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 Diapherodes angulata (Fabricius, 1793) rev. stat. ( Figs. 100–108 , 346 , 365 , 381 ) Mantis angulata Fabricius, 1793: 13 . Type (s), ♀ (♀): Insula Guadeloupe Dom. De Badier (not traced—presumed lost). [Unnecessary replacement name for Mantis gigas Drury, 1773 ]. NT ( by present designation), ♀: Guadeloupe Le Moult [MNCN]. Phasma angulata , Fabricius, 1798: 187 . Phasma angulatum , Lichtenstein , 1802: 11 . Cyphocrana angulata , Audinet-Serville, 1831: 61 . Diapherodes angulata , Burmeister, 1838: 547 . Rehn, 1903: 136. Brock, 1998c: 33. Haplopus grayi Kaup, 1871: 36 , pl. 2: 1 (♀) & 20, pl. 1: 20 (eggs). HT , ♀: Haplopus Grayi Kaup Mol. v. Rosenberg, Holotypus [HLDH]. n. syn. Moxey, 1971: 99 (in litt.). [Listed as a synonym of Diapherodes gigas (Drury, 1773) — in error ] Diapherodes grayi , Kirby, 1904a: 363 . Otte & Brock, 2003: 305. Diapherodes gigantea , Redtenbacher, 1908: 434 (in part). Moxey, 1972: 99 (in litt.; in part). Diapherodes gigas , Moxey, 1971: 98 (in litt.; in part—only specimens from Guadeloupe ). Langlois & Lelong, 1997: 43, figs. 20a (♀) & 20 b (♂). Lelong & Langlois, 2001: 242, figs. 1 (♀), 2 (♂). [ Not: Phasma angulata Stoll, 1813: 61 , pl. 21: 77 (♀), = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825: 445] Further material [14 ♀♀, 21 ♂♂, 6 nymphs, eggs]: GUADELOUPE : 5 ♀♀, 5 ♂♂, 2 ♂♂ (nymphs): Guadeloupe Le Moult [ MNCN ]; 1 ♀, 2 ♂♂: Guadeloupe Trois Rivières; D.Vitrac Dedit.; Collection A. Finot, Diapherodes gigas , Drury [ MNHN , coll. Finot, box no. 294]; 1 ♀: Guadeloupe Trois Rivières; D.Vitrac Dedit.; Collection A. Finot, Diapherodes gigas , Drury [ MNHN ]; Eggs (ex abdomen): Diapherodes gigas Drury, Trois Rivières , Guadeloupe , 69.321 [ MNHN ]; 1 ♀, 1 ♂ , 1 ♂ (penultimate instar), 2 ♀♀ n5: Guadeloupe , Gourbeyre 1896 Diapherodes gigas Drury [ MNHN ]; 1 ♂ : Vernou, 15–30-VI- 19771; Museum Paris Guadeloupe , A. Villiers [ MNHN ]; 5 ♂♂: M. Delfosse Elevage ♂ 1993; Diapherodes gigantea gigantea (Gmélin, 1789) , E. Delfosse Dét. 23/02/03 [ MNHN ]; 1 ♀: M. Delfosse, Guadeloupe ♀ Elevage 1994; Diapherodes gigantea gigantea [ MNHN ]; 1 ♂ : M. Delfosse, Basse-Terre, Guadeloupe 90; Diapherodes gigantea gigantea 04/05 [ MNHN ]; 1 ♀: M. Delfosse, Elevage, Guadeloupe , 1993; Diapherodes gigantea gigantea (Gmélin, 1789) E. Delfosse Dét. 04/05 [ MNHN ]; 2 ♂♂: M. Delfosse Guadeloupe , 6/1993; Elevage, Diapherodes gigantea [ MNHN ]; 1 ♂ : M. Delfosse, Guadeloupe , 07/1993 [ MNHN ]; 2 ♂♂: M. Delfosse Guadeloupe ♂ 04/1993; Elevage, Diapherodes gigantea [ MHNH ]; 1 ♀: Guadeloupe , Antilles ♀; Diapherodes gigas Drury [ MHNG ]; 1 ♀: Antilles; Diapherodes gigas Drury [ MHNG ]; 1 ♀: 3/33 Guadeloupe , Mr. H. d. Sauss.; Diapherodes gigas Drury [ MHNG ]; 1 ♀ (nymph): Larve, Guadeloupe , Mr. H. d. Sauss; Diapherodes gigas Drury [ MHNG ]. NO DATA : 1 ♀: no data [ NHMUK ]; 1 ♂ : no data [ MNHN ]. FIGURES 100–102. Diapherodes angulata (Fabricius, 1793) . 100. ♀: Guadeloupe, Trois Rivères [MNHN]; 101. ♂: Guadeloupe [MNHN]; 102. ♂: Guadeloupe [MNHN]. Diagnosis: Distinguished from the type-species D. gigantea gigantea (Gmélin, 1789) by: the smaller size and prominently bi-spinose vertex of both sexes; more numerous but smaller spiniform tubercles of the mesonotum; shorter and more numerous spines of the marginal row of the meso- and metapleurae; more slender legs and much less prominently dentate, not considerably expanded carinae of the mid and hind legs of ♀♀ ( Fig. 106 ). From D. dominicae (Rehn & Hebard, 1938) ♀♀ differ by: the larger size; more robust legs; much more prominent cephalad pair of spines ( Fig. 107 ); less numerous but more pronounced tubercles of the mesonotum; larger anterior pair of spines on the pronotum; larger and more pointed spines of the meso- and metapleurae ( Fig. 100 ) and larger, broader tegmina which overlap interiorly ( Fig. 100 ). ♂♂ are at once distinguished from all other representatives of the gigantea species-group by the green body, tegmina and costal region of the alae and very sparsely granulose mesonotum ( Figs. 101–102 ). FIGURES 103–108. Diapherodes angulata (Fabricius, 1793) . 