Acyphoderes Audinet-Serville, 1833

Clarke, Robin O. S., 2015, Revision of the genus Acyphoderes Audinet-Serville, 1833, with a brief synopsis of the genus Bromiades Thomson, 1864 (Coleoptera, Cerambycidae), Insecta Mundi 2015 (401), pp. 1-92 : 10-17

Acyphoderes Audinet-Serville, 1833

( Fig. 7-10 , 13-21 )

Type species. Necydalis aurulenta Kirby, 1818 , Newman (1840) designation.

Acyphoderes Audinet-Serville, 1833: 549 . Monné 2005: 453 (cat.).

Diagnosis. Prothorax Type 1A in Acyphoderes Groups 1 and 2 species (only shared with Bromiades , see under this genus for diagnostics); or Type 1B in Group 3 species (shared with Anomaloderes and Brachyphoderes , see under these genera for diagnostics). Elytra subtype 3A only found in Acyphoderes Group 1 species; or subtype 3B in Acyphoderes Groups 2 and 3 species (shared with Acyderophes and Anomaloderes , see under these genera for diagnostics). Male abdomen Type 2A in all species of Acyphoderes (shared with Acyderophes , Amerispheca , Anomaloderes and Bromiades , see under these genera for diagnostics; and shared with Ameriphoderes amoena ), but in Acyphoderes apical margin of urosternite V with broad projection at middle, or slightly bisinuate, or almost truncate, and inclination of abdominal process varies from flat to 30° (in Ameriphoderes amoena apical margin of urosternite V deeply excavate, and abdominal process with 45° inclination). Hind leg EITHER subtype 1A in Acyphoderes Group 1 species (only shared with Bromiades , see under this genera for diagnostics); OR subtypes 2A and 2B, the first unique to Acyphoderes auricapilla , the second unique to Acyphoderes crinita and A. rubrohirsutotibialis ; OR subtype 3Ba in Acyphoderes hirtipes (shared with Anomaloderes and Forficuladeres , see under these genera for diagnostics); or subtype 3Bc in Acyphoderes amboroensis and A. carinicollis (shared with Ameriphoderes bayanicus , A. cribricollis and A. prolixa ), but in the two species of Acyphoderes hind leg is 1.7-2.0 longer than front leg (in the three species of Ameriphoderes hind leg 2.3-2.5 longer than front leg). Aedeagus with Type 2 tegmen (shared with Acyderophes and Bromiades , see under these genera for diagnostics).

Description of genus. Medium sized (14.5-25.0 mm), compact species (Group 1 and 2 species), or moderately compact (Group 3 species); forebody 0.87-1.17 length of abdomen.

Head with eyes distinctly narrower than width of prothorax (width prothorax/head 1.24-1.54). Rostrum long, 1.7-2.1 wider than long (Group2), or shorter, 2.4-2.75 (in Group 3 species), or short, 2.55-3.15 (Group 1 species). Maxillary palps short (Group 2 species) or long (Groups 1 and 3 species), apical palpomeres cylindrical with truncate apices; galea moderately long and narrow, or long and narrow (Group 1 species, and in A. hirtipes ). Labrum moderately large, not strongly transverse, EITHER about 1.3 wider than long (in Group 1 species), OR 1.5 wider than long (in A. auricapilla , A. crinita and A. rubrohirsutotibialis ), OR about two times wider than long (in Group 3 species, and in A. hirtipes ); front margin emarginate, slightly declivous; sides rounded (most Group 2 and 3 species, and in A. aurulenta ), or sides straight and convergent (in A. abdominalis and A. crinita ). Clypeus usually flat, but moderately convex (in A. carinicollis ), slightly inclined to labrum, and hardly wider; EITHER planar with frons (in A. auricapilla , A. crinita and A. rubrohirsutotibialis ), OR slightly declivous (in A. hirtipes ), OR strongly declivous (in Group 1 species). Frontal suture in female short, as sulcus or raised line, not reaching anterior margin of inferior lobes (in Groups 1 and 3 species, and in A. auricapilla , A. crinita and A. rubrohirsutotibialis ), or nearly reaching clypeus (in A. hirtipes ).

