Elateriformia (Coleoptera) Larvae

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

Character List

#1. <Synonymy>/

#2. <Classification - suborder: series/superfamily>/

1. Archostemata/

2. Myxophaga/

3. Adephaga/

4. Polyphaga: Eucinetoidea/

5. Polyphaga: Hydrophiloidea/

6. Polyphaga: Staphylinoidea/

7. Polyphaga: Scarabaeoidea/

8. Polyphaga: Elateriformia/

9. Polyphaga: Derodontoidea/

10. Polyphaga: Bostrichoidea/

11. Polyphaga: Lymexyloidea/

12. Polyphaga: Cleroidea/

13. Polyphaga: Cucujoidea/

14. Polyphaga: Tenebrionoidea/

15. Polyphaga: Chrysomeloidea/

16. Polyphaga: Curculionoidea/

17. Placement Uncertain/

#3. <Geographical distribution>/

#4. <Biology and comments>/

#5. <References>/

Body #6. Length/

1. less than 3 mm/

2. 3 to 15 mm/

3. more than 15 mm/

#7. Body <whether disc-like>/

1. not broadly ovate and strongly flattened/

2. broadly ovate, strongly flattened and disc-like/

#8. Body <whether C-shaped>/

1. relatively straight or only slightly curved ventrally/

2. strongly curved ventrally (c-shaped)/

#9. Body <cross-section>/

1. circular in cross-section/

2. slightly flattened/

3. strongly flattened/

#10. Body, as viewed from above <shape>/

1. elongate and more or less parallel-sided/

2. oblong to ovate, not parallel-sided/

#11. Vestiture <type>/

1. consisting of fine hairs or setae only/

2. including bristles, scales, expanded or complex hairs/

#12. Dorsal surfaces <pigmentation or sclerotization>/

1. more or less heavily pigmented or sclerotized/

2. very lightly pigmented or sclerotized/

#13. Dorsal surfaces <sculpturing>/

1. generally smooth/

2. generally granulate or tuberculate/

3. generally spinose or complexly sculptured/

#14. Ventral surfaces <pigmentation>/

1. more or less heavily pigmented/

2. very lightly pigmented/

Head #15. Head <orientation>/

1. prognathous or slightly declined/

2. moderately to strongly declined (hypognathous)/

#16. Head <whether retracted>/

1. protracted or slightly retracted/

2. strongly retracted/

#17. Head <whether concealed by prothorax>/

1. not concealed from above by prothorax/

2. completely concealed from above by prothorax/

#18. Head <whether broader than thorax>/

1. narrower than thorax/

2. broader than thorax/

#19. Head <whether forming sclerotized, serrate, wedge-like plate>/

1. not forming sclerotized, serrate, wedge-like plate/

2. forming sclerotized, serrate, wedge-like plate/

#20. Head capsule <whether depressed and wedge-like>/

1. not strongly depressed and wedge-like anteriorly/

2. strongly depressed and wedge-like anteriorly/

#21. Posterior edge of head capsule <whether emarginate>/

1. not or only slightly emarginate/

2. distinctly emarginate/

#22. Median epicranial region <whether with longitudinal furrow>/

1. without longitudinal furrow/

2. with longitudinal furrow (sometimes obscuring epicranial stem) for attachment of retractor muscle/

#23. Epicranial stem <presence>/

1. absent/

2. present/

The ‘epicranial stem’ or coronal suture is the basal or posterior portion of the Y-shaped ecdysial line known as the ‘epicranial suture’. It may be very faint or well marked and sometimes accentuated by the presence of an internal ridge or ‘endocarina’ (see ‘median endocarina’ characters). Occasionally the epicranial stem is located within a basal longitudinal furrow housing the retractor muscle of the head (see ‘median eppicranial region’ character).

#24. Epicranial stem <length>/

1. absent or very short/

2. moderately long/

The ‘epicranial stem’ or coronal suture is the basal or posterior portion of the Y-shaped ecdysial line known as the ‘epicranial suture’. It may be very faint or well marked and sometimes accentuated by the presence of an internal ridge or ‘endocarina’ (see ‘median endocarina’ characters). Occasionally the epicranial stem is located within a basal longitudinal furrow housing the retractor muscle of the head (see ‘median eppicranial region’ character).

#25. Epicranial stem <whether bent to the left>/

1. straight, not bent to the left/

2. bent to the left/

The ‘epicranial stem’ or coronal suture is the basal or posterior portion of the Y-shaped ecdysial line known as the ‘epicranial suture’. It may be very faint or well marked and sometimes accentuated by the presence of an internal ridge or ‘endocarina’ (see ‘median endocarina’ characters). Occasionally the epicranial stem is located within a basal longitudinal furrow housing the retractor muscle of the head (see ‘median eppicranial region’ character).

#26. Frontal arms <presence>/

1. absent/

2. present/

The ‘frontal arms’ represent the distal or anterior portions of the ecdysial line known as the ‘epicranial suture’. These lines may be faint or absent in some instars, but sometimes they are reinforced by paired ‘endocarinae’ or the anterior portions of a Y-shaped endocarina (see ‘endocarina’ characters).

#27. Frontal arms <shape>/

1. lyriform/

2. V-shaped or U-shaped/

The ‘frontal arms’ represent the distal or anterior portions of the ecdysial line known as the ‘epicranial suture’. These lines may be faint or absent in some instars, but sometimes they are reinforced by paired ‘endocarinae’ or the anterior portions of a Y-shaped endocarina (see ‘endocarina’ characters).

#28. Frontal arms <whether joined anteriorly>/

1. not joined anteriorly by transverse line/

2. joined anteriorly by transverse line/

The ‘frontal arms’ represent the distal or anterior portions of the ecdysial line known as the ‘epicranial suture’. These lines may be faint or absent in some instars, but sometimes they are reinforced by paired ‘endocarinae’ or the anterior portions of a Y-shaped endocarina (see ‘endocarina’ characters).

