Studies on Neotropical Phasmatodea XXV: Revision of Pterinoxylus Serville, 1838, with the descriptions of two new species from Costa Rica. (Phasmatodea: Oriophasmata: Cladomorphinae: Pterinoxylini) Author Hennemann, Frank H. Author Conle, Oskar V. Author Valero, Pablo 0000-0002-5471-3458 Author Nishida, Kenji 0000-0002-6547-4704 kenji.nishida@gmail.com text Zootaxa 2022 2022-11-14 5208 1 1 72 http://dx.doi.org/10.11646/zootaxa.5208.1.1 journal article 188712 10.11646/zootaxa.5208.1.1 0121aac5-c21f-46c1-a222-6917ee5e2e6f 1175-5326 7318962 FDBFF270-AF6B-45ED-9995-BB8D77DD372D Pterinoxylus eucnemis ( Burmeister, 1838 ) Figs. 7–9 , 21C , 22C–D , 28 , 29A–B , 32 Haplopus eucnemis Burmeister, 1838: 577 . HT , ♀ : 805, eucnemis Burm. *, difformipes Serv., Westw. *, Brasil . Virm. [MNHU]. Burmeister, 1840: 37 . Phasma (Haplopus) eucnemis , Haan, 1842: 127 . Pterinoxylus eucnemis , Kirby, 1904: 362 . Rehn, 1904: 61 . Otte & Brock, 2005: 294 . Zompro, 2005: 261 . Bellanger, Lelong & Jourdan, 2018: 274 . Conle, Bellanger, Lelong, Jourdan & Valero, 2020: 125. Pterinoxylus difformipes Serville, 1838: 227 . HT , ♀ : Amérique méridionale [MNHN – valde defectum]. Westwood, 1859: 90 , pl. 36: 1 ( ♀ ). Kirby, 1904: 362 . [As a synonym of P. eucnemis Burmeister, 1838 ] Redtenbacher, 1908: 428 (in part – only the records from Brazil and Suriname ). Shelford, 1909: 365 (in part – only the records from Brazil and Suriname ). Chopard, 1911: 348 . Rehn, 1957: 184 , pl. 21: 2 ( ♀ ). Roubaud & Lelong, 1993: 12, fig. 5 ( ♀ ). Clark–Sellick, 1998: 221, fig. 30h & i. [Egg and internal micropylar plate]. Otte & Brock, 2005: 294 . [As a synonym of P. eucnemis Burmeister, 1838 ] [ Not: Pterinoxylus difformipes , Rehn, 1904: 61 . Misidentification – nymphs from Costa Rica are P. spinulosus Redtenbacher, 1908 ] [ Not: Pterinoxylus difformipes , Zompro, 1997: 180 , figs. 2a & 2b. Misidentification relating to Pterinoxylus spinulosus Redtenbacher, 1908 ] Material examined ( 15 ♂♂ , 6 ♀♀ , 5 nymphs, eggs): BRAZIL : 2 ♀♀ : Brazil : Para, A. Miles Moss Coll., B.M. 1947–453 [ NHMUK ]; 1 ♂ : Obidos, s/data, Diversos 1 [MZUSP]; 1 ♀ : Manaos, Brazil , II– III .43 [AMNH]. FRENCH GUIANA : 1 ♂ : Guyane Francaise, Nouveau Chantier, Collection Le Moult [ MNHN ]; 1 ♂ : Guyane Francaise, St–Jean du Maroni, Collection Le Moult, Mai, Museum Paris, Collection Lucien Chopard, 1914 [ MNHN ]; 1 ♀ : Guyane Francaise, St–Jean du Maroni, Collection Le Moult [ MNHN ]; 1 nymph (n3): Guyane, Febrere, Roubaud rec. [ MNHN ]; 2 nymphs (n3): Guyane, A 1 53 N 1, 10. VII .93, Roubaud rec. [ MNHN ]; 1 ♂ : Französisch Guyana : Commune de Roura, Montagne des Chevaux, RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , S.E.A.G [coll. OC, No. 0269–1]; 1 ♂ : Französisch Guyana : Commune de Mana Laussat (Ouest), 05°28'31.6N – 053°35'07.3W , P3 Sable Blanc [coll. OC, No. 0269–2]; 1 ♂ : Französisch Guyana : Commune de Régina, Nouragues, Saut Pararé, 4°02'N – 52°41'W , S.E.A.G [coll. OC, No. 0269–3]; 1 ♂ : Französisch Guyana : Commune de Mana Laussat (westlich), 05°28'31.6N – 053°35'07.3W , P3 Sable Blanc [coll. OC., No. 0269–4]; 2 ♂♂ : Französisch Guyana : Commune de Roura, Montagne des Chevaux, RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , S.E.A.G [coll. OC., No. 0269–5 to 6)]; 1 ♂ : Französisch Guyana : Commune de Camopi, Bergmassiv Sommet Tabulaire, Mount Itoupé, N 03°01’23’’ W 053°05’44’’ , 600m , Pente ouest, S.E.A.G [coll. OC, No. 0269–7]; 3 ♂♂ : Französisch Guyana : Commune de Roura, Montagne des Chevaux, RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , S.E.A.G, Lichtfang [coll. OC, No’s. 0269–8 to 10]; 1 ♂ : Französisch Guyana : Commune de Roura, Montagne des Chevaux, RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , S.E.A.G, Malaise–trap [coll. OC, No. 0269–11]. SURINAME : 1 ♂ : Coll. Br. v. W., Surinam ?, Deyrolle; det. Br. v. W. Pterinoxylus difformipes , 46.; 6371, 6371 [ NHMW , No. 824]; 1 ♀ (penultimate instar): Suriname , Coeroeni–Eiland, 17.IX.1959 , St. Legoni [RMNH]; 1 ♀ (nymph): Suriname , Coeroeni Eiland, 3–X–1959 , D. C. Geyskes [RMNH]. BOLIVIA : 1 ♀ : Bolivia , Mamon River, A. E. Mc. Dougall, B. M. 