Inferior lobes of eyes in males large (especially in A. crinita ) and round (in Groups 2 and 3 species), or distinctly wider than long (in Group 1 species), in female flatter and smaller; subcontiguous, in male width of one lobe 2.34-4.40 interocular distance (in female 1.14-1.60); their distal margins lying on frons in male, closer to gena in female, proximal margins usually transverse in males and weakly oblique in females.

Superior lobes moderately large and wide, and narrowed laterally (to 0.40-0.67 their mesal width, except female A. auricapilla 0.8); interocular distance/width of one lobe 1.9-2.5 (in Groups 2 and 3 species), or 2.4-2.8 (in Group 1 species); with fine ommatidia, in males arranged in approximately 13-15 rows (in Group 3 species), or 17-20 rows (in Group 2 species), or 21-23 rows (in Gp.1 species); in females, 13-18 (in Group 2 species), or 19 rows (in Group 3 species), or 23-24 (in Group 1 species).

Mentum-submentum represented by, EITHER quadrate area in male, subquadrate in female (in A. crinita and A. rubrohirsutotibialis ), OR transverse area (in Groups 1 and 3 species, and in A. auricapilla and A. hirtipes ); multicarinate (the carinas confused with small elliptical areas), and sparsely punctured (the punctures moderately small to large, alveolate, and usually situated in these ellipsoids).

Antennal tubercles rather small, rounded, weakly prominent (in female A. auricapilla somewhat prominent); the distance between them/width of scape 1.80-2.10 (in males of Group 2 species), or 2.00- 2.10 (in males of Group 3 species), or 2.20-2.27 (in both sexes of Group 1 species, and in females of Group 2 and 3 species).

Antennae short, EITHER just reaching apex of metepisternum (in Group 1 females), OR apex of metasternum (in Group 1 males), OR rather short, just passing metacoxae (in Group 2 males), OR reaching from metacoxae to near middle of urosternite I (in Groups 2 and 3 females); EITHER subfiliform (in Group 3 species), OR subcrassate (in A. abdominalis , A. crinita and A. rubrohirsutotibialis , and least so in A. auricapilla ), OR nearly crassate (in female A. aurulenta and in both sexes of A. hirtipes ); scape subcylindrical (in most Group 2 species, and in both Group 3 species), or subpyriform (in A. hirtipes ), or pyriform (in Group 1 species); antennomere III filiform, EITHER shorter than scape (0.7-0.8 length of scape in Group 1 species), OR longer than scape (1.13-1.19 in Group 3 species), OR much longer than scape (1.29-1.46 in most Group 2 species, and even longer in male A. hirtipes 1.67); IV subcylindrical, short (0.40-0.66 length of III (in Groups 2 and 3 species), or moderately long (0.8-0.9 length of III in Group 1 species), shorter than V (in Groups 2 and 3 species, and in A. aurulenta ), equal in length to V (in A. abdominalis ); V subcylindrical (in Groups 2 and 3 species), or strongly widened at apex and serrate (in Group 1 species, but weakly serrate in females of A. aurulenta and A. carinicollis ), and slightly longer than VI (in Groups 2 and 3 species), or subequal to VI (in Group 1 species); VI widened at apex and strongly serrate, equal to VII or slightly longer; VII-X incrementally shorter, widened at apex and strongly serrate; XI longer than X, with moderately small cone, the latter narrow (in A. aurulenta , A. crinita , A. rubrohirsutotibialis , and males of A. hirtipes and A. amboroensis ), and in some of these species rendering this segment somewhat subserrate in appearance.

Prothorax Type 1. Subtype 1A (in Groups 1 and 2 species), or subtype 1B (in Group 3 species). Subtype 1A. Prothorax discoid (length/width 0.90-1.08). Central depression less ample and not as deep in A. auricapilla ; lateral calli do not project beyond sides of pronotum in A. auricapilla and some females of A. hirtipes ; lateral calli crimped at middle (in most species). Subtype 1B. Prothorax obovate (length/ width 1.03-1.12).

