Ichnea Laporte, 1836

Opitz, Weston, 2010, Classification, Natural History, and Evolution of the Epiphloeinae (Coleoptera: Cleridae). Part VIII. The Genera Acanthocollum Opitz, Stegnoclava Opitz, and Ichnea Laporte, The Coleopterists Bulletin (mo 9) 64, pp. 1-65 : 17-24

Ichnea Laporte, 1836

Ichnea Laporte, 1836: 55 . Type species: Ichnea lycoides Laporte, 1836: 55 . By monotypy. Spinola 1841: 75, 1844: 20; Lacordaire 1857: 476; Chenu and Desmarest 1857: 265; Gemminger and Harold 1869: 1751; Chevrolat 1876: 7; Gorham 1877: 409, 1883: 178; Lohde 1900: 100; Schenkling 1903: 101, 1910: 122; Gahan 1910: 68; Corporaal 1942: 142, 1950: 270; Blackwelder 1945: 389; Ekis 1975: 49; Opitz 1997: 57, 2002: 278.

Diagnosis. The synapotypic character that distinguishes members of this epiphloeine genus is a triangular, depigmented mark on the pygidium ( Fig. 127 ). Other characteristics that may be useful in distinguishing these beetles involve the antenna ( Fig. 45 ), eyes ( Figs. 63, 64 ), color patterns ( Figs. 52–60), and form of the elytra. The scape is short, the funicular antennomeres are transverse, the antennal capitulum is greatly extended ( Fig. 45 ), and the eyes are prominent and broadly notched in front ( Fig. 63 ). The elytra are finely punctate with punctations seriate in elytral basal half. Moreover, they are profusely setose and together with the forebody approximate a triangular or narrow subovoid body form.

Description. Size: Length 4.5–12.0 mm; width 1.7–4.0 mm. Form: Narrowly triangular or oblong subovoid; elytra gradually widened to posterior fourth or subconvex to posterior forth. Integumental color: Antennae unicolored or bicolored ( Figs. 52–60), if unicolored dark brown, if bicolored most antennomeres dark brown but apex yellow; cranium usually bicolored, yellow with dark brown infuscations, rarely unicolored, when unicolored yellow; pronotum usually bicolored, most often disc with dark medial stripe, yellow at upper sides, and dark in lower sides, rarely completely yellow or roseate; elytral disk most often with combination of yellow and dark maculae or dark streaks, sometimes entirely yellow and nearly entirely black or dark brown, with or without pale

40) Stegnoclava zorropsis ; 41) Ichnea acanthomelina 42) Ichnea atra ; 43–44) Ichnea aequinoctialis .

line along epipleural and sutural margins; legs usually bicolored, with basal half of femora yellow and rest of legs black or dark brown, or femora yellow and rest of legs brown; pterothorax and abdomen piceous or testaceous. Vestiture: Integument highly setose, cranium and pronotum often densely vested with decumbent setae, elytra densely vested with short secondary setae and less profusely distributed primary setae, latter particularly prominent along epipleural margin and elytral distal fourth.

59) Ichnea gremia ; 60) Ichnea divisa .

64) Head of I. marginella .

