Studies on Neotropical Phasmatodea XXVI: Taxonomic review of Cladomorformia tax. n., a lineage of Diapheromerinae stick insects, with the descriptions of seven new genera and 41 new species (Phasmatodea: Occidophasmata: Diapheromerinae) Author Hennemann, Frank H. 0000-0002-6547-4704 Tannenwaldallee 53, 61348 Bad Homburg, Germany. hennemann @ phasmatodea. com; http: // zoobank. org / urn: lsid: zoobank. org: author: 651 FCCFA- 271 B- 48 A 3 - A 58 E-A 30 FDC 739493 https: // orcid. org / 0000 - 0002 - 6547 - 4704; Website: www. phasmatodea. com hennemann@phasmatodea.com Author Conle, Oskar V. 0000-0002-6547-4704 Tannenwaldallee 53, 61348 Bad Homburg, Germany. hennemann @ phasmatodea. com; http: // zoobank. org / urn: lsid: zoobank. org: author: 651 FCCFA- 271 B- 48 A 3 - A 58 E-A 30 FDC 739493 https: // orcid. org / 0000 - 0002 - 6547 - 4704; Website: www. phasmatodea. com & Am Freischütz 16, 47058 Duisburg, Germany. conle @ phasmatodea. de; http: // zoobank. org / urn: lsid: zoobank. org: author: D 2712 C 02 - 7973 - 4 FAA-A 186 - 5 F 8540 A 66691 https: // orcid. org / 0000 - 0001 - 5609 - 5107; Website: www. phasmatodea. com * Corresponding author & Tannenwaldallee 53, 61348 Bad Homburg, Germany. hennemann @ phasmatodea. com; http: // zoobank. org / urn: lsid: zoobank. org: author: 651 FCCFA- 271 B- 48 A 3 - A 58 E-A 30 FDC 739493 https: // orcid. org / 0000 - 0002 - 6547 - 4704; Website: www. phasmatodea. com hennemann@phasmatodea.com text Zootaxa 2024 2024-04-26 5444 1 1 454 http://dx.doi.org/10.11646/zootaxa.5444.1.1 journal article 10.11646/zootaxa.5444.1.1 1175-5334 11071464 5DE4A9DD-99F7-4E23-AD50-58DC491BB75E Phanocles falcatus sp. n. urn:lsid:zoobank.org:act: 81A0ABFB-FBBA-4391-BAAC-AAF6CA4BFC41 ( Figs. 52B , 55B , 57H–J , 59J , 63K–M , 88O , 93C , 97A–B , 99O ) HT , ♂ : Jungle vegetation; Ecuador , Napo Province , Coca , II.1982 , G. Onore , BM 1985–61 [ NHMUK ] . PT , 1 ♂ , 2 ♀♀ : Via Cupidos : forêt de plateau lisiéres; Prov. Napo , entres rios Rumiyacu et Tiputini Pindo , 04 / 16 III 1995 ; Museum Paris , Ecuateur, S . Poulain réc. [ MNHN ] . PT , ♀ : forêt de plateau lisiéres; Prov. Napo , stat. Rech. “Yasuni”, cflt ríos Tiputini / Tivacuno , 05 / 15 II 1995 ; Museum Paris , Ecuateur, S . Poulain réc. [ MNHN ] . PT , ♀ : Prov. Napo , Puerto Napo / Ahuano , 450 m . 16 VIII / 06 IX 1991 ; Museum Paris , Ecuateur , Amedegnato / Poulain rec. [ MNHN ] . PT , ♂ : Pr. Sucumbios , Coca / L. Agrio , S. P. de los Cofanes , 415 m . 22/ 23 IX 1997 , 77°52’W , 0°8’S ; Museum Paris , Ecuateur , Amédéganto / Poulain rec. [ MNHN ] . PT , ♂ : Umbria , upper Putumayo , R. Narino , Colombia , IX.’30 [ ANSP ] . PT , ♂ : Dept. du Loreto , Rio Yubineto , 22.I–1978 ; Pérou , S. Poulain rec. [ MNHN ] . PT , ♂ : Dept. du Loreto , Rio Yubineto , V–1978 ; Pérou , S. Poulain rec.; la colpa [ MNHN ] . PT, ♀ : Macas , Ecuador or.; 409 [ USNM ] . PT , 2 ♀♀ , 2 ♀♀ (penultimate instar), 2 ♂♂ , 1 egg : Ecuador : Provinz Napo , Oriente , Misahualli , 400 m , jungle vegetation, eg. Heinz van Herwaarden 1989 [ FH , No’s 0919–1 to 6 & E] . Diagnosis. Males of this new species strongly resemble those of the two Ecuadorian species Ph. aequatorialis ( Redtenbacher, 1908 ) comb. n. and Ph. ecuadoricus sp. n. but differ from both by the more prominent, pointed cephalad horns ( Figs. 59J ), slightly shorter alae (at least reaching to posterior margin of abdominal segment III in the other two species) and the very large, elongated and distinctly arcuate and falcate vomer ( Figs. 65D , 97A ). From ♂♂ of the first species they may also be distinguished by the larger size (body length <100.0 mm in aequatorialis ), more globose head, less pronounced medio-longitudinal carina of the mesonotum, longer cerci (notably shorter than anal segment in aequatorialis ), presence of a spiniform central protuberance of the poculum ( Fig. 65D ) and having the