Prothorax subtypes 1A and 1B. Front margin (f) slightly narrower, to wider, than hind margin (h), in male f/h 0.85-1.10, in female 0.84-1.05. Apical constriction weak; basal constriction EITHER moderately broad, and to either side of midline connected to central depression (in Group 2 and 3 species), OR moderately broad and deep laterally, narrower and separated from deepest part of central depression by slight ridge towards midline (in Group 1 species). Sides EITHER widest close to middle, prothoracic quotient 1.97-2.13 (in Group 1 males and some females of A. hirtipes ), OR before middle, prothoracic quotient 2.21-2.33 (in Group 2 males and female A. hirtipes , and in Group 1 females), OR well before middle, prothoracic quotient 2.58-2.96 (in most Group 2 females, in Group 3 males, and in female A. carinicollis ). Surface of pronotum uneven; and details may be obscured by rugose puncturation (in all Group 2 species). Disc of pronotum EITHER with moderately broad, long callus at midline (in Group 1 species, and A. carinicollis ), OR this callus mostly evanescent (in most Group 2 species, and in A. amboroensis ), OR absent (in A. auricapilla ); and pair of calli to either side of midline (anterior callus narrow, the posterior callus broad), these connected to form an inverted comma-shaped crescent delimiting basal two-thirds of pronotal disc, the crescent abrupt (in Group 1 species), prominent, but rounded (in Groups 2 and 3 species).

Prosternum EITHER declivous for apical third (in male A. hirtipes ); OR apical half (in females of A. hirtipes ), OR hardly declivous (in Groups 1 and 3 species, and females of A. auricapilla and A. rubrohirsutotibialis , and both sexes of A. crinita ); surface of prosternum EITHER inclined to prosternal process (in Group 1 species, and some Group 2 species), OR rather flat (in Group 3 males, and in females of A. auricapilla , and in both sexes of A. hirtipes ), OR tumid (in both sexes of A. crinita , and females of A. carinicollis and A. rubrohirsutotibialis ); and the surface adjacent to base of prosternal process raised into large conical tubercle (in Group 1 species). Prosternal process flat, or exceptionally arced (in male A. amboroensis and female A. auricapilla ); base exceptionally broad in male, 1.87-2.17 times narrower than width of procoxal cavity (in males of Group 1 species, and 1.56 in females); or moderately narrow in both sexes, 2.44-8.00 times narrower than width of procoxal cavity (in Groups 2 and 3 species; narrowest in A. amboroensis , widest in female A. crinita ). Apex of process strongly declivous across apical third; EI- THER characteristically, axe-shaped (in Gp.1 species), OR more typically, large and trapezoidal (in most Group 2 species, and both Group 3 species), OR somewhat triangular and not declivous (in female A. auricapilla ). Procoxal cavities plugged at sides, closed behind.

Mesosternum abrupt and deep. Base of mesosternal process EITHER wider than, or nearly as wide as, mesocoxal cavity, coxal cavity 0.92-1.09 width of process (in Group 1 females), OR narrower than coxal cavity, coxal cavity 1.39-1.41 wider than process (in Group 1 males), OR 1.31-2.20 wider than process (in Group 3 females, and Group 2 species, narrowest in male A. hirtipes , widest in females of A. hirtipes and A. crinita ), OR 2.13-3.00 wider than process (in Group 3 males) ; apex of process bilobed, the lobes hardly divergent, each one rather broad and rounded at apex. Mesocoxal cavity moderately open to epimeron.

Length of mesosternum/metasternum 0.60-0.87.

Elytra Type 3. Subulate; and subfissate for apical half (in Group 1 species), or for apical two-thirds (in Groups 2 and 3 species; but A. carinicollis shows considerable variation) .

Subtype 3A (Group 1 species). Relatively short, in male 1.98-2.00 longer than width of humeri, in female 1.70-1.91 longer; apex reaching from middle to apical third of urosternite II (in A. abdominalis ), from basal quarter to basal third of II (in A. aurulenta ); apical margin acuminate.

Subtype 3B (in Groups 2 and 3 species). Moderately long, in males, 2.60-2.70 longer than width of humeri, in females shorter, 2.30-2.61; in males apex reaching from middle of urosternite III to middle of IV, in females basal third of III to base of IV; apical margin narrow and blunt (subacuminate).

Subtypes 3A and 3B. Each elytron with well demarcated translucent panel (less so at base in A. aurulenta ); regularly narrowed from humeri to apex (in Group 1 species), or regularly narrowed from humeri to apical two-thirds, almost parallel to apex (in Groups 2 and 3 species). Humeri not or hardly projecting (projecting in A. amboroensis ); almost square and hardly prominent (in Group 1 species, and in A. carinicollis , A. auricapilla , A. crinita and A. rubrohirsutotibialis ), or well rounded and moderately prominent (in A. amboroensis and A. hirtipes ).