1922–351 [ NHMUK ]. NO/WRONG DATA: 1 ♀ : Amboina, Phasma Angustata [RMNH – possibly from Suriname ]. FIGURE 7: Pterinoxylus eucnemis ( Burmeister, 1838 ) ♀♀. A. HT from Brazil, dorsal view [MNHU]; B. Specimen from French Guiana (Saint Jean du Maroni) in dorsal view [MNHN]; C. Specimen from French Guiana (Saint Jean du Maroni) in dorsolateral view [MNHN]; D. Specimen from French Guiana in lateral view [MNHN]. Diagnosis: Females are very similar to those of P. spinulosus ( Redtenbacher, 1908 ) , with which they share the flattened basal portion of the subgenital plate and distinct dorsal apical lobe of the meso- and metatibiae. They can however be distinguished by: on average more pronounced cephalic tubercles; more broadened subgenital plate, which has the lateral margins more prominently deflexed and arcuate over the entire length of the plate and the apex obtusely rounded to very gently biconcave (more decidedly tri-dentate in spinulosus ); more prominent praeopercular organ, as well as the much more prominently deflexed and unevenly lobate/undulate posteroventral carina of the protibiae ( Fig. 9G ) and notably more distinct sub-basal dorsal lobe of the mesotibiae. Males differ from those of P. spinulosus by: the somewhat more robust body, relatively shorter mesothorax; prominent lateral lobes of abdominal tergum VII ( Figs. 9E–F ); lack of spines on the mesonotum; considerably more prominent cephalic tubercles; more developed and broader lobes of the protibiae and generally more distinct armature of the extremities as well as the paler grey anal region of the alae. Also the eggs ( Figs. 22C–D ) strongly resemble those of P. spinulosus and may only differentiated by the more distinct posterior constriction just above the polar extension. Description: In addition to preserved specimens, the colouration is described from pictures of a live individual from French Guiana taken by Philippe Lelong ( France ). FIGURE 8: Pterinoxylus eucnemis ( Burmeister, 1838 ) ♂♂. A. Specimen from French Guiana, dorsolateral view [coll. OC]; B. Specimen from French Guiana, dorsolateral view [coll. OC]; C. Specimen from French Guiana, dorsal view [coll. OC]. ♀♀ ( Figs. 7 , 9A–C , 28 , 29A–B ). Medium to large for the genus (body length including subgenital plate 142.0– 174.0 mm), form twig –like and moderately stocky with the body surface partly and unevenly sculptured, the anterior legs very strongly undulate and lobate. General colouration variable and ranging from almost blackish brown over various shades of greyish or ochraceous mid brown to buff; either almost plain but more often irregularly flecked with combinations of these colours and/or with lichenose areas. Abdominal tergum VIII usually with an ocelliform black spot anterolaterally and terga II–VI with an aggregation of dark spots in posterior portion. Legs in particular irregularly mottled with paler and darker tones of brown. Head with two conspicuous triangular, sharply defined black markings between the eyes. Antennae ochre to dull straw with all antennomeres brown apically. Tegmina and costal region of alae roughly of same colour as body; alae generally paler although. Basal portion of costal region of alae with an oval, sub–basal, slightly pinkish area. Anal region of alae slightly transparent dark brown to dark grey with all anal veins boldly marked with deep black (figs. 7A, 26). Head: Oval in cross-section, parallel-sided and about 1.6x longer than wide. Between the eyes with two transverse swellings. Vertex with two bluntly conical swellings in centre that each bear two to three low tubercles; a further pair of more distinct, conical tubercles at posterior margin. Otherwise sparsely granulose. Eyes circular in outline and their diameter contained about 2.3x in that of genae. Antennae almost reaching to tip of protarsi and laid back about reaching two-thirds the way along mesothorax; consisting of 22–23 antennomeres. Scapus flattened dorsoventrally, strongly deflexed laterally and sub-circular in outline with the base narrowed. Pedicellus subcylindrical and about three-fifths the length of scapus. Third antennomere slightly longer