Haplopus Burmeister 1838
Creators: name: Frank H. Hennemann
name: Oskar V. Conle
name: Daniel E. Perez-Gelabert
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Phasmida
Phasmatidae
Haplopus

description: Haplopodini Günther, 1953 rev. stat.
Type-genus: Haplopus Burmeister, 1838: 576.
Haplopodini Günther, 1953: 557 (in part).
Bacteridae (Sectio: Bacteriae), Brunner v. Wattenwyl, 1893: 80, 83 (in part). Bacteriinae, Kirby, 1904a: 431 (in part).
Bacterinae, Rehn, 1904: 61 (in part).
Cladoxerinae, Karny, 1923: 130 (in part).
Cranidiini, Zompro, 2005: 135 (in part). Otte & Brock, 2005: 32 (in part).
Hesperophasmatini Bradley & Galil, 1977: 188 (in part). Phibalosomini (Sectio: Phibalosomata), Redtenbacher, 1908: 399. Phibalosominae, Shelford, 1908: 343, 355 (in part). Phibalosomatinae, Moxey, 1971: 44 (in part).
Description: ♀♀, ♂♂: Small to very large (body lengths: ♀♀ incl. subgenital plate 50.0–197.0 mm, ♂♂ 60.0–126.0 mm), moderately slender to very robust and massive (♀♀ in particular) Cladomorphinae with long but thickened antennae. ♀♀ either various shades of brown and grey or bright green, ♂♂ often very colourful insects. Body cylindrical in ♂♂ and cylindrical to oval in ♀♀; in one case triangular in cross section with a prominently raised longitudinal median keel along entire dorsal body surface. Body often ± glabrous (♂♂ in particular). ♀♀ mostly brachypterous or with vestigial wings, rarely apterous, ♂♂ with scale-like tegmina and alae either well developed or shortened but rarely apterous. Head indistinctly longer than wide with the vertex ± convex and bispinose or bi-cornute; rarely flat and unarmed. No gula. No ocelli. Antennae long and projecting over mesothorax (♀♀) or median segment (♂♂); cylindrical and consisting of 55–80 segments. Antennomeres shortened indistinctly longer than wide and increasingly perlamorph; scapus slightly compressed dorsoventrally. Pro- and mesothorax to a variable degree armed with granules, tubercles or spines, rarely unarmed. Pronotum longer than wide and narrower than head. Probasisternum or profurcasternum without sensory areas. Mesothorax elongate and cylindrical (♂♂) or ± widened towards the posterior (♀♀). Mesonotum often with a cluster of tubercles or spines near anterior margin; in one case with a prominently semi-circularly raised longitudinal median keel. Mesopleurae of ♀♀ often ± expanded and armed with a longitudinal marginal row of tubercles, teeth or spines. Mesosternum simple. Median segment longer than metanotum (only in one case slightly shorter than metanotum). Abdomen longer than head and thorax combined. Tergites mostly unarmed but II–IV in ♀♀ may bear longitudinal carinae or single tooth- or crest-like lobes. Abdomen of ♀♀ ± swollen and broadened basally or pre-medially, segments II–VII of ± uniform width in ♂♂. Praeopercular organ of ♀♀ indistinct and formed by a longitudinal ridge or tubercle near posterior margin of sternum VII; in one case formed by a cluster of three granules (Figs. 326–343). Anal segment of ♂♂ slightly tectiform with the posterior margin rounded or roundly indented medially; posterior margin ventro-medially with a variable number of small in-curving denticles. Cerci at best equal in length to anal segment, cylindrical to oval in cross-section and ranging from straight to slightly in-curving. Vomer well developed and sclerotized, either roundly triangular or with the base semi-circular and the single terminal hook strongly elongated and papillate (Figs. 344–364). Poculum of ♂♂ convex, cup-like to conical and often with a ± distinct basal swelling or spiniform projection. Gonapophyses VIII of ♀♀ short and fully hidden under anal segment; not or hardly longer than gonapophyses IX. Gonoplacs lacking. Epiproct small, either hidden under anal segment or slightly projecting; shape ranging from triangular to semi-circular. Subgenital plate of ♀♀ ± projecting over apex of abdomen, often lanceolate or naviculate and keeled longitudinally. Legs rather short and stocky (♀♀) to moderately long and slender (♂♂), at least with small sub-apical ventral spines on the meso- and metafemora but often with additional teeth, lobes or expansions. Profemora curved and compressed basally; cross-section slightly triangular with the anterodorsal carina ± raised. Meso- and metafemora and all tibiae trapezoidal in cross-section. Medioventral carina of meso- and metafemora distinct and usually armed with a longitudinal row of spines; slightly misplaced towards anteroventral carina in profemora. Tibiae without an area apicalis. Tarsi simple and slender, probasitarsus occasionally with a rounded lobe dorsally and with the two dorsal carinae melted with another. Meso- and metabasitarsus sulacte dorsally with the two dorsal carinae well separated. All basitarsi at best as long as following three tarsomeres combined.
