most Carabidae), or a weak, oblique, setose impression or dense patch of hairs (some Cerambycidae). Antenna cleaners occur only in taxa with filiform antennae. SEE "outer subapical edge of mesotibia (whether with antenna cleaner)".

#286. Protibial spurs <whether single or absent>/

1. double/

2. single/

3. absent/

Protibial spurs are often reduced to 1 or absent in those taxa having the tibial apex modified for either digging or grasping.

#287. Articulations of protibial spurs <whether located in different planes>/

1. located in same plane/

2. located in different planes, one being subapical/

The articulation of the one of the protibial spurs in most Carabidae has been displaced in connection with the development of the antenna cleaner.

#288. Hind legs <whether with swimming hairs>/

1. without swimming hairs/

2. with swimming hairs/

The swimming hairs on the mid and-or hind legs of some water beetles (Hygrobiidae, most Dytiscidae and many Hydrophilidae) usually occur on the tarsi but may be present on the tibiae as well.

#289. Preapical surfaces of metatibia <whether with ridges or combs>/

1. without ridges or combs/

2. with transverse or oblique ridges or combs/

A taxon coded as state 2 has a series of short, transverse or oblique ridges, often crenulate or comb-like and darkly pigmented, but sometimes bearing spines.

#290. Metatibial articular area <whether broadly expanded and flattened>/

1. not to only moderately expanded, narrowly oval or oblique/

2. greatly expanded, broadly oval to circular and flattened/

The apex of the metatibia may become greatly expanded in burrowing beetles, so that the apical surface forms a broadly oval, more or less flattened area housing the tibial spurs and tarsal articulation.

#291. Metatibial spurs <whether single or absent>/

1. double/

2. single/

3. absent/

#292. Metatibial spurs <whether differing in length and-or form>/

1. subequal in length and form/

2. differing distinctly in length/

3. differing distinctly in form/

Metatibial spurs may be very different in length and-or form in some burrowing scarabaeoids.

Abdomen #293. Number of abdominal ventrites/

Abdominal ventrites are visible abdominal sternites. The first ventrite is usually sternite 3, but may be sternite 2, and the last ventrite is usually sternite 7 but may be sternite 8. In counting ventrites, the male genital capsule (sternite 9) is not included, although it occasionally projects beyond the apex of the last ventrite. Because the male genital capsule is excluded, normal adults never have more than 8 ventrites; however larviform females may have 9 or 10.

#294. Number of basal ventrites connate/

1. none/

2. two/

3. three/

4. four/

5. five or six/

Connate ventrites are more or less solidly fused together or at least incapable of being moved independently of one another. There is a gradation between slightly connate ventrites and those which are soldered together and separated by a weak or incomplete suture. Differences between basal joints and those between the more apical ventrites often suggest that the basal ventrites are connate. For instance, basal ventrites may have no membrane between them while apical ones do. A taxon is coded as state 1 when the basal two ventrites are free, even though two or more of the more apical ones are may be connate. SEE "ventrites 4 (whether articulated with 3 but connate with 5)".

#295. Abdominal sternite 2 <whether concealed or apparently absent>/

1. visible/

2. present as concealed or partly exposed lateral sclerites/

3. apparently absent/

Abdominal sternite 2 is usually concealed beneath the metacoxae or entirely fused to sternite 3 (state 3); in this case the basal ventrite is sternite 3. In some groups, however, there may be a complete 2nd ventrite present and in others this ventrite is divided into 2 or 3 parts by the metacoxae. SEE "first ventrite (whether completely divided by metacoxae)". State 2 refers to the condition in which sternite 2 is represented by lateral sclerites usually concealed beneath the elytra and visible only in dissections.

#296. First ventrite <whether completely divided by metacoxae>/

1. not completely divided by metacoxae/

2. divided into 2 or 3 parts by metacoxae/

The complete division of ventrite 1 into 2 or occasionally 3 parts characterizes the suborder Adephaga; however the suture between the first two ventrites may be incomplete or obscure, in which case ventrite 1 appears to be very large and the coxae do not appear to completely subdivide it. A divided first ventrite may also occur in some in Polyphaga, where the metacoxae are particularly large and the first ventrite reduced. It is particularly likely to occur in Staphyliniformia or Scarabaeoidea, where separated portions of sternite 2, usually concealed beneath the elytral bases, are partly exposed.