103. ♀ NT, apex of abdomen (lateral view) [MNCN]; 104. ♀ NT, apex of abdomen (dorsal view) [MNCN]; 105. Apex of ♂ abdomen (lateral view) [MNHN]; 106. Right hind leg of ♀ NT (posterolateral view) [MNHN]; 107. Head and pronotum of ♀ NT [MNHN]; 108. Head and pronotum of ♂ [MNHN]. Description: ♀ ( Fig. 100 ). Medium-sized to large (body length including subgenital plate 141.0– 162.5 mm ) and moderately broad for the genus. General colouration usually plain pale to mid green, the ventral body surface with a whitish wash; occasionally olive or pale to mid brown specimens may occur. Ventral surface of meso- and metapleurae white, spines of the longitudinal marginal row dull orange to very dark red. Dorsal spines of the head and thorax orange with ochre bases. Tegmina and alae of same colouration as body. Antennae pale brown, the two basal segments greenish. Tarsi greenish pale to mid brown. Eyes dull orange to reddish mid brown. Head: Globose, vertex strongly convex and armed with two prominent, blunt to fairly acute spines; a few small granules are present in the posterior portion ( Fig. 107 ). Eyes circular, convex and of moderate size; their length contained about 2.3x in that of cheeks. Between the bases of the antennae with two rather distinct, oval impressions. Antennae longer than fore legs and laid back, projecting over posterior margin of median segment; with more than 60 segments. Scapus dorsoventrally flattened, 2x longer than wide and gently narrowed towards the base. Pedicellus cylindrical, slightly constricted towards the apex and about half as long as scapus. Thorax: Pronotum slightly shorter and narrower than the head; indistinctly longer than wide and gently widened towards the posterior. In front of the very prominent and well defined, slightly curved transverse median sulcus armed with a large pair of rather long, blunt spines; posterior half with a more or less decided pair of tubercles close to posterior margin and one small tubercle at each posterolateral angle ( Fig. 107 ). Mesothorax 1.6–1.7x longer than head and pronotum combined, and decidedly constricted anteriorly. Mesonotum dorsally armed with two pairs of spiniform tubercles close to anterior margin ( Fig. 107 ) and two or three further pairs of slightly enlarged tubercles in the anterior half of the dorsal surface; otherwise irregularly set with a few small tubercles. Along lateral margins with a longitudinal row of small, rounded granules. Metanotum unarmed, about 2/ 5 the length of mesonotum and about as long as wide. Spines of the meso- and metapleurae very distinct and acute; the longest being about 2x as long as its basal width. Mesopleurae with 17–22, metapleurae with 13–17 spines. Meso- and metasternum each with a variable number of pointed tubercles, which are more decided and larger on the metasternum. Tegmina rather large for the genus and usually slightly projecting over posterior of metanotum (length 11.9–13.0 mm) with posterior margin roundly angulate; overlapping or leaving a small space (up to 3 mm ) inbetween them. Alae very small, projecting underneath tegmina by only about 0.5–2.0 mm. Abdomen: Median segment slightly longer than metanotum and trapezoidal, being gently narrowed towards the anterior . Segments II–V roughly of equal length, VI and VII slightly shorter. II–IV about 1.6–1.7x wider than long, VI about as long as wide. VII narrower and slightly longer than VI, parallel-sided. Praeopercular organ formed by an elongate, longitudinal swelling near