but considerably narrower than pedicellus. Thorax: Pronotum about as long but slightly narrower than head, almost rectangular and with a slight constriction medially; transverse median sulcus distinct, gently curved and expanding over entire width of segment. Median line slightly impressed in anterior portion; the surface set with several granules and small tubercles. Mesothorax about 3.8x longer than pronotum, oval in cross-section and very slightly swollen pre-medially. Mesonotum with a fine but irregularly median carina, surface irregularly tuberculose to nodose and about one-thirds off the anterior margin with two more or less prominent knots or swellings of irregularly strumose rugulae. Meso- and metapleurae rugulose and tuberculose. Prosternum with a small, rough anterolateral sensory area at lateral margins; a further much larger and oval central sensory area on probasitsternum. Meso- and metasternum irregularly and to a variable degree set with small nodes or tubercles. Tegmina sub-oval with texture similar to that of body and with a fairly prominent, rounded hump in centre; slightly projecting over posterior margin of metanotum. Alae slightly projecting over posterior margin of median segment. Abdomen: Median segment almost 2x longer than metanotum and notably longer than abdominal segment II; 1.5x longer than wide and smooth. Segments II–VII almost uniform in length; on average 1.4x longer than wide and sub-rectangular. Segments II and III very slightly widening, IV and V widest segment, VI narrower than preceding. Tergum VII with lateral margins in posterior half of segment dilated into a prominent, bluntly angular or foliaceous, somewhat dorsal directed lobe which laterally extends by at least half of the body width; surface of lobe strongly wrinkled and rugulose ( Fig. 9B–C ). Terga VIII–X considerably narrower than all preceding and roughly uniform in width. Complete surface of all terga rugulose and granulose; all with a fine longitudinal carina and two irregular, sub-parallel rugulae. These rugulae posteriorly terminating in a more or less crenulate or foliaceous lobe on tergum III. The median carina terminating in an obtuse posterior swelling on VII–IX. Sterna II–VII unevenly rugulose and each with a pair of short, obtuse, converging ridges near posterior margin. Sternum VII with praeopercular organ formed by a small posteromedian hump ( Fig. 9C ). VIII almost two-thirds the length of VII strongly convex and almost 2x longer than wide; IX three-quarters the length of VIII, rectangular, slightly tectiform and about 1.5x longer than wide. Anal segment notably shorter than IX, flattened and slightly narrowed in posterior portion; posterior margin sub–truncate with an almost semi-circular median excavation and the outer angles bluntly rounded and somewhat labiate ( Fig. 9B ). Epiproct sub-truncate, shield-shaped and extending beyond posterior margin of anal segment ( Fig. 9B ). Cerci very small, subcylindrical, tapered towards a fairly pointed tip and just reaching posterior margin of anal segment. Subgenital plate extending over apex of abdomen by at least the length of the two terminal terga combined; uniformly canaliculate longitudinally, scaphiform in dorsal aspect with the lateral margins strongly arcuate but gradually lowering towards the bluntly sub-truncate posterior margin ( Figs. 9A–C ). Legs: All relatively short and stocky; profemora two-thirds the length of mesothorax, mesofemora shorter than metathorax and hind legs reaching about halfway along abdominal segment VI. Anterodorsal carina of profemora strongly raised, deflexed and undulate. Posteroventral carina slightly dilated and forming two or three rounded lobes; the terminal one being the largest and somewhat foliaceous. Medioventral carina indistinct. Anterodorsal carina of protibiae with several large, rounded to bluntly triangular and tooth-like foliaceous lobes of variable sizes; the posteroventral carina very prominently dilated and lamellate with the margin very irregularly and unevenly undulate; anteroventral carina less distinctly dilated and gently wavy ( Fig. 9G ). Posterodorsal carina