Pterinoxylus spinulosus Redtenbacher, 1908

Hennemann, Frank H., Conle, Oskar V., Valero, Pablo & Nishida, Kenji, 2022, Studies on Neotropical Phasmatodea XXV: Revision of Pterinoxylus Serville, 1838, with the descriptions of two new species from Costa Rica. (Phasmatodea: Oriophasmata: Cladomorphinae: Pterinoxylini), Zootaxa 5208 (1), pp. 1-72 : 44-54

Pterinoxylus spinulosus Redtenbacher, 1908

Figs. 17–19 , 20B , 21F , 22G–H , 26–27 , 33

Pterinoxylus spinulosus Redtenbacher, 1908: 428 , pl. 20: 3 (♂). LT, ♂: Coll. Br. v. W, Chiriqui ( Panama) Staudinger; det. Br. v. W. Pterinoxylus spinosus ; 10.295 [NHMW, No. 825]. (Not: PLT, ♂ (penultimate instar nymph): Syntype; V. de Chiriqui, below 4000 ft., Champion; 89; Godman–Salvin Coll. 1908.–168.; B.C.A. Orth. II, Pterinoxylus spinulosus Redt. ; Pterinoxylus sp. larva; BMNH(E) #844939 [NHMUK]. This is P. perarmatus ( Redtenbacher, 1908)) Shelford, 1909: 365 . Hebard, 1923: 361. Rehn, 1957: 188, pl. 21: 1 (♀). [Description of ♀] Robinson, 1968: 195, figs. 1–4. [Notes on defensive behaviour] Robinson, 1970: 52, fig. 1. [Notes on defensive behaviour] Hogue, 1993: 169. [Notes on defensive behaviour] Brock, 1997: 21. [Type data in NHMUK] Brock, 1998: 59. Clark–Sellick, 1998: 221. [Micropylar plate] Otte & Brock, 2005: 294. Conle, Hennemann & Gutiérrez, 2011: 64. [Lectotype designation] Brock, Marshall, Beccaloni & Harman, 2016: 194 . [Type data in NHMUK]

Pterinoxylus difformipes, Rehn, 1904: 61 . [Nymphs from Costa Rica – misidentification]. Redtenbacher, 1908: 428 (in part – only specimen from Costa Rica in MNHN). Shelford, 1909: 365 (in part – only records from Costa Rica and Panama). Zompro, 1997: 180, fig. 2a & b (egg). [Description of egg from Costa Rica – misidentification].

Material examined (35 ♂♂, 46 ♀♀, 8 nymphs, eggs):

PANAMA:

1 ♀: Barro Colorado Isl., Panama, July29 1933, J.D. and H.Hood; Rehn 1957 figured; Measured specimen Rehn 1957; Pterinoxylus spinulosus Det. Rehn 1957 A.N.S.P. [ANSP]; 1 ♀: Barro Colorado Island, C. Z., Panama, I.12.1956, Berthe Ruud; Measured specimen Rehn 1957; Pterinoxylus spinulosus Det. Rehn 1957 A.N.S.P. [ANSP]; 1 ♀: Bruja Point, Canal Zone, Panama, X.21.1929 (J. Zetek); Measured specimen Rehn 1957; Pterinoxylus spinulosus Det. Rehn 1957 A.N.S.P. [ANSP]; 1 ♂: Pan.-Barro Col. Is. C.Z.–coll. Rettmeyer, VI.- VIII.1957 [ANSP]; 1 ♀: Barro Colo Is; CZ, VI.1939; Jas Zetek, No 4997; Lot No 39–15883 [USNM]; 1 ♀: Panama, Pearl Is, San Jose; Morrison JPE, Feb. 18.1944 [USNM]; 1 ♂: Muséum Paris, Barro Colorado, Z. Gregoire, 1954; 214; Pterinoxylus difformipes Serv. [ MNHN]; 3 nymphs: V. de Chiriqui, below 4000 ft., Champion; Pterinoxylus difformipes Serv., Godman – Salvin Coll. 1908.–168. [ NHMUK]; 2 eggs: Laid 11/67 Balboas; Panama; Pterinoxylus spinulosus, Brit. Mus. 1974 –118 [ NHMUK]; 1 ♂: Gamboa, Ex.-Z. Rov. Pmá, Colón, 3 Feb 1980, col. R. Angulo, Victor Leon [ MIUP]; 1 ♂: Panama: Panamá Pr., Arraijun, 20–Jun–2001, W.I. Louiza [ MIUP]; 1 ♀: Panamá, Colón, Cerro Botija, 23 julio 2012, col. David Correa [ MIUP]; 1 ♀: Panama: Los Santos, R.F. La Tronosá, El Cortezo, 6 may 2006, A. Santos [ MIUP]; 1 ♀: Panama: El Valle de Anton, IV–30–1945, T. Patiño [AMNH].