posterior margin of sternum VII ( Fig. 346 ). Tergum VIII slightly about 2/3 the length and narrower than VII, decidedly more convex than previous and gently widened towards the posterior. IX about half as long as VIII and slightly wider than long. Anal segment longer than IX, gently narrowed towards the posterior and with a faint longitudinal median keel dorsally ( Fig. 104 ). Posterior margin broadly rounded and with a very shallow median indentation, the lateral margins with a concave excavation at bases of cerci. Epiproct very small and hidden under anal segment. Subgenital plate long and gradually narrowed towards a pointed tip; projecting over apex of abdomen by at least the combined length of tergites IX and X ( Figs. 103–104 ). Legs: All of moderate length, profemora about as long as mesothorax and metatibiae reaching about halfway along abdominal tergum VIII. All carinae densely and sharply but very minutely granulate or denticulate (more decided on mid and hind legs). Mid and hind legs decidedly carinate ( Fig. 106 ). Medioventral carina of meso- and metafemora blunt but low and with 4–6 stout spines which decrease in size towards the base of femur. Posteroventral carina with one, anteroventral carina with two rather small pointed sub-apical spines. Anterodorsal carina of meso- and metatibiae very weakly rounded apically. Basitarsus about 1.3x longer than second tarsomere. ♂ ( Figs. 101–102 ). Medium-sized to large (body length 93.5–110.0 mm) and stocky for the genus, with long alae (53.0–56.0 mm) and a moderately, dorsally sparingly granulose mesothorax. General colouration pale yellowish green to mid green, the tegminal and costal region of the alae with distinctly darker longitudinal veins. Ventral thoracal segments and abdominal sternum II dull green, abdominal sternites III–VII whitish. Vent ral surfaces of meso- and metapleurae white. Largest tubercles of the head and dorsal surface of the thorax tipped with dull orange. Tegmina and costal region of same colouration as body, the tegmina with a broad longitudinal and very pale creamish green stripe along the anterior margin which is continued in the basal quarter of the alae. Bases of alae pale red, anal region hyaline. Antennae pale to mid brown and becoming gradually darkler brown towards the apex, two basal segments greenish. Eyes dark orange to reddish brown. Head: Generally as in ♀♀. Vertex strongly convex and smooth except for two blunt but well decided, conical tubercles; posterior portion usually with two small granules ( Fig. 108 ). Eyes large, cylindrical and projecting hemispherically; their length contained about 2x in that of cheeks. Antennae reaching halfway along abdominal tergum III, otherwise structured like in ♀♀. About 70 antennomeres. Thorax: Pronotum shorter and slightly narrower than head, about as long as wide with the anterior margin slightly widened. Otherwise generally as in ♀♀ but armature less distinct ( Fig. 108 ). Mesothorax rather elongate, about 2x longer than head and pronotum combined; mesonotum almost 6x longer than wide. Anterior of mesonotum with a well decided pair of blunt tubercles, dorsal surface otherwise irregularly set with a few low tubercles and granules; a longitudinal row of small granules along lateral margins. Meso- and metapleurae each with a longitudinal row of minute granules. Meso- and metasternum sparsely granulose. Alae reaching halfway along tergum VI. Abdomen: Tergum VII with posterolateral angles very gently expanded and rounded. Tergum VIII 2 /3 of VII and gradually widened towards the posterior, broadest segment. IX ¾ the length of VIII and very weakly narrowed towards the posterior. Posterior half of tergum VII, as well as VIII and IX with two parallel, longitudinal carinae dorsomedially. Anal segment with a very faint longitudinal