Pterinoxylus Serville, 1838

Hennemann, Frank H., Conle, Oskar V., Valero, Pablo & Nishida, Kenji, 2022, Studies on Neotropical Phasmatodea XXV: Revision of Pterinoxylus Serville, 1838, with the descriptions of two new species from Costa Rica. (Phasmatodea: Oriophasmata: Cladomorphinae: Pterinoxylini), Zootaxa 5208 (1), pp. 1-72 : 5-9

Pterinoxylus Serville, 1838

Type –species: Pterinoxylus difformipes Serville, 1838: 227 (= Haplopus eucnemis Burmeister, 1838: 577 ), by original monotypy.

Pterinoxylus Serville, 1838: 226 .

Westwood, 1859: 89, pl. 36: 1.

Kirby, 1889: 502.

Kirby, 1904: 362.

Rehn, 1904: 61.

Redtenbacher, 1908: 427.

Shelford, 1909: 365.

Rehn, 1957: 185, pl. 21.

Zompro, 1997: 180, fig. 2a–b.

Bragg, 2001: 642.

Langlois, Lelong, Rastel, Polidori, & Dorel, 2000: 50, figs. 20a–g & photos 19–22.

Lelong, Langlois, Rastel, & Dorel, 2003: 75, figs. 26a–g & photos 35–38.

Zompro, 2004: 320.

Lelong & Langlois, 2005: 268, figs. 25–30 & 43–46.

Otte & Brock, 2005: 294.

Conle, Hennemann & Gutiérrez, 2011: 64.

Hennemann, Conle & Perez-Gelabert, 2016: 8, 28, figs 37–44, 46.

Haplopus, Burmeister, 1838: 577 (in part).

Haan, 1842: 127 (in part). [Erroneous synonymy of Pterinoxylus ].

Redtenbacher, 1908: 429 (in part).

Description: ♀♀ / ♂♂. Medium sized to large (body lengths: ♀♀ incl. subgenital plate 116.0–196.0 mm, ♂♂ 82.0– 118.0 mm), elongate and moderately slender, stick-like Cladomorphinae with very prominent sexual dimorphism. ♂♂ with scale-like tegmina and alae exceeding abdominal tergum IV. ♀♀ much larger and much stockier, brachypterous but alae projecting over posterior margin of median segment. Alae with a tympanal area (= stridulatory organ) in the basal portion of the costal region, that is positioned between the radial and medial veins ( Figs. 20A–B); radial vein notably thickened, domed and with rough dorsal texture. Body of ♂♂ cylindrical, of ♀♀ subcylindrical. Entire body surface ± prominently rugulose and wrinkled, thorax partly granulose, tuberculose or spinose. Head longer than wide, dorsoventrally flattened and ± parallel-sided. Vertex with ± prominent tubercles or spines and occasionally convex. No ocelli. A slender and transverse gula present. Antennae moderately robust and not reaching ( ♀♀) or slightly projecting over posterior margin of mesothorax; consisting of 27–33 segments. Scapus dorsoventrally flattened and ± roundly deflexed laterally, pedicellus subcylindrical. Following antennomeres with a longitudinal median keel or bulge ventrally. Pronotum rectangular and distinctly longer than wide. Probasisternum with two rough sensory areas near lateral excavations of procoxae ( Figs. 1B–C); profurcasternum with a raised, oval central sensory area ( Figs. 1B, 1D). Mesothorax elongate, at least 3x longer than pronotum (comparatively longer in relation in ♂♂), parallelsided. Metanotum quadrate to slightly transverse. Meso- and metasternum as well as pleurae simple; granulose, rugulose and/or tuberculose. Tegmina ovate and scale-like with a rounded central hump; slightly projecting over posterior margin of metanotum. Anal region of alae dark grey to black with ± distinct transparent spots, markings or radially arranged stripes; in ♀♀ of a conspicuous convex contour when opened. Abdomen longer than head and thorax combined. Median segment about 2x longer than metanotum. Segments II–VI parallel-sided and either more than 2x longer than wide ( ♂♂) or just indistinctly longer than wide ( ♀♀). Tergum VII parallel-sided to ± laterally deflexed and sometimes with a lateral lobe ( ♀♀ in particular). VIII–X slightly narrower ( ♀♀) or a little broader than previous segments ( ♂♂). Praeopercular organ of ♀♀ indistinct and usually formed by a median tubercle close to posterior margin of sternum VII. Anal segment quadrate to slightly transverse, the posterior margin rounded with a small median indentation ( ♂♂) or broadly emarginated ( ♀♀). Epiproct small and with a longitudinal median carina, mostly concealed by anal segment in ♂♂. Vomer of ♂♂ well developed, sclerotized and broadly triangular with a single, fairly short, upcurved terminal hook ( Figs. 21A–F). Cerci very small, compressed laterally, ± carinate dorsally and ventrally and slightly incurved. Gonapophyses VIII of ♀♀ not considerably elongated or extending beyond anal segment but notably longer than gonaphophyses IX ( Fig. 1A). No gonoplacs. Gonangulum present. Subgenital plate of ♀♀ elongate, irregularly naviculate or spatulate and distinctly projecting over apex of anal segment; longitudinally keeled basally but broadened and flattened towards a rounded to angular apex. Poculum (= subgenital plate) of ♂♂ gently convex, cup-like and with a median carina; posterior margin notched medially. Legs short and robust ( ♀♀ in particular) with more or less prominently expanded or lobed carinae. Tibiae notably shorter than corresponding femora. Protibiae with broad foliaceous lobes and lamellate expansions ( ♀♀ in particular). Meso- and metafemora and corresponding tibiae with a ± distinct sub-apical dorsal lobe or tooth. Profemora shorter than mesothorax, metafemora at best reaching half way along abdominal tergum IV. Profemora compressed and curved basally and distinctly triangular in cross-section with the anterodorsal carina conspicuously raised and ± lamellate; medioventral carina indistinct and slightly displaced towards anteroventral carina. Meso- and metafemora and all tibiae trapezoidal in cross-section, the two anterior carinae strongly approaching each other. Medioventral carina of meso- and metafemora indistinct and unarmed or at best with a few minute spinules. Meso- and metatibiae shorter than corresponding femora. Tibiae without an area apicalis. Medioventral carina of meso- and metatibiae conspicuously displaced towards the anteroventral carina in the median portion. Tarsi short and stout, probasitarsus with a distinct rounded dorsal lobe, that is formed by the melted dorsal carinae. Meso- and metabasitarsi simple, furcate dorsally with the two dorsal carinae strongly approaching each other but well separated, and no longer than following two tarsomeres combined.

