Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “ Anareolatae ”: Phasmatidae: Cladomorphinae) Author Frank H. Hennemann Author Oskar V. Conle Author Daniel E. Perez-Gelabert text Zootaxa 2016 4128 1 1 211 journal article 38706 10.11646/zootaxa.4128.1.1 553faca2-0799-4bbe-8b54-92960421d9c9 1175-5326 271800 B4D2CD84-8994-4CEF-B647-3539C16B6502 5.5. Genus Haplopus Burmeister, 1838 Type-species: Cyphocrana micropterus St. Fargeau & Audinet-Serville, 1828: 445 , by monotypy. Haplopus Burmeister, 1838: 560 , 576. [Emendation and replacement name for Aplopus Gray, 1835 ] De Haan, 1842: 107, 127. Westwood, 1859: 85 (in part). Saussure, 1871–1872 : 192. Stål, 1875: 31. Bolivar, 1888: 140. Kirby, 1904a: 363 (in part). Redtenbacher, 1908: 429 (in part). Zompro, 2005a: 30. Otte & Brock, 2005: 150 (in part). Aplopus Gray 1835: 34 . [Preoccupied by Aplopus Dejean, 1821 ( Coleoptera : Curculionidae )] Rehn, 1904: 63 (in part). Caudell, 1905: 83. Rehn, 1909: 200. Karny, 1923: 240. Rehn & Hebard, 1938: 52. Adams & Adams, 1982: 263. Brock, 1998c: 33. . Fargeau & Audinet-Serville , 1828 : 445 . Diapherodes Gray, 1835 : 33 (in part) . Moxey, 1972: 85 (in litt.; in part). [As a synonym of Aplopus Gray, 1835 in error] Phasma , Stoll, 1813: 61 & 1788: pl. 21: 77 (♀). [ Not: Haplopus , Shelford, 1908: 355 , 365. → Pterinoxylus Audinet-Serville, 1838 ] Description: ♂♂, ♀♀. Medium-sized to large (body length ♀♀ (including subgenital plate) 115.5–172.0 mm, ♂♂ 82.0–103.0 mm), slender and elongate Haplopodini, ♀♀ brachypterous, ♂♂ brachypterous or with well developed alae. Body ± cylindrical in cross-section. Colouration of ♀♀ various shades of brown, straw and grey, rarely greenish. ♂♂ rather colourful insects and sometimes multi-colorous but mostly green or greenish brown; body surface glabrous. Head indistinctly longer than wide and globose, vertex ± convex and distinctly bi-spinose or bicornute; the dextral spine larger than the sinistral. Antennae filiform in ♀♀ and considerably thickened and robust in ♂♂; in ♀♀ almost as long as head and complete thorax combined, in ♂♂ reaching at least to abdominal segment III. Pronotum about as long as head and with a ± distinct pair of anterior tubercles or spines; otherwise ± tuberculose. Mesothorax elongate and parallel-sided, at least 2x longer than head and pronotum combined. Mesonotum in ♀♀ rather irregularly and to a variable degree armed with granules or spiniform tubercles, in ♂♂ armed with a variable number of distinct spines. Mesopleurae smooth in ♂♂ and with a longitudinal row of tubercles in ♀♀. Meso- and metasternum irregularly set with a variable number of granules or tubercles; metasternum may be smooth in ♂♂. Tegmina broadly oval and with a moderately distinct central hump in ♂♂; ± reaching posterior margin of metanotum. Alae of ♀♀ ± as long as tegmina; of ♂♂ variable, either slightly shorter than tegmina or reaching as far as abdominal segment VI. Anal region reticulate with distinct brown to black radial and transverse veins, translucent pink or red in ♂♂. Abdomen considerably longer than head and thorax combined. Median segment longer than metanotum, smooth. Segments II–VII distinctly longer than wide. Tergites unarmed in ♂♂, II–V often with a ± distinct pair of posterior spines in ♀♀. VII parallel-sided, with lateral margins gently rounded posteriorly, or with a small posterolateral lobe or tooth. VIII and IX narrower (♀♀) or slightly wider (♂♂) than previous segments. Sternites II–VII smooth. Praeopercular organ of ♀♀ formed by a ± distinct, elongate and longitudinal wart-like median tubercle close to posterior margin of sternum VII ( Figs. 288–289 ). Anal segment with a longitudinal median carina and the posterior margin with a ± distinct median notch; tapered in ♀♀. Epiproct very small in ♂♂, triangular or shield-shaped and ± projecting beyond anal segment in ♀♀. Vomer of ♂♂ well developed, broadened basally and with an elongate, ± papillate terminal hook ( Figs. 304–310 ). Cerci straight, small and tapered in ♀♀, obtuse, about as long as anal segment and cylindrical in ♂♂. Poculum of ♂♂ convex, cup-like, often longitudinally carinate and with a blunt hump or spine basally. Subgenital plate of ♀♀ long, ± lanceolate with the apex narrow and ± pointed; projecting over apex of abdomen by at least the combined length of tergites IX and X. Legs of moderate length, pro- and mesofemora shorter than mesothorax in ♀♀, about as long or a little longer in ♂♂. Hind legs ± reaching (♂♂) or clearly not reaching (♀♀) to the apex of abdomen. Two outer ventral carinae of meso- and metafemora each with 1–3 sub-apical spines, the medioventral carina of these femora with a longitudinal row of 4–7 ± distinct spines. Dorsal carinae of all femora unarmed, those of