Eggs: Medium-sized to large, capsule ovoid or barrel-shaped. Capsule surface finely punctured to strongly coriaceous, rugulose or tuberculose. Micropylar plate at best ½ as long as capsule, shape variable. Internal micropylar plate open and with a widely triangular posteromedian notch, no median line (Fig. 45). Operculum either flat and with a circular rim of tubercles or ± conically raised to knob-like.
Differentiation (Table 2): Closely related to Pterinoxylini n. trib. and Hesperophasmatini with which it shares the lack of gonoplacs and the short gonapophyses VIII of ♀♀, both characters of which are likely synapomorphies of these three tribes. On this basis and as discussed above Haplopodini possibly forms a monophyletic clade together with these two tribes within present Cladomorphinae sensu lato. The lack of sensory areas on the probasisternum and profurcasternum, increasingly shortened but greater number (55–80) of antennomeres and increasingly thickened, perlamorph antennae suggest Haplopodini is the sister-taxon of Pterinoxylini n. trib. + Hesperophasmatini (Fig. 409). While the lack of sensory areas most certainly is a plesiomorphic character, the morphology of the antennae is likely to be an autapomorphy of Haplopodini. The lack of a gula is here estimated as a synapomorphy of Haplopodini + Hesperophasmatini. A confirmed definition of these character states however deserves a comprehensive phylogenetic analysis of the subfamily Cladomorphinae and supposedly closely related taxa.
Furthermore, Haplopodini differs from both tribes by the distinct medioventral carina of the profemora which is slightly displaced towards the anteroventral carina, distinct and often spinose medioventral carina of the meso- and metafemora of both sexes, as well as the ± glabrous body surface, mostly pretty colouration and transparent orange to pink anal region of the alae of ♂♂ (either plain or reticulate).
From Pterinoxylini n. trib. it also differs by: the lack of a tympanal region in the basal portion of the alae (= stridulatory organ) of ♀♀; less distinctly triangular cross-section of the profemora; not dilated or lamellate carinae of the protibiae and ovoid to barrel-shaped, much smaller eggs which lack hollow, peripheral extensions on the polar-area and operculum (→ Table 2).
From Hesperophasmatinim, with which they share the lack of a gula, members of Haplopodini may also be distinguished by the more decidedly triangular cross-section of the profemora, longer often lanceolate or naviculate subgenital plate of ♀♀ and lack of conspicuous hairy structures in the eggs. Also, Haplopodini on average are larger and less stocky than members of Hesperophasmatini.
Comments: Günther (1953: 557) originally established the tribe Haplopodini to include some of the neotropical genera of Redtenbacher's Phibalosomatini: Sectio Phibalosomata (1908: 399) and Phibalosomatini: Sectio Eurycanthae (1908: 339), this is: Haplopus Burmeister, 1838, Diapherodes Gray, 1835, Pterinoxylus Audinet-Serville, 1838, Hesperophasma Rehn, 1904 and Lamponius Stål, 1875. Bradley & Galil (1977: 188) renamed Günther's tribe Hesperophasmatini and added some not closely related genera (Agamemnon Moxey, 1971, Taraxippus Moxey, 1971 and Aploploides Rehn & Hebard, 1838). Zompro (2004: 139) transferred Hesperophasmatini from Cladomorphinae to Pseudophasmatidae: Xerosomatinae and placed the genera Haplopus, Aploploides, Diapherodes and Paracranidium in the distinctive South American tribe Cranidiini Günther, 1953. However, these four Antillean genera differ significantly from Cranidium Westwood, 1843, the type-genus of Cranidiini, and are here removed from that tribe (→ 4.2.3, Table 1). Haplopodini Günther, 1953 is therefore reinstated to contain Haplopus, Aploploides, Diapherodes and Paracranidium as well as four newly described genera (rev. stat.). These share a good number of morphological characters, which sufficiently distinguish them from Cranidiini, Hesperophasmatini or Pterinoxylini n. trib. and obviously form a monophyletic clade, the tribe Haplopodini (→ Table 2).