Head ( Figs. 46 , 63, 64 ): Cranial setose punctures small, frons indented ( Fig. 76 ) or not; eyes prominently bulged, finely faceted ( Fig. 98 ) and deeply, broadly incised along frontal margin; cranial postocular setose punctures with globose elevations at base ( Fig. 99 ); antenna ( Fig. 45 ) capitate, inserted at lower angle of eye incision (= ocular notch, Fig. 63 ), 10-segmented, scape shorter than length of funicular antennomeres (3–7) combined ( Fig. 66 ), antennomeres 3–7 expanded laterally, antennomere 8–10 form a large capitulum about three-fourths length of antenna, capitular antennomeres with well developed sensilla chaetica and sensilla basonica ( Fig. 100 ); labrum ( Fig. 83 ) deeply incised, transverse tormal processes horizontal and contiguous; mandible ( Fig. 86 ) falciform, dens well developed, mandibular penicillus reduced; maxilla ( Fig. 87 ) distinctly developed, maxillary palpomere 4 tapered distally, laterolacinia present; labium ( Fig. 84 ) distinctly developed, palpomere 3 tapered distally; gula ( Fig. 46 ) triangular; cranium notably indented near posterior margin of eye ( Figs. 98 , 112 ). Thorax: Pronotum usually slightly elongate ( Fig. 62 ), slightly transverse ( Fig. 47 ) or quadrate ( Fig. 131 ), discal setose punctures small, lateral margins subparallel ( Fig. 102 ) or sinuous ( Fig. 61 ); anterior transverse depression absent, posterior transverse depression prominent, pronotal collar very narrow, discal and lateral trichobothria ( Figs. 49 , 77 , 113 ) prominent, bothria domed, pronotal projection ( Fig. 48 ) only slightly extended towards the middle, lateral carina not extended to anterior margin of pronotum and posteriorly confluent with pronotal hem ( Figs. 49– 51 ); elytra broadened to distal four-fifths, then gradually rounded to apex, about 5 times longer than pronotum, 2–6 times longer than broad, epipleuron well developed ( Fig. 79 ), notably explanate, punctations small, subseriate at basal half, much smaller than width of interstitial spaces ( Fig. 105 ); primary and secondary elytral setae present ( Fig. 101 ); mesoscutellum subglobose; protibial anterior margin with 9–14 spines ( Fig. 78 ); tibial spur formula 0-1-1; tarsal pulvillus formula 3-3-1; metathoracic wing as in Fig. 80 ; metendosternite without furcal lamina ( Fig. 82 ). Abdomen: Six visible sterna; pygidium with triangular depigmented notch, posterior margin evenly convex in females, slightly incised in males. Male genitalia: Aedeagus narrow, long ( Fig. 176 ), parameres usually reduced, phallobasic and phallic apodemes expanded distally; spicular fork apodeme divided in anterior half, interspicular plate slender and bifid distally. Female genitalia: Ovipositor of moderate length; dorsal lamina comprised of five lobes; ventral lamina trilobed. Alimentary canal ( Fig. 88 ): Proventriculus moderately developed; ventriculus tapered at extremities, with poorly developed papillae; stomodaeal valve ( Fig. 81 ) with four primary lobes, dorsal and ventral lobes short, dorsal lobe very broad; four cryptonephridial malpighian tubules. Male mesodermal internal reproductive organs ( Fig. 114 ): Two pairs of accessory glands; testis comprised of 12–22 follicles. Female mesodermal internal reproductive organs ( Fig. 115 ): Spermathecal capsule not visibly sclerotized; spermathecal gland attached near apex of spermathecal capsule; bursa copulatrix saccular.

Distribution. The members of this genus range from southwestern United States to northeastern Argentina.

KEY TO THE SPECIES- GROUPS AND SPECIES

OF ICHNEA

1. Body form narrowly triangular ( Fig. 44); usually pronotum and elytral disc with various combinations of yellowish and black colorations....................................................2

1′. Body form not narrowly triangular, elytral epipleural margins parallel or slightly convex, more suboval ( Figs. 42, 58); pronotum and elytral disc brownish or various degrees of pale yellow, and elytral sutural and epipleural margins pale ..................................................11

2. Pronotum oblong ( Fig. 121 )........................3

2′. Pronotum broad trapezoidal or quadrate ( Fig. 122 ) ( lycoides species-group) .............9

3. Eyes nearly contiguous in dorsal view ( Fig. 126 ) ( aequinoctialis species-group)...4

3′. Eyes widely separated in dorsal view ( Fig. 125 ) ( aterrima species-group) ............5

4. Pronotum with triangular dark mark that extends from pronotal base, but does not extend to pronotal anterior margin ( Fig. 44) ( Honduras, Panamá, Colombia, Venezuela, Guyana, French Guiana, Peru, Ecuador, Bolivia, Brazil, Paraguay, Argentina) ( Figs.43, 44, 67, 68) ............................ Ichnea aequinoctialis Spinola

4′. Pronotum without triangular dark mark, broad dark vertical line extends from pronotal base to anterior pronotal margin ( Colombia, Bolivia, Brazil, Peru) ( Figs. 69, 70)................. .............. Ichnea callanga Opitz , new species

5. Basal half of elytral disc black, distal half yellowish ( Colombia) ( Fig. 75) ..................... .................... Ichnea dimidiatipennis Spinola

5′. Elytral color not as above............................ 6

6. Pronotum strongly constricted in anterior half ( Fig. 52).......................................................7

6′. Pronotum moderately constricted in anterior half ( Fig. 71)................................................8

7. Basal antennomere of antennal capitulum shorter than combined length of funicular antennomeres ( Peru) ( Fig. 52) ......................... ................ Ichnea gregata Opitz , new species

7′. Basal antennomere of antennal capitulum as long as or longer than combined length of funicular antennomeres ( Brazil, Argentina) ( Figs. 53–55) .......... Ichnea praeusta (Klug)