posterior margin of the poculum entire (indented medially and bidentate in fallax ). From those of ecuadoricus sp. n. these ♂♂ can also be distinguished by the more globose head; distinctly black lateral margins of the pronotum ( Fig. 59J ), relatively more elongate anal segment, that is gradually narrowed towards the posterior ( Fig. 65E ; with posterior margin broadly rounded in ecuadoricus ), more distinct and acute central protuberance of the poculum ( Fig. 65D ) as well as the slender basitarsi which merely have a faint median dorsal deflexion (dorsal carina notably raised and rounded in ecuadoricus ). In many aspects (e. g. the general shape, morphology of the head and shape of the large lobes of the praeopercular organ and dorsal crests of the basitarsi) ♀♀ are very similar to those of the Colombian Ph. significans ( Redtenbacher, 1908 ) . They can however readily be separated by the more anteriad directed cephalad horns ( Figs. 57H–J ) and distinct lateral lobe of abdominal tergum VII ( Figs. 63K–M ; no lateral lobes in significans ). Etymology. The name ( falcatus lat. = falcate, falciform) refers to the very large, elongate and strongly arched vomer of this new species ( Fig. 97A ). Description. ♀♀ ( Fig. 52B ): Large to very large (body length including subgenital plate 189.0–212.0 mm) and slender species with a sub-spherical and prominently bi-cornute head, smooth thorax and a short subgenital plate, that projects beyond the tip of the abdomen by less than the length of the anal segment. Body surface wholly unarmed with the thoracic segment sometimes very minutely and sparsely granulose. General colouration variable and ranging from pale drab over various shades of brown to blackish dark brown; occasionally all over flecked with dark brown and/or light grey and somewhat lichenose in appearance or with a weakly indicated medio-longitudinal streak along dorsal body surface. Outer margin of the cephalad horns black.Antennae drab dorsally and dark greyish brown ventrally in the basal one-third. Head ( Figs. 57H–J ): Sub-spherical and scarcely longer than wide with the vertex roundly convex, smooth and armed with a prominent pair of distinctly forward directed cephalad horns; these oval at the base and increasingly compressed laterally and carinate towards the apex, variable in shape and size, with the anterior margins sometimes dentate and at least projecting by half of capsule height. Frons with two small but distinct impressions between the bases of the antennae and an obtuse pair of shallow swellings between the eyes. Eyes moderately projecting and almost circular in outline with the anterior margin somewhat angular, their diameter contained 2.2x in length of genae.Antennae ± reaching half the way along abdominal segment III. Scapus compressed dorsoventrally, somewhat narrowed towards the base and with the lateral margins gently rounded; 1.3x longer than wide. Pedicellus round in cross-section, slightly narrowed apically and roughly half the length of scapus. Thorax: Pronotum about as long but noticeably narrower than head, 1.7x longer than wide and roundly rectangular in dorsal aspect with the anterior angles somewhat widened and the posterior margin rounded. Entire surface with a moderately impressed medio-longitudinal furrow, the transverse median sulcus distinct, gently rounded and almost reaching to lateral margins of segment ( Figs. 57H–J ). Mesothorax 7x longer than prothorax, slightly constricted anteriorly and the posterior portion just weakly expanded. Metanotum somewhat less than one-third the length of mesonotum, with a slightly narrowing pre-medially and about 3.2x longer than wide. Meso- and metanotum and pleurae wholly smooth but the nota with a weakly indicated medio-longitudinal line and occasionally with a few minute scattered granules. Meso- and metasternum minutely granulose and