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 aequatorialis ( Redtenbacher, 1908 ) comb. n. ( Figs. 54B, 54F , 58A , 60A , 61K–M , 66H , 99A , 111C , 118C ) Bacteria aequatorialis Redtenbacher, 1908: 419 . Brock, 1998: 12. Phanocloidea aequatorialis , Zompro, 2001: 196 . Otte & Brock, 2005: 262 . Brock & Büscher, 2022: 514 . Further material examined [ 12 ♀♀ , 10 ♂♂ , eggs]: 1 ♀ : Prov. Esmeraldas , 20 km . N.-O. Lita , 250 m . 12.IV.1990 ; Museum Paris , Equateur , S. Poulain rec. [ MNHN ] ; 2 ♂♂ , 3 ♀♀ , eggs: ex Zucht F. Hennemann 2009/10, Herkunft : NW-Ecuador, Prov. Esmeraldas , Durango , leg. H. Kaech [ FH , No’s 0659–1 to 5, E] ; 8 ♂♂ , 8 ♀♀ : Ex Zucht B. Kneubühler 2018, F1, Ecuador : Prov. Esmeraldas , Durango , leg. H. Kaech [coll. OC –0550–1 to 16] . Diagnosis. Females of this small species ( Fig. 54B ) most closely resemble those of the Panamanian Ph. ploiaria ( Westwood, 1859 ) but may be separated by the smaller size, less globose head, smaller but more acute cephalad horns ( Fig. 58A ), more rounded and node-like tubercles of the mesonotum, much smaller lobes of the praeopercular organ ( Fig. 61M ), somewhat more delicate legs and lack of a rounded apical lobe on the posterodorsal carina of the meso- and metatibiae. Males ( Fig. 54F ) resemble those of the two Ecuadorian species Ph. ecuadoricus sp. n. and Ph. falcatus sp. n. , but readily differ from both by the considerably smaller size (body length <100.0 mm), more elongate and rather flattened head ( Fig. 60A ), presence of some small, node-like tubercles on the mesonotum and mesopleurae (wholly smooth in ecuadoricus and falcatus ), reddish brown legs and different morphology of the three terminal abdominal segments, which includes a notably shorter and broader, basically triangular vomer. From those of ecuadoricus they may furthermore be distinguished by the slender basitarsi, while they differ from falcatus also by the much smaller cephalad tubercles and somewhat longer alae. Description. The colouration is described from photos of live captive reared specimens. ♀♀ ( Fig. 54B ): Fairly small (body length including subgenital plate 138.1–162.2 mm ) and slender species with a moderately globose, bi-cornute head, a sparsely but distinctly tubercular and nodulose meso- and metathorax and a rather short subgenital plate. General colouration either various shades of brown and grey or dull to mid green. Tubercles of the mesothorax pale orange in green and dull ochre in brown specimens. Eyes pale yellowish with brown mottling. Meso- and metafemora with a faint pale transverse band in brown specimens. Antennae except for scapus and pedicellus reddish mid brown dorsally and slightly darker ventrally. Head ( Fig. 58A ): Moderately globose, about 1.3x longer than wide with the vertex weakly convex. Vertex supplied with a few minute granules and a pair of small ± acute conical horns just behind the eyes; these slightly directed laterad and cephalad and extending by less than one-quarter the height of head capsule. Between these horns often a narrowly placed pair of node-like tubercles. Frons with a distinct, oval impression between the bases of the antennae that has a granule at each outer corner. Eyes small but strongly projecting, circular in outline and their diameter contained about 2.2x in length of genae. Antennae ± reaching to posterior margin of abdominal segment II. Scapus compressed dorsoventrally, very slightly constricted towards the base but with the lateral margins straight; 1.6x longer than wide. Pedicellus round in cross-section, slightly constricted apically and less than half the length of scapus. Antennomere III scarcely longer than pedicellus. Thorax: Pronotum slightly shorter and noticeably narrower than head, 1.7x longer than width of posterior margin with the anterior half notably narrowed and the lateral margins very gently rounded in posterior half. Surface supplied with some node-like granules, which are roughly arranged in two converging rows in front and two sub-parallel and closely placed longitudinal rows posterior of the shallow and gently curved transverse median sulcus; each row consisting of three granules. A few further scattered granules in lateral portions of segment, the anterior margin somewhat raised and with four small granules in the median portion. Mesothorax 8x longer than prothorax and uniform in diameter with only a slight widening posteriorly. Mesonotum unevenly set with a variable number of differently sized node-like to conical tubercles and small granules in between but the armature much less pronounced in the very posterior portion; a longitudinal row of granules indicated close to lateral margins. Metanotum scarcely more than one-quarter the length of mesonotum and some 1.8x longer than wide; surface sparsely granulose and with a few small, conical tubercles only towards the lateral margins. Meso- and metapleurae with a longitudinal marginal row of moderately sized tubercles and some further but much smaller, irregularly