of mesofemora with a prominent, rounded sub-apical lobe and 2–3 much smaller, occasionally toothed lobes in basal half; anterodorsal carina smooth except or a prominent, rounded sub-apical lobe. Antero- and posteroventral carinae sparsely and minutely denticulate, with a slight rounded expansion post-medially and an enlarged, triangular tooth sub-apically. Dorsal carinae of metafemora each with a prominent, obtusely triangular sub–apical lobe, which is notably more pronounced on the posterior carina; antero- and posteroventral carinae with 4–9 indistinct, blunt teeth which gradually increase in size towards the apex of femur. The medioventral carina of meso- and metafemora very indistinct. Ventral carinae of meso- and metatibiae smooth; the anterodorsal carina with a prominent toothed lobe near the apex and a narrower, rounded lobe sub-basally; the latter much less pronounced on metatibiae. Probasitarsus with dorsal carina strongly raised and bearing a distinct denticulate or crenulate lobe; posteroventral carina somewhat rounded; second tarsomere with a similar but much smaller dorsal lobe. Meso- and metabasitarsus slightly longer than following tarsomere and only with a small triangular, bi-dentate dorsal lobe. ♂♂ ( Figs. 8 , 9D–F, 9H , 21G ). Medium sized for the genus (body length 89.3–105.5 mm ), form moderately slender and stick-like with well-developed alae (length 50.7–51.8 mm ) and distinctly undulate protibiae. General colouration of body different shades of straw over grey and mid to dark or almost blackish brown; usually irregularly flecked with combinations of these tones and occasionally lichenose. Head with a pair of conspicuous, triangular black markings between the eyes. Pronotum in anterior half with a bold back longitudinal median streak and often with two black spots at posterior margin. Anterior margin of mesonotum sometimes with a small V-shaped black marking. Abdominal terga II–IV with two minute black spots in posterior third of segment. The largest cephalad and thoracic nodes and tubercles often buff, brown or ochre. Tegmina and costal region of same colour as body; in lighter brown specimens the alae with several irregular dark brown to black markings along the longitudinal veins. Anal region of alae translucent grey with numerous smaller and larger transparent patches; all anal veins dark greyish brown with interruptions at the transparent patches. Antennae straw to reddish pale brown. Head: Generally as in ♀♀ , but eyes more prominent and projecting hemispherically from head capsule with their diameter contained less than 2x in length of genae. All tubercles smaller but more defined and node-like. Antennae reaching posterior margin of tegmina and with 28–30 segments; otherwise as in ♀♀ but scapus less prominently dilated. Thorax: Pronotum as in ♀♀ but tubercles less prominent; a prominent pair of spiniform tubercles present near posterior margin and the lateral margins with a deep but narrow semi-circular excavation pre-medially. Mesothorax elongate, slender, cylindrical and 4x longer than pronotum; complete surface slightly and unevenly rugulose. Mesonotum with a faint longitudinal median carina and set with a variable number of distinct nodes to spiniform tubercles, mostly in anterior two-thirds; one pre-medial pair usually larger than all others. Meso- and metapleurae with a longitudinal row of small nodes. Mesosternum with prominent longitudinal median rugulae in anterior half; metasternum with two irregular and partly interrupted, sub-parallel longitudinal median carinae and a few small nodules. Tegmina sub-oval in outline, slightly projecting over posterior margin of metanotum and strongly convex with a prominent, rounded central hump. Alae reaching about one third to half way along abdominal tergum VI. Abdomen: Median segment 1.5x longer than metanotum, 3x longer than wide and smooth. Segment II shorter than median segment, II to VII slightly decreasing in length; II about 4x, VII only 3x longer than wide. All with a faint longitudinal median carina; II–VI parallel-sided and of uniform width. Tergum VII with lateral margins in posterior half