#297. Suture between ventrites 1 and 2 <whether incomplete or vaguely indicated>/

1. distinct/

2. incomplete or vaguely indicated/

In some taxa with state 2 the line dividing ventrites 1 and 2 is more or less completely obliterated, so that there seems to be one less ventrite present. In taxa where this is likely to occur, the characters "abdominal ventrites (number)" and "basal ventrites connate (number)" are coded for both the actual and apparent number of ventrites.

#298. Suture between ventrites 2 and 3 <whether incomplete or vaguely indicated>/

1. distinct/

2. incomplete or vaguely indicated/

In some taxa with state 2 the line dividing ventrites 2 and 3 is more or less completely obliterated, so that there seems to be one less ventrite present. In taxa where this is likely to occur, the characters "abdominal ventrites (number)" and "basal ventrites connate (number)" are coded for both the actual and apparent number of ventrites.

#299. Ventrite 4 <whether articulated with 3 but connate with 5>/

1. articulated with or connate with both 3 and 5/

2. articulated with 3 but connate with 5/

State 2 refers to a condition in which the basal ventrites are freely articulated, but the last 2 ventrites (4 and 5) are solidly fused together.

#300. Postcoxal lines on ventrite 1 <presence and type>/

1. absent/

2. 1 pair, strongly curved or recurved/

3. 1 pair, straight or slightly curved/

4. 2 pairs/

Postcoxal lines on ventrite 1 are similar in structure and function to those on the metaventrite. SEE characters "paired postcoxal lines of metaventrite (whether present)" and "postcoxal lines of metaventrite (type)".

#301. Ventrite 1 <whether much longer than 2>/

1. not much longer than 2/

2. much longer than 2/

#302. Ventrite 1 at middle of metacoxa <whether much shorter than 2>/

1. about as long as or longer than ventrite 2/

2. distinctly shorter than ventrite 2/

This character has been coded for all taxa of Curculionoidea, and serves to distinguish typical Platypodinae from other members of the superfamily; elsewhere it has been coded only sporadically.

#303. Abdominal process <shape>/

1. acute or narrowly rounded/

2. broadly rounded or angulate/

3. truncate or slightly emarginate/

4. absent/

The abdominal process is an anterior extension of the first ventrite, which extends in between the two metacoxal cavities. It is formed only when the base of the abdomen is countersunk to form cavities for housing the coxae. In those taxa in which the metacoxae project below the abdomen (and thus overlap the base of the abdomen when viewed from below), the cavities are usually absent, as is the abdominal process.

#304. Ventrite 5 in female <whether with circular depression>/

1. without circular depression/

2. with distinct circular depression/

#305. Posterior edge of ventrite 5 <whether crenulate>/

1. not crenulate/

2. crenulate/

#306. Last visible tergite and-or sternite (7 or 8) <whether forming terminal spine>/

1. not forming terminal spine/

2. produced forming terminal spine/

#307. Tergite and sternite 7 <whether fused>/

1. separated by membrane or distinct suture/

2. solidly fused, without or with incomplete suture between them/

#308. Subapical abdominal luminous organ <whether present>/

1. absent/

2. present/

The subapical luminous organ is visible as one or more yellowish-white patches which may occupy an entire segment or more than one segment near the apex of the abdomen. Taxa coded as state 1 may still have smaller luminous spots located elsewhere on the body.

#309. Ventrites <whether with setose patches or foveae>/

1. without setose patches or foveae/

2. with 1 or more setose patches/

3. with 1 or more setose foveae/

Setose patches (state 2) or setose foveae (state 3) are usually median, but are sometimes paired. They are most likely to occur on the basal ventrites. These are sometimes referred to as sex patches and apparently produce pheromones. The term "trichobothria" has also been applied to this type of structure in weevils, but that term normally refers to a specific type of sensory hair.

#310. Functional spiracles on abdominal segment 8 <whether absent>/

1. present/

2. absent/

A functional spiracle is one which has a distinct opening and a well-developed tracheal connection; it may also have a spiracular closing apparatus. There may be a pair of non-functional, spiracular remnants with no distinct opening and with the tracheal connection absent or narrow with a blocked lumen; in this case the specimen is considered to have state 2. Segment 8 is often lightly sclerotized and telescoped within segment 7; however in some groups with 6 ventrites (or visible sternites) it is well sclerotized and forms the visible abdominal apex. Distally it is attached to the genital capsule (9–10 complex) in the male or to the ovipositor in the female.

