Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley &amp; Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “Anareolatae”: Phasmatidae: Cladomorphinae)
Authors: HENNEMANN FRANK H. , CONLE OSKAR V. , PEREZ-GELABERT DANIEL E. 
Published: June 2016

The anareolate New World subfamily Cladomorphinae Bradley & Galil, 1977 is reviewed and keys to the six tribes currently included are presented; these are: Cladomorphini Bradley & Galil, 1977, Cladoxerini Karny, 1923, Cranidiini Günther, 1953, Pterinoxylini n. trib., Hesperophasmatini Bradley & Galil, 1977 and Haplopodini Günther, 1953 rev. stat.. New diagnoses are presented for all these tribes and possible relationships within Cladomorphinae are discusssed. Morphology of the genitalia and egg-structures indicate Cladomorphinae as presently treated to be polyphyletic. Two subordinate groups are recognized within present Cladomorphinae, which differ considerably in numerous morphological characters of the insects and eggs. The first group and here regarded as Cladomorphinae sensu stricto is formed by the mostly South American Cladomorphini + Cranidiini + Cladoxerini, while the second group is formed by the predominantly Caribbean Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini. Members of the first group (= Cladomorphini sensu stricto) share the dorsally carinate basitarsus in which the two dorsal carinae are melted with another, increasingly elongated gonapophyses VIII of females which are noticeably longer than gonapophyses IX and lamellate as well as strongly displaced medioventral carina of the profemora. Cranidiini + Cladomorphini share the strongly elongated and filiform gonapophyses VIII and presence of gonoplacs in the females, specialized poculum of males and presence of a median line in the eggs. Cranidiini differs from all other tribes of Cladomorphinae by the entirely unarmed legs of both sexes, distinctly broadened and leaf-like body and prominent longitudinal keel of the mesosternum of females, prominently enlarged poculum and spinulose phallus of males as well as the conspicuous narrowing of the posteromedian gap of the internal micropylar plate of the eggs and noticeably separated median line. Cladomorphini is characteristic for the specialized vomer and poculum of males and distinct opercular structures of the eggs. Certain representatives of Cladomorphini indicate relationships to the “Phanocles-group” of Diapheromerinae: Diapheromerini, hence Cladomorphini as presently treated may be paraphyletic. The exclusively South American Cladoxerini (= Baculini n. syn.) differs from the other two tribes of Cladomorphinae sensu stricto by the distinctly serrate profemora of both sexes and conspicuously shortened antennae of females, which consist of less than 30 segments and are much shorter than the profemora in females. Genital morphology, such as the elongated gonapophyses VIII and presence of gonoplacs in females, as well as the lamellate medioventral carina of the profemora indicate close relation to Cladomorphini. Cranidiini appears to be the sister-taxon of Cladomorphini + Cladoxerini. The tribe Baculini Günther, 1953 is synonymised with Cladoxerini (n. syn.), on the basis that the type-genera of both tribes are congeneric, with Baculum Saussure, 1861 being a junior synonym of Cladoxerus St. Fargeau & Audinet-Serville, 1827 (n. syn.). The genus Tersomia Kirby, 1904 is removed from Hesperophasmatini and transferred to Cladoxerini. Wattenwylia Toledo Piza, 1938 is removed from Pachymorphinae: Gratidiini and transferred to Cladoxerini. A detailed new diagnosis is presented for Cranidiini along with a detailed differentiation and the tribe is shown to be monotypical, only containing its type-genus Cranidium Westwood, 1843. All Caribbean genera subsequently added to Cranidiini are removed and transferred to Haplopodini rev. stat.. The three tribes Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini rev. stat. are closely related and might form a monophyletic clade within Cladomorphinae sensu lato. They differ from Cladomorphinae sensu stricto by the short gonapophyses VIII and reduced gonoplacs of females, unspecialized poculum of males and lack of a