8. Pronotum usually with a centrally located broad yellow vertical line flanked by black, or pronotum entire black ( USA, Mexico, Guatemala, Belize, Honduras, Nicaragua, Costa Rica, Panama, Venezuela, Colombia, Ecuador) ( Figs. 71–74)............. Ichnea aterrima (Klug)

8′. Pronotum with centrally located broad black vertical line that may be incised by yellow streak at anterior pronotal margin ( Peru, Bolivia) ( Fig. 56) ........................................... .......................... Ichnea procera Schenkling

9. Pronotum not strongly trapezoidal, nearly quadrate ( Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Peru, Ecuador, Bolivia, Brazil) ( Figs. 90–94) ........................................ ............................. Ichnea frenata (Erichson)

9′. Pronotum strongly trapezoidal...................10

10. Basal antennomere of antennal capitulum about as long as combined length of funicular antennomeres ( Fig. 169 ); elytral disc with yellow fascia at middle ( Brazil) ( Figs. 106 , 133 ). ................................. Ichnea mimica Gorham

10′. Basal antennomere of antennal capitulum shorter than combined length of funicular antennomeres ( Fig. 170 ); elytral disc without yellow fascia at middle ( Bolivia, Brazil) ( Figs. 95–97)......... Ichnea lycoides Laporte

11. Frons planar, not indented with a deep pit .......................................................................12

11′. Frons not planar, widely concave or indented with a deep pit...........................................15

12. Eyes nearly contiguous in dorsal view, interocular width narrower than width of scape ( marginella species-group) ( Mexico, Guatemala, Belize, Honduras, Costa Rica, Panama, Colombia, Venezuela, Trinidad, Guyana, Suriname, French Guiana, Ecuador, Brazil, Peru) ( Fig. 107 )..................................... ............................... Ichnea marginella (Klug)

12′. Eyes not nearly contiguous in dorsal view, interocular width as broad as or broader than width of scape............................................13

13. Interocular width in dorsal view much broader than width of scape ( plumbea species-group) ( Panama, Trinidad, Colombia, Ecuador, Peru, Bolivia, Brazil) ( Figs. 108, 109 )............ ............................... Ichnea plumbea Gorham

13′. Interocular width in dorsal view about as wide as width of scape ( digna species-group)....14

14. Sutural and epipleural margins only briefly yellow near middle of elytral disc ( Ecuador) ( Fig. 59) ......................................................... .............. Ichnea gremia Opitz , new species

14′. Sutural and epipleural margins yellow throughout their length, or elytral disc entirely dark brown ( Costa Rica, Panama, Colombia) ( Figs. 57, 58).................. Ichnea digna Wolcott

15. Pronotum roseate.......................................16

15′. Pronotum not roseate.................................17

16. Broad suboval species; pronotum strongly transverse; pronotum with pink lateral spots ( Guyana, French Guiana, Brazil) ( Fig. 89) ... .................................... Ichnea opaca (Klug)

16′. Slender species; pronotum quadrate; pronotum without pink lateral spots ( Panama, Colombia, Venezuela, Bolivia, Ecuador, Peru, Paraguay. Brazil) ( Fig. 60)....... Ichnea divisa Chevrolat

17. Pronotum yellow or sides yellow and disc infuscated ( Brazil) ( incerta species-group) ( Figs. 110, 111 ) ...... Ichnea incerta Gorham

17′. Pronotum mostly black ( acanthomelina species-group)............................................18

18. Pronotum densely vested with golden setae ( Brazil) ( Fig. 41) ............................................ ... Ichnea acanthomelina Opitz , new species

18′. Pronotum not densely vested with golden setae ( Guyana, Brazil, Suriname) ( Fig. 42)... ................... Ichnea atra Opitz , new species

CLAVE PARA LOS GRUPOS DE ESPECIES Y LAS

ESPECIES DE ICHNEA

1. Cuerpo de forma triangular ( Fig. 44); pronoto y discos elitrales generalmente con alguna combinación de negro y amarillo.................2

1′. Cuerpo de forma más ovalada ( Fig. 58); pronoto y discos elitrales generalmente café o con variaciones de amarillo pálido, sutura elitral y márgenes epipleurales pálidas.... 11

2. Pronoto ovalado ( Fig. 121 )..........................3

2′. Pronoto trapezoidal o cuadrado ( Fig. 122 ) (grupo de especies lycoides )........................9

3. Ojos casi contiguos en vista dorsal ( Fig. 126 ) (grupo de especies aequinoctialis )................. 4

3′. Ojos bien separados en vista dorsal ( Fig. 125 ) (grupo de especies aterrima ) ......................... 5