median keel, the posterior portion abruptly narrowed and with a broadly triangular median indentation. Sternum VIII with an acute longitudinal median carina. Vomer triangular with basal portion strongly broadened and almost equal to overall length, terminal hook short and slender ( Fig. 365 ). Poculum granulose, convex and cup-like ( Fig. 105 ) with the posterior margin broadly triangular ( Fig. 365 ); indistinctly projecting over posterior margin of tergum IX. Legs: All of moderate length for the genus, profemora slightly longer than mesonotum, metatibiae almost reaching posterior margin of abdominal tergum VII. All carinae densely but very minutely granulate to denticulate. Posteroventral carina of meso- and metafemora with one, anteroventral carina with two, slender and pointed subapical spines. Medioventral carina broad and flat, armed with 5–6 short but strong spines which decrease in size towards the base of femur. Tarsi rather stout, slightly less than half the length of corresponding tibia. Basitarsi about 1.3x longer than second tarsomere. Comments: This species was first described by Fabricius (1793: 13) who stated the type locality to be “ Insula Guadeloupe ”. The original ♀ type specimen(s) is not traced in ZMUC nor in any other likely collection and hence believed lost (Brock, 1998c: 33). Fabricius (1793) based his description on a specimen(s) from Guadeloupe and introduced Mantis angulata as a replacement name for Mantis gigas Drury, 1773 , which had however already been replaced by Mantis gigantea Gmélin, 1798 . The name of Fabricius is therefore an unnecessary replacement name and a junior objective homonym of Mantis gigantea Gmélin. Diapherodes gigantea (Gmélin, 1789) is endemic to Grenada and Saint Vincent and detailed examination of several specimens, including ♂♂, from Guadeloupe in MNCN and MNHN has shown these to be a distinct species. As the type specimen(s) on which Fabricius (1793: 13) based his description of Mantis angulata were from Guadeloupe and represent a distinct species, Mantis angulata Fabricius is not a synonym of Mantis gigantea and becomes an available name ( rev. stat. ). Therefore the specific name of Fabricius (1793) is here re-introduced and used for the species from Guadeloupe . As the original type-specimen(s) is lost, an almost perfect ♀ in MNCN from a series of specimens with the data “ Guadeloupe Le Moult” is here selected as the neotype of Mantis angulata Fabricius, 1793 in order to ensure stability of the name. Further specimens from the same source are in MNHN. Kaup (1871: 36, pl. 2: 1) described and figured Haplopus grayi from a ♀ wrongly labelled “ Molukken ”, but there can however be no doubt his specimen originated from the Lesser Antilles. Examination of Kaup's holotype in HLDH has proven this to be clearly distinct from Diapherodes gigantea but to match very well with typical ♀♀ of D. angulata from Guadeloupe . Comparison has proven the synonymy of Kaup's species and suggests this specimen to have originated from Guadeloupe . Moxey (1971: 99) erroneously synonymised Haplopus grayi with Diapherodes gigantea . Diapherodes angulata appears to have been moderately abundant in certain localities in Guadeloupe , which is seen in the good series of specimens collected by Le Moult (now in MNCN and MNHN). It has however become apparently rare and was not encountered during recent extensive collections conducted on the island in the late 1990's (Langlois & Lelong, 1997: 43). D. angulata is endemic in Guadeloupe and all records of D. gigantea from this island are based on misidentifications. Distribution ( Fig. 381 ): Guadeloupe (Basse-Terre [MNHN]; Trois Riviéres [MNHN]), endemic. Number of specimens examined: 43 TABLE 11. Measurements of Diapherodes angulata (Fabricius, 1793) [mm] ♀, NT ♀♀ ♂♂ [ MNCN ] [ MNCN ] [ MNCN ]