#29. Frontal arms <whether meeting mandibular articulations>/

1. not extending anteriorly to meet mandibular articulations, either incomplete or ending at antennal sockets/

2. extending anteriorly to meet mandibular articulations/

The ‘frontal arms’ represent the distal or anterior portions of the ecdysial line known as the ‘epicranial suture’. These lines may be faint or absent in some instars, but sometimes they are reinforced by paired ‘endocarinae’ or the anterior portions of a Y-shaped endocarina (see ‘endocarina’ characters).

#30. Bases of frontal arms <whether separated>/

1. contiguous/

2. distinctly separated/

The ‘frontal arms’ represent the distal or anterior portions of the ecdysial line known as the ‘epicranial suture’. These lines may be faint or absent in some instars, but sometimes they are reinforced by paired ‘endocarinae’ or the anterior portions of a Y-shaped endocarina (see ‘endocarina’ characters).

#31. Median endocarina <presence>/

1. absent/

2. present/

‘Endocarinae’ are internal ridges, which usually show up as distinct lines when light passes through the cuticle. The term is used here only for those internal ridges on the dorsal part of the head capsule. Ventral internal ridges include the ‘hypostomal rods’ (see ‘hypostomal rod’ characters) and sometimes paired structures associated with the tentorium. Median or paired endocarinae are not always pigmented and they may appear to be thickened portions of the epicranial suture.

#32. Median endocarina <position>/

1. absent or coincident with epicranial stem/

2. Y-shaped, coincident with epicranial stem and frontal arms/

3. moderately to very long, extending anterad of epicranial stem/

4. short or long, extending between frontal arms (epicranial stem absent)/

5. short or long, epicranial suture not evident/

‘Endocarinae’ are internal ridges, which usually show up as distinct lines when light passes through the cuticle. The term is used here only for those internal ridges on the dorsal part of the head capsule. Ventral internal ridges include the ‘hypostomal rods’ (see ‘hypostomal rod’ characters) and sometimes paired structures associated with the tentorium. Median or paired endocarinae are not always pigmented and they may appear to be thickened portions of the epicranial suture.

#33. Paired endocarinae <presence>/

1. absent/

2. present/

‘Endocarinae’ are internal ridges, which usually show up as distinct lines when light passes through the cuticle. The term is used here only for those internal ridges on the dorsal part of the head capsule. Ventral internal ridges include the ‘hypostomal rods’ (see ‘hypostomal rod’ characters) and sometimes paired structures associated with the tentorium. Median or paired endocarinae are not always pigmented and they may appear to be thickened portions of the epicranial suture.

#34. Paired endocarinae <position>/

1. absent or coincident with frontal arms/

2. located mesad of frontal arms/

‘Endocarinae’ are internal ridges, which usually show up as distinct lines when light passes through the cuticle. The term is used here only for those internal ridges on the dorsal part of the head capsule. Ventral internal ridges include the ‘hypostomal rods’ (see ‘hypostomal rod’ characters) and sometimes paired structures associated with the tentorium. Median or paired endocarinae are not always pigmented and they may appear to be thickened portions of the epicranial suture.

#35. Stemmata <presence>/

1. absent/

2. present/

In Coleoptera, there are 6 or fewer larval eyes or ‘stemmata’ (sometimes called ‘ocelli’), which may be large and strongly convex or reduced to eye spots. Stemmata should always be looked for in the living or preserved specimen before the head has been macerated and cleared, since the pigment often disappears during the clearing process.

#36. Number of stemmata/

1. one/

2. two/

3. three/

4. four/

5. five/

6. six/

7. zero/

In Coleoptera, there are 6 or fewer larval eyes or ‘stemmata’ (sometimes called ‘ocelli’), which may be large and strongly convex or reduced to eye spots. Stemmata should always be looked for in the living or preserved specimen before the head has been macerated and cleared, since the pigment often disappears during the clearing process.

#37. Number of stemmata <whether 6>/

1. fewer than six/

2. six/

In Coleoptera, there are 6 or fewer larval eyes or ‘stemmata’ (sometimes called ‘ocelli’), which may be large and strongly convex or reduced to eye spots. Stemmata should always be looked for in the living or preserved specimen before the head has been macerated and cleared, since the pigment often disappears during the clearing process.

#38. Number of antennal segments/

1. one/

2. two/

3. three/

4. four/

5. five or more/

Mistakes can be made when counting the number of antennal segments in a larva. The antenna is often borne on a membranous prominence which may be mistaken for a basal segment. Also the penultimate segment in beetle antennae bears a large sensory structure known as a ‘sensorium’ (see ‘sensorium’ characters), which may be mistaken for a terminal segment when the true terminal segment is absent or fused to that preceding it. Both the basal membrane and the sensorium differ from a true segment in lacking pigmentation, setae or sensilla. In some groups of weevils (Curculionoidea), the antenna consists of a membranous elevation (usually accompanied by a few setae and sensilla and a sensorium); this is considered to be 1-segmented.

#39. Ratio of antennal length to head width/

1. less than 0.15/

2. 0.15 to 0.50/

3. more than 0.50/

#40. Sensorium(a) on preapical antennal segment <presence>/

1. absent/

2. present/

3. apparently absent in antennae (usually 1- or 2-segmented) with apical sensorium/

The ‘sensorium’ is an enlarged sensillum usually located on the preapical segment of the antenna. It is a transparent vesicle, which varies in shape from conical or palpiform to flat and dome-like. When the apical segment is highly reduced or absent the sensorium appears to be located at the antennal apex. In this case, the specimen is coded as having state 3. The sensorium never bears setae or sensilla on its surface and thus may be distinguished from an apical antennal segment. See images for the other ‘sensorium’ characters.