Eggs ( Figs. 22A–H): Large to very large (overall length> 6.0 mm), alveolar, at least 3x longer than wide and cylindrical to oval in cross-section. Entire capsule surface minutely punctured and strongly sculptured, with prominent raised ridges and wrinkles. Polar-area with a more or less prominent, hollow peripheral extension. Operculum circular and with a peripheral extension on outer margin similar to that of the polar-area. Micropylar plate elongate, oval and pointed anteriorly; covering about one-quarter of capsule length. Micropylar cup distinct. Internal micropylar plate open with a wide, triangular posteromedian gap. No median line.

Comments: The genus Pterinoxylus was originally established by Serville (1838: 227) for his Pterinoxylus difformipes , described from a ♀ from “Amérique méridionale” (= South America). As this was the only species included in the genus when it was described, P. difformipes Serville is the type-species of Pterinoxylus by original monotypy. Shortly before in the same year, Burmeister (1838: 577) described Haplopus eucnemis based on a ♀ from the State of Pará, Brazil. Westwood (1859: 89) was correct in recognizing that these two taxa were conspecific and synonymised P. eucnemis with P. difformipes . This author also provided a beautiful figure of Burmeisters's holotype as well as a redescription in his famous 1859 catalogue ( Westwood, 1859, plate 36: 1). However, as Burmeister's publication predates that of Serville, P. difformipes must be regarded a synonym of P. eucnemis . Haan (1842: 127) erroneously treated Pterinoxylus as a synonym of Haplopus Burmeister, 1838 .

A third species, Pterinoxylus crassus , was described by Kirby (1889: 502) based on a ♀ from Dominica, one of the Lesser Antillean islands. Redtenbacher (1908: 428) described a fourth species, Pterinoxylus spinulosus , based on a ♂ and nymph from Panama and provided a description of the ♂ of Burmeister's P. eucnemis . A fifth but generically misplaced species, Haplopus perarmatus Redtenbacher, 1908 from Nicaragua was transferred to Pterinoxylus by Zompro (2005: 275).

Rehn (1957) briefly discussed the genus and supplied a description and illustration of the so far unknown ♀♀ of P. spinulosus Redtenbacher. The egg of P. spinulosus was described by Zompro (1997: 180) from material collected in Costa Rica and erroneously identified as P. difformipes Serville, 1838 . Langlois et al. (2000), Lelong et al. (2003) and Lelong & Langlois (2005) provided descriptions and illustrations of both sexes and the eggs of P. crassus Kirby, 1889 along with notes on the habitats and biology in Dominica and Martinique. Robinson (1970) presented information on camouflage and active defensive behaviour of P. spinulosus .

Distribution ( Figs. 31–33): Central America, northern South America and Lesser Antilles ( Dominica, Martinique & Saint Lucia).

Species included:

1. Pterinoxylus cocoense n. sp.

Distribution: Costa Rica ( Isla del Coco).

2. Pterinoxylus crassus Kirby, 1889: 502 .

Distribution: Lesser Antilles ( Dominica, Martinique & Saint Lucia).

3. Pterinoxylus eucnemis Burmeister, 1838: 577 .

= Pterinoxylus difformipes Serville, 1838: 227 .