4. Pronoto con una marca triangular oscura que se extiende desde la base del pronoto y no alcanza el margen anterior del pronoto ( Fig. 44) ( Honduras, Panamá, Colombia, Venezuela, Guyana, Guyana Francesa, Peru, Ecuador, Bolivia, Brasil, Paraguay, Argentina) ( Figs. 43, 44, 67, 68)......................................... ......................... Ichnea aequinoctialis Spinola

4′. Pronoto sin marca triangular oscura, con una banda oscura que se extiende desde la base del pronoto hasta el margen anterior ( Colombia, Bolivia, Brasil, Perú) ( Figs. 69, 70)............... ......... Ichnea callanga Opitz , especie nueva

5. Mitad basal del disco elitral negra, mitad distal amarillenta ( Colombia) ( Fig. 75).............. .................... Ichnea dimidiatipennis Spinola

5′. Coloración de los élitros diferente..............6

6. Mitad anterior del pronoto más angosta que mitad posterior ( Fig. 61 )..............................7

6′. Mitad anterior del pronoto más o menos de igual ancho que mitad posterior ( Fig. 124 ).........................................................................8

7. Antenomero basal de la maza antenal más corto que la suma de los antenomeros funiculares ( Perú) ( Fig. 52).................................................. ............. Ichnea gregata Opitz , especie nueva

7′. Antenomero basal de la maza antenal igual o más largo que la suma de los antenomeros funiculares ( Brasil, Argentina) ( Figs. 53–55) ............................... Ichnea praeusta (Klug)

8. Pronoto con una banda longitudinal central ancha y amarilla, flanqueada de negro, o pronoto totalmente negro (EEUU, México, Guatemala, Belice, Honduras, Nicaragua, Costa Rica, Panamá, Venezuela, Colombia, Ecuador) ( Figs. 71–74)........... Ichnea aterrima (Klug)

8′. Pronoto con una banda longitudinal central ancha y negra, que puede presentar un poco de amarillo cerca del margen pronotal anterior ( Perú, Bolivia) ( Fig. 56) ......................... .......................... Ichnea procera Schenkling

9. Pronoto levemente trapezoidal, casi cuadrado ( Nicaragua, Costa Rica, Panamá, Colombia, Venezuela, Perú, Ecuador, Brasil) ( Figs. 90– 94)........................ Ichnea frenata (Erichson)

9′. Pronoto fuertemente trapezoidal................10

10. Antenomero basal de la maza antenal aproximadamente igual a lo largo combinado de los antenomeros funiculares ( Fig. 169 ); disco elitral con una banda amarilla en el medio ( Brasil) ( Figs. 106 , 133 ) ..................... ................................ Ichnea mimica Gorham

10′. Antenomero basal de la maza antenal más corto que lo largo combinado de los antenomeros funiculares ( Fig. 170 ); disco elitral sin banda amarilla en el medio ( Bolivia, Brasil) ( Figs. 95– 97) .............................. Ichnea lycoides Laporte

11. Frente plano, sin depresión........................12

11′. Frente muy cóncavo o con una depresión profunda.....................................................15

12. Ojos casi contiguos en vista dorsal, ancho interocular menor del ancho del escapo (grupo de especies marginella ) ( México, Guatemala, Belice, Honduras, Costa Rica, Panamá, Colombia, Venezuela, Trinidad, Guyana, Suriname, Guyana Francesa, Ecuador, Bolivia, Perú, Brasil, Paraguay) ( Fig. 107 )........ .................................. Ichnea marginella (Klug)

12′. Ojos no contiguos en vista dorsal, ancho interocular igual o más ancho que el ancho del escapo ........................................................... 13

13. Ancho interocular en vista dorsal mucho más ancho que el escapo (grupo de especies plumbea ) ( Panamá, Colombia, Ecuador, Perú, Bolivia, Brasil) ( Figs. 108, 109 ) .................... .............................. Ichnea plumbea Gorham

13′. Ancho interocular en vista dorsal aproximadamente igual al ancho del escapo (grupo de especies digna )......................................14

14. Márgenes suturales y epipleurales con solo un poco de amarillo cerca de la mitad del disco elitral ( Ecuador) ( Fig. 59) .................... ........... Ichnea gremia Opitz , especie nueva

14′. Márgenes suturales y epipleurales con amarillo en todo su longitud, o disco elitral completamente café oscuro ( Costa Rica, Panamá, Colombia) ( Figs. 57, 58) .............. .................................. Ichnea digna Wolcott

15. Pronoto rosado...........................................16

15′. Pronoto no rosado......................................17

16. Forma ancha subovalada; pronoto transverso; pronoto con manchas laterales rosadas ( Guyana, Guyana Francesa, Brasil) ( Fig. 89).... ......................................... Ichnea opaca (Klug)