COSTA RICA:

1 ♀, 1 egg (ex ovipositor): Costa Rica, Farm Hamburg, a. Reventazon 28.10.1927 a. Madero negro. Eing. 1928 No. 1, Ferd. Nevermann leg. [ ZMUH]; 1 ♂: Costa Rica, Guapiles 30.5.33, Eing. Nr. 69, 1936; PHA 73 Zoologisches Museum Hamburg [ ZMUH]; 1 ♀ (nymph): Turrialba, Costa Rica; Coll. Schild & Burgdorf [ANSP]; 1 ♀ (nymphs): Tucurrique, Costa Rica; Coll. Schild & Burgdorf; Pterinoxylus eucnemis (Burm.) ? Det. Rehn [USNM]; 1 ♀: 26–53; 15–11–53, I.I. CA, Turrialba, C. R.; Viale; Pterinoxylus spinulosus Redt. , det. A.B. Gurney 1961 [USNM]; 1 ♀: Cachi, C. R., C.H.L. [ NHMUK]; 1 ♂: Costa Rica, Prov. Limón, Veragua Rainforest, Restaurant, 400–440m, 7 SEP 2008, J. Mara, Colecta Libre, L_N_212220_625230 #94928; MNCR –A004170413 MNCR-ACRI Costa Rica [ MNCR-A]; 1 ♀: Est. Sirena, 0–100m, P.N. Corcovado, Prov. Punt., Costa Rica, G. Fonseca, May 1991, L–S– 270500, 508300; Costa Rica MNCR –A CRI000 598337 [ MNCR-A]; 1 ♀: Est. Pistilla, 700m, 9km S Sta. Cocilla, Prov. Guan., Costa Rica, P. Ríos & C. Moraga, Oct 1990, L–N–830200, 880200; Costa Rica, MNCR-A CRI000 226284 [ MNCR-A]; 1 ♂: Costa Rica, Prov. Heredia, Sarapiqui, Z.P. La Selva, El Ceibo, 500–600m, 19 AGO 2005, M. Ballestero, I. Chavez, Colecta Libre, L_N_256615_527735, #94220; INB004154644, MNCR-ACRI Costa Rica [ MNCR-A]; 1 ♀: Terraba (Pacifique) H. Pittier, Oa Kúnjiga = expulgadera del diablo; Pterinoxylus difformipes Serv. [ MHNG].

GUATEMALA:

1 ♀: Polochic River, Guatemala, Oct. 07 A.P. Coll.; Pterinoxylus eucnemis Burm. ♀, A.N.C. [USNM]; 1 ♀ (nymph, n3): Chacoj, R. Polochic, Guatemala, Champion, 91, B.C.A. Orth. II, Pterinoxylus difformipes Serv. [ NHMUK].

BELIZE:

1 ♀: ex Zucht: M. Rotter; Belize, Green Hills nr. Belmopan 2020 [coll. FH, No. 1238–1]; 2 ♀♀, 1 ♀ n4, 42 eggs: ex Zucht: F. Hennemann; Belize, Green Hills nr. Belmopan 2020 [coll. FH, No. 1238–2 to 4, E2]; 1 ♀, 4 ♂♂: Ex Zucht: B. Kneubühler 2018, F1. Belize: Belmopan, Green Hills Butterfly Ranch, leg. J. Meerman 08.2016 [coll. OC, No. 0291–2 to 6]. 14 ♀♀, 16 ♂♂, 11 eggs: Ex Zucht: B. Kneubühler 2019, F2. Belize: Belmopan, Green Hills Butterfly Ranch, leg. J. Meerman 08.2016 [coll. OC, No. 0291–7 to 37].

HONDURAS:

1 ♀: Honduras 1923, Rio Paulaya: Barranco, IV.16, 407; T.H. Hubbell; Pterinoxylus sp. cf. spinulosus Redt. ♀ Det. T.H. Hubbell 1949; Pterinoxylus sp. near spinulosus ; UMMZI–178790 [UMMZ]; 1 ♂: Honduras 1923, Tela: V –5 Guaimas district, 529 T.H. Hubbell; Pterinoxylus sp. near spinulosus Redt. ♂ Det. T.H. Hubbell 1949; Pterinoxylus sp. near spinulosus ; UMMZI–178789 [UMMZ].

