Ptychoderes Schoenherr, 1823
Mermudes, José Ricardo M. & Napp, Dilma Solange, 2006, Revision and cladistic analysis of the genus: Ptychoderes Schoenherr, 1823 (Coleoptera, Anthribidae, Anthribinae, Ptychoderini), Zootaxa 1182 (1), pp. 1-130: 14-20
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|Ptychoderes Schoenherr, 1823|
Ptychoderes Schoenherr, 1823: 1135 (non Ptychoderes sensu Schoenherr, 1833 ); Schoenherr, 1833: 3, 120 (description); Dejean, 1837: 256 (cat.); Schoenherr, 1839: 156; Labram & Imhoff, 1842: fasc. 10; Jekel, 1849: 10 (list.); Jekel, 1855: 41; Lacordaire, 1866: 488; Gemminger & Harold, 1872: 2722 (cat.); Bovie, 1906: 219 (cat.); Jordan, 1906: 302; Wolfrum, 1929: 5 (cat.), 1953: 3 (cat. supl.); Blackwelder, 1947: 765 (cat.); AlonsoZarazaga & Lyal, 1999: 33 (cat.); Rheinheimer, 2004: 10 (cat.).
Anthribus (Ptychoderes) ; Schoenherr, 1826: 34.
Type species: Macrocephalus nebulosus Olivier, 1795 , by original designation.
Size variable (7–24 mm). Body elongate, about three times as long as width across humeri; dorsally depressed, height at metathorax level slightly less than humeral width.
Integument black to reddishbrown, antennae and legs concolorous or slightly lighter. Dorsal vestiture: dorsum of rostrum, frons, pronotum and elytra with a wide, longitudinal median vitta formed by dense, moderately long, cylindricaldepressed scales truncate at apex; dorsal vitta usually lighter than sides of pronotum and epipleura, and suddenly narrowed just behind the middle of elytra ( Figs. 208–219 View FIGURES 208–214 View FIGURES 215–220 , 221–227 View FIGURES 221–227 ) (except P. bivittatus , Fig. 220 View FIGURES 215–220 ). Blackishbrown scales forming: two irregular vittae, one at either side of vertex extending to basal third of pronotum, another from posterior third of pronotum reaching to elytral base; numerous spots at sides of pronotum and elytra, intermingled predominantly with lightbrown scales (except P. brevis , P. viridanus , P. jekeli , P. callosus and P. jordani , with prevalenct green scales). Pygidium with suberect scales, not forming vittae. Sides of rostrum with sparse, blackishbrown, suberects scales. Ventral vestiture: at sides with scales similar to dorsal scales, but yellowishwhite to greyishwhite, always dense, with setigerous punctures each bearing a semidecumbent scale coloured otherwise; medially finely pubescent, pubescence usually dense, concolorous with the lateral scales (except for P. elongatus ). Sides of mentum densely hairy within depressions or grooves. Antennae with fine, whitish pubescence, denser at apex of segments; club with dense, shining, blackishbrown pubescence. Legs with vestiture similar to that of the sides of ventral surface intermingled with blackishbrown or pale scales (except for P. bivittatus ). Outer surface of meso and metafemora either with two dark spots or with a dark, not ringlike, postmedian strip; profemora with a single spot or lacking both spots and stripes. Anterior side of all tibiae with two spots formed by dense blackishbrown pubescence, one near base, another postmedian, both not ringlike. Apical half of protibiae with a dense strip of brown bristles extending along inner side of tarsi III. Tarsal pads of tarsi III bicolorous, with dark bristles on inner side, and pale bristles on outer side; III with compact yellowish pilosity, with a golden tint.
