Azotoctla, Cardona-Duque & Franz, 2012

Cardona-Duque, Juliana & Franz, Nico M., 2012, Description and phylogeny of a new Neotropical genus of Acalyptini (Coleoptera: Curculionidae: Curculioninae) associated with the staminodes of Cyclanthaceae, Zoological Journal of the Linnean Society 166 (3), pp. 559-623 : 561-569

publication ID 10.1111/j.1096-3642.2012.00851.x

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Diagnosis: Azotoctla resembles other genera of Acalyptini (= Derelomini Lacordaire sensu Franz, 2006 ; as corrected in Alonso-Zarazaga, 2007) that key to Phyllotrox Schoenherr in Kuschel’s key to the ‘Petalochilinae’ Pascoe ( Kuschel, 1952: 271–273) – now separated from the Acalyptini and placed in the Molytinae (see, e.g. Franz, 2003a, 2006). However, Azotoctla may be distinguished from all remaining acalyptine genera by the combination of the following diagnostic traits: a compact and rounded body shape (which motivated the genus name ‘turtle’; see Etymology), ventrally with a more flattened surface; a posteroventral marginal area of the prothorax that is inflected (concave) in relation to surrounding lateral regions; a short to long and dorsally glabrous (noncarinate) rostrum that is tumescent above the antennal insertion in males; incomplete abdominal tergites, connected via a mesal longitudinal membrane; and triangularly shaped hemisternites of sternum 8 in males. In addition, Azotoctla shares all phylogenetically relevant characteristics of the Staminodeina as provided in Franz (2006: 277); viz. a labial prementum with two triangular projections, two-segmented labial palps that lack setae, a well-developed anal lobe of the wing, and a specialized and primarily parasitic association with the staminodes of Cyclanthaceae . Azotoctla is therefore readily separated from most members of the Phyllotrogina by, for example, the absence of a perpendicularly orientated pygidium (in relation to body axis) in males and generally less convex elytra; from the Acalyptina by the fully developed elytra, which cover all tergites; and from the Notolomina by its association with Cyclanthaceae as well as numerous mouthpart characters. Azotoctla is furthermore differentiated from the putative sister genus Staminodeus (see Franz, 2001) by a dorsally rounded (as opposed to carinate) rostrum, simple (as opposed to armed) profemora and protibiae in males, an even (as opposed to spinulose to spinose) frons in females, and the absence of specialized female staminode-transporting and ritualized male- male fighting behaviours (see Franz, 2003b).

Description: Male ( Fig. 1A View Figure 1 ): small, length 1.4–3.0 mm, width 0.6–1.6 mm, shape in dorsal view oval, l/w = 1.6–2.3, greatest width near central region of elytra; in lateral view, dorsally nearly flattened to convex ( Figs 6A View Figure 6 , 13A View Figure 13 ), ventrally nearly flat; colour variously light yellowish to dark brown; sculpture nearly even to punctulate; vestiture short to long, sparse to dense, appressed to slightly erect, golden.

Mouthparts: mandible ( Fig. 1B View Figure 1 ) with two teeth similar in length: outer tooth wider, slightly superposed, outer margin rounded, protruded; inner tooth slightly narrower, inner margin rounded, concave; inner surface with a small cutting edge; pharyngeal process inconspicuous. Maxilla ( Fig. 1C View Figure 1 ) with cardo basally bilobed, arcuate, apically widened (subtriangular); stipes broad, triangular, apicodorsal margin oblique, apicoventral margin straight, with one long ventral seta; galea + lacinia + palpiger fused, outer margin apically with one long ventral seta, two thirds of inner surface setose, with six lacinial teeth (sensu Ting, 1936); maxillary palps three-segmented, extending beyond margin of palpiger; palpomeres I and II similar in length, both transverse; III nearly two times length of II, equilateral, papillate. Labium ( Fig. 1D View Figure 1 ) with prementum trapezoidal (apical margin nearly two times length of basal margin), nearly as large as the mandible, with two large, apicolateral, ventrally inserted setae; apicolateral margins rounded, with two stout marginal setae; apical margin with two triangular projections laterad of labial palps; basal margin nearly straight; ligula sclerotized, subtriangular, not extending beyond basal two thirds of prementum, bearing densely arranged dorsal setae; labial palps two-segmented, small, extending beyond apex of triangular projections of prementum, similar in length; palpomere I slightly transverse, slightly clavate; II equilateral, apically papillate. Postmentum elongate.