micropylar line in the eggs. Haplopodini Günther, 1953 is re-established (rev. stat.) and comprises almost exclusively Caribbean genera previously placed in Hesperophasmatini by Bradley & Galil (1977) or Cranidiini by Zompro, (2004). Aploploides Rehn & Hebard, 1938, Diapherodes Gray, 1835, Haplopus Burmeister, 1838 and Paracranidium Brock, 1998 were misplaced in Cranidiini and are transferred to Haplopodini. On the basis of numerous morphological characters of the insects and eggs Hesperophasmatini is removed from Pseudophasmatidae: Xerosomatinae and re-transferred to its previous position in the subfamily Cladomorphinae sensu lato. A detailed newdiagnosis of Hesperophasmatini is presented, but is only provisional since the true diversity is as yet only fractionally known. The lack of a gula distinguishes Hesperophasmatini from all other tribes. The genus Laciphorus Redtenbacher, 1908 is removed from Hesperophasmatini and transferred to Diapheromeridae: Diapheromerinae: Diapheromerini. The new tribe Pterinoxylini n. trib. is established to contain only the type-genus Pterinoxylus Audinet-Serville, 1838. It is closely related and perhaps the sister taxon of Hesperophasmatini, with which it shares the presence of rough sensory areas on the probasisternum and profurcasternum. It differs from Hesperophasmatini and Haplopodini by the presence of a tympanal region (= stridulatory organ) in the alae of females and the alveolar eggs, which possess peripheral opercular and polar structures. Haplopodini is likely to be the sister group of Pterinoxylini n. trib. + Hesperophasmatini. The tribe Haplopodini rev. stat. is revised at the species level and comprises eight almost exclusively Caribbean genera, four of which are newly described. All eight genera now contained in Haplopodini are described in detail, differentiated from their closest relatives and their relationships and systematic position within Haplopodini are discussed. Keys and maps showing their distributions are presented along with a discussion of the distributional patterns. Detailed descriptions, differential diagnoses, synonymic listings, illustrations, material listings and measurements are given of all 26 currently known species and subspecies of Haplopodini. Four new genera are described within Haplopodini. The monotypical Apteroplopus n. gen. (type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907) from Honduras is the only taxon of the tribe represented in Central America. It is only known from the male which differs from all other genera by being entirely apterous. Cephaloplopus n. gen. (type-species: Cephaloplopus pulchellus n. sp.) and Parhaplopus n. gen. (type-species: Haplopus cubensis Saussure, 1868) occur only on Hispaniola and Cuba. Both are closely related to Haplopus Burmeister, 1838 but in addition to having noticeably different eggs, both genera differ from Haplopus in several morphological characters. The monotypical Venupherodes n. gen. (type-species: Platycrana venustula Audinet-Serville, 1838) is endemic to Cuba, and in females being apterous resembles the second exclusively Cuban genus Aploploides Rehn & Hebard, 1938. It however differs from all other members of Haplopodini by the laterally expanded mesonotum of females, which overlaps the mesopleurae, as well as the morphology of the eggs. Two species-groups are recognized within Diapherodes Gray, 1835. The gigantea species-group comprises the species from the Lesser Antilles, which are: D. angulata (Fabricius, 1793), Diapherodes dominicae (Rehn & Hebard, 1938), D. gigantea gigantea (Gmélin, 1789), D. gigantea saintluciae n. ssp. and Diapherodes martinicensis Lelong & Langlois, 2005. The three species of the jamaicensis species-group, which are D. achalus (Rehn, 1904), D. jamaicensis (Drury, 1773) and D. laevicollis Redtenbacher, 1906, are restricted to the two Greater Antillean islands Jamaica and Puerto Rico. Haplopus Burmeister, 1838 is the most widely distributed genus being represented on all islands of the Greater Antilles except Jamaica, and also in the Virgin Islands, Bahamas, Florida Keys, Dry Tortugas and as far southwest as the Cayman Islands and Swan Islands. Nine new species