COLOMBIA:

1 ♀: Colombia, Andagoya, R. Condoto, Choco, H.G.F. Spurrell, 1916–273 [ NHMUK]; 1 ♂: fecha: I–05/69; loc: Anserma, Hosp: Maleza, Col: J.G.B. [UCA]; 1 ♂: Orden: Phasmatodea , Familia: Phasmatidae, Localidad : SJ. Putm., Fecha: [...] 2005 [MHN-UC]; 1 ♂: Colombia, Valle, Punta Soldado, En.15/2001, leg. Salazar, F. V. ♂; Colombia, Universidad Caldas – Centro de Museo Historia Natura, CI–01–044– [MHN-UC]; 1 ♀: Colombia, Valle, Punta Soldado, En. 28/2001, leg. Salazar, J.C. Vargas ♀; Colombia, Universidad Caldas – Centro de Museo Historia Natura, CI–01–044– [MHN-UC]; 1 ♀ (penultimate instar nymph): Colombia, Valle, Punta Soldado, Dic. 30/2000, leg. Salazar, J.C. Vargas ♂; Colombia, Universidad Caldas – Centro de Museo Historia Natura, CI–01–044–6 [MHNUC]; 1 ♂: CO05. San Luis, Vereda La Tebaida, Intradomita, Techo, 3– III –2005, CEUA [CEUA]; 1 ♂: Colombia: Antioquia, Porce Lat. N 6°54'38'' Long. W75°4'49'' Alt. m.s.n.m. 1500, Maleza, Enero 1983; Raúl Vélez; MEFLG N.C. 12013 [MEFLG]; 1 ♀: Colombia, Valle del Cauca, Buenaventura, Bajo Calima, Noviembre 1961, Francisco Gallego; MEFLG N.C. 14464 [MEFLG]; 1 ♂: Kolumbien: Dept. Tolima, Hamburgo, Río Magdalena, 800, Ex ZMHB [coll. OC, No. 0291–1].

ECUADOR:

1 ♂: ECUADOR – Esmeraldas Montalvo, La Mayronga 95m 14/ 17.XI.2004, N00º53'27.2'' W079º13'02.5'', leg. F. M. Buzzetti & G. Carotti [coll. OC, No. 0291–38].

Diagnosis: Females are very similar to those of the type –species P. eucnemis ( Burmeister, 1838) , with which they share the flattened basal portion of the subgenital plate and distinct dorsal apical lobe of the meso– and metatibiae. They are however easily distinguished by: the averaging smaller and more obtuse cephalic tubercles; more slender subgenital plate, which has the lateral margins less distinctly deflexed and narrowing towards the apex ( Fig. 19A ) and the apex more or less distinctly tri–dentate (obtusely angular to slightly biconcave in eucnemis ); less distinct praeopercular organ, as well as the less prominently deflexed and more even and less lobate/undulate carinae of the protibiae that is more or less gradually widening towards the apex of the tibia ( Fig. 19K ) and much less notable to obsolete sub-basal dorsal lobe of the mesotibiae ( Fig. 19L ). Males differ from those of P. spinulosus by: the somewhat slenderer body, relatively longer mesothorax; parallel-sided abdominal tergum VII ( Fig. 19G ); spinose mesonotum ( Figs. 19I–J ); considerably less pronounced and obtuse cephalic tubercles; notably less deflexed carinae of the protibiae and generally less distinct armature of the extremities as well as the darker grey anal region of the alae. The eggs are very similar and almost indistinguishable from those of P. eucnemis . They may only be differentiated by the on average less elongate overall shape and less distinct posterior constriction of the capsule.

Description: ♀♀ ( Figs. 17 , 19A–E, 19K–L , 26A , 27 ). Medium to large for the genus (body length including subgenital plate 137.0–189.0 mm), form twig –like and fairly slender (maximum body width at mesothorax 7.0– 8.5 mm) with the body surface partly and unevenly sculptured, the anterior legs strongly undulate and lobate. Colouration very variable and ranging from dark 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 white or creamy white areas. Abdominal tergum VIII with an ocelliform black spot anterolaterally (sometimes also present on tergum IX). Legs like body irregularly mottled with paler and darker tones of brown. Head with two blackish markings between the eyes. The larger cephalad and thoracic tubercles may be ochre to dull orange. Antennae ranging from greyish ochre to reddish mid brown. Tegmina and costal region of alae roughly of same colour as body; the basal portion of costal region of alae that is covered by the tegmina contrastingly red. Anal region of alae slightly transparent dark brown to dark grey with all anal veins boldly marked with deep black.