rugose along the mid-line and each with a fine but acute longitudinal carina parallel to lateral margins. Abdomen: Median segment 1.2x longer than metanotum, distinctly narrowed medially and about 4.3x longer than width of anterior margin. Segment II scarcely shorter than median segment and 2.7x longer than wide. II–V slightly increasing in length, V longest segment and 3.3x longer than wide, VI–VII decreasing in length and VII notably shorter than II. Terga I–VII with a ± distinct, obtusely bi-lobed posteromedian projection and an obtuse longitudinal carina laterally. Tergum VII narrower than all preceding, somewhat widening towards the posterior and only about three-fifths the length of VI, lateral margins posteriorly deflexed to form a distinct, rounded lobe, that may extend laterally by as much as on-third the width of the segment ( Figs. 63K–M ). Sterna smooth to very sparsely and minutely granulose and with a fine longitudinal carina laterally. Praeopercular organ on sternum VII very prominent and formed by a pair of large, oval lobes, that are larger than the posterolateral lobe of the tergum and strongly project beyond the posterior margin; these lobes formed by the longitudinal lateral carinae, which are in general strongly pronounced on this sternum ( Figs. 63K, M , 93C ). Tergum VIII about half the length and narrower than VII, slightly narrowed at posterior margin and 1.8x longer than wide. IX scarcely longer than wide, about half as long as VII and narrowed anteriorly. Anal segment a little longer than IX, carinate medio-longitudinally with the carina increasingly pronounced towards the apex, the lateral margins slightly widened and gently rounded basally and the posterior margin obtusely angular ( Fig. 63L ). Epiproct minute and wholly concealed under anal segment. Cerci small and tapering towards a narrow apex; roughly reaching to posterior margin of anal segment. Gonapophyses VIII very elongate, distinctly upcurved with the apex pointed and projecting over tip of abdomen by almost the length of anal segment (occasionally also projecting beyond tip of subgenital plate, Figs. 63K–L ). Subgenital plate obtusely keeled longitudinally in the apical half, with the apex obtusely triangular and projecting over the tip of abdomen by less than length of anal segment ( Figs. 63K–M ); basal half with two prominent, uneven lateral longitudinal lamellae. Legs:All long and moderately stocky. Profemora slightly shorter than mesothorax, mesofemora almost reaching to posterior margin of abdominal segment II, metafemora projecting over posterior margin of abdominal segment IV and metatarsi slightly projecting beyond apex of abdomen. Two outer ventral carinae of mesofemora with a small, ± distinct, rounded deflexion sub-basally. Meso- and metatibiae with the two outer ventral carinae ± deflexed and rounded sub-apically. Probasitarsus with the dorsal carina lamellate and gradually increasing in height with the apical portion rounded; almost as long as remaining tarsomeres combined. Meso- and metabasitarsi with a prominent, roundly triangular dorsal crest; all slightly longer than corresponding following three tarsomeres taken together. ♂♂ ( Fig. 55B ): Medium-sized (body length 108.5–121.0 mm) and slender for the genus with moderately developed alae, that reach ± half the way along abdominal tergum III (31.5–43.0 mm), a globose and bi-tuberculate to bi-spinose head and a very weakly rounded dorsal carina of the basitarsi. General colour buff to ochre mid brown or drab. Mesopleurae pale yellow posteriorly. Mesosternum with the medio-longitudinal carina pale green in the anterior portion. Head with a pair of black spots between the eyes, two ± defined blackish longitudinal markings or streaks in posterior portion of vertex and a broad blackish postocular streak; lower portion of the genae pale yellow and pair of cephalic