Rostrum ( Fig. 2A View Figure 2 ): in lateral view short to long, 0.3–0.9 mm; slightly shorter to longer than pronotum r/p = 0.9–1.4, yellowish brown to dark reddish brown; in lateral view usually dorsally arcuate, ventrally subrectate; in cross-section subcircular, usually slightly tumescent above antennal insertion, apex dorsally flattened (distad of antennal insertion), glabrous; in dorsal view apically slightly or distinctly widened; surface glabrous to pilose throughout; even, rugulose or punctulate between antennal insertion and eyes, dorsolateral margins smooth; two shallow gular sutures usually extending to basal third of rostrum, subparallel ( Figs 2B View Figure 2 , 22B View Figure 22 ); antennal insertion positioned near apical one third to one quarter (generally visible in dorsal view), scrobe subrectate or arcuate, ventral margin acutely projected beyond antennal socket to apex, positioned laterally at apex, extending ventrad of eye, well defined, shallow or deep. Antenna ( Figs 2A View Figure 2 , 23C View Figure 23 ): in repose extending beyond anterior margin of pronotum, almost reaching posterior margin. Antennal socket large ( Fig. 26C View Figure 26 ). Scape extending to base of rostrum, apex reaching eye, slightly shorter or similar in length to funicle + club, slightly to distinctly clavate; funicle pilose (setae generally long); segment I shorter to longer than II + III, clavate, apex nearly as wide to wider than apex of scape; II as long as III to slightly longer than III + IV, clavate; III- VII small, similar in length, progressing from subcircular to transverse; club oval to elongate, similar in length or slightly longer than funicular segments III- VII jointly, apically rounded to slightly pointed, with dense, short pubescence, darker than funicle; segment I similar in length to II to slightly longer than two times length of II + III; II similar in length to III or two times length of III.

Head ( Fig. 1A View Figure 1 ): small, globular, colour yellowish to dark reddish brown; ventrally glabrous or scarcely pilose; dorsally barely to distinctly pilose, setae golden, directed towards midpoint; frons pilose, setae dense, golden, directed to midline; area posterior to eyes rugulose to even, pilose ( Fig. 2C View Figure 2 ). Eyes ( Figs 1A View Figure 1 , 2C View Figure 2 ) small, subcircular, protruded, distant from ante- rolateral margin of pronotum by one quarter to three quarters¥ the diameter; interocular distance usually similar to interantennal distance at insertion point.