Distribution (Fig. 52): West Indies (Greater and Lesser Antilles, Bahamas), Florida Keys, Cayman Islands, Santanilla & Central America (Honduras).
Most of the Haplopodini are distributed throughout the West Indies, with only one genus (Apteroplopus n. gen., Fig. 377) represented in Central America (eastern Honduras), hence this is a principally Antillean taxon. Haplopus is restricted to the Greater Antilles, Virgin Islands, St. Thomas, St. Croix, Florida Keys, Dry Tortugas and Bahamas and one species is found also on the Cayman Islands and Swan Islands (= Santanilla, Fig. 376). Parhaplopus n. gen. only contains three species, one of which is endemic to Cuba and the other two known from Hispaniola and Gonaive Island (Fig. 378). Cephaloplopus n. gen. is represented by three species on Hispaniola and has one species on Cuba (Fig. 377). Diapherodes is distributed throughout almost the complete Lesser and Greater Antilles, but has not yet been recorded from Hispaniola, Cuba or the Bahamas (Figs. 375, 381). Paracranidium is restricted to the mountainous regions of East-Jamaica (Fig. 378) and the two monotypical genera Aploploides and Venupherodes n. gen. are endemic to Cuba (Fig. 374).
Genera included:
1. Aploploides Rehn & Hebard, 1938: 49. Type-species: Aploploides stenocephalum Rehn & Hebard, 1938: 49, by original designation of Rehn & Hebard, 1938: 49.
2. Apteroplopus n. gen. Type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907: 323, by present designation.
3. Cephaloplopus n. gen. Type-species: Cephaloplopus pulchellus n. sp., by present designation.
4. Diapherodes Gray, 1835: 13, 33. Type-species: Mantis gigantea Gmélin, 1789: 2055, by subsequent designation of Kirby, 1904a: 362.
5. Haplopus Burmeister, 1838: 576. Type-species: Cyphocrana micropterus St. Fargeau & Audinet-Serville, 1825: 445, by monotypy. = Aplopus Gray, 1835: 13, 34.
6. Paracranidium Brock, 1998: 28. Type-species: Diapherodes (Cranidium) pumilio Westwoood, 1843: 50, pl. 61: 2, by original designation of Brock, 1998: 28.
7. Parhaplopus n. gen. Type-species: Haplopus cubensis Saussure, 1868: 68, by present designation.
8. Venupherodes n. gen. Type-species: Platycrana venustula Audinet-Serville, 1838: 242, by present designation.
Keys to the genera of Haplopodini Günther, 1953
♀♀*
1. Wings present........................................................................................ 4
- Apterous............................................................................................ 2
2. Body length> 6 cm; dorsal body surface without a prominent keel nor tectiform; Cuba .............................. 3
- Small insects (body <6 cm); dorsal body surface strongly tectiform; mesonotum with a prominent, semicircularly raised longitudinal median keel (Fig. 287); Jamaica (Fig. 378)................................................ Paracranidium
3. Massive insects with thorax and abdomen considerably broadened (Fig. 314); head globose and indistinctly longer than wide; vertex bi-tuberculate (Fig. 318)........................................................... Venupherodes n. gen.
- Slender insects (Fig. 53); head elongate and about 2x longer than wide; vertex flat and unarmed (Fig. 58)....... Aploploides
4. Slender insects with body cylindrical in cross-section; colour mostly brown or grey; mesothorax> 2.0x longer than head and pronotum combined; anal region of alae reticulate, with dark radial and transverse veins; not Lesser Antilles or Jamaica .... 5
- Robust to very massive insects, with meso-, metathorax and basal portion of abdomen broadened; mostly bright green; body oval in cross-section; mesothorax <2x longer than head and pronotum combined; anal region of alae plain pink or violet; Greater & Lesser Antilles (Fig. 375).............................................................. Diapherodes
5. Large insects (body length> 10 cm); pair of spines on vertex extending <2/3 the height of head capsule and with a single point; abdominal tergum VII at best gently expanded or with a small posterolateral lobe.............................. 6
- Smaller insects (body <10 cm); pair of spines of the vertex extending by> 2/3 the height of head capsule, crenate or multituberculate; abdominal tergum VII prominently dilated, lateral lobes extending> ½ of body width; Cuba & Hispaniola (Fig. 377)............................................................................... Cephaloplopus n. gen.