Head longer than wide; vertex convex or almost flat; surface punctaterugose at sides behind eyes, punctures dull, interstices shining; posteroventral margin corrugated. Eyes lateral ( Figs. 9 View FIGURES 1–11 , 12, 14 View FIGURES 12–15 ), almost rounded and weakly prominent, anterior margin obliquely truncate; ommatidia moderately fine, weakly prominent; subocular pit between lower eye margin and distal area of extension of scrobe present ( Figs. 9 View FIGURES 1–11 , 18 View FIGURES 16–18 , 22 View FIGURES 22–23 ). Gula almost dull. Frons slightly longer than wide, oblique, convex or almost flat, slightly narrower near base of rostrum, with a short carina on each side of median carina (except for P. bivittatus ) slightly depressed near ocular margin. Rostrum ( Figs. 7, 9 View FIGURES 1–11 , 12–23 View FIGURES 12–15 View FIGURES 16–18 View FIGURES 19–21 View FIGURES 22–23 ) cylindrical, thickened, suboblique, dorsally depressed, narrower than head at base, widest at apex, 1.2–2.0 times as long as wide across the base, height at base subequal to basal width. Dorsal carinae three, elongate, thickened, rostrum strongly impressed at base between carinae; median carina extending from apical notch to beyond middle of frons, flattened near apical notch and weakly prominent on frons; lateral carinae slightly thicker, slightly curved and convergent, far from each other at base, extending from anterior margin of eyes to scrobes. Apex ( Fig. 13 View FIGURES 12–15 ) with wide, deep, almost round notch; dorsally, strongly impressed on each side of the notch, with short, weakly prominent carina between impressed areas and dorsal margin of scrobes. Sides of rostrum ( Figs. 9 View FIGURES 1–11 , 14 View FIGURES 12–15 ) bicarinate and strongly impressed between carinae (except males of P. virgatus ); looking tricarinate in some species because of the careniform upper margin of scrobes. Scrobes ( Figs. 9 View FIGURES 1–11 , 17 View FIGURES 16–18 ) latero apical, not prominent; anterior region, where the condyle of scape fits, narrow and shallow; posterior region conspicuously wider and deep, with a ventrally directed oblique extension, and with a distal pit for accommodation of apex of 2 nd antennomere. Mentum ( Figs. 11 View FIGURES 1–11 , 19–21 View FIGURES 19–21 ) with wide, angulate emargination, lateral lobes well developed; groove of anterior margin shallow or vestigial; median region prominent, broad, dull, glabrous, impressed or grooved on each side; maxilar sinus deep. Labrum ( Fig. 24 View FIGURES 24–36 ) convex, transverse, apical margin almost rounded. Mandibles ( Figs.25–27 View FIGURES 24–36 ) stout, wider than long, widest at base, weakly, acutely projecting at apex; inner cutting edge with a blunt tooth more developed in left mandible (except P. mixtus ); mola developed, strongly concave, dull. Maxillae ( Fig. 28 View FIGURES 24–36 ): galea cylindrical reaching apical segment of palp, with a basal, weakly sclerotized ring; inner edge densely hairy; apex rounded with long, spatulate setae; lacinia narrow, elongate, rounded at apex, inner margin densely hairy, hairs thickener and shorter apically, apex with spatulate setae; basal segment of maxillary palps short, slightly sclerotized; 2 nd