Distribution: NE– Brazil, French Guiana & Suriname.

4. Pterinoxylus perarmatus Redtenbacher, 1908: 431 .

Distribution: Guatemala, Nicaragua, Costa Rica & Panama.

5. Pterinoxylus speciosus n. sp.

Distribution: Costa Rica.

6. Pterinoxylus spinulosus Redtenbacher, 1908: 428 , pl. 20: 3.

Distribution: Belize, Guatemala, Costa Rica, Panama, Colombia & NW– Ecuador.

Keys to the species of Pterinoxylus :

♀♀

1. Basal portion of subgenital plate convex and bulgy............................................................2

– Basal portion of subgenital plate flattened.................................................................. 4

2. Mesothorax slender and ± uniform in width; without spines.................................................... 3

– Mesothorax notably swollen medially and heavily spinose ( Fig. 16E); Costa Rica...................... speciosus n. sp.

3. Vertex with a pair of prominent, blunt, carinate horns ( Figs. 12I–J); subgenital plate extending beyondcapex of abdomen by more than combined length of two terminal terga ( Figs. 12C–D); Central America.......................... perarmatus

– Vertex multi–spinose ( Fig. 6G); subgenital plate extending beyond apex of abdomen by less than combined length of two terminal terga ( Figs. 6A–B); Lesser Antilles............................................................. .. crassus

4. Large insects (body length incl. subgenital plate> 13 cm); alae projecting considerably over posterior margin of median segment; carinae of protibiae strongly deflexed and lamellate......................................................5

– Smaller insects (body length incl. subgenital plate < 14 cm); alae hardly reaching to posterior margin of median segment ( Figs. 2A–C); carinae of protibiae just weakly expanded ( Fig. 2A); Cocos Island............................. cocoense n. sp.

5. Subgenital plate with lateral margins strongly deflexed and arcuate over entire length of plate ( Fig. 9A); posteroventral carina of protibiae very strongly deflexed and unevenly undulate ( Fig. 9G); mesotibiae with a distinct sub–basal dorsal lobe; NE–South America....................................................................................... eucnemis

– Subgenital plate with lateral margins narrowing and declining towards apex of plate ( Fig. 9A); posteroventral carina of protibiae more even in outline and widening towards apex of tibia ( Fig. 19K); sub–basal dorsalclobe of mesotibiae small to obsolete ( Fig. 19L); Central America & Colombia........................................................... spinulosus

♂♂

1. Large insects (body length> 9 cm); vertex armed; legs dentate, lobulate and/or lamellate............................. 2

– Small insects (body length < 9 cm); vertex unarmed; all legs slender, simple and destitute of teeth, lobes or lamellae ( Figs. 3A–C); Costa Rica (Cocos Island)............................................................. cocoense n. sp.

2. Vertex tuberculose, spinulose or multi–spinose...............................................................3

– Vertex armed with a pair of large, laterally compressed and tuberculose lobes ( Fig. 12H); Central America...... perarmatus

3. Mesothorax> 4x longer than pronotum, rugulose, tuberculose or spinose; no green stripe along costal margin of alae; femora brown and unicolourous.................................................................................4

– Mesothorax <3.5x longer than pronotum, heavily armed with large, blunt spines ( Fig. 15G); costal region of alae with a green longitudinal stripe in basal half; basal ¾ of femora green; Costa Rica................................ speciosus n. sp.

4. Abdominal tergum VII parallel–sided; protibiae with carinae just gently lamellate.................................. 5

– Abdominal tergum VII with lateral margins deflexed and rounded in posterior half ( Figs. 9E–F); protibiae with carinae strongly deflexed, lamellate and notably expanded apically; NE–South America..................................... eucnemis

5. Mesonotum distinctly spinulose to spinose ( Figs. 10I–J); meso– and metatibiae with a broad, dentate dorsal apical lobe ( Fig. 19M); Central America & Colombia................................................................ spinulosus

– Mesonotum ± tuberculose ( Fig. 6H); no apical lobe on meso– and metatibiae; Lesser Antilles.................... crassus

Eggs*

1. Capsule tube–like and ± round in cross–section............................................................. 2 – Capsule compressed laterally with dorsal surface strongly raised and tectiform longitudinally ( Figs. 22E–F)..... perarmatus 2. Capsule surface punctured and with ± distinctly raised ridges.................................................. 3 – Capsule surface densely rugulose/tuberculose; polar extension shorter than micropylar plate ( Fig. 3G)....... cocoense n. sp. 3. Ridges of capsule shallow; polar and opercular extensions close................................................ 4 4. Ridges of capsule very prominently raised; polar and opercular extensions open and net–like ( Figs. 22A–B)......... crassus 4. Capsule with a posterior constriction before polar extension............................................. eucnemis – Capsule without a distinct posterior constriction.................................................... ... spinulosus * Eggs of P. speciosus n. sp. are not known