tubercles brown to black. Eyes ochre with darker mottling. Pronotum with lateral margins and most of epimerum black. Tegmina drab to mid to dark brown with the central portion somewhat darker than the rest and the anterior margin broadly pale yellow. Costal region of alae alike, the anal fan transparent pale brown with all veins brown. Two terminal abdominal terga often with a slight greenish was. Cerci straw to ochre with the apex greyish green. Vomer drab basally, greenish towards the apex and the terminal hook reddish brown ( Fig. 97B ). All coxae with two blackish sub-parallel longitudinal stripes. All femora with three and tibiae with two faint pale yellowish transverse bands. Antennae reddish brown dorsally and greyish brown ventrally. Head ( Fig. 57J ): Shape basically as in ♀♀ but vertex somewhat less convex, broadest at the eyes and narrowing towards the posterior with the cephalad horns much less pronounced and only represented by a pair of obtuse swelling or short, conical spines. Between the bases of the antennae with a transverse ridge that is followed by two distinct and deep impressions and two obtuse, low swellings between the eyes. Eyes large, almost circular in outline and projecting hemispherical from head capsule, their diameter contained 1.5x in length of genae. Antennae reaching to abdominal segment V. Scapus and pedicellus basically as in ♀♀ . Thorax: Pronotum slightly longer and considerably narrower than head, roundly rectangular in dorsal aspect with the lateral margins slightly sub-parallel and somewhat converging posteriorly and notably expanded anteriorly; roughly 2.2x longer than wide. The transverse median sulcus weakly W-shaped and not reaching to lateral margins of segment ( Fig. 57J ). Mesothorax 10x longer than and a little narrower than prothorax. Mesonotum smooth except for a weakly developed irregularly granulose longitudinal carina parallel to lateral margins; meso- and metapleurae smooth but with a fine medio-longitudinal carina. Mesosternum with a rather obtuse longitudinal median carina that is scarce in the median portion of the segment ( Fig. 88O ), metasternum with the medio-longitudinal carina weakly indicated. Tegmina elongate, oval and gradually narrowed in the anterior portion; the central protuberance small and obtusely rounded. Alae somewhat variable in length and reaching ± half the way along abdominal tergum III. Abdomen: Median segment over 4x longer than metanotum and gently narrowing towards the posterior. Abdominal segment II scarcely more than half the length of median segment. II–VI slightly sub-equal in length and on average 5.5x longer than wide, all constricted medially. VII notably shorter than all preceding segments, only about two-thirds the length of VI and widened in the posterior portion; 3.2x longer than width at anterior margin. Tergum VIII about three-fifths the length of VII, gradually widening towards the posterior, trapezoidal in dorsal aspect and the posterior margin about 1.6x wider than anterior margin. IX scarcely shorter than VIII, constricted medially and with the anterior portion slightly deflexed and notably wider than posterior half; lateral margins straight Fig. 65D ). Anal segment very slightly longer than IX, 1.6x longer than width at anterior margin, the posterior half gradually narrowing ( Fig. 65E ), the lateral margins with an obtuse, roundly triangular tooth medially and the apex rounded ( Fig. 65D ); dorsal surface with an obtuse medio-longitudinal bulge that gets increasingly pronounced towards the posterior. Ventral surface of entire outer margin of posterior half of segment unevenly set with minute denticles. Cerci of moderate length, slender, very gently incurved and tapered towards a slightly club-like apex; projecting scarcely beyond apex of anal segment. Vomer very large, elongate, slender, gradually narrowing