Thorax: prothorax rounded, slightly flattened l/w = 0.6–0.9; pronotum yellowish brown to dark brown, in dorsal view subcircular, semicircular or suboval, anterior margin 0.3–0.8¥ width of posterior margin, lateral margins either rounded or anteriorly parallel, greatest width at varying positions, slightly to distinctly punctulate, with short to long vestiture, setae usually orientated toward a longitudinal midline; anterior margin rounded, posterior margin slightly to distinctly bisinuate, rounded or nearly straight; in lateral view subrectangular ( Fig. 1A View Figure 1 ), subquadrate ( Fig. 7A View Figure 7 ), or conical ( Fig. 18A View Figure 18 ), posteroventral marginal area inflected (concave) in relation to surrounding lateral regions ( Figs 18A View Figure 18 , 22A View Figure 22 ). Epipleura ( Fig. 18A View Figure 18 ) with mesanepisternum equilaterally triangular, anterior marginal area inflected from lateral plane ( Figs 18A View Figure 18 , 22A View Figure 22 ), scarcely to distinctly pilose; mesepimeron rhomboidal-elongate, scarcely pilose, ventrally slightly to distinctly projected; anterior margin of metanepisternum slightly widened, curved, meeting ventral margin at a sharp angle, posteriorly narrowed; sclerolepidia (cf. Lyal, Douglas & Hine, 2006; Fig. 2E, F View Figure 2 ) present along metanepisternal sutures except for anterior one sixth where suture is curved: consisting of small, posterodorsally directed scales, longer than wide, with weak longitudinal lamellae, somewhat dentate apically, as described by Lyal et al. (2006) as the squamiform 2B type; metepimeron suboval or subtriangular to elongate, extending above posterior one fifth to one sixth of metanepisternum or positioned posteriad to metanepisternum. Sterna ( Fig. 2D View Figure 2 ) with prosternum glabrous or scarcely pilose, anterior and posterior margins rounded; procoxal cavities usually contiguous, inserted between middle and posterior three quarters; prosternal process acute to variously rounded; mesoventrite retracted from ventral plane, mesoventral process rounded, narrowly emarginate or acute (mesocoxae narrowly separated), slightly protruded; metaventrite transverse, anteriorly or entirely pilose, punctulate or even, laterally convex, mesally nearly flat, metathoracic discrimen usually running along entire length, slightly impressed, anterior and posterior margins bisinuate, mesally widely emarginate, posterolateral area with a depressed, glabrous region adjacent to metacoxa, usually preceded by a ridge; metacoxal cavities separated by distance similar to one to two times the mesocoxal diameter. Venter usually even. Metendosternite ( Fig. 3A View Figure 3 ): stalk in ventral view equilaterally triangular, posterior margin rounded, mesally narrowly emarginate; ventral flange in lateral view slender (stalk resembling a coplanar structure); sheaths quadrangular, concave, nearly 1.5¥ length of stalk (cf. Velázquez de Castro, 1998); hemiducts half as long as stalk, clubbed, truncated; furcal arms apically bifurcated, pointed.

Legs ( Figs 6A View Figure 6 , 11A View Figure 11 , 22A View Figure 22 ): prothoracic legs usually similar in length to meso- and metathoracic legs, light yellowish brown to reddish brown; procoxa conical, inner margin with one subapical foveola ( Fig. 2D View Figure 2 ), glabrous to pilose; trochanter oblique; profemur moderately stout, similar in length to meso- and metafemur, f/p = 1.1–1.5, slightly sinuate, compressed, widest near apex, with golden setae (pilosity often denser on apical half); protibia slightly shorter than profemur t/f = 0.7–1.0, slender, apically slightly widened, anteroventrally pubescent along apical third, anteroapical margin roundly oblique, unarmed, with a row of stout setae on pro-, meso-, and metatibiae; protarsus ( Fig. 22C View Figure 22 ) nearly three quarters times the protibial length, dorsally with long setae, ventrally with dense vestiture; tarsomere I similar in length to two times length of II, each clavate; III varying shorter than or similar in length to I + II, lobes slender, apically slightly diverging; IV between one third and three quarters times length of II; V as long as III to II + III; tarsal claws paired, with two to four setae. Meso- and metathoracic legs similar in length; mesocoxa rounded, glabrous or slightly pilose; meso- and metafemora similar in length and width to profemur; meso- and metatibiae anteroventrally pubescent along apical one third to one quarter.

Scutellum: exposed, small, elongate, triangular or pentagonal, covered with setae, yellowish brown to brown.

Elytra: semicircular to elongate, l/w = 1.2–1.7, greatest width near central region, anterior margins rounded or sinuate, jointly wider than posterior margin of pronotum; humeri rounded, sometimes slightly protruded [marking a lateral elytral curvature ( Fig. 22A View Figure 22 )]; lateral margins in dorsal view subparallel along anterior half to two thirds, thereafter uniformly converging (nearly straight or evenly rounded), posteriorly rounded, subcontiguous; in lateral view dorsally flattened along anterior half to three quarters or evenly convex, lateral margins nearly straight; ten-striate, striae narrower or subequal to intervals, shallow or deep, distinct; III- VI merging towards apex, IX merging with X along lateral margin; strial punctures small to large, brown to dark brown, subcircular or suboval; intervals light yellowish brown to dark reddish brown, vestiture short or long, dense, appressed, directed posteriad.