Head: Oval in cross-section, parallel-sided and about 1.6x longer than wide. Between the eyes with a low transverse swelling, that is indented medially. Vertex gently rounded and unevenly tuberculose, usually with two pairs of rounded swellings in centre and a further pair of enlarged, obtuse tubercles at posterior margin. Eyes circular in outline and their diameter contained about 2.3x in that of genae; strongly projecting. Antennae almost reaching tip of protarsi and laid back about three-quarters the way along mesothorax; consisting of 28–29 antennomeres. Scapus flattened dorsoventrally, strongly deflexed laterally and strongly narrowed basally. Pedicellus subcylindrical and almost three-quarters the length of scapus. Third antennomere slightly longer but considerably narrower than pedicellus.

Thorax: Pronotum longer and slightly narrower than head, sub–rectangular with anterior somewhat narrower than posterior margin; transverse median sulcus U-shaped, moderately distinct and not reaching lateral margins of segment. Median line slightly impressed in anterior portion; the surface set with several granules and small tubercles. Occasionally there is a conspicuously enlarged pair of spiniform tubercles near posterior margin. Mesothorax variable in shape and length, ranging from 3.7x to 4.3x the length of pronotum and ranging from fully parallel-sided to being slightly swollen pre-medially. Mesonotum with a fine but irregularly median carina, surface texturing very variable and ranging from irregularly rugulose over sparsely nodose to prominently tuberculose; about one-thirds off the anterior margin with two more or less prominent knots or swellings of irregularly strumose rugulae or blunt tubercles. Meso- and metapleurae rugulose and more or less prominently tuberculose. Sensory areas at lateral margins of prosternum weakly developed and indistinct; the central sensory area of probasitsternum distinct and notably swollen. 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 moderately prominent, rounded hump in centre; slightly projecting over posterior margin of metanotum. Alae at least reaching one-thirds the way along abdominal segment II.

Abdomen: Median segment almost 2x longer than metanotum and considerably longer than abdominal segment II; 1.6x longer than wide and smooth. Segments II–VII almost uniform in length; on average 1.5–1.8x longer than wide and sub-rectangular. Segments II widening towards the posterior, III widest and the following in general very slightly gradually narrowing with VIII–X almost uniform in width and narrowest segments. Tergum VII with lateral margins in posterior half of segment dilated into a prominent, bluntly angular but irregularly shaped lobe, which laterally extends by at least one-thirds of the body width ( Figs. 19B–E ). Complete surface of all terga rugulose and granulose; all with a closely placed pair of fine longitudinal carinae that vary in emphasis and usually terminate in a nodule or small swelling posteriorly on each tergum. Surface furthermore with two irregular, sub-parallel and sinuate rugulae, which posteriorly terminate in a more or less crenulate, bi- or trifid lobe on III and IV; the size and shape of these lobes very variable. Sterna II–VII unevenly rugulose with the rugulae longitudinally directed and each with a pair of short, obtuse, converging ridges near posterior margin; otherwise sparsely granulose. Sternum VII with praeopercular organ formed by a small conspicuously node-like posteromedian swelling ( Fig. 19D–E ). Tergum VIII widened anteriorly, narrowed pre-medially and almost two-thirds the length of VII; strongly convex and almost 2x longer than wide. IX about three-quarters the length of VIII, rectangular and about 1.5x longer than wide. Anal segment somewhat shorter than IX, flattened and slightly narrowed in posterior portion; the posterior margin sub-truncate with a wide, slightly angular median excavation and the outer angles bluntly triangular. Epiproct small shieldshaped and extending beyond posterior margin of anal segment; with an acute longitudinal median carina dorsally ( Figs. 19B–C ). Cerci very small, oval in cross–section, compressed laterally, tapered towards a fairly pointed tip and just reaching posterior margin of anal segment; the apex somewhat incurved. Subgenital plate variable in length and extending over apex of abdomen by at least the length of the two terminal terga combined; uniformly canaliculate longitudinally ( Fig. 19A ), narrowly scaphiform in dorsal aspect with the lateral margins sub-parallel, more or less arcuate and gradually lowering towards the sub-truncate and roughly tri-dentate posterior margin ( Figs. 19B–E ).