#41. Sensorium(a) on preapical antennal segment <relative length>/

1. shorter than apical segment/

2. as long as or longer than apical segment/

#42. Sensorium(a) on preapical (or apical) antennal segment <shape>/

1. conical or palpiform/

2. dome-like/

3. bifid or complex/

#43. Apex of preapical antennal segment <whether oblique>/

1. truncate, so that sensorium and apical segment arise together/

2. oblique, so that sensorium arises proximad of apical segment/

#44. Antenna separated from mandibular articulation <distance>/

1. by strip of membrane or cuticle narrower than antennal socket/

2. by strip of cuticle between 1 and 2 times as wide as antennal socket/

3. by strip of cuticle at least 2 times as wide as antennal socket/

#45. Antennal base <whether with eversible lobe>/

1. without eversible membranous lobe/

2. with large, eversible, membranous lobe/

#46. Frontoclypeal suture <presence>/

1. absent or vaguely indicated/

2. distinct/

The ‘frontoclypeal suture’ (sometimes called the ‘epistomal suture’) is an impressed line between the frons and clypeus, which marks the position of an invagination called the ‘epistomal ridge’. Usually this ridge shows up as a dark line in cleared specimens, so that there appears to be a suture between the frons and clypeus. If the specimen is examined at an angle the true surface features may be ascertained. Taxa have been coded as 48,2 only when there is a distinct, externally visible impression.

#47. Labrum <attachment>/

1. separated from head capsule by complete suture/

2. partly fused to head capsule (suture incomplete)/

3. completely fused to head capsule (suture absent)/

#48. Labrum <whether with crenulate emargination>/

1. without crenulate emargination/

2. with crenulate emargination/

#49. Epipharyngeal rods <presence>/

1. absent/

2. present/

‘Epipharyngeal rods’, also known as ‘labral rods’, are paired longitudinal paramedian sclerites lying just beneath the surface of the epipharynx but easily seen in cleared specimens. They should not be confused with the ‘tormae’, which project posteriorly from the posterolateral corners of the labrum-epipharynx.

#50. Mouthparts <whether forming sucking tube>/

1. not forming sucking tube/

2. forming sucking tube containing stylets/

#51. Mandibles <whether asymmetrical>/

1. symmetrical/

2. asymmetrical/

Asymmetry here refers to obvious differences, especially with respect to the molar area. Some mandibles with minor differences in the positions of teeth on the incisor edge are coded as symmetrical.

#52. Mandibles <whether non-opposable>/

1. opposable, at least slightly curving mesally at apex/

2. non-opposable, straight or curving laterally at apex/

#53. Mandibles <whether forming diverging blades or stylets>/

1. not approximate, diverging and longitudinally divided/

2. approximate at base and divided longitudinally so that 2 pairs of diverging blades or stylets are formed/

#54. Mandibles <shape>/

1. absent or fused to head capsule/

2. narrow and falcate/

3. flattened and more or less triangular/

4. broad at base and narrow at apex/

5. broad and stout or more or less wedge-like/

6. stylet-like/

7. elongate, deeply cleft and with apical teeth directed both laterally and mesally/

#55. Mandibles <whether grooved or perforate>/

1. without groove or perforation/

2. with open or partly closed groove/

3. with internal perforation/

#56. Apex of mandible <orientation>/

1. narrow when viewed from in front (parallel or oblique to plane of movement)/

2. broad when viewed from in front (perpendicular to plane of movement)/

#57. Apex of mandible <number of lobes or teeth>/

1. with single lobe or tooth/

2. bilobed or bidentate/

3. trilobed or tridentate/

4. multilobed or multidentate/

#58. Accessory ventral process of mandible <presence>/

1. absent/

2. present/

This process is present only in mandibles with a mola and is located on the ventral surface of the mandibular base, just laterad of the molar area.

#59. Incisor edge of mandible <whether simple>/

1. simple/

2. with 1 or 2 subapical teeth/

3. serrate (sometimes in addition to retinaculum)/

4. with 1 or 2 heavily sclerotized retinacula/

5. with subapical pseudomola/

A ‘retinaculum’ is a relatively large and heavily sclerotized tooth or lobe on the cutting edge of the mandible; this is usually found only on those mandibles lacking a mola, but in Dascillidae and Scarabaeoidea a mola coexists with one or two heavy teeth coded here as retinacula (and sometimes known as ‘scissorial teeth’).

A ’pseudomola’ is a mola-like structure which is not considered to be homologus with the true basal mola. Obviously, the user of this key is not expected to make decisions on the homology of structures, and so the term is here restricted to a few obvious cases of mola-like structures located near the apex of the mandible.

#60. Mesal surface of mandibular base <whether simple>/

1. simple or slightly expanded/

2. with brush of hairs or spines/

3. with articulated process/

4. with fixed, rigid, hylaline process, sometimes partly sclerotized/

5. with 2 to 5 hyaline processes, sometimes joined basally/

6. with membranous lobe/

7. with mola or basal processing area/

8. with brush of hairs and fixed, hyaline process/

9. with brush of hairs and articulated process/

10. with articulated process and membranous lobe/

11. with finely pubescent sclerotized lobe/

A ‘mola’ is an enlargement and modification of the mesal surface of the mandibular base which meets a similar structure on the opposite mandible for the purpose of ingesting, breaking down or compacting food material. Some larvae may have a slightly expanded but otherwise unmodified mandibular base, which is not considered to be a mola.

#61. Mandibular mola <presence>/

1. absent/

2. present/

A ‘mola’ is an enlargement and modification of the mesal surface of the mandibular base which meets a similar structure on the opposite mandible for the purpose of ingesting, breaking down or compacting food material. Some larvae may have a slightly expanded but otherwise unmodified mandibular base, which is not considered to be a mola.

#62. Mola <armature at base>/

1. without hyaline lobe or brush at base/

2. with hyaline lobe at base/

3. with brush or hairs or spines at base/

4. with hyaline lobe and brush of hairs at base/

A ‘mola’ is an enlargement and modification of the mesal surface of the mandibular base which meets a similar structure on the opposite mandible for the purpose of ingesting, breaking down or compacting food material. Some larvae may have a slightly expanded but otherwise unmodified mandibular base, which is not considered to be a mola.

#63. Molar surface <whether simple>/

1. simple/

2. tuberculate or asperate/

3. with numerous fine ridges/

4. with few coarse ridges/

A ‘mola’ is an enlargement and modification of the mesal surface of the mandibular base which meets a similar structure on the opposite mandible for the purpose of ingesting, breaking down or compacting food material. Some larvae may have a slightly expanded but otherwise unmodified mandibular base, which is not considered to be a mola.