conspicuously thickened, slightly longer than apical segment; 3 rd conical; apical segment elongate, attenuate at apex, 1/3 longer than 3 rd. Labium ( Fig. 29 View FIGURES 24–36 ): prementum with apical third of palpiger free; labial palps with long, cylindrical segments, 1 st longest, 3 rd about 1/3 longer than 2 nd, rounded truncate at apex; ligula deeply notched, lateral lobes fingerlike, densely hairy, reaching apex of basal segment of palps; inner surface with sclerotized median line. Antennae stout, 11segmented, sexually dimorphic, greatly variable in length in males; scape short, cylindrical, thickened, slightly depressed dorsally;segments IIVIII swelling at apex; II short; club distinct, formed by segments IX – XI; IX almost as long as VIII (except P. crustatus ), segments X– XI conspicuously shorter; XI narrowed at apical half, rounded at apex. Prothorax ( Figs. 32–33 View FIGURES 24–36 , 37–46 View FIGURES 37–40 View FIGURES 41–42 View FIGURES 43–46 , 241–250 View FIGURES 239–250 ) from wider than long to slightly longer than wide, declivous behind antebasal carina; sides declivous, slightly convergent anteriorly from antebasal carina (except P. jordani , P. crustatus and P. longicollis ). Disk of pronotum with wide medial depression (except P. bivittatus , Fig. 248 View FIGURES 239–250 ) tuberculate at middle. Center of pronotum usually with transverse rugae ( Fig. 38 View FIGURES 37–40 ) not reaching anterior and posterior margins. Antebasal carina ( Figs. 37, 39 View FIGURES 37–40 , 41 View FIGURES 41–42 ) interrupted at middle. Lateral carina ( Fig. 43 View FIGURES 43–46 ) elongate, almost reaching anterior margin of prothorax. Secondary carina ( Figs. 37, 39–40 View FIGURES 37–40 ) elongate, smooth, and laterobasal carina ( Figs. 43–44 View FIGURES 43–46 ), conspicuously prominent. Antebasal, lateral and laterobasal carinae with bract units, more evident in the latter ( Figs. 41–42 View FIGURES 41–42 , 44–45 View FIGURES 43–46 ). Posterodorsal margin prominent, with setiferous punctures ( Figs. 40 View FIGURES 37–40 , 44 View FIGURES 43–46 ). Sides of prosternum with deep setiferous punctures, each provided with a colored scale ( Fig. 46 View FIGURES 43–46 ). Intercoxal process of prosternum triangular, reaching the middle of anterior coxal cavities; obliquely elevated in relation to prosternum (except P. nebulosus and P. elongatus ). Antecoxal prosternal furrow ( Figs. 253–262 View FIGURES 251–262 ) variable (vestigial in P. nebulosus and P. elongatus , Figs. 261, 262 View FIGURES 251–262 ). Procoxae globular, not prominent. Proendosternite ( Fig. 30 View FIGURES 24–36 ) developed, lateral projections far from each other, upwards directed, somewhat expanded externally at apex. Mesoscutum ( Fig. 34 View FIGURES 24–36 ) not raised at anterior and lateral margins, the former rounded between lateral projections, these short and tapered; disc minutely, densely granulated. Scutellum reduced, quadrangular to suboval. Mesosternum ( Figs. 35, 36 View FIGURES 24–36 ) convex, almost dull, with a glabrous vitta on each side, depressed near anterior margin and irregulary impressed near base of intercoxal process (except P. nebulosus and P. elongatus ). Mesepisternum developed ( Figs. 35, 36 View FIGURES 24–36 ). Intercoxal process of mesosternum ( Figs. 35, 36 View FIGURES 24–36 ) projected, 2/3 as wide as mesocoxa. Mesocoxae globular. Mesendosternite ( Figs. 47 View FIGURES 47–54 ) elongate, directed towards mesepimeron, slender apically, with well developed projections for tendon insertion. Metasternum ( Figs. 35, 36 View FIGURES 24–36 ) flat, slightly longer than mesosternum, grooved along anterior margin, groove deeper at middle, shallower on sides (except P. elongatus ). Metepisternum ( Fig. 36 View FIGURES 24–36 ) almost flat, wide and depressed anteriorly. Metendosternite ( Figs. 48–50 View FIGURES 47–54 ) with a narrow, elongate peduncle, lateral lamina perpendicular to peduncle, slightly longer than lateral arms which are divergent, widely separated and oblique to lateral lamina.