dispersed granules. Meso- and metasternum just sparsely granulose and tubercular. Abdomen: Median segment 1.6x longer than metanotum, gently constricted medially with the posterior margin broader than anterior margin and about 5.3x longer than wide. Segment II somewhat shorter than median segment and 3x longer than wide. II–V slightly increasing and VI–VII decreasing in length; VI longest and 3.6x longer than wide; all roughly uniform in diameter. Tergum VI with a ± distinct bi-lobed excrescence posteromedially. VII shorter than all preceding segments with the lateral margins just very weakly deflexed and rounded posteriorly. Sterna just very weakly and sparsely granulose to almost smooth. Praeopercular organ indistinct and only represented by a dark marking and two small and shallow posteromedian protuberances close to posterior margin of sternum VII ( Fig. 61M ). Tergum VIII slightly less than half the length and somewhat narrower than VII, about 1.3x longer than wide, rectangular in dorsal aspect and strongly convex longitudinally. IX scarcely shorter than VIII and rectangular dorsal aspect. Anal segment a little longer than IX with the posterior portion somewhat widened, carinate medio-longitudinally, flattened towards the posterior and the posterior margin almost straight with the outer angles roundly deflexed ( Fig. 61L ). Cerci very small and tapered towards a blunt apex. Gonapophyses VIII elongate, upcurved, notably projecting over posterior margin anal segment and ± reaching apex of subgenital plate ( Figs. 61K–L ). Subgenital plate canaliculate in the posterior three-fifths and with a prominent, lamellate longitudinal lateral carina in basal two-fifths with the apex obtusely rounded and projecting beyond apex of abdomen by ± the length of anal segment ( Figs. 61K–M ). Legs: All moderately long and slender. Profemora about equal in length to mesothorax, mesofemora slightly longer than metathorax, metafemora almost reaching to posterior margin of abdominal segment IV and metatarsi not reaching apex of abdomen. Two outer ventral carinae of meso- and metafemora with a ± distinct, roundly triangular lobe about one quarter off the base of femur (less distinct on metafemora). Legs otherwise unarmed. All basitarsi with a prominent, roundly triangular dorsal crest; all slightly longer than corresponding following three tarsomeres combined. ♂♂ ( Fig. 54F ): Small (body length 89.0– 100.2 mm ) and slender for the genus with fairly well-developed alae, that roughly reach to posterior margin of abdominal segment III ( 30.2–32.8 mm ), a rather flattened and unarmed head, a sparsely granulose mesonotum and distinctly white lower portion of the genae. General colour dark greyish brown, the mesothorax with a slight dull greyish green wash, the abdomen rather ochreous mid brown basally and becoming darker brown towards the apex. Abdominal terga IX and X with an ochre to creamy white medio-dorsal marking. Head brown with a washed black postocular streak and the entire lower portion of the genae distinctly white. Eyes as in ♀♀ . Pronotum with the lateral portions blackish brown. Tegmina and costal region of alae pale brown with darker brown veins; both with the anterior margin broadly white to pale cream coloured (only in the basal portion of the alae although). Anal region of alae transparent pale brown. Vomer dark brown. Legs reddish mid brown and all femora and tibiae with three faint straw transverse annulae. Antennae greyish dark brown dorsally and slightly darker ventrally. Head ( Fig. 60A ): Oval and broadest at the eyes with the vertex flattened and smooth except for an impressed coronal line in the posterior half; only a pair of shallow and obtuse swellings just behind the eyes. Frons with a very distinct oval impression between bases of antennae. Eyes large, almost circular in outline and projecting hemispherical from head capsule; their diameter contained only 1.5x in length of genae. Antennae ± reaching half the way along abdominal segment VI. Scapus and pedicellus as in ♀♀ . Thorax: Pronotum almost as long but noticeably narrower than head and basically as in ♀♀ but destitute of granules. The transverse median sulcus somewhat more pronounced and almost reaching lateral margins of segment ( Fig. 60A ). Mesothorax 7.5x longer and a little narrower than prothorax, uniform in diameter with only a slight widening posteriorly. Mesonotum supplied with a very few scattered node-like granules and a longitudinal row of minute granules laterally. Mesopleurae with about ten node-like granules and the mesosternum acutely carinate medio-longitudinally but destitute of granules. Metapleurae and metasternum smooth. Tegmina elongate, slender, ovate-spatulate in shape and gradually narrowed in the anterior portion; central protuberance moderate and roundly conical. Alae ± reaching to posterior margin of abdominal segment III. Abdomen: All segments smooth dorsally and ventrally. Median segment over 4x longer than metanotum, very gently