#64. Prostheca <presence>/

1. absent/

2. present/

A ‘prostheca’ is defined as a structure of variable form lying just distal to the mandibular mola, but not including projections of the mola itself or teeth or serrations lying on the cutting edge of the mandible just before the apex. The prostheca may be membranous, hyaline or partly sclerotized, or it may consist of a group of processes or hairs. By definition, the prostheca cannot be present unless a mola is also present. A basal mandibular structure in a mandible lacking a mola is often referred to as a ‘lacinia mobilis’; although such structures may resemble prosthecae, they are coded as separate features (see ‘mesal surface of mandibular base’ character).

#65. Prostheca <type>/

1. absent/

2. a rigid hyaline process, fixed or articulated, sometimes partly sclerotized/

3. complex, consisting of 2 or more hyaline processes, a brush of simple or complex hairs, or a series of fringed membranes/

4. a simple, membranous lobe/

5. a membranous lobe fringed with simple or complex hairs/

6. a fixed, rigid process and brush of hairs/

A ‘prostheca’ is defined as a structure of variable form lying just distal to the mandibular mola, but not including projections of the mola itself or teeth or serrations lying on the cutting edge of the mandible just before the apex. The prostheca may be membranous, hyaline or partly sclerotized, or it may consist of a group of processes or hairs. By definition, the prostheca cannot be present unless a mola is also present. A basal mandibular structure in a mandible lacking a mola is often referred to as a ‘lacinia mobilis’; although such structures may resemble prosthecae, they are coded as separate features (see ‘mesal surface of mandibular base’ character).

#66. Prostheca or basal mandibular process <shape>/

1. narrow with acute or narrowly rounded apex/

2. broad with obtusely angulate apex/

3. broad and rounded/

4. narrow with blunt apex/

5. narrow with bifid or multidentate apex/

6. broad at base and abruptly narrowed to acute apex/

A ‘prostheca’ is defined as a structure of variable form lying just distal to the mandibular mola, but not including projections of the mola itself or teeth or serrations lying on the cutting edge of the mandible just before the apex. The prostheca may be membranous, hyaline or partly sclerotized, or it may consist of a group of processes or hairs. By definition, the prostheca cannot be present unless a mola is also present. A basal mandibular structure in a mandible lacking a mola is often referred to as a ‘lacinia mobilis’; although such structures may resemble prosthecae, they are coded as separate features (see ‘mesal surface of mandibular base’ character).

#67. Prostheca or basal mandibular process <vestiture>/

1. glabrous/

2. generally pubescent/

3. pubescent or serrate along one edge/

4. pubescent at apex only/

A ‘prostheca’ is defined as a structure of variable form lying just distal to the mandibular mola, but not including projections of the mola itself or teeth or serrations lying on the cutting edge of the mandible just before the apex. The prostheca may be membranous, hyaline or partly sclerotized, or it may consist of a group of processes or hairs. By definition, the prostheca cannot be present unless a mola is also present. A basal mandibular structure in a mandible lacking a mola is often referred to as a ‘lacinia mobilis’; although such structures may resemble prosthecae, they are coded as separate features (see ‘mesal surface of mandibular base’ character).

#68. Ventral mouthparts <whether protracted>/

1. retracted/

2. strongly protracted/

It is the bases of the ventral mouthparts (cardines and postmentum or submentum) which are retracted or protracted and not necessarily the apices; sometimes the bases are retracted while the apices of the maxillae and labium appear to be protracted, due to an elongation of the stipes or a change in the orientation of the cardines (to strongly oblique or longitudinal).

#69. Cardo <presence>/

1. absent, indistinct or membranous/

2. distinct and sclerotized/

#70. Cardo <whether divided>/

1. undivided/

2. divided by internal ridge/

3. externally divided/

The cardo often appears to be divided into 2 parts (state 3) when it is not; this is caused by an internal division by a sclerotized ridge (state 2).

#71. Cardo <orientation>/

1. transverse to slightly oblique/

2. strongly oblique or longitudinal/

When the cardines are strongly oblique, the entire maxillolabial complex may be pushed forward and appear to be protracted (see ‘ventral mouthparts’ character).

#72. Cardines or maxillary bases <whether contiguous or fused together>/

1. separated from each other by labium/

2. closely approximate or contiguous, not separated by labium/

3. completely fused or combined with labium to form single plate/

#73. Stipes <length-width ratio>/

1. longer than wide/

2. wider than long/

#74. Maxillary articulating area <presence>/

1. absent/

2. present/

#75. Maxilla <number and articulation of lobes>/

1. without apical lobes/

2. with single, fixed mala/

3. with single, articulated mala/

4. with articulated galea and fixed lacinia/

5. with fixed galea and lacinia/

6. with articulated galea and lacinia/

The apical lobe or lobes of the maxilla begin at the point where the palp arises and extend distally (anteriorly) from that point. There may be two lobes: an outer (lateral) galea and an inner (mesal) lacinia; however when one of these is absent or when they are fused together, the resulting single lobe is called a ‘mala’. Rarely, there is no apical lobe at all, and the first palp segment arises from the end of the maxilla.

#76. Mala or galea <whether 2-segmented>/

1. consisting of a single segment/

2. 2-segmented/

#77. Apex of mala <whether cleft>/

1. simple, not cleft/

2. cleft or distinctly emarginate/

#78. Apex of mala or galea <shape>/

1. rounded or truncate/

2. falciform/

3. stylet-like/

4. trilobed/

#79. Apex of mala or galea <armature-vestiture>/

1. glabrous/

2. bearing simple seta or spine/

3. bearing single spur/

4. setose or spinose/

5. bearing comb-hairs or spatulate hairs/

6. bearing 2 or 3 spurs or bifid spur/

7. setose with 2 or 3 spurs or bifid spur/

#80. Inner apical angle of mala <whether with teeth or uncus>/

1. simple/

2. with one to several small teeth/

3. with distinct lobe or uncus/

An ’uncus’ is not always easy to distinguish from one or more teeth at the inner apical angle of the mala. In general, the uncus is larger than a tooth and separated from the rest of the apex by an emargination.