16′. Forma estrecha; pronoto cuadrado; pronoto sin manchas laterales rosadas ( Panamá, Colombia, Venezuela, Bolivia, Ecuador, Perú, Paraguay, Brasil) ( Fig. 60) ............................... ................................. Ichnea divisa Chevrolat

17. Pronoto principalmente amarillo, sea totalmente amarillo o amarillo con disco cafesoso ( Brasil) (grupo de especies incerta ) ( Figs. 110, 111 )............................ Ichnea incerta Gorham

17′. Pronoto casi totalmente negro (grupo de especies acanthomelina ) ............................ 18

18. Pronoto densamente revestido de setas doradas ( Brasil) ( Fig. 41) .................................................. ... Ichnea acanthomelina Opitz , especie nueva

18′. Pronoto no densamente revestido de setas doradas ( Guyana, Brasil, Suriname) ( Fig. 42) ................ Ichnea atra Opitz , especie nueva

acanthomelina species-group

Among representatives of species characterized by a suboval body form, these beetles can be distinguished by the almost entirely dark brown coloration of the pronotum. This group, which consists of two species, ranges from the coastal northern environs of South America to the central highlands of Brazil.

Figures

Figs. 121–129. Ichnea spp. 121) Pronotum of I. aequinoctialis; 122) Pronotum of I. lycoides; 123) Pronotum of I. praeusta; 124) Pronotum of I. aterrima; 125) Head of I. aterrima; 126) Head of I. aequinoctialis; 127) Pygidium of I. marginella; 128) Aedeagus of I. praeusta; 129) Aedeagus of I. atra.

Figs. 45–51. Ichnea frenata. 45) Antenna; 46) Head, ventral view; 47) Pronotum, dorsal view; 48) Pronotum, ventral view; 49) Pronotum, lateral view; 50) Pronotum, dorsoventral view; 51) Pronotum, posterior view.

Figs. 61–64. Ichnea spp. 61) Forebody of I. praeusta; 62) Forebody of I. aequinoctialis; 63) Head of I. frenata;

Figs. 76–79. Ichnea spp. 76) Forebody of Ichnea marginella; 77) Pronotal trichobothrium of I. aequinoctialis;

Figs. 98–101. Ichnea frenata. 98) Postocular region; 99) Postocular setae; 100) Last antennomere; 101) Elytral setae.

Figs. 65–66. Ichnea frenata. 65) Mouthparts; 66) Funicular antennomeres.

Figs. 80–88. Ichnea spp. 80) Metathoracic wing of I. frenata; 81) Stomodaeal valve of I. frenata, internal view; 82) Metendosternite of I. frenata; 83) Labrum of I. frenata; 84) Labium of I. frenata; 85) Mesoscutellum of I. frenata; 86) Mandible of I. frenata; 87) Maxilla of I. frenata; 88) Alimentary canal of I. divisa.

Figs. 112–113. Ichnea frenata. 112) Postocular region; 113) Pronotal trichobothrium.

Figs. 130–132. Line habiti of 130) Stegnoclava zorropsis; 131) Stegnoclava velatis; 132) Ichnea gregata.

Figs. 102–105. Ichnea spp. 102) Forebody of I. frenata; 103) pronotal trichobothrium of I. divisa; 104) Metatibia and metatarsus of I. praeusta; 105) Section of elytron of I. praeusta.

Figs. 176–190. Aedeagi of Ichnea spp. 176) I. plumbea; 177) I. gremia; 178) I. digna; 179) I. callanga; 180) I. frenata; 181) I. procera; 182) I. opaca; 183) I. divisa; 184) I. incerta; 185) I. aequinoctialis; 186) I. gregata; 187) I. acanthomelina; 188) I. marginella; 189) I. aterrima; 190) I. lycoides.

Figs. 114–119. Mesodermal reproductive organs of Ichnea spp. 114) I. divisa, male; 115) I. divisa, female; 116) I. marginella, male; 117) I. marginella, female; 118) I. frenata, male; 119) I. frenata, female.

Figs. 169–175. Antennae of Ichnea spp. 169) I. mimica; 170) I. lycoides; 171) I. frenata; 172) I. opaca; 173) I. praeusta; 174) I. procera; 175) I. callanga.

Figs. 106–111. Habiti of 106) Ichnea mimica; 107) Ichnea marginella; 108–109) Ichnea plumbea; 110–111) Ichnea incerta.

Fig. 133. Habitus of Ichnea mimica.