dilated into a gently rounded lobe, that laterally extends by about one-thirds the width of segment ( Figs. 9E–F ). Terga II–IV smooth, V–VII and all sterna rugulose; sterna II–VIII sparsely nodose and each with an irregular longitudinal lateral carina. Tergum VIII three-quarters the length of VII, strongly convex and very slightly widening towards the posterior. IX three-quarters the length of VIII, almost 2x longer than wide and parallel-sided; both with a blunt longitudinal median carina. Anal segment two-thirds the length of IX, sub-rectangular in dorsal aspect, slightly narrowed and weakly tectinate in posterior portion; the posterior margin with a slight median indention and the outer portions broadly rounded; ventral surface of outer portions of posterior margins armed with a few small denticles. Epiproct very small and rounded. Cerci small, distinctly oval in cross-section, gradually tapered towards an obtuse apex and slightly projecting over posterior margin of anal segment; the apical portion slightly incurved. Vomer triangular in shape, somewhat longer than breadth of base, the terminal hook posterior directed, short but acutely triangular and strongly upcurved ( Fig. 20C ). Poculum weakly convex with a bluntly conical basal hump and roughly reaching to posterior of abdominal tergum IX ( Fig. 9D ), the posterior half carinate longitudinally ( Fig. 9F ) and the posterior margin broadly bi-lobed with a fairly small median indention ( Fig. 20C ). Legs: Relatively longer and slenderer than in ♀♀ ; profemora about as long as mesothorax, mesofemora threequarters the length of mesothorax and hind legs reaching about half way along abdominal segment VI. Anterodorsal and posteroventral carina of profemora moderately lamellate, the former irregularly lobate with the lobes bluntly triangular, the latter slightly wavy with a few small teeth, the most apical ones of which are largest. Anterodorsal carina of protibiae raised and irregularly undulate, occasionally with a prominent rounded lobe near the apex; posteroventral carina strongly lamellate with the margin gently wavy to undulate. Armature of mid and hind legs generally as in ♀♀ but much less pronounced. Probasitarsus as long as following two tarsomeres combined and with a distinct (sometimes dentate) triangular dorsal lobe; second tarsomere with a much smaller triangular dorsal lobe. Meso- and metabasitarsus almost as long as following two tarsomeres combined, dorsal carina slightly raised and rounded. Eggs ( Figs. 22G–H ): Very large, alveolar, capsule more than 3x longer than wide, almost cylindrical in crosssection. Dorsal surface more convex than ventral and lateral surfaces. Capsule surface minutely but very densely granulose, distinctly pitted and with several irregular and obtusely raised longitudinal ridges. A distinct, obtuse, longitudinal swelling below micropylar plate. Polar-area with a hollow, crest-like and slightly lamellar excrescence; the outer margin irregularly crenate. Micropylar plate small, covering less than one-thirds of capsule, ovoid to spear-shaped in outline and narrowed towards anterior end. Micropylar cup small and near posterior margin of plate. Operculum almost circular and with a minute central tubercle. Outer margin with a prominent, hollow crest-like protuberance, which has the anterior margin irregularly crenate; height over one-quarter of capsule length (not fully developed or artificially deformed in the illustrated example, Figs. 22G–H ). General colour plain chestnut brown. A second example from Réserve Naturelle Nationale de la Trinité examined from photographs is considerably darker brown in colour and shows and much more pronounced posterior constriction of the capsule just above the polar extensions (seen in the ovipositor, Fig. 9B ). Measurements [mm]: Overall length 11.8, capsule length 10.0, width 3.3, height 3.5, length of micropylar plate 2.6, height of operculum 1.8. FIGURE 9: Pterinoxylus eucnemis ( Burmeister, 1838 ) . A. Apex of abdomen of ♀ from French Guiana in lateral view (Photo by Philippe Lelong) [coll. PL]; B. Apex of abdomen of same ♀ as in figure A in dorsal