#81. Apex of lacinia <shape>/

1. rounded or truncate/

2. falciform/

#82. Apex of lacinia <armature-vestiture>/

1. simple and glabrous/

2. bearing single seta/

3. bearing single spur/

4. bearing 2 or 3 spurs or bifid spur/

5. setose or spinose/

#83. Number of segments in maxillary palp/

1. one/

2. two/

3. three/

4. four/

5. five/

6. zero/

Errors in determining the number of maxillary palp segments usually occur because the palp-bearing process or ‘palpifer’ is mistaken for a basal palp segment. As a rule, the palpifer lacks a line of demarcation at its base and has no inner edge.

#84. First segment of maxillary palp <whether with digitiform appendage>/

1. without digitiform appendage/

2. with digitiform appendage/

The appendage on the first palp segment in hydrophilids is often considered to be a displaced galea, but this interpretation is not used here.

#85. Labium <subdivision>/

1. consisting of single plate/

2. consisting of prementum and postmentum/

3. consisting of prementum, mentum and submentum/

#86. Mentum, postmentum or labial plate <degree of freedom>/

1. completely free or basally connate with maxillae/

2. completely or almost completely connate with maxillae/

3. partly or completely fused with maxillae (sutures incomplete or absent)/

#87. Postmentum or labial plate <whether subtriangular>/

1. subquadrate or trapezoidal, with blunt posterior edge/

2. subtriangular, with acute posterior edge/

#88. Postmentum or labial plate <whether divided longitudinally>/

1. not divided longitudinally/

2. divided longitudinally into 3 parts/

#89. Ligula <relative length>/

1. absent/

2. shorter than labial palp/

3. as long as or longer than labial palp/

The ‘ligula’ is a prominence or elevation of the prementum in between the labial palps. A prominence lying behind the palps and separated from them by a short distance is not considered to be a ligula, but is usually termed a ‘prehypopharynx.’ In those cases where the palps are located posterior to the apex of the prementum, the entire apical area is considered to be the ligula.

#90. Ligula <whether bilobed>/

1. simple, not bilobed/

2. bilobed/

The ‘ligula’ is a prominence or elevation of the prementum in between the labial palps. A prominence lying behind the palps and separated from them by a short distance is not considered to be a ligula, but is usually termed a ‘prehypopharynx.’ In those cases where the palps are located posterior to the apex of the prementum, the entire apical area is considered to be the ligula.

#91. Ligula <whether with sclerome>/

1. without tooth-like sclerome/

2. with tooth-like sclerome/

The ‘ligula’ is a prominence or elevation of the prementum in between the labial palps. A prominence lying behind the palps and separated from them by a short distance is not considered to be a ligula, but is usually termed a ‘prehypopharynx.’ In those cases where the palps are located posterior to the apex of the prementum, the entire apical area is considered to be the ligula.

#92. Number of segments in labial palp/

1. one/

2. two/

3. three/

4. four/

5. zero/

Errors in counting the number of labial palp segments usually occur because the palp-bearing process or ‘palpiger’ is mistaken for a basal palp segment. As a rule, the palpiger lacks a line of demarcation at its base and has no inner edge.

#93. Labial palps <degree of separation>/

1. contiguous or separated by less than width of first palp segment/

2. separated by more than width of first palp segment/

#94. Hypopharyngeal bracon <presence>/

1. absent/

2. present/

The ‘hypopharyngeal bracon’ is a narrow cuticular bar (often lightly pigmented) which extends between the lower mandibular articulations and behind the labium to which is also attached. It is most prominent in larvae with retracted ventral mouthparts for which it forms a brace, but it is noticeably absent in members of the Chrysomelidae.

#95. Hypopharyngeal sclerome/

1. absent/

2. consisting of transverse bar only/

3. consisting of a single sclerite shaped like a molar tooth (usually irregularly concave)/

4. divided into 2 parts or consisting of more than 1 sclerite/

The ‘hypopharyngeal sclerome’ is not always easy to see, because it lies within the mouth cavity. Sometimes removing the labrum-epipharyx allows the sclerome to be seen from above. It is almost always present in some form when the mandibles have well-developed molae and rarely present (and then only as a sclerotized bar) when the molae are absent.

#96. Hypostomal region <presence>/

1. absent (labium contiguous with thoracic membrane)/

2. present (separating labium from thorax)/

The ‘hypostomal region’ includes the gula when the latter is delimited by distinct sutures.

#97. Hypostomal rods <presence>/

1. absent/

2. present (1 pair)/

3. present (2 pairs)/

‘Hypostomal rods’ are sclerotizations extending posteriorly from the posterolateral portions of the hypostomal cavity (just lateral to the attachments of the cardines). These rods usually diverge posteriorly, but they may be parallel and are rarely convergent.

#98. Hypostomal rods <length>/

1. very short/

2. moderately long but not extending to posterior edge of head/

3. extending to posterior edge of head/

4. long with 1 pair extending to posterior edge of head/

‘Hypostomal rods’ are sclerotizations extending posteriorly from the posterolateral portions of the hypostomal cavity (just lateral to the attachments of the cardines). These rods usually diverge posteriorly, but they may be parallel and are rarely convergent.

#99. Hypostomal rods <direction>/

1. subparallel/

2. diverging posteriorly/

3. converging posteriorly/

4. with 1 pair diverging and another subparallel or slightly converging/

‘Hypostomal rods’ are sclerotizations extending posteriorly from the posterolateral portions of the hypostomal cavity (just lateral to the attachments of the cardines). These rods usually diverge posteriorly, but they may be parallel and are rarely convergent.

#100. Ventral epicranial ridges <presence>/

1. absent/

2. present/

‘Ventral epicranial ridges’ extend posteriorly from the ventral mandibular articulations and tend to form a housing for the maxillolabial complex. They should not be confused with ‘hypostomal rods’, which are sclerotizations, not ridges, and begin posteriorly near the attachments of the cardines.