#311. Functional spiracles on abdominal segment 7 <whether absent>/

1. present/

2. absent/

A functional spiracle is one which has a distinct opening and a well-developed tracheal connection; it may also have a spiracular closing apparatus. There may be a pair of non-functional, spiracular remnants with no distinct opening and with the tracheal connection absent or narrow with a blocked lumen; in this case the specimen is considered to have state 2. Segment 7 often forms the visible abdominal apex, enclosing segment 8 and the genital capsule or ovipositor; however in some groups with 6 or more ventrites (or visible sternites) it and precedes the last visible segment.

#312. Functional spiracles on abdominal segment 6 <whether absent>/

1. present/

2. absent/

A functional spiracle is one which has a distinct opening and a well-developed tracheal connection; it may also have a spiracular closing apparatus. There may be a pair of non-functional, spiracular remnants with no distinct opening and with the tracheal connection absent or narrow with a blocked lumen; in this case the specimen is considered to have state 2. Segment 6 is associated with the 4th or 5th ventrite, depending upon whether the true second sternite is visible at the base of the abdomen.

#313. Functional spiracles on abdominal segment 5 <whether absent>/

1. present/

2. absent/

A functional spiracle is one which has a distinct opening and a well-developed tracheal connection; it may also have a spiracular closing apparatus. There may be a pair of non-functional, spiracular remnants with no distinct opening and with the tracheal connection absent or narrow with a blocked lumen; in this case the specimen is considered to have state 2. Segment 5 is associated with the 3rd or 4th ventrite, depending upon whether the true second sternite is visible at the base of the abdomen.

#314. 7th abdominal spiracles <position>/

1. located in pleural membrane/

2. located on sternite/

3. located on tergite/

4. located on tergosternal suture or area of fusion/

A specimen is considered to have state 2 when the functional spiracle on segment 7 is located within a distinct notch in the sternite or state 3 when in a distinct notch in the tergite. State 4 refers to those cases in which the 7th tergite is connate with or fused to the sternite and the spiracle is located either on the suture or in the area of fusion when the suture is absent.

#315. 6th abdominal spiracles <position>/

1. located in pleural membrane/

2. located in sternite/

3. located in tergite/

A specimen is considered to have state 2 when the functional spiracle on segment 6 is located within a distinct notch in the sternite or state 3 when in a distinct notch in the tergite.

#316. 5th abdominal spiracles <position>/

1. located in pleural membrane/

2. located in sternite/

3. located in tergite/

A specimen is considered to have state 2 when the functional spiracle on segment 5 is located within a distinct notch in the sternite or state 3 when in a distinct notch in the tergite.

#317. Anterior edge of sternite 8 in male <whether with median strut>/

1. without median strut/

2. with median strut/

A taxon is coded as state 2 when the mesal portion of the anterior edge of sternite 8 is strongly produced to form a narrow, rod-like strut. If the anterolateral corners of the sternite are produced to form a pair of narrow struts, the taxon is still coded as having state 1.

#318. Pygidium (sclerotized tergite 7 or 8) <orientation>/

1. more or less horizontal/

2. moderately to strongly oblique/

3. vertical or deflexed/

The pygidium is a well sclerotized, terminal tergite, which usually represents segments 7 or 8. When exposed the tergite may be oblique (state 2) or vertical to deflexed (state 3). In the last case it is not visible from above and may appear to be a ventrite.

#319. Anterior edge of sternite 9 in male <whether with median strut>/

1. without median strut/

2. with median strut (spiculum gastrale)/

A taxon is coded as state 2 when the mesal portion of the anterior edge of sternite 9 is strongly produced to form a narrow, rod-like strut, the spiculum gastrale. A state 1 9th sternite may be produced to form a broad anterior process, rounded or acute at apex.