Metathorax broad; sides EITHER subparallel with apical margin oblique (in females of A. carinicollis and A. auricapilla ), OR sides weakly rounded and converging towards apex, and apical margin moderately oblique (in A. abdominalis , A. amboroensis , male A. carinicollis and A. hirtipes ), OR apical margin almost truncate (in A. aurulenta ), OR strongly rounded from base to middle of apex (in A. crinita and A. rubrohirsutotibialis ). Metasternum tumid, but broadly flattened on disc; more or less planar with mesocoxae, but well below level of mesocoxae (in males and some females of A. hirtipes ); longitudinal suture moderately shallow to moderately deep, but always long. Metepisternum cuneate, broad at base, subacuminate at apex.

Male abdomen Type 2. Cylindrical or subcylindrical; apical segments not distinctly wider than basal segments; urosternite II not conical.

Subtype 2A in all three Groups (Fig. 72, 74, 77, 79, 82, 84). Widest at base of urosternite I, or middle of 1/base of II, hardly narrowed to apex, or slightly wider towards apex (in A. amboroensis ); urosternites I-IV parallel-sided, but slightly contracted at base and apex; urosternite I quadrate or elongate, II-IV transverse. Male urosternite V EITHER cylindrical (in Group 1 species), OR trapezoidal (in Group 3 species, and in A. crinita ), OR quadrate (in A. hirtipes ); soleate depression EITHER shallow without raised sides (in Group 1 species), OR with weakly raised sides (in A. hirtipes ), OR soleate depression deep with narrow sides, these increasingly projecting towards base (in A. amboroensis , A. carinicollis and A. crinita ), and terminating in pair of acute tubercles (in A. carinicollis and A. crinita ); apical margin EITHER with abrupt, broad projection at middle (in Group 1 species, and in A. carinicollis and A. hirtipes ), OR slightly bisinuate (in A. amboroensis ), OR almost truncate (in A. crinita ).

Female abdomen (Fig. 73, 75, 76, 78, 80, 81, 83) fusiform, rather wide and flat (less so in A. abdominalis , more so in A. aurulenta ). Urosternite V EITHER trapezoidal or subconical, and rather short to moderately long (in Groups 1 and 3 species, and in A. crinita , A. hirtipes and A. rubrohirsutotibialis ), OR very short and transverse (in A. auricapilla ); the surface undifferentiated and weakly declivous across middle (in Groups 1 and 3 species, and in A. auricapilla , A. crinita and A. rubrohirsutotibialis ), OR characteristic and male-like, with rounded soleate depression between basal and apical thirds, the depression Ushaped and demarcated by narrow, raised sides (in A. hirtipes ); apical margin simply rounded to subacuminate (in Groups 1 and 3 species, and in A. auricapilla , A. crinita and A. rubrohirsutotibialis ), or apical third of segment narrowed and parallel-sided, and truncate at apex (in A. hirtipes ).

Male and female abdominal process broadly triangular, with rather blunt apex (in Group 1 species, most Group 2 species, and in female A. carinicollis ), or narrowly triangular with acuminate apex in (in Group 3 males, and in male A. hirtipes ); in most species EITHER planar with abdomen, or almost so (in most females, in male A. hirtipes , and in both sexes of A. carinicollis ); OR with 10-15° slope (in Group 1 males, male A. crinita , and some female A. hirtipes ), OR with 30° slope (in male A. amboroensis ).

Apical tergite in male cylindrical to subcylindrical, rather short to moderately long, broad, strongly convex, and rounded at apex. In female apical tergite subconical to subcylindrical, long, narrow and flat, apex truncate, rounded or subacuminate (in A. abdominalis , and in Groups 2 and 3 species), or somewhat trapezoidal and short, but with narrow, excavate-sided extension to rounded apex (in A. aurulenta ); and midline occupied by narrow raised line (in Group 2 species, becoming wide and strongly raised in A. hirtipes ; and in Group 3 species).

Legs. Ratio lengths front/middle/hind leg 1.0:1.1:1.7-1.9 (in Group 1 species), or 1.0:1.1-1.2:1.7-2.0 (in Group 3 species), or 1.0:1.1-1.4:2.0-2.4 (in Group 2 species).