narrowed towards the posterior. Segment II about 0.7x the length of median segment. II–VII uniform in diameter but slightly decreasing in length with II over 5x and VII hardly 4x longer than wide; VII gently constricted medially. Tergum VIII 3/5 the length of VII, gradually widening towards the posterior, trapezoidal in dorsal aspect and the posterior margin about 1.3x wider than anterior margin. IX about 1.3x longer than VIII, constricted medially and with the anterior portion slightly widened; lateral margins straight but somewhat deflexed posteriorly. Anal segment noticeably shorter than IX ( Fig. 66H ), roundly rectangular in dorsal aspect, scarcely longer than wide and slightly narrowed apically; posterior margin with a very shallow median indention and the outer angles rounded. Ventral surface of posterior margin set with a few minute black teeth. Cerci shorter than anal segment, very gently incurved and slightly thickened at the apex. Vomer basically triangular in shape and gradually narrowing towards a single, acute and upcurved terminal hook; lateral margins straight and moderately inflated, ventral surface Vshaped, moderately lowered and irregularly furrowed transversely. Poculum moderately bulgy and convex, roundly rectangular in lateral aspect ( Fig. 66H ) with a fairly pronounced medio-longitudinal carina that is most prominent and somewhat lamellate at the angle; almost reaching to posterior margin of tergum IX and with posterior margin obtusely triangular. Legs:All long, slender and wholly unarmed. Profemora a little longer than head, pro- and mesothorax combined, mesofemora slightly longer than mesothorax, metafemora almost reaching posterior margin of abdominal segment V and metatibiae projecting considerably over apex of abdomen. All basitarsi slender and noticeably longer than remaining tarsomeres taken together. Variability. In addition to the colour, some variability is seen in the size of the cephalic horns and degree of the meso- and metathoracic nodes and tubercles of the ♀♀ . The immature holotype and specimen from near Lita in the collection of MNHN have the cephalic horns notably larger than the captive reared specimens from Durango in the authors collections. Both these specimens also have the thoracic armature somewhat more pronounced . Egg ( Fig. 99A ). Fairly small but typical for the genus. Shape angularly ovoid with the polar-area flattened and the dorsal egg surface strongly convex in the anterior portion; oval in cross-section. Capsule surface deeply and densely punctured, the pits of moderate size and well defined. Micropylar plate large, elongate-oval and gradually narrowing towards the anterior, slightly more than two-thirds the length of capsule and only about 2.5x longer than wide; sculptured like capsule but with the pits smaller and the entire surface gently convex longitudinally. Micropylar cup a distinct elongate swelling with a cup-like protuberance at posterior end of plate. Median line indistinct, short and not reaching to posterior pole of capsule. Operculum oval and with a moderately sized, hollow and raised structure that is formed by excrescences of the outer margin; shape roundly conical, flattened and pitted dorsally and with the lateral surface almost smooth basally; height only corresponding to about one-fifth the length of capsule. General colour plain grey with a slight ochre wash, mostly with a faint and weakly defined dark grey to black marking laterally. Centre of polar area and opercular collar brown, the micropylar cup blackish brown. Operculum reddish ochre. Measurements [mm]: Length including operculum 3.4–3.5, length 2.8–2.9, width 1.9–2.0, height 2.5–2.6, length of micropylar plate 2.0. Comments. The unique holotype in the collection of NHMW is a fairly damaged penultimate instar ♀ nymph, that lacks both antennae, the right front leg, left mid and hind leg and terminal four abdominal segments. Comparison with the fresh material at hand leaves no doubt the latter specimens represent this fairly distinctive and rather small species. Culture stock that originated from Durango in the Esmeraldas Province in north-western Ecuador was imported to Europe in 2007 and first successfully reared in captivity by Bruno Kneubühler ( Luzern , Switzerland ). It was since then distributed as Phanocles sp. “Durango” but the culture could only be maintained for few generations. In captivity in Europe bramble ( Rubus spp. , Rosaceae ), oak ( Quercus robur , Fagaceae ) and salal ( Gaultheria shallon , Ericaceae ) were accepted as alternative food plants. Unfortunately, the natural host plants are not known. The incubation time of eggs was 4–5 months at temperatures of 20–25°C and at the same climatic circumstances ♂♂ reached maturity after about 12 weeks, while ♀♀ on average took some 14 weeks to reach maturity. Females laid 3– 5 eggs per day, which they simply flicked away by an abrupt movement of the abdomen. Table 30: Measurements of Phanocles aequatorialis ( Redtenbacher, 1908 ) comb. n.