#320. Tergite 9 in male <whether emarginate or divided>/

1. truncate/

2. slightly to moderately emarginate/

3. deeply emarginate/

4. almost completely divided into two parts/

5. completely fused to tergite 10/

An apically truncate tergite 9 (state 1) is separated from 10 by a straight suture. In states 2 and 3, tergite 9 is slightly to strongly emarginate and separated from 10 by a curved suture, which is sometimes obliterated in the middle due to a partial fusion of 9 and 10. SEE state 2 of character "tergite 10 in male (whether fused to 9)". In state 4, the 9th tergite appears to be completely divided into 2 parts, although there is a membranous connection at the base, and 10 may lie in between these two paratergites. State 5 refers to a variety of conditions in which tergite 10 is either membranous or completely fused to 9 and thus not visible as a distinct sclerite. SEE state 3 of character "tergite 10 in male (whether fused to 9)".

#321. Tergite 10 in male <whether fused to 9>/

1. well developed and free/

2. partly fused to tergite 9/

3. completely membranous or fused to tergite 9/

#322. Aedeagus <type>/

1. trilobate/

2. bilobate/

3. histeroid/

4. buprestoid/

5. adephagan/

6. staphylinoid/

7. bostrichoid/

8. cucujiform/

9. tenebrionoid/

10. chrysomeloid/

11. curculionoid/

12. unclassified/

The aedeagus is the term used by most coleopterists for the male external genitalia (for an alternative use of this term, see end of note). The basic, trilobate aedeagus (state 1) consists of a pair of free parameres (or lateral lobes) attached to the apex of a phallobase (or basal piece) and lying ventral to the penis (or median lobe). This type occurs in various basal Polyphaga, and in most Elateriformia. A series of modified types (states 2 to 11) have been described, but some aedeagi do not fall readily into any group (state 12). The bilobate type of aedeagus (state 2) is one in which there is a well-developed phallobase and parameres, but the penis has become membranous, except for endophallic sclerotizations and sometimes paired basal struts; it is characteristic of many Lucanidae, Geotrupidae. Hybosoridae and Scarabaeidae.

The histeroid aedeagus (state 3) is a modified trilobate type in which the phallobase is large, the parameres are more or less fused together, and the two form a tube around the penis. The buprestoid aedeagus (state 4) is also tubular, but the phallobase is either reduced in size or fused to the parameral tube and the latter is flattened and often more of less expanded apically.

In the adephagan aedeagus (state 5), the phallobase is absent and the parameres are articulated directly to the base of the penis and-or to each other; the parameres are often asymmetrical. This type of aedeagus occurs throughout Archostemata, Myxophaga and Adephaga. This is referred to as the pseudotrilobate by Crowson (1981) and the articulate type by d'Hotman and Scholtz (1990).In the staphylinoid type (state 6), the parameres are fused to the highly reduced phallobase, which is usually represented by a narrow, transverse strip of cuticle joining the parameres on each side of it. In some this is not distinguishable from the adephagan type (state 5).

The bostrichoid type of aedeagus (state 7), the phallobase broadly overlaps the parameres, which are basally united with the penis and may also be joined by a narrow bridge. In the cucujiform type of aedeagus (state 8), the phallobase is enlarged and forms a sheath or ring around the penis, usually with a well-developed posterior, dorsal part joined on each side to a ventral, anterior part, often drawn out forming a strut. This strut, usually called a manubrium, may be articulated or fixed at its base. Parameres are often reduced and may be articulated or fused to the posterior end of the sheath. When the parameres and phallobase are completely fused together, the resulting structure is called the tegmen. These aedeagi are often referred to as cleroid (sheath or vaginate) or cucujoid (ring or annulate), depending upon whether a broad sheath or narrower ring is formed, but the two tend to grade into one another. In a modification called the "double tegmen" the anterior end of the sheath has a pair of dorsal struts in addition to the ventral strut. The cucujiform aedeagus occurs not only in cleroids and cucujoids but also in basal groups of Chrysomeloidea and Curculionoidea.

The tenebrionoid type (state 9) is also sheath-like, but the tegmen lies enitrely on one side of the penis (normally dorsally, but sometimes ventrally due to torsion), and does not enclose or surround it. Articulated parameres are not present, although the apex of the tegmen may be cleft or bilobed. The tegmen may be divided into a basal and apical part, which are apparently homologous to phallobase and fused paramreres, respectively, but they are usually called the basale and apicale. The apicale in some tenebrionoids has a pair of articulated appendages usually located near the base; these are called accessory lobes.