Front and middle legs moderately long to long (body length/length of legs 2.2-2.8 and 1.8-2.4 respectively); moderately strongly pedunculate-clavate (in Group 3 species), or strongly pedunculate-clavate (least so in A. auricapilla ).

Front Leg. Profemoral clave tumid mesally, pro- and mesofemoral claves broad and abrupt. Protibiae shorter or longer than profemora; narrow at base, EITHER gradually widening to apex (in Groups 1 and 3 species), OR gradually widening to middle (in some female Group 2 species), OR somewhat abruptly widening to middle (in females of A. hirtipes ), and parallel-sided to apex, OR very abruptly widened to middle, and narrowing to apex (in male A. hirtipes ); apical margin oblique.

Middle Leg. Mesofemur longer in males than in females, in males 1.29-1.45 longer than mesotibia (shortest in male A. abdominalis , longest in male A. hirtipes ), in females 1.22-1.32 longer than mesotibia (shortest in female A. abdominalis , longest in female A. hirtipes ). Mesotibia moderately short and robust (in Group 1 species and in A. hirtipes ), or moderately short and slender (in the remaining species); narrow at base, EITHER gradually widening to apex (in Groups 1 and Group 3 species, and in A. crinita ), OR parallel-sided for apical half (in male A. hirtipes ), OR parallel-sided for apical two-thirds (in female A. hirtipes , and in remaining Group 2 species).

Hind leg (Fig. 101, 103, 104, 106, 108) subtype 1A (in Group 1 species), or Types 2 or 3 (in Groups 2 and 3 species). EITHER long, body length 1.0-1.2 longer than legs (in Group 2 species, and in A. carinicollis ), OR moderately long, body length 1.3-1.4 longer than legs (in Group 1 species), OR short, body length 1.6 longer than legs (in A. amboroensis ).

Hind leg subtype 1A (Fig. 101) in A. abdominalis and A. aurulenta . Metafemoral peduncle moderately short (length clave/peduncle 1.35-1.80); apical half of metatibia with rather short, compact, dense, rufous, or rufous and black brush.

Hind leg Type 2 (Fig. 103, 104) in A. auricapilla , A. crinita and A. rubrohirsutotibialis ). Metafemoral peduncle flattened, and moderately short to long (length clave/peduncle 0.85-1.75); apical half of metatibia with rather long, but not very wide, black, or yellow to rufous brush.

Subtype 2A (Fig. 103) in A. auricapilla . Legs moderately robust; apex of clave reaching middle of urosternite IV.

Subtype 2B (Fig. 104) in A. crinita and A. rubrohirsutotibialis . Legs slender; apex of clave reaching from base to apex of urosternite V.

Hind leg Type 3 (in A. amboroensis , A. carinicollis and A. hirtipes ). Peduncle flattened, narrow, and moderately short to very long (length clave/peduncle 0.85-1.30); metatibia with or without brush; metatarsomere I cylindrical or subcylindrical, I/II+III (1.0-1.1).

Subtype 3Ba (Fig. 106) in A. hirtipes . Apex of clave reaching apex of urosternite IV to apex of V; apical half of metatibia with rather long, but not very wide, black, or rufous brush.

Subtype 3Bc (Fig. 108) in A. amboroensis and A. carinicollis . Apex of clave reaching from apex of urosternite III to middle of urosternite IV; metatibia without brush.

Hind leg Types 1, 2 and 3. Metafemoral apex (when viewed laterally) weakly narrowed (in Group 1 species and most Group 2 species), or moderately strongly narrowed (in Group 3 species, and some female A. hirtipes ). Metatibia (when viewed from above) EITHER almost straight (in A. carinicollis ), OR bent inwards for apical third (in Group 1 species), OR bent inwards for apical half (in Group 2 species, and in A. amboroensis ); shorter than metafemora; robust (in Group 1 species), or elegant (in Groups 2 and 3 species); and gradually widening to apex. Metatarsus moderately short, less than half, to half length of metatibia (in both sexes of A. carinicollis , in male of A. abdominalis and females of A. auricapilla and A. hirtipes ), or long, more than half length of metatibia (in female A. abdominalis , male A. amboroensis , and both sexes of A. aurulenta , and in A. crinita , male A. hirtipes and female A. rubrohirsutotibialis ). Metatarsomere I trapezoidal (in some Group 1 species) or cylindrical (in some Groups 1 and 2 species); II subtrapezoidal; III moderately long, with weakly divergent lobes.