6. Mesothorax slender and ± parallel-sided; mesonotum granulose or tuberculose; apex of subgenital plate narrow and ± pointed; probasitarsus slender; Greater Antilles, Dry Tortugas, Florida Keys, Bahamas, Cayman Islands and Swan Islands (Fig. 376)............................................................................................... Haplopus
- Mesothorax swollen medially; mesonotum armed with large, blunt spine-like tubercles; apex of subgenital plate rounded or broadly truncate; probasitarsus with a lobe dorsally; only Cuba & Hispaniola (Fig. 378).............. Parhaplopus n. gen.
*♀♀ of Apteroplopus n. gen. are not known
♂♂*
1. Pterous or brachypterous; median segment longer than metanotum; West Indies.................................... 2
- Apterous (Fig. 53); median segment less than half the length of the metanotum; Honduras (Fig. 377)... Apteroplopus n. gen.
2. Head globose and indistinctly longer than wide; vertex armed.................................................. 3
- Head elongate, about 2x longer than wide; vertex flat and unarmed; Cuba (Fig. 374)....................... Aploploides
3. Spines of vertex extending <2/3 the height of head capsule, with a single point; abdominal tergum VII parallel-sided or with a small posterolateral lobe................................................................................ 4
- Spines of the vertex extending by> 2/3 the height of head capsule, often crenate or multi-tuberculate; abdominal tergum VII strongly expanded with lateral lobe extending by> ½ of body width; Cuba & Hispaniola (Fig. 377)... Cephaloplopus n. gen.
4. Mesothorax> 2x longer than head and pronotum combined; not Lesser Antilles or Jamaica ........................... 5
- Mesothorax <2x longer than head and pronotum combined; Greater & Lesser Antilles (Figs. 367)............ Diapherodes
5. Body surface dull; cerci slender, distinctly shorter than anal segment and laterally compressed; anal region of alae plain.... 6
- Body surface glabrous; cerci obtuse, ± as long as anal segment and ± cylindrical in cross-section; anal region of alae reticulate (Fig. 247); Greater Antilles, Dry Tortugas, Florida Keys, Bahamas, Cayman Islands and Swan Islands (Fig. 376)... Haplopus
6. Body length <7 cm; head globose; pronotum with a pair of anterior spines (Fig. 319); mesonotum only with 2–4 paired spines near anterior margin; Cuba (Fig. 374)...................................................... Venupherodes n. gen.
- Body length> 8 cm; head with vertex at best gently rounded; pronotum unarmed; complete mesonotum armed with spines; Cuba & Hispaniola (Fig. 378)............................................................ Parhaplopus n. gen.
*♂♂ of Paracranidium Brock, 1998 are not known
Eggs*
1. Capsule surface strongly sculptured; no raised central capitulum................................................ 2
- Capsule surface minutely punctured or rugulose; operculum with a raised conical or knob-like central capitulum... Haplopus
2. Micropylar plate <1/3 the length of capsule................................................................. 3
- Micropylar plate> 1/3 the length of capsule................................................................. 4
3. Micropylar plate longer than wide; heart or drop-shaped; capsule irregularly granulose and covered with fine, raised ridges (Figs. 312-313)....................................................................... Parhaplopus n. gen.
- Micropylar plate wider than long; shield-shaped; capsule very prominently and enevenly sculptured (Figs. 324–325)............................................................................................ Venupherodes n. gen.
4. Large (capsule length> 3.5 mm)................................................................. Diapherodes
- Small (capsule length <3.5 mm)........................................................ Cephaloplopus n. gen.
* eggs of Aploploides Rehn & Hebard, 1938, Apteroplopus n. gen. and Paracranidium Brock, 1998 are not known
description: Published as part of Frank H. Hennemann, Oskar V. Conle & Daniel E. Perez-Gelabert, 2016, 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), pp. 1-211 in Zootaxa 4128 (1) on pages 33-37, DOI: 10.11646/zootaxa.4128.1.1, http://zenodo.org/record/271800