#101. Gula <presence, length-width ratio>/

1. absent/

2. wider than long/

3. longer than wide/

#102. Gular sutures <presence>/

1. absent/

2. separate/

3. partly or completely fused/

#103. Gula or gular region <whether fused to labium>/

1. fused to labium/

2. separated from labium by suture/

#104. Occipital foramen <whether divided>/

1. not divided by tentorial bridge/

2. divided into 2 parts by tentorial bridge/

The occipital foramen appears to be divided by the tentorial bridge because the latter lies almost on the same plane with it, rather than being well within the head capsule.

Thorax #105. Prothorax <whether wider than abdomen>/

1. not or only slightly wider than abdomen/

2. much wider than abdomen/

#106. Prothorax <whether longer than meso- and metathorax combined>/

1. not longer than meso- and metathorax combined/

2. longer than meso- and metathorax combined/

#107. Thoracic terga <whether with patches of asperities>/

1. without patches of asperities/

2. with patch(es) of asperities on metathorax only/

3. with patch(es) of asperities on meso- and metathorax/

4. with patch(es) of asperities on prothorax only/

5. with patches of asperities on all segments/

#108. Thoracic terga <whether with rows of asperities>/

1. without rows of asperities or carinae/

2. with transverse row(s) of asperities (sometimes forming carinae) on metathorax only/

3. with transverse row(s) of asperities (sometimes forming carinae) on meso- and metathorax/

4. with transverse row(s) of asperities (sometimes forming carinae) on prothorax only/

#109. Protergum <whether with sclerotized plates>/

1. without sclerotized plates/

2. with 1 or more sclerotized plates/

#110. Meso- and metaterga <whether with sclerotized plates>/

1. without sclerotized plates/

2. with 1 or more sclerotized plates/

#111. Thorax <whether with lateral processes>/

1. without well-developed lateral tergal processes/

2. with well-developed lateral tergal processes/

#112. Prosternum <armature>/

1. without armature/

2. with patch of asperities/

3. with 2 or more patches of asperities/

4. with single, median tooth/

5. with rod-like sclerotization/

6. with patch of asperities and rod-like sclerotisation/

7. with transverse rows of asperities or carinae/

#113. Mesocoxae or coxal lobes <degree of separation>/

1. separated by less than 1 basal coxal diameter/

2. separated by 1 to 2 basal coxal diameters/

3. separated by more than 2 basal coxal diameters/

#114. Thoracic legs <presence>/

1. absent or represented by non-articulated lobes or processes/

2. articulated/

In a few instances the thoracic legs are highly reduced and only visible under high magnification; however, these legs are obviously segmented and usually end in a claw. In some curculionoids there are rounded, unsegmented prominences, which have a basal line of demarcation and are thus considered to be 1-segmented legs.

#115. Number of segments in mesothoracic leg/

1. one/

2. two/

3. three/

4. four/

5. five including pretarsus (claw)/

6. six including pretarsus (claw or claws)/

7. zero/

The ‘pretarsus’ or ‘claw’ in the 5-segmented legs of Myxophaga and most Polyphaga is considered by many workers to be a ‘tarsungulus’ (fused tarsus and pretarsus). In this case, the preceding segment would be considered a ‘tibia’, rather than a ‘tibiotarsus’(fused tibia and tarsus).

#116. Number of segments in mesothoracic leg <whether 6>/

1. five or fewer including pretarsus (claw)/

2. six including pretarsus (claw or claws)/

The ‘pretarsus’ or ‘claw’ in the 5-segmented legs of Myxophaga and most Polyphaga is considered by many workers to be a ‘tarsungulus’ (fused tarsus and pretarsus). In this case, the preceding segment would be considered a ‘tibia’, rather than a ‘tibiotarsus’(fused tibia and tarsus).

#117. Legs <whether highly reduced>/

1. not highly reduced, more than 0.2 times as long as head width/

2. highly reduced, less than 0.2 times as long as head width/

#118. Metathoracic leg <whether reduced relative to mesothoracic leg>/

1. about same size as mesothoracic leg/

2. much smaller than mesothoracic leg/

#119. Meso- and metathoracic legs <whether forming stridulatory organ>/

1. not forming stridulatory organ/

2. forming stridulatory organ/

#120. Apical segment of leg <whether rounded>/

1. rounded or truncate, not acute and claw-like/

2. acute and claw-like (or consisting of 2 claws)/

#121. Number of movable pretarsal claws/

1. one/

2. two/

#122. Number of setae on pretarsus/

1. zero/

2. one/

3. two/

4. three/

5. four or more/

#123. Pretarsal setae <disposition>/

1. lying side by side or obliquely situated/

2. lying in row, one distal to the other/

#124. Thoracic spiracles <type>/

1. annular/

2. annular-uniforous/

3. annular-biforous/

4. biforous or bilabiate/

5. cribriform/

6. undulate/

7. reduced and non-functional or absent/

8. annular-multiforous/

The ‘annular’ spiracle (state 1) has a simple opening, the edge of which (‘peritreme’) may be somewhat scalloped or uneven, but is never lined with a series of small accessory chambers as in the ‘annular-multiforous’ type (state 8). The ‘annular-uniforous’ and (‘unicameral’) and ‘annular-biforous’ (‘bicameral’) spiracles (states 2 and 3) have a simple main opening, with either one or two parallel and contiguous accessory openings connected to it. Sometimes these accessory openings are very short and can be missed under lower magnifications. In other cases, the openings are very long and almost as wide as the main opening; they then resemble ‘biforous’ spiracles (state 4) in which the main opening is completely blocked by a pair of narrow slit-like openings. In this last type there is always a spiracular scar lying near one end of the paired openings. In the ‘cribriform’ type of spiracle (state 5), the opening is partly or completely blocked by a ‘sieve plate’ containing a number of small openings. The undulate type of spiracle (state 6) is merely a variation of the biforous type in which the paired openings are undulate.