Male genitalia. AedeagusType 2 (Fig. 112-115, 118)

Tegmen Type 2. Lateral lobes (length/width 4.4-7.1) paddle-shaped apically; base moderately broad; divergent at base; towards apex more so, or subparallel, or convergent; densely chitinized; twisted; and basalpiece short.

Subtype 2a (Fig. 112, 113) in A.abdominalis and A.crinita . Lateral lobes (length/width 5.5-7.1) strongly divergent from base to apex; apices non abrupt, elongate paddles; extreme apex and lateral side of paddle setose.

Subtype 2b (Fig. 114, 115) in A. aurulenta and A. hirtipes . Lateral lobes (length/width 4.8-5.3) divergent at base, convergent towards apex (but widely separated throughout); apices abrupt, rounded paddles; hardly setose.

Subtype 2d (Fig. 118) in A.amboroensis and A. carinicollis ). Lateral lobes (length/width 4.4) subparallel to slightly divergent; apices weakly abrupt, elongate, narrow paddles; apex and sides of paddle setose.

Median lobe Type 1. Length of median lobe 2.3 mm (in Group 3 species), 2.7 mm (in Group 2 species), 2.8-3.0 mm (in Group 1 species).

Surface ornamentation/pubescence. On upper side notable pubescence (discrete areas of dense, recumbent, metallic or brightly colored pubescence) only present on frons between eyes, most of pronotum, and scutellum; the pubescence recumbent (in A. aurulenta pale golden in color, in male A. hirtipes brassy and ochraceous, in females of A. hirtipes bright coppery), or replaced by notably dense, erect, pubescence on pronotum (in A. abdominalis mostly dark golden in color, in A. auricapilla , A. crinita and A. rubrohirsutotibialis black, and mixed with longer, setose-like hairs). Elytra almost glabrous; but some semirecumbent hairs between humeri; and translucent panels with rows of fine punctures, each with a short, erect, black hair.

Antennal scape, and antennomeres III (in A. abdominalis and A. aurulenta ), III and IV (in A. crinita and A. rubrohirsutotibialis ), III-V (in Group 3 species), III-VI (in A. auricapilla and A. hirtipes ) with fringe of black setae on meso-ventral surface. Pedicel and apex of III-VI with 2-4 longer, thicker setae at apex; VII-X usually with single, incrementally smaller seta at apex.

On underside notable pubescence limited to prosternum (apical third glabrous in Group 1 species, and most of prosternum in female A. carinicollis ). EITHER with very long, often untidy, semi-erect hairs (in Group 3 males semi-erect, ashy colored hairs), OR recumbent hairs (in male A. hirtipes ashy colored hairs; in male A. aurulenta very long and brassy in color; in female A. aurulenta shorter, but denser, rufous on prosternum, golden on prosternal process; in male A. abdominalis shorter and untidy, brassy and black in color; in female A. abdominalis short, erect, and black in color); mesepimeron and adjacent sides of mesosternum (in Group 1 species golden colored hairs; in male A. hirtipes brassy hairs, in female ochraceous hairs; in Group 3 species ashy-white to silver hairs; in some female A. hirtipes silver colored hairs; in A. crinita with whitish hairs, but not extending to sides of mesosternum in male); and all of metasternum, and much of metepisternum (in A. aurulenta and in males of A. crinita and A. hirtipes with brassy hairs, in females somewhat dusky hairs), or basal half of metasternum and extreme apex of metepisternum (in A. abdominalis with brassy hairs, and in Group 3 species with silver hairs).

Less notable pubescence long, semi-erect, moderately dense hairs (usually untidy and sometimes rather sparser and shorter) supplement the underside pubescence at centre of prosternum (in both sexes of A. crinita , and females of A. auricapilla and A. rubrohirsutotibialis with dusky hairs; and in females of A. hirtipes with ochraceous hairs); sides of meso- and metasternum (in Group 1 species with pale golden hairs; in A. amboroensis with brownish hairs); apical half of metasternum and most of metepisternum (in Group 3 species with silver colored pubescence); OR dense, short, and recumbent pubescence uniformly covering most of meso- and metasterna (in females of A. auricapilla with rufous hairs, and A. rubrohirsutotibialis with rufous and black hairs).