The chrysomeloid type (state 10) is another modification of the cucujoid type in which the parameres are reduced and fused or absent, and the anterior strut may be laterally compressed, forming vertical plate; in the extreme type of chrysomeloid aedeagus, the dorsal portion of the ring is membranous, so that only a forked ventral anterior strut remains. The curculionoid aedeagus (state 11) is similar to the chrysomeloid type, in that the parameres are reduced, fused to the ring and often absent, and the entire dorsal portion of the ring may be membranous; the manubrium is often short and never vertically compressed.

The term aedeagus is used here to refer to the external genitalia, including the intromittent organ (penis) and associated structures (phallobase and parameres, or tegmen) but excluding modifications of apical or pregenital abdominal segments. Some coleopterists restrict the term aedeagus to the intromitten organ alone (penis above), and the term penis is used for this structure combined with phallobase and parameres or tegmen (aedeagus above). Some authors refer to the phallobase as fused gonocoxites, the parameres as gonostyli, and the two together as gonoforceps; the penis is considered to represent fused penis valves. For more information on terms for insect genitalia (including other systems of nomenclature), SEE Crowson (1984), D'Hotman and Scholtz (1990), Gilbert (1952), Iablokoff-Khnzorian (1980), Jeannel (1955), Jeannel and Paulian (1944), Lindroth (1957), Matsuda (1976), Nichols (1989), Tuxen (1970) and Wood (1952) in General Bibliography.

#323. Aedeagus <whether asymmetrical>/

1. symmetrical/

2. asymmetrical/

A taxon is coded as state 2 when the parameres are of a different length or form (the right differing from the left) or when the phallobase and-or the penis are asymmetrical. If the endophallic armature is asymmetrically arranged, this character is still coded as state 1.

#324. Anterior edge of tegmen or phallobase <whether with strut(s)>/

1. without struts/

2. with single strut/

3. with single strut and opposing paired struts/

A taxon is still coded as state 1 if the tegmen or phallobase is broadly emarginate at base so it appears to have paired struts. The paired struts in a state 3 tegmen are located opposite median strut (usually dorsally, but ventrally in an inverted aedeagus).

#325. Apicale or apical portion of tegmen <whether with accessory lobes>/

1. without accessory lobes/

2. with accessory lobes/

The accessory lobes are relatively narrow, articulated processes arising from the apical piece or apicale (fused parameres) of a tenebrionoid tegmen; they are usually sub-basal in position.

#326. Parameres <whether fused to phallobase, fused together or absent>/

1. individually articulated to phallobase or base of penis/

2. fused to phallobase or base of penis but free from one another/

3. partly or entirely fused together but articulated to phallobase/

4. partly or entirely fused together and to phallobase, or absent/

In state 1, the parameres may have a narrow basal connection to one another or with the phallobase.

#327. Parameres <whether outwardly hooked>/

1. not outwardly hooked/

2. outwardly hooked/

#328. Penis <whether with dorsal and ventral lobes>/

1. without dorsal and ventral lobes/

2. with dorsal and ventral lobes/

In some taxa coded as state 1 there is a longitudinal strut located in membrane beneath the main body of the penis. A state 2 penis must have distinct dorsal and ventral sclerotized lobes. These lobes are not necessarily homologous in all groups. The ventral lobe in dascilloids, psephenoids and some scirtids is smaller than the dorsal one, whereas in some Chrysomeloidea and Curculionoidea, the dorsal lobe (tectum) is smaller than the ventral one (pedon).

#329. Anterior edge of penis <whether with strut(s)>/

1. without struts/

2. with single strut/

3. with paired struts/

A taxon may be coded as state 1 when there is a broad anterior process articulated with main body of the penis. In state 2, there is a narrow, rod-like anterior strut, which may or may not be articulated with the main body of the penis. A state 3 penis has paired anterior struts.

Taxonomic level #330. <Taxonomic level>/

1. family/

2. lowest level/


The interactive key provides access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting specified attributes, and summaries of attributes within groups of taxa.

Cite this publication as: ‘Lawrence, J.F., Hastings, A.M., Dallwitz, M.J., Paine, T.A., and Zurcher, E.J. 2000 onwards. Elateriformia (Coleoptera): descriptions, illustrations, identification, and information retrieval for families and subfamilies. Version: 9th October 2005. http://delta-intkey.com’.

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