Wings ( Fig. 3B View Figure 3 ): 2.1¥ length of elytra, l/w = 2.5–3.3 (N = 3), transparent, anal angle obliterated, anal lobe large, with setae along its distal half. Anterior margin nearly straight to evenly curved along basal three quarters, apical quarter rounded, with groups of short setae along costa (C) vein; posterior margin with setae throughout (except just in the region of the anal fold), setae shortened to distal portion of posterior margin; apex widely rounded; C merging with subcosta (Sc), strongly sclerotized along basal half to one third; posthumeral patch (pp) extending posteriorly over Sc; radial (R) ascending, merging with C + Sc near their midlength; stigmal patch (sp) subtriangular to elongate (posteriorly irregular), longer than wide, rounded at apex, apically with two setae (cf. Zherikhin & Gratshev, 1995); radial stripe (rst) narrow, continuous with sp; radial fold (rf) distinct, vein-like, shortened before sp or reaching sp; two radial sclerite (rs) present, distad to apex of sp, proximal semilunar to subtriangular, distal rs subtriangular; stigmal fold inconspicuous; radial sclerotization (rsc) rounded, anteriorly slightly to distinctly emarginate (hook-like); anterior part of postradial stripe (app) weak, fused with posterior part of postradial stripe (ppp); ppp distinct, wide near midpoint, laterally narrowed; medial stripe (mst) short to long, conspicuous near posterior margin, slightly hooked at apex; cubital (Cu) curved, widened at inflection point; Cu 1 narrow, continuous with apical fold (af), directed posteriad-distad; anal cell opened; 3 anal (A) present, not reaching posterior margin.

Abdomen ( Fig. 3C View Figure 3 ): venter usually even. Sternites 1–5 jointly 1.5–2.5¥ length of lateral margin of metaventrite, even or distinctly punctulate, vestiture variable, sometimes sparser anteriad (near mesal concavity), similar to or denser than on metaventrite; sternites 1 and 2 mesally fused, nearly flat to concave, laterally convex; 1 similar in length or slightly longer than 2; 2 similar in length to 3 + 4; 3 and 6 similar in length; 5 as long as 2; posterior margin rounded. Tergites ( Fig. 3D View Figure 3 ) 1–6 variously incomplete or complete, with a mesal longitudinal membrane (of the common type sensu Morimoto, 1962); tergites 4–6 posteriorly or posterolaterally and tergite 7 anterolaterally with conspicuous strigate-sculptured regions (presence and position variable), each covered with densely arranged, appressed setae, rarely simple ( Fig. 3D, E View Figure 3 ); tergite 7 covered by elytra. Tergite 8 (pygidium) partly to mostly covered by elytra; in dorsal view subrectangular, slightly to distinctly wider than long, posterior margin slightly rounded or straight, plicate, laterally rounded; posterior half to one third covered with medium to large, circular setal sockets, setae long; in lateral view medium to wide, orientated along the same plane of tergites.

Terminalia: sternum 8 ( Fig. 4A View Figure 4 ) consisting of two equilaterally triangular sclerites (hemisternites) connected by a narrow membrane, without apodeme (= spiculum relictum sensu Thompson, 1992); distal angle glabrous or with three to eight large setae. Sternum 9 (spiculum gastrale) ( Fig. 4B View Figure 4 ) with basal plate divided in an entire, variously shaped basal portion and a subquadrate, plicate, mesally projected apical portion, each portion separated yet connected via a membrane; apodeme similar throughout, 1.0– 2.5¥ width of aedeagal apodemes, apically widened. Tegmen ( Fig. 4C View Figure 4 ) with basal piece usually Y-shaped (tegminal plate reduced), if developed (basal piece O-shaped), then dorsally not closed ( Fig. 11D View Figure 11 ); tegminal apodeme subrectate to deflexed, between one quarter and one times length of aedeagus. Aedeagus ( Fig. 4D View Figure 4 ) in dorsal view with lateral margins subparallel throughout, short to long, l/w = 1.7–5.0, with paired, lateral, longitudinally orientated, variously shaped plates, basal margin narrowly or widely rounded, apex wide rounded, conical or subrectate ( Fig. 11C View Figure 11 ), simple ( Fig. 24B View Figure 24 ), mesally projected (projection short to medium, Figs 4D View Figure 4 , 13C View Figure 13 ) or opened ( Fig. 30B View Figure 30 ); ostium subapical; tectum usually membranous, simple or mesally with a weakly sclerotized tissue constituted of dense papillae or sparsely arranged spines; endophallus with one to numerous variously orientated and positioned, usually heavily sclerotized sclerites, sometimes extended basad of aedeagus (in resting position); in lateral view slender ( Fig. 18C View Figure 18 ) or wide ( Fig. 4D View Figure 4 ), dorsal margin curved, ventral margin distinctly deflexed, or subrectate and slightly sinuate, apex variously shaped; aedeagal apodemes in lateral view basally deflexed, apically slightly to distinctly sinuate.