Elytra ( Figs. 208–227 View FIGURES 208–214 View FIGURES 215–220 View FIGURES 221–227 ) elongate, about 1.2–2.5 times as long as wide across humeri, subconvex, either slightly impressed along interstriae 2 or strongly depressed between suture and interstriae 3; basal margin prominent and thickened. Humeri prominent and rounded. Surface transversely rugose, punctures organized in rows. Epipleura declivous, deeply grooved at basal third (except P. elongatus ), outer margin sinuous, prominent ( Fig. 230 View FIGURES 228–238 ). Apex almost rounded (except males of P. depressus , P. elongatus and P. nebulosus , Fig. 229 View FIGURES 228–238 ), apical margin prominent (except P. viridanus ); apical declivity variable, tubercles present or absent, or interstriae 3, 5, 7 and 9 swollen ( Figs. 229–230, 232–234 View FIGURES 228–238 ). Wings ( Fig. 51 View FIGURES 47–54 ) developed, 3 times as long as width at base; apical part, from apex of Rs to apex of wing, about half as long as the basal part from the base of wing to apex of Rs; anal lobe poorly defined; subcosta (Sc) moderately short, about 1/3 as long as the radial ( RP); radial cell (Rc), if present, rectangular, poorly developed; radial sector (Rs) developed; transverse radialmedia (rm) sinuous, weakly sclerotized or vestigial; media (M) developed; 1 st branch of 1 st anal (1A 1) at least 1/3 longer than 2 nd branch (1A 2); 2 nd anal (2A) elongate; 3 rd anal (3A) about half as long as 2A, weakly sclerotized or vestigial; 4 th anal (4A) missing; jugal (J) vestigial; anal cell missing.
Legs ( Figs. 55 View FIGURES 55–57 , 58 View FIGURES 58–60 , 61 View FIGURES 61–63 , 69 View FIGURES 64–69 , 228–230 View FIGURES 228–238 ) short. Femora subclavate, club of profemora stronger. Mesofemora slightly longer than pro and metafemora. Apical outer half of femora with glabrous areas ( Figs. 55–63 View FIGURES 55–57 View FIGURES 58–60 View FIGURES 61–63 ): somewhat rounded or oval, finely punctate on profemora (except P. brevis and P. jekeli ), oval to fusiform, densely, finely punctate on meso and metafemora (except P. jekeli ). Inner surface of femora with narrow, elongate, glabrous area with a row of fine punctures. Tibiae from weakly to strongly depressed, slightly enlarged apically. Tarsomeres I and II subequal in length. Claws appendiculate, three times as long as the internal process.
Abdomen. Ventrite I slightly shorter than II; intercoxal process triangular, grooved along anterior margin (except P. elongatus , P. crustatus e P. brevis ), this prominent and thickened, groove deeper at middle, shallow and sinuous at sides. Setiferous sex patches of males, if present, variable in shape ( Figs. 236–238 View FIGURES 228–238 ). Ventrites II – III subequal in length. Ventrite V in males slightly shorter than IV, in females, always longer; lateroapical angles not projected (except females of P.brevis ); apex slightly emarginate or truncate. Pygidium ( Figs. 52–54 View FIGURES 47–54 , 235 View FIGURES 228–238 ) subvertical, subconvex before middle or depressed lenghtwise, truncate or rounded at apex; impressed at middle near apex in females (except for P. brevis ). Hindgut with sclerotized rectal plates ( Figs. 198–201 View FIGURES 196–201 ): dorsal longest; lateral slightly shorter, ventral conspicuously shorter; rectal loop oblique and straight, either conspicuously projected dorsally or not.
Male terminalia ( Figs. 71–126 View FIGURES 70–84 View FIGURES 85–112 View FIGURES 113–126 , 130–184 View FIGURES 127–141 View FIGURES 142–157 View FIGURES 158–176 View FIGURES 177–195 ). Tergite VIII as wide as to slightly wider than long, more sclerotized at sides and apex; sides subparallel or convergent. Sternite VIII membranous, transverse, with apical lobes poorly to well defined, pigmented and with long, dense hairs; apodeme triangular, short or very much reduced (vestigial). Sternite IX with apodeme (= spiculum gastrale) variable in length. Tegmen with apodeme about as long as the sclerotized ring, this emarginate before apex; parameres fused, wider at base, variable at apex. Aedeagus elongate, slightly curved, body shorter than half length of apodemes; arc between apodemes present or absent, bridge missing; tectum twice as long as wide, barely shorter and slender than body of pedon; lateral membrane lobed and evenly pigmented; pedon continuous with apodemes, its sides gradually convergent to rounded apex; apodemes weakly enlarged basally. Internal sac variable in length; distal 2/3 spiculated in variable areas; sclerotized pieces from developed to absent; proximal 1/3 spiculated, with minute, weakly pigmented areas.