view with an egg held in the ovipositor (Photo by Philippe Lelong) [coll. PL]; C. Apex of abdomen of ♀ from French Guiana in ventral view [MNHN]; D. Apex of abdomen of ♂ from French Guiana in lateral view [coll. OC]; E. Apex of abdomen of ♀ from French Guiana in dorsal view [coll. OC]; F. Apex of abdomen of ♂ from French Guiana in ventral view [coll. OC]; G. Right protibia of ♀ from French Guiana in extero–lateral view, arrows indicate dorsal and apical direction [MNHN]; H. Ventral view of head, pro– and mesothorax of ♂ from French Guiana, showing sensory areas [coll. OC]. Comments: Burmeister (1838: 577) described Haplopus eucnemis based on a ♀ from “Aus dem Innern Brasiliens” (= Inner Brazil ) in the collection of MNHU ( Fig. 7A ). A full-sized figure and redescription of the specimen were presented by Westwood (1859: 90 , pl. 36: 1). The original description of P. difformipes was based on a single ♀ specimen from “Amérique méridionale” (= South America) which Serville had bought from the collection of P. A. Latreille. As the holotype has since not been traced and due to great parts of Latreille's collection being housed in MNHN, extensive investigation to locate Serville's type–specimen was conducted by the authors in 2002. This revealed a strongly mutilated, but certainly very old ♀ specimen without collecting data but with an old green determination label stating “ P. difformipes Serville ”. The green label is typical for specimens of that author and indicates the possible type-status of the concerned specimen. It lacks the complete head, pro- and mesothorax as well as the fore and mid legs as well as parts of the subgenital plate. Nevertheless, it is quite certainly Serville's holotype of P. difformipes . Westwood (1859: 90) synonymised Haplopus eucnemis Burmeister with P. difformipes Serville. Although this synonymy is correct in the aspect that these two taxa are conspecific, Westwood used the wrong name. Serville's work was published in late December 1838 (often erroneously cited as 1839) while Burmeister’s “Handbuch der Entomologie” part 2 section 1 was published earlier in 1838 (Bragg, 2001: 620). Hence, Burmeister's publication pre–dates Serville's work and P. difformipes Serville must be regarded a junior synonym, with H. eucnemis Burmeister being the senior name. The ♂ was subsequently described by Redtenbacher (1908: 428) based on a specimen from Surinam in the collection of NHMW. Zompro (1997: 180 , figs. 2a & b) described and figured the egg of what the author believed to be P. difformipes , based on an example which was extracted from the ovipositor of a ♀ from Costa Rica in ZMUH. Examination of the specimen however clearly shows this to be misidentified and to represent P. spinulosus Redtenbacher, 1908 (à see below). P. eucnemis appears to be widely distributed in the northern half of South America, although only rarely encountered. Redtenbacher (1908: 428) recorded it from Brazil and Surinam but also from Guatemala and Costa Rica . The last two Central American records however are based on misidentified material which represents P. perarmatus ( Redtenbacher, 1908 ) and P. spinulosus Redtenbacher, 1908 . Chopard (1911: 348) recorded two ♂♂ from French Guiana in MNHN but obviously overlooked a ♀ also from French Guiana and contained in the collection of Le Moult in MNHN ( Figs. 7B–C ). The collection of NHMUK contains a ♀ from as far south as the Mamoré River in NE– Bolivia , which is far away from all other currently known records of P. eucnemis . As the record is unique and no further specimens have been recorded from nearby localities subsequently it must currently be regarded doubtful. Distribution ( Fig. 32 ): NE– Brazil (Est. Pará: Óbidos; Manaos). French Guiana (Nouveau Chantier; St. Jean du Maroni & Rue Nationale 1; Réserve Naturelle Nationale de la Trinité ( Bellanger, Lelong & Jourdan, 2018: 274 ); Commune de Roura, Montagne de Chevaux; Commune de Mana Laussat, Sable Blanc; Commune de Régina, Nourages, Saut Pararé; Commune de Camopi, Sommet Tabulaire, Mount Itoupé). Suriname ( Sipaliwini District , Res. Coeroeni). NE– Bolivia (Est. Beni , Río Mamoré).