Female: Length 1.7–3.0 mm, width 0.8–1.4 mm, l/w = 1.7–2.3. Rostrum ( Fig. 31A View Figure 31 ) slightly longer than in males, 0.4–0.7 mm; r/p = 1.1–1.7, stout, dorsally and ventrally slightly arcuate, simple above antennal insertion (as opposed to slightly tumescent in males), apex continually curved (distad of antennal insertion), basally pilose (setae short), glabrous distad of antennal insertion, punctulate, dorsolateral margins smooth, parallel; gular sutures extending along basal two thirds of rostrum ( Fig. 23B View Figure 23 ); antennal insertion positioned near midpoint, scrobe nearly straight, either acutely or not projected to apex of rostrum, basally reaching eye. Antennae usually as in males. Head yellowish to dark brown, frons setae converging towards midpoint. Eyes rounded, distant from anterior margin of pronotum by nearly half of their diameter. Pronotum as in male, yellowish brown to dark brown, l/w = 0.6–0.8, anterior margin 0.5–0.8¥ width of posterior margin; lateral margins rounded. Prothorax laterally pilose, either conical, subquadrate, or subrectangular, procoxal cavities contiguous; mesoventrite glabrous, mesoventral process rounded. Prothoracic legs similar in length to meso- and metathoracic legs, f/p = 1.1–1.6; protibia slightly shorter than profemur, t/f = 0.7–0.9. Elytra as in males, l/w = 1.3–1.8. Wings as in males. Abdominal sternites 1 and 2 fused (suture slight), convex; sternites 1–5 pilose, sternite 5 slightly shorter to longer than 3 + 4, posterior margin rounded. Tergites 1–6 variously incomplete; tergites 4-7 with strigate-sculptured regions as in males (or tergite 7 simple); tergite 8 ( Fig. 5A View Figure 5 ) entirely covered by elytra, semilunar, semicircular, or subrectangular ( Fig. 19A View Figure 19 ), even (as opposed to punctulate); posterior margin rounded, plicate, or simple, setose; anterior margin widely rounded or emarginate; lateral margin simple or folded, rounded.

Terminalia: s ternum 8 ( Figs 5B View Figure 5 , 26A View Figure 26 , 33B View Figure 33 ) with lamina typically bifurcated, mesally projected ( Fig. 5B View Figure 5 ) or emarginate ( Fig. 33B View Figure 33 ) when bifurcated; furcal arms explanate, diverging, distally widened, distal margin usually concave, inner and outer corners of furcal arms rounded and projected, inner corners usually with one to 20 setae (cf. Velázquez de Castro et al., 2007); apodeme slender, subrectate and usually long. Coxites ( Figs 5C View Figure 5 , 12B View Figure 12 ) slightly shorter to slightly longer than apodeme of sternum 8, elongate, anteriorly widened, posteriorly gradually narrowed, styli slender and elongate, apically with one to four long setae. Spermatheca ( Fig. 5D View Figure 5 ) C-shaped, deflexed; corpus slightly to distinctly swollen ( Figs 5D View Figure 5 , 33C View Figure 33 ) or tubular ( Fig. 19C View Figure 19 ); apex of cornu narrowly or widely rounded; ramus (opening to spermathecal gland) and collum (opening to spermathecal duct) narrowly to widely separated, protruded or not.

Etymology: The name Azotoctla is used as a noun in nominative singular. It originates from the Aztec word ‘Azotochtli’, meaning turtle-rabbit and referring to armadillos ( McBee & Baker, 1982). This name is chosen in reference to the compact and convex habitus of the adult weevils, which in addition share a Neotropical origin with their mammalian namesakes ( Patterson & Pascual, 1972).

Type species: Azotoctla gomezi , by present designation.