and one new subspecies are described: Cephaloplopus alope n. sp. and Haplopus sobrinus n. sp. from Cuba, Cephaloplopus euchlorus n. sp., Cephaloplopus laetus n. sp., Cephaloplopus pulchellus n. sp., Haplopus brachypterus n. sp., Haplopus intermedius n. sp. and Parhaplopus navarroi n. sp. from Hispaniola, Haplopus woodruffi n. sp. from Cayman Brac (Cayman Islands) and Diapherodes gigantea saintluciae n. ssp. from Saint Lucia. Seven of these are described from both sexes but Cephaloplopus alope n. sp. and Haplopus sobrinus n. sp. are only known from the females and Cephaloplopus laetus n. sp. only from the males. The previously unknown males of Diapherodes angulata (Fabricius, 1793), Diapherodes laevicollis Redtenbacher, 1908, Haplopus bicuspidatus de Haan, 1842 and Parhaplopus cubensis (Saussure, 1868) as well as the previously unknown female of Parhaplopus evadne (Westwood, 1859) n. comb. are described and illustrated for the first time. Descriptions and illustrations of the eggs of eleven species are presented: Cephaloplopus euchlorus n. sp., Cephaloplopus pulchellus n. sp., Diapherodes achalus (Rehn, 1904), Diapherodes dominicae (Rehn & Hebard, 1938), Diapherodes gigantea gigantea (Gmélin, 1789), Diapherodes martinicensis Lelong & Langlois, 2005, Diapherodes jamaicensis (Drury, 1773), Haplopus bicuspidatus de Haan, 1842, Haplopus micropterus St. Fargeau & Audinet-Serville, 1825, Parhaplopus navarroi n. sp. and Venupherodes venustula (Audinet-Seville, 1838) n. comb.. Type specimens of the newly described taxa are deposited in the collections of ANSP, NHMUK, IIBZ, FSCA, MCZC, MNHN and USNM. Six species are transferred to other genera (n. comb.): Bacteria grossetuberculata (Brunner v. Wattenwyl, 1907) to Apteroplopus n. gen.; Haplopus cubensis Saussure, 1868 and Haplopus evadne Westwood, 1859 to Parhaplopus n. gen.; Diapherodes venustula (Audinet-Serville, 1838) to Venupherodes n. gen.; Haplopus jamaicensis (Drury, 1773) and Haplopus achalus Rehn, 1904 to Diapherodes Gray, 1835. Mantis angulata Fabricius, 1793 and Diapherodes gigantea dominicae Rehn & Hebard, 1938 are removed from synonymy with D. gigantea (Gmélin, 1789) and shown to be valid species (n. stat.). Fifteen new synonymies are revealed amongst the species studied: Diapherodes longiscapha Redtenbacher, 1908 = Diapherodes achalus (Rehn, 1904) n. syn.; Haplopus grayi Kaup, 1871 = Diapherodes angulata (Fabricius, 1793) n. syn.; Diapherodes glabricollis Gray, 1835 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Diapherodes pulverulentus Gray, 1835 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Diapherodes christopheri Westwood, 1859 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Haplopus murinus Redtenbacher, 1908 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Haplopus bituberculatum de Haan, 1842 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus cythereus Westwood, 1859 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus ligiolus Redtenbacher, 1908 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus ligia Westwood, 1859 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus mayeri Caudell, 1905 = Haplopus scabricollis (Gray, 1835) n. syn.; Aplopus similis Rehn, 1904 = Haplopus scabricollis Gray, 1835 n. syn.; Diapherodes spinipes Gray, 1835 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus obtusus Redtenbacher, 1908 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn. and Haplopus juvenis Redtenbacher, 1908 = Venupherodes venustula (Audinet-Serville, 1838) n. syn.. The previously presumed lost holotype of Cyprocrana microptera St. Fargeau & Audinet-Serville, 1825 (= Phasma angulata Stoll, 1813), was traced in the collection of RMNH. The designation of a neotype has become necessary for Mantis angulata Fabricius, 1793 (in MNCN) and Platycrana venustula Audinet-Serville, 1838 (in MNHU). Lectotypes are designated for eight species: Diapherodes longiscapha Redtenbacher, 1908; Diapherodes scabricollis Gray, 1835; Haplopus christopheri Westwood, 1859; Haplopus cytherea Westwood, 1859; Haplopus juvenis Redtenbacher, 1908; Haplopus ligiolus Redtenbacher, 1908; Haplopus ligia Westwood, 1859 and Haplopus murinus Redtenbacher, 1908.