towards the terminal hook and strongly arcuate with the apical portion upcurved, almost at a right angle to the basal portion and weakly sinistral directed; shape sickle-like in lateral aspect ( Figs. 65D , 97A ), the outer portions strongly inflated and the basal two-thirds with a deep medio-longitudinal furrow ( Figs. 65F , 97B ). Poculum bulgy, strongly convex, angular and cup-shaped in lateral aspect with a small and obtuse protuberance at the angle ( Fig. 65D ) and the vertical posterior portion carinate medio-longitudinally; the posterior margin obtusely triangular and roughly reaching one-third along the anal segment. Legs:All very long, slender and wholly unarmed except for an indistinct, weakly rounded sub-basal deflexion of the two outer ventral carinae of the mesofemora. Profemora about as long as head, pro- and mesothorax combined, mesofemora roughly as long as mesothorax, metafemora reaching to posterior margin of abdominal segment V and metatibiae projecting greatly beyond apex of abdomen. All basitarsi noticeably longer than remaining tarsomeres taken together, the dorsal carina of the probasitarsi lamellate and very weakly rounded, that of the meso- and metabasitarsi almost uniform in height. Variability. Females show noteworthy variability in size, colour, shape and size of the cephalic horns, size of the posteromedian protrusion of abdominal terga I–VII and length of the gonapophyses VIII. Slight variability is also seen in the development of the sub-basal lobe on the two outer ventral carinae of the mesofemora as well as the size and shape of dorsal crest of the basitarsi. No variability other than concerning to the overall colouration, size and development of the cephalic pair of spines can be observed in ♂♂ . The latter are least developed and only represented as obtuse swellings in the ♂ from the Rio Nariño, Colombia in the collection of ANSP and the Ecuadorian examples in the collection of MNHN but most prominent in the two specimens in the first author’s collection (coll. FH ). Since , the latter two specimens and those in the collection of MNHN are from fairly nearby localities in the Napo Province in north-eastern Ecuador , this trait is believed to be not peculiar to certain localities, which is in accordance to the variability of the cephalic armature seen in the ♀♀ at hand . Eggs ( Fig. 99O ). One fully developed egg could be removed from the ovipositor of one of the ♀ paratypes in the first author’s collection (coll. FH ) . Moderately sized for the genus, shape ovoid with the polar-area somewhat flattened and the dorsal egg surface strongly convex, but in the region of the micropylar plate in particular; oval in cross-section. Capsule surface almost smooth and all over but rather sparsely covered with irregularly dispersed, fairly small pits; surface otherwise very slightly glossy. Micropylar plate moderately elongate, almost two-thirds as long as capsule, parallel-sided and with both ends rounded; 2.6x longer than wide. Plate wholly convex and projecting from surface of capsule, the surface with same puncturing as capsule and the outer margin obtusely and slightly inflated. Micropylar cup small, node-like. Median line obtuse, short and not reaching to polar-area. Operculum oval and with a moderately sized, hollow and raised structure that is formed by excrescences of the outer margin; shape roundly conical with the lateral surface almost smooth basally and crenate apically; height only corresponding to about 0.6x the length of capsule. Colour of capsule plain ochre, the micropylar plate reddish mid brown in the outer portions and otherwise like capsule, opercular structure reddish brown with the base darker in colour. Measurements [mm]: length including operculum 4.4, length 3.7, width 2.8, height 3.0, length of micropylar plate 2.4. Table 38: Measurements of Phanocles falcatus sp. n.