Abdomen lacking notable pubescence (except in Group 3 species), and urosternite I often rather glabrous; urosternites II-IV EITHER mostly glabrous with scattered patches of recumbent, fine, short pubescence, and longer, sparser semi-erect hairs supplementing these patches (in Group 1 males); OR II- IV almost uniformly clothed with, recumbent, moderately short and dense hairs (in females of A. abdominalis and A. auricapilla with ashy colored hairs, in A. rubrohirsutotibialis and some A. hirtipes with black hairs, in A. aurulenta with black and rufous hairs, in some females A. hirtipes with rufous hairs; and in both sexes of Group 3 species with denser silver colored hairs of more notable appearance, supplemented by ocellate patches in males); OR II-IV with rather sparse, very short, somewhat setoselike hairs (in A. crinita with ashy and yellowish hairs, in male A. hirtipes with testaceous hairs).

Legs. Femora rather densely clothed with fine, long, pale hairs; recumbent on profemora; suberect on meso- and metafemora (in A. abdominalis and A. amboroensis ), or femora clothed with thick, rather stifflooking, black hairs (in Group 2 species, and in A. aurulenta and A. carinicollis ). Metatibiae with moderately long haired, dense brush (lacking brush in Group 3 species), occupying apical two-thirds (in Group 2 species), or apical half (in Group 1 species), the hairs EITHER mostly yellowish (in female A. auricapilla ), OR rufous (in A. abdominalis , some A. crinita , and females of A. hirtipes and A. rubrohirsutotibialis ), OR rufous and black (in A. aurulenta ), OR black (in some A. crinita ). Metatarsi EITHER rather densely covered by thick, long hairs (in A. aurulenta , in male A. crinita with black hairs, in both sexes of A. hirtipes with yellow hairs), OR rather short hairs (in females of A. crinita and A. rubrohirsutotibialis with black hairs), OR metatarsi covered by rather fine hairs (in Group 3 species black hairs, in A. abdominalis rufous hairs, in female A. auricapilla yellow hairs).

Surface ornamentation/puncturation. On upper side frons and vertex of head densely punctate, the punctures deep and alveolate. Pronotum EITHER entirely and densely, rugosely punctured, the punctures rather small, alveolate and deep; and embedded in matrix of dense micropunctures (in Group 1 species, A. auricapilla , A. crinita and A. rubrohirsutotibialis ); and on apices of the broad, basal calli the punctures infiltrated by network of smooth carinas (in Group 1 species); OR entirely and densely covered with alveolate punctures (with smooth impunctate areas on calli), otherwise the punctures uniformly distributed and all of small size (in A. amboroensis ), or larger on, and adjacent to, calli, denser, smaller, and variable in size in depressed areas (in both sexes of A. carinicollis ).

Antennal scape with small rounded punctures; moderately dense (in Group 3 species; in A. aurulenta , A. auricapilla , A. crinita and A. rubrohirsutotibialis ), or moderately sparse (in A. abdominalis and A. hirtipes ).

Elytra rather densely punctate (except on translucent panels), the punctures alveolate or simple, on the following. Across base subcontiguous, on humeri denser (in Group 1 species somewhat evanescent) and beveled; a single row of very small punctures adjacent to sutural border; epipleur with 2-5 rows of larger and smaller, deep and confluent punctures; and rather disperse rows of very small, shallow, setose punctures on translucent panels.