Female terminalia ( Figs. 185–192 View FIGURES 177–195 ). Tergite IX elongate, barely convergent apically; lateral rods 3 times as long as the ovipositor, strongly narrowed and sclerotized, slightly enlarged at base. Tergite VIII from barely to conspicuously longer than wide; sternite VIII shorter than or about as long as apodeme. Ovipositor with body distinct from lateral rods; apex strongly sclerotized forming a toothed plate, with 3–4 teeth; stylus short, cylindrical; lateroventral lobes membranous at apex; ventral median lobe bilobate or acuminate; median rods narrow or thickened, fused along proximal half; bursa copulatrix developed, ventral lobe present or absent, insertion of spermathecal duct close to median oviduct, insertion region not sclerotized; spermatheca reniform, with short or elongate tubular gland.
Based on the cladistic analysis, Ptychoderes is here defined by the following synapomorphies (character number and states are discussed below in the section "characters"): presence of a pale vitta formed by whitish scales intermingled with differently coloured scales, extending dorsomedially along the rostrum, prothorax and elytra (1 1); anterior side of the tibiae with two not ringlike spots, one near base and another postmedian, formed by dense, dark brown pilosity (6 2); rostrum with a prominent, thickened medial carina, extending from the apical notch, where it is flat, to beyond the middle of frons (9 1); dorsolateral carinae of rostrum thickened, rounded and convergent, far from medial carina at the base of rostrum, slightly curved and closer to medial carina near scrobes (10 2); extension of the scrobe from shallow to deep in both sexes (11 2); mentum without sexual dimorphism, medially widely raised, dull and glabrous, either grooved or impressed at sides (12 1); frons with short carinae (13 1); central depression of pronotum deeper behind middle (18 3); tubercle inside the depression transverse (20 2); prothorax with a prominent second lateral carina (22 2); prosternum deeply grooved in front of procoxae, the anterior margin almost straight or sinuous (23 3); elytra impressed along the interstria 2 (27 1); ovipositor quadridentate, two distal teeth, convex, developed and similarly sized, one proximal dorsolateral moderately developed, and another proximal, ventrolateral, minute (47 1); rectal loop oblique, straight, weakly projected dorsally (50 1).
Ptychoderes is the sister group of Unanthribus Mermudes, 2003 . They differ in the following features (condition present in Unanthribus indicated in parenthesis): 1) frons near the base of rostrum almost as wide as the maximum width of eye (narrower); 2) rostrum impressed at base between the carinae and at sides, the median carina thickened (weakly impressed only between carinae, median carina narrow); 3) antennae of males slightly longer than body, exceeding the elytral apex by two segments (X+XI), segments II–V cylindrical (twice as long as body, segments II–V thickened and depressed); 4) pronotal depression deep (shallow); 5) prosternal furrow near procoxae (far from procoxae); 6) setigerous punctures at sides of prosternum coarse, as well as those beyond pleural suture (fine beyond pleural suture); 7) elytra depressed dorsally, not raised at base (elytra conspicuously more convex, raised at base); 8) lateral membrane of the internal sac uniformly pigmented (pigmented only at margins).
Sexual dimorphism. In males, the rostrum is always longer than the width across the base; the antennae are longer, reaching at least, the elytral basal third, at most exceeding the elytral apex; the elytra more evidently depressed; and ventrite V shorter than IV. In addition, males of P. rugicollis , P. callosus , P. viridanus , P. antiquus , P. nebulosus , P. elongatus , P. jordani and P.mixtus with setiferous sex patches on ventrite I.
In females, the rostrum is shorter, the distance between scrobe and eye less than the maximum eye width; antennae shorter, segments II–VIII conical, club conspicuously thickener; prothorax from slightly longer than wide to wider than long; elytra more convex; and ventrite V longer than IV. Also, the pygidium is strongly impressed at middle near apex (except P. brevis ), and ventrite V expanded at sides in P. antiquus , P. nebulosus and P. elongatus .