the meso- and metafemora often forming a shallow to acutely triangular sub-apical tooth or lobe. Tibiae unarmed except for a few small teeth in the apical portion of the medioventral carina; anterodorsal carina in ♀♀ sometimes gently elevated sub-basally and sub-apically. All basitarsi slender and rather elongate, longer than following two tarsomeres combined. Eggs: Medium-sized (capsule length 3.1–4.4 mm ), capsule ovoid, round in cross-section and> 1.5x longer than wide, polar area rounded. Capsule surface minutely granulose or rugulose. Micropylar plate elongately heartshaped, the anterior end narrowed and the widened posterior end with a prominent median notch;> 2/3 as long as capsule. No median line ( Fig. 40 ). Operculum with a prominent, raised, squamiform, hat or knob-like capitulum. Colouration of capsule variable, often prettily mottled in various shades of brown, straw, cream, grey and orange. Differentiation: Very close to Parhaplopus n. gen. but differing by: the convex and distinctly bi-cornute vertex and anterior pair of tubercles or spines on the pronotum of both sexes; slender mesothorax and slender probasitarsus of ♀♀, as well as the reticulate anal region of the alae; larger and conspicuously thickened cerci and more convex poculum with a ± distint central spine of ♂♂. The eggs clearly differ from those of Parhaplopus n. gen. by having the capsule surface much less sculptured and possessing a ± distinct conical, hat or knob-like capitulum. For a more detailed differentiation see Table 29 . Comments: Burmeister (1838: 576) established Haplopus as an amendment to Aplopus Gray, 1835 based on the original Greek pronounciation. This was however an unjustified emendation (ICZN, 1999: Article 32.5) and hence Haplopus is a junior objective synonym of Aplopus Gray (ICZN, 1999: Article 33.2.3). Aplopus Gray, 1835 however is preoccupied and a junior homonym of Aplopus Dejean, 1821 ( Coleoptera : Curculionidae ) and therefore replaced by Haplopus Burmeister, 1838 (Zompro, 2005a: 30) , which is an available name (ICZN, 1999: Article 19.1). In the composition it was used by most former authors (e.g. Kirby, 1904a; Redtenbacher, 1908 or Moxey, 1972 in litt.) Haplopus Burmeister, 1838 was not a natural grouping, which had already been suggested by Rehn & Hebard (1938: 52). This is obviously seen in two clearly distinct types of eggs and several obvious distinctive features of the insects. Eggs of typical Haplopus exhibit a more or less prominently conical, knob or hat-like capitulum and have a rather faintly coriaceous capsule surface, whereas those of the second group lack a central capitulum and have the capsule surface prominently sculptured with raised ridges and humps. Species belonging to the second group are here transferred to the “ jamaicensis species-group” of Diapherodes Gray, 1835 . One species formerly attributed to Haplopus , i.e. Haplopus dubius (Gray, 1835) , is definitely misplaced and here removed from the genus. The exact systematic position however remains uncertain (→ 5.9). Distribution ( Fig. 376 ): Bahamas , Florida Keys, Dry Tortugas , Greater Antilles (Hispaniola and Puerto Rico ), Gonaive, Mona Island east of Hispaniola, Navassa Island west of Hispaniola, Virgin Islands (St. Croix, St. Thomas, St. John , Anegada , Guana, Little Thatch, Moskito and Tortola ), Cayman Islands and Swan Islands. Although no definite records are available so far, the genus is most certainly also represented on the Caicos Islands north of Hispaniola ( Fig. 312 ). Species included: 1. Haplopus bicuspidatus de Haan, 1842: 128 . [Distribution: Hispaniola] 2. Haplopus brachypterus n. sp. [Distribution: Hispaniola] 3. Haplopus intermedius n. sp. [Distribution: Hispaniola] 4. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828: 445) [ Cyphocrana ]. = Phasma angulata Stoll, 1813: 61 , pl. 21: 77 (♀). = Haplopus bituberculatum de Haan, 1842: 128 . n. syn. = Haplopus cythereus Westwood, 1859: 86 , pl. 18: 5, 5a & b (♂). n. syn. = Haplopus ligiolus Redtenbacher, 1908: 432 . n. syn. = Haplopus ligia Westwood, 1859: 89 , pl. 1: 1, 1a, 1b (♂) & 2, 2a (♀). n. syn. = Haplopus obtusus Redtenbacher, 1908: 431 . n. syn. = Diapherodes spinipes Gray, 1835: 34 . n. syn. [Distribution: Hispaniola, Mona Id., Puerto Rico , St. Croix & Virgin Islands ] 5. Haplopus scabricollis (Gray, 1835: 34) [ Diapherodes ]. = Aplopus mayeri Caudell, 1905: 83 . n. syn. = Aplopus similis Caudell, 1904: 65 . n. syn. [Distribution: Bahamas , S-Florida, Florida Keys, Dry Tortugas , Navassa Id., Cayman Ids. & Swan Ids.] 6. Haplopus sobrinus n. sp. [Distribution: Cuba ] 7. Haplopus woodruffi n. sp. [Distribution: Cayman Islands ( Cayman Brac)] Keys to the species of Haplopus ♀♀ 1. Vertex convex; alae as long as or longer than tegmina and covering> 2/3 of median segment.......................... 