On underside puncturation partially hidden by pubescence. Prosternal puncturation variable, EI- THER with smooth surface, the punctures mostly contiguous (but in some areas becoming confluent), alveolate, relatively small to moderately large, and deep (in A. amboroensis , in A. crinita , in females of A. hirtipes and A. rubrohirsutotibialis ), OR the punctures confluent, and the surface rugose (in female A. auricapilla the surface dull, in A. carinicollis the surface shining), OR surface not rugose, but dull and reticulate, all punctures contiguous, alveolate, and small (and of similar size), embedded in matrix of micropunctures (in male A. hirtipes ); OR apical third usually finely carinate, and almost impunctate (in female A. abdominalis ), OR moderately closely punctured (in A. aurulenta ), the punctures alveolate and mostly small; basal two-thirds densely punctate (in female A. abdominalis the punctures a mixture of contiguous and confluent ones; in both sexes of A. aurulenta somewhat rugosely punctate; in male A. abdominalis uniformly, very densely and minutely punctate). Mesosternum at midline EITHER surface smooth, with fine, round, non-alveolate punctures of mixed size (in Group 2 species); OR midline, including mesosternal process, rugosely punctured, the punctures embedded in matrix of micropunctures (in Groups 1 and 3 species); towards sides EITHER front and hind margins of declivity with moderately small punctures embedded in matrix of micropunctures, and middle of declivity smooth and sparsely punctured (in A. hirtipes ), OR almost whole surface with moderately small punctures embedded in matrix of micropunctures (in A. auricapilla , A. crinita and A. rubrohirsutotibialis ), OR adjacent to midline smooth, shining and sparsely punctured, the punctures moderately large to very small, rather shallow and non-alveolate, and towards sides hidden by dense pubescence, but appears to be somewhat rugosely punctured (in Groups 1 and 3 species). Metasternum with EITHER entire surface rather dull and micropunctate with scattered larger, shallow punctures below dense pubescence (in Groups 1 and 3 species, A. crinita and A. rubrohirsutotibialis ), OR only basal half of metasternum similarly punctured (in A. auricapilla and A. hirtipes ); and on glabrous areas of apical half the surface EITHER smooth, and impunctate with group of thirty or so, small tubercles (in male A. hirtipes ), OR the surface sculpted, with tracery of slender, arced carinas creating rows of very dense microsculpture, among which small, scattered punctures occur, these shallow and alveolate (in females of A. auricapilla and A. hirtipes ). Metepisternum usually similar to pubescent areas of metasternum.

Distribution of the abdominal puncturation mirrors the state of its pubescence (see above); the punctures in all species small to very small; rather shallow and beveled (in Group 2 species); and in matrix of micropunctures (in Groups 1 and 3 species).

Species groups. The genus is divided into three species groups (see the list of species of Acyphoderes ), with the following diagnoses based on males unless otherwise stated.

Group 1. Rostrum short (width/length 2.8-3.2); antennae do not pass apex of metasternum, and in both sexes antennomere III shorter than scape (lengths antennomere III/scape 0.7-0.8); prothorax subtype 1A, transverse in females; elytra subtype 3A; hind leg subtype 1A, apical half of metatibia with brush. Aedeagus Type 2a or 2b.

Group 2. Rostrum long (width/length 1.7-1.8); antennae just pass metacoxae, and in both sexes antennomere III distinctly longer than scape (lengths antennomere III/scape 1.3-1.7); prothorax subtype 1A, usually elongate in females; elytra subtype 3B; hind leg subtypes 2A, 2B or 3Ba, apical two-thirds of metatibia with brush. Aedeagus Type 2a or 2b.

Group 3. Rostrum moderately short (width/length 2.4-2.5); antennae reach apical third of urosternite I, and in both sexes antennomere III not much longer than scape (lengths antennomere III/scape 1.1-1.2); prothorax subtype 1B, elongate in females; elytra subtype 3B; hind leg subtype 3Bc, metatibia without brush. Aedeagus Type 2d.

Figures

Figures 7–12. Species habitus in the genera Acyphoderes and Bromiades. Figures 7–8. Acyphoderes aurulenta (Kirby, 1818). 7) male. 8) female. Figures 9–10. Acyphoderes abdominalis (Olivier, 1795). 9) male. 10) female. Figures 11–12. Bromiades brachyptera (Chevrolat in Guérin-Meneville, 1838). 11) male. 12) female.

Figures 13–18. Species habitus in the genus Acyphoderes. Figures 13–15. Acyphoderes hirtipes (Klug, 1825). 13) male. 14) female and 15) female Acyphoderes rufofemorata Fisher, 1930 (junior synonym). 16) Acyphoderes crinita (Klug, 1825), female. 17) Acyphoderes rubrohirsutotibialis Tippmann, 1953, female. 18) Acyphoderes auricapilla Fisher, 1947, female paratype.

Figures 19–24. Species habitus in the genera Acyphoderes and Acyderophes. gen. nov. Figures 19–20. Acyphoderes carinicollis Bates, 1873. 19) male. 20) female. 21) Acyphoderes amboroensis, 2013, male holotype. Figures 22–24. Acyderophes fulgida (Chemsak and Linsley, 1979). 22) paratype. 23) female paratype, Costa Rica. 24) male Mexico (VC).