#125. Thoracic spiracles <whether at ends of tubes>/

1. not placed at ends of spiracular tubes/

2. placed at ends of lateral spiracular tubes/

3. placed at ends of dorsal spiracular tubes/

Abdomen #126. Ratio of length of abdomen (without appendages) to length of thorax/

1. less than 1.2/

2. 1.2 to 2.0/

3. more than 2.0/

#127. Number of visible abdominal segments/

1. ten/

2. nine/

3. eight/

4. seven or fewer/

Remember, this is the total number of abdominal segments which can be seen from any angle, not the number seen from above. States 2 to 4 only occur when two or more of the terminal segments are fused together or when the terminal segment is completely membranous and concealed by the meeting of the tergum and sternum of the preceding segment.

#128. Abdominal terga <whether extending laterally beyond sterna>/

1. not extending laterally beyond edges of sterna/

2. extending laterally beyond edges of sterna/

#129. Abdominal terga <whether with patches of asperities>/

1. without patches of asperities/

2. with patches of asperities on 1 or more segments/

#130. Abdominal terga <whether with rows of asperities>/

1. without rows of asperities/

2. with rows of asperities on 1 or more segments/

#131. Rows of abdominal tergal asperities <orientation>/

1. transverse/

2. longitudinal/

#132. Rows of abdominal tergal asperities <type>/

1. single/

2. double/

3. forming open or closed circles/

#133. Rows of abdominal tergal asperities <number per segment>/

1. one per segment/

2. two per segment, one on each side of midline/

3. two per segment, anterior and posterior/

#134. Abdomen <whether with lateral processes>/

1. without well-developed lateral processes/

2. with 1 pair of lateral processes on most segments/

3. with 2 pairs of lateral processes (dorsal and ventral) on most segments/

#135. Abdominal tergum 3 <whether with dorsal transverse folds or lobes>/

1. without transverse dorsal folds or lobes/

2. with 2 or more transverse dorsal folds or lobes/

A ‘fold’, as used here, refers to a convexity, not a groove or furrow. Thus an abdominal segment either has no folds, two folds separated by a furrow, 3 separated by 2 furrows etc.

#136. Abdominal tergum 3 <number of transverse folds or lobes>/

1. simple, not divided into transverse dorsal folds or lobes/

2. divided into 2 transverse dorsal folds or lobes/

3. divided into 3 transverse dorsal folds or lobes/

4. divided into 4 or more transverse dorsal folds or lobes/

A ‘fold’, as used here, refers to a convexity, not a groove or furrow. Thus an abdominal segment either has no folds, two folds separated by a furrow, 3 separated by 2 furrows etc.

#137. Paired abdominal glands (openings or porous plates) <presence>/

1. absent/

2. present on segments 1 to 7/

3. present on segments 1 and 8/

4. present on segments 1 to 8 or 9/

5. present on apical or subapical segment only/

#138. Dorsal, balloon-like tracheal gills <presence>/

1. absent/

2. present on prothorax and abdominal segments 1 and 8/

3. present on abdominal segments 1 to 8/

#139. Long and narrow, lateral gills <presence>/

1. absent/

2. present on first 7 abdominal segments/

3. present on first 8 abdominal segments/

4. present on first 9 abdominal segments/

#140. Abdominal sterna <whether with patches of asperities>/

1. without patches of asperities/

2. with patches of asperities on 1 or more segments/

#141. Abdominal sterna <whether with rows of asperities>/

1. without rows of asperities/

2. with rows of asperities on 1 or more segments/

#142. Paired ventral prolegs (or asperity-bearing ampullae) <presence>/

1. absent/

2. present on abdominal sterna 3 and 4, 2 to 4 or 2 to 5/

3. present on abdominal sterna 3 to 7/

#143. Ventral abdominal gill tufts <presence>/

1. absent/

2. present on segments 1 to 7/

3. present on segments 2 to 6/

4. present on segments 1 to 4/

5. present on segments 1 to 3/

#144. Abdominal tergum 8 <special armature>/

1. without special armature/

2. with single median process, simple at apex/

3. with single median process bearing spiracles at apex/

4. with paired processes, simple at apex/

5. with paired processes, each bearing spiracle at apex/

6. with respiratory chamber/

The spiracular openings within the respiratory chamber (state 6) may not be visible externally, but in cleared specimens the enlarged tracheal trunks leading to them are fairly obvious. See image for ‘ventral abdominal gill tufts’ character.

#145. Abdominal apex <presence of respiratory chamber>/

1. without respiratory chamber/

2. with respiratory chamber formed from 8th and 9th terga and enclosing pair of enlarged spiracles/

The spiracular openings within the respiratory chamber may not be visible externally, but in cleared specimens the enlarged tracheal trunks leading to them are fairly obvious.

#146. Abdominal segment 9 excluding appendages <length relative to 8>/

1. shorter than segment 8/

2. as long as or slightly longer than segment 8/

3. much longer than segment 8/

#147. Abdominal tergum 9 <whether forming articulated plate>/

1. not forming articulated plate/

2. forming articulated plate/

#148. Abdominal tergum 9 <whether extending onto ventral surface>/

1. completely dorsal/

2. extending onto ventral surface/

3. completely ventral/

#149. Abdominal tergum 9 <whether with paired processes or urogomphi>/

1. without paired processes or urogomphi/

2. with paired processes or urogomphi/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#150. Abdominal tergum 9 <whether with pits between urogomphi>/

1. without pit between urogomphi/

2. with single pit between urogomphi/

3. with 2 pits between urogomphi/

#151. Abdominal tergum 9 <whether with median process or terminal disc>/

1. without median process or terminal disc/

2. with median process/

3. with terminal disc/

In those scraptiids which have a median process (large and rounded), it often breaks off. In this case the apex of tergum 9 is rather abruptly truncated.