4 - Vertex gently rounded; alae at best equal in length to tegmina and covering at best ½ of median segment................ 2 2. Spines of the vertex blunt or obsolete; anal region of alae not red................................................ 3 - Spines of the vertex acute ( Fig. 268 ); alae much shorter than tegmina and covering only 1/3 of median segment, anal region red; epiproct distinct, shield-shaped ( Fig. 273 ); Bahamas , Florida Keys, Dry Tortugas , Navassa Island , Cayman Islands & Swan Islands................................................................................. scabricollis 3. Spines of the vertex obsolete ( Fig. 277 ); tegmina distinctly shorter than alae and covering only 1/3 of median segment ( Fig. 278 ), anal region reticulate; epiproct acutely triangular and keeled dorsally; Cuba ........................ sobrinus n. sp. - Spines of the vertex blunt ( Fig. 281 ); alae subequal in length to tegmina and covering about ½ of median segment, anal region pale transparent pink; epiproct very small ( Fig. 283 ); Cayman Brac.................................. woodruffi n. sp. 4. Abdominal tergum VII parallel-sided; epiproct small, triangular; only Hispaniola................................... 5 - Abdominal tergum VII ± broadened / expanded, or with a lobe posterolaterally; epiproct distinct....................... 6 5. Epiproct very small; anal segment with a shallow concave excavation posteromedially ( Fig. 208 ); profurcasternum tuberculate..................................................................................... brachypterus n. sp. - Epiproct acutely triangular; anal segment with a distinct, triangular posteromedian incision ( Fig. 197 ); profurcasternum smooth.................................................................................... bicuspidatus 6. Epiproct roundly triangular to semi-circular, ± shield-shaped ( Figs. 241–242 ); mesonotum <2.3x longer than head and pronotum combined; alae covering> ¾ of median segment; antennae red ventrally............................. micropterus - Supraanal plate acutely triangular ( Fig. 215 ); mesonotum 2.4x longer than head and pronotum combined; alae covering <¾ of median segment; antennae brown........................................................... intermedius n. sp. ♂♂ 1. Brachypterous, alae not reaching to apex of median segment................................................... 2 - Alae well-developed, exceeding abdominal tergum IV........................................................ 5 2. Large (body> 80 mm ); horns of the head prominent and acute; three terminal abdominal segment scarcely broader than previous; tergum VII with a posterolateral tooth or lobe........................................................... 3 - Small (body < 80 mm ); horns of the head very small and blunt ( Fig. 282 ); three terminal abdominal segments strongly swollen and club-like, distinctly broader than previous ( Fig. 285 ); tergum VII slender; Cayman Brac.............. woodruffi n. sp. 3. Alae scarcely shorter than tegmina and covering roughly ½ of median segment, anal region reticulate; no white markings on pronotum and median segment;.......................................................................... 4 - Alae considerably shorter than tegmina and covering only 1/3 of metanotum, anal region red; lateral margins of pronotum white; median segment with a longitudinal white marking; Bahamas ; Florida Keys; Dry Tortugas , Navassa Island , Cayman Islands & Swan Islands......................................................................... scabricollis 4. Abdominal tergum VII slightly expanded posteriorly ( Fig. 210 ); dorsal carinae of meso- and metafemora smooth; vomer with base large and terminal hook only about 1.2x longer than basal portion ( Fig. 350 ).................... brachypterus n. sp. - Abdominal tergum VII with a distinct triangular posterolateral lobe ( Fig. 217 ); dorsal carinae of meso- and metafemora with a low triangular tooth sub-apically; vomer with base fairly small and the terminal hook papillate and almost 2x longer than basal portion ( Fig. 351 )....................................................................... intermedius n. sp. 5. Head with cheeks white ( Figs. 231–233 ); alae reaching to abdominal segment VI; anterior of pronotum with a distinct pair of spines ( Figs. 231–233 )........................................................................ micropterus - Head uniformly green (sometimes with a faint yellow marking on frons, Fig. 194 ); alae at best reaching half way along abdominal segment V; anterior of pronotum merely with a pair of small tubercles ( Fig. 194 )....................... bicuspidatus