The variability in both body size and proportional antennal length in males of Ptychoderes has been reported by several authors ( Jekel 1855; Lacordaire 1866; Jordan 1906). The present work corroborates these authors mainly with reference to the antennae. Medium sized males with moderately long antennae have the antennal segments slightly shorter, but their proportion (except for the scape) is similar to the observed in large males. Otherwise, the antennae in small males are slightly longer than in females, but the length of the antennal segments is similar to females. Also, some species may have a transverse carina between the lower lateral carina of rostrum and the extension of scrobe. When present, this carina may be entire or incomplete.
In the Neotropical region, from Mexico to Argentina, reaching Ecuador to the west, and Brazilian Atlantic coast to the east.
The biology of the species of Ptychoderes is poorly known. Costa et al. (1988) described and illustrated larva and pupa of Ptychoderes elongatus ( Germar, 1824) . Larvae, pupae and adults of this species were found inside vertical, cylindrical galleries (diameter about 7 mm) excavated in the duramen of newly fallen trunks. The last instar larvae make a thin, pergaminaceous cocoon and the pupal chambers are located underneath the bark. The pupae remain vertically inside the cocoon with the pronotal sclerites directed upwards. Well sclerotized and pigmented adults were found inside the pupal chambers.
Howden (1992) observed females of Ptychoderes rugicollis Jordan, 1895 ovipositing inside vertical holes of 10 mm depth, which had been previously bored with the toothed plate of ovipositor. She also reported that the males, before mating, apparently clean the sawdust remains from the females’ abdomen with the protarsi pads. They also protect the females during oviposition, placing the antennae or the rostrum on the females’ back.
Gombauld & Duranton (1996) recorded Loreya arborescens , Miconia sp. ( Melastomataceae ), Eperua rubiginosa ( Caesalpiniaceae ), Licania alba ( Chrysobalanaceae ), Ocotea rubra ( Lauraceae ) e Tapirira guianensis ( Anacardiaceae ) as host plants of P. viridanus Boheman, 1833 . According to Maes (pers. comm.), he collected females of P. bivittatus that emerged from decayed stems of Ficus insipida ( Moraceae ) previously cut by Acrocinus longimanus ( Cerambycidae , Lamiinae), and observed adults of that species on branches while they were green.
Magno (pers. comm.) colected adults of P. magnus , P. callosus , P. virgatus , P. viridanus , P. jordani and P. nebulosus on trunks of newly cut down trees, in Amapá state, northern Brazil. O’Brien (1998) noticed different ways of collecting specimens of Ptychoderes : P. bivittatus at night, P. tricostifrons attracted by UV and HgV light, and P. rugicollis , with beating umbrella. According to label data, a male of P. nebulosus was collected on Virola ( Myristicaceae ), a male of P. antiquus , on a dead avocado tree ( Persea , Lauraceae ), and some specimens of P.crustatus , P. mixtus , P.callosus , P. rugicollis and P. antiquus , in light traps.
Royal British Columbia Museum - Herbarium
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Ptychoderes Schoenherr, 1823
|Mermudes, José Ricardo M. & Napp, Dilma Solange 2006|
|Schoenherr, C. J. 1826: 34|
|Rheinheimer, J. 2004: 10|
|Alonso-Zarazaga, M. A. & Lyal, C. H. C. 1999: 33|
|Blackwelder, R. E. 1947: 765|
|Wolfrum, P. 1929: 5|
|Bovie, A. 1906: 219|
|Jordan, K. 1906: 302|
|Gemminger, M. & Harold, E. 1872: 2722|
|Lacordaire, T. 1866: 488|
|Jekel, H. 1855: 41|
|Jekel, H. 1849: 10|
|Schoenherr, C. J. 1839: 156|
|Dejean, P. F. 1837: 256|
|Schoenherr, C. J. 1833: 3|
|Schoenherr, C. J. 1823: 1135|