#152. Median process on tergum 9 <shape>/

1. acute or narrowly truncate/

2. large, round and dehiscent/

3. bifurcate/

#153. Ratio of length of urogomphus to basal width of tergum 9/

1. less than 0.2/

2. between 0.2 and 1.0/

3. more than 1.0/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#154. Urogomphi <whether articulated at base>/

1. fixed at base/

2. articulated at base/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#155. Urogomphi <whether segmented>/

1. unsegmented/

2. segmented/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#156. Urogomphi <whether bifurcate or with accessory processes>/

1. not bifurcate and without accessory processes/

2. bifurcate or with accessory processes/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#157. Urogomphi <orientation>/

1. posteriorly oriented/

2. dorsally or posterodorsally oriented/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#158. Urogomphi separated by <distance>/

1. less than 1 basal width/

2. between 1 and 2 basal widths/

3. more than 2 basal widths/

‘Urogomphi’ are paired processes located on the 9th tergum, or the most obvious pair when there are several pairs. They are not all considered to be homologous structures, and they represent numerous parallel developments. When smaller paired process occur in front of the urogomphi, they are called ‘pregomphi’.

Occasionally tergum 8 has a pair of long processes which appear to be urogomph but actually belong to the preceding segment (see ‘paired ventral prolegs’ character). The abdominal segments should be counted to be sure.

#159. Abdominal sternum 9 <whether concealed or apparently absent>/

1. completely concealed or apparently absent/

2. partly or entirely exposed/

#160. Abdominal sternum 9 <armature>/

1. simple or apparently absent/

2. with row of apical asperities/

3. with 2 apicomesal asperities/

4. with 1 basal asperity on each side/

5. with 2 to 6 basal asperities on each side/

6. with more than 6 basal asperities on each side/

7. U-shaped with apicolateral horns or teeth/

8. with 2 basal and at least 2 apical asperities/

#161. Abdominal sternum 9 <whether enclosed by sternum 8>/

1. not enclosed by sternum 8/

2. partly or almost completely enclosed by sternum 8/

#162. Row of basal asperities on sternum 9 <whether curved posteriorly>/

1. straight or very slightly curved posteriorly/

2. strongly curved posteriorly at sides/

3. strongly, doubly curved/

#163. Row of basal asperities on sternum 9 <whether interrupted>/

1. more or less continuous or very narrowly interrupted at middle/

2. broadly interrupted at middle/

#164. Abdominal apex <whether with hinged operculum>/

1. without hinged operculum/

2. with dorsally hinged operculum/

3. with ventrally hinged operculum/

#165. Abdominal segment 10 <whether with oval lobes>/

1. without oval lobes separated by longitudinal groove/

2. with pair of oval lobes separated by longitudinal groove/

#166. Abdominal segment 10 <development>/

1. concealed from above, sometimes membranous or more or less fused to segment 9/

2. distinct and visible from above/

#167. Abdominal segment 10 <whether with asperated holdfast organs>/

1. without asperated, tubular holdfast organs/

2. with several asperated, tubular, protrusible holdfast organs/

#168. Paired pygopods on segment 10 <presence and elongation>/

1. absent/

2. shorter than wide/

3. longer than wide/

#169. Anal region <orientation>/

1. posteriorly or terminally oriented/

2. posteroventrally oriented/

3. ventrally oriented/

#170. Anal gill tufts <presence>/

1. absent/

2. present/

An ‘anal gill tuft’ consists of several fine gill filaments joined to a common base. They may be confused with a group of anal papillae, each of which is a single membranous evagination.

#171. Anal hooks <number>/

1. absent/

2. 1 on each side/

3. 2 on each side/

4. 3 or more on each side/

#172. Anterior abdominal spiracles <type>/

1. annular/

2. annular-uniforous/

3. annular-biforous/

4. biforous or bilabiate/

5. cribriform/

6. undulate/

7. reduced and non-functional or absent/

8. annular-multiforous/

The ‘annular’ spiracle (state 1) has a simple opening, the edge of which (‘peritreme’) may be somewhat scalloped or uneven, but is never lined with a series of small accessory chambers as in the ‘annular-multiforous’ type (state 8). The ‘annular-uniforous’ and (‘unicameral’) and ‘annular-biforous’ (‘bicameral’) spiracles (states 2 and 3) have a simple main opening, with either one or two parallel and contiguous accessory openings connected to it. Sometimes these accessory openings are very short and can be missed under lower magnifications. In other cases, the openings are very long and almost as wide as the main opening; they then resemble ‘biforous’ spiracles (state 4) in which the main opening is completely blocked by a pair of narrow slit-like openings. In this last type there is always a spiracular scar lying near one end of the paired openings. In the ‘cribriform’ type of spiracle (state 5), the opening is partly or completely blocked by a ‘sieve plate’ containing a number of small openings. The undulate type of spiracle (state 6) is merely a variation of the biforous type in which the paired openings are undulate.

#173. Abdominal spiracles <whether at ends of tubes>/

1. not placed at ends of spiracular tubes/

2. all placed at ends of lateral spiracular tubes/

3. on segment 8 placed at ends of lateral spiracular tubes/

4. on segments 1 and 8 placed at ends of lateral spiracular tubes/

#174. Spiracular tubes or processes on abdominal segment 3 <elongation>/

1. shorter than wide/

2. between 1 and 2 times as long as wide/

3. more than 2 times as long as wide/

#175. Eighth abdominal spiracles <whether reduced, absent or enlarged>/

1. functional and about the same size as others on abdomen/

2. reduced or absent/

3. much larger than others on abdomen/

4. only functional pair on abdomen/

#176. Abdominal spiracle 8 <position>/

1. lateral or dorsolateral, facing laterally/

2. dorsal, often near midline, facing posteriorly/

Distribution #177. Geographical distribution <whether including America north of Mexico>/

1. not including America north of Mexico/

2. including America north of Mexico/

#178. Geographical distribution <whether including Australia>/

1. not including Australia/

2. including Australia/

#179. Geographic distribution <whether including New Zealand>/

1. not including New Zealand/

2. including New Zealand/

#182. <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. 1995 onwards. Elateriformia (Coleoptera) larvae: descriptions, illustrations, identification, and information retrieval for families and subfamilies. Version: 9th October 2005. http://delta-intkey.com’.

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