Legs: All relatively short and stocky; profemora two-thirds the length of mesothorax, mesofemora shorter than metathorax and hind legs reaching about half way along abdominal segment VI. Anterodorsal carina of profemora strongly raised, deflexed and undulate. Posteroventral carina much dilated with margin wavy and with a large, roughly triangular lobe sub-apically. Medioventral carina indistinct. Anterodorsal carina of protibiae with several roundly triangular teeth-like lobes and a somewhat enlarged, denticulate lobe apically; the posteroventral carina more or less gradually dilated and lamellate towards the apex of tibia with the margin somewhat undulate; the apical portion usually somewhat arched towards the anterior and forming a more or less distinct sub-trigonal lobe ( Fig. 19K ). Posterodorsal carina of mesofemora with a prominent, almost semi-circular sub-apical lobe ( Fig. 19L ) and 2–3 much smaller, occasionally toothed lobes in median portion; armature of anterordorsal carina similar but less pronounced. Antero- and posteroventral carinae very sparsely and minutely denticulate, supplied with a tooth-like expansion post-medially and an enlarged, triangular tooth sub-apically. Posterodorsal carina of metafemora with a very prominent, upright and narrow triangular sub-apical lobe, which has the apical margin dentate; antero- and posteroventral carinae supplied with several small teeth and notably enlarged, flat but broad tooth sub-apically. The medioventral carina of meso- and metafemora very indistinct. Ventral carinae of meso- and metatibiae smooth; the anterodorsal carina with a fairly small tooth sub-basally and a prominent more or less rounded and toothed lobe in the apical portion ( Fig. 19L ). Probasitarsus with dorsal carina strongly raised and bearing a distinct denticulate, roughly triangular lobe; second tarsomere with a similar but much smaller dorsal lobe. Meso- and metabasitarsus hardly longer than following tarsomere and with the dorsal carina just slightly raised and rounded.

♂♂ ( Figs. 18 , 19F–J, 19M , 21F ). Medium sized to large (body length 96.0– 104.8 mm), form slender and stick-like with well-developed alae (length 39.5–46.0 mm) and a distinct apical dorsal lobe on all tibiae. General colouration of body different shades of ochre, grey or mid to dark brown more rarely with a greenish hue especially on thoracic segments and front legs; usually irregularly flecked with combinations of these tones. Head with a pair of small blackish markings between the eyes ( Fig. 19J ). The largest thoracic tubercles and spines ochre to yellowish straw. Metapleurae often greenish. Tegmina and costal region of same colour as body; the tegmina often with a white diagonal stripe in posterior half, forming a white V-shaped marking when the tegmina are closed; the costal region of the alae with the portion posterior to the radial vein more or less distinctly red. Anal region of alae translucent greyish brown with numerous smaller and larger transparent patches; all anal veins marked with darker greyish brown and with interruptions at the transparent patches.Antennae dark straw to reddish pale brown; apex of all segments brown.

Head: Generally as in ♀♀, but eyes more prominent and projecting hemispherically from head capsule with their diameter contained a little less than 2x in length of genae. All cephalad tubercles and tubercles averaging more prominent, more pointed and spiniform ( Figs. 19I–J ). Antennae almost reaching posterior margin of median segment and with about 30 segments; otherwise as in ♀♀ but with scapus less prominently dilated.

Thorax: Pronotum as in ♀♀ but somewhat longer in relation; near posterior margin with a prominent pair of spiniform tubercles or spines and the lateral margins distinctly concave ( Figs. 19I–J ). Mesothorax elongate, slen- der, cylindrical and somewhat more than 4x longer than pronotum; complete surface very sparsely and unevenly granulose. Mesonotum with a very faint longitudinal median carina and all over set with a very variable number of spiniform tubercles to spines of variable sizes ( Fig. 19I ), which range from obtuse to slender and more or less acutely pointed in shape; size and number of spines decreasing towards the posterior. A marginal row of smaller tubercles along lateral margins. Meso- and metapleurae with a longitudinal row of nodes. Meso- and metasternum with some irregularly placed nodes; the mesosternum with some irregular rugulae in anterior portion. Tegmina oval in outline, slightly projecting over posterior margin of metanotum and moderately convex with a fairly prominent, rounded central hump. Alae ± reaching to posterior margin abdominal tergum VI.

Abdomen: Median segment 1.6x longer than metanotum, 2.8x longer than wide and smooth. Segment II longer than median segment, II to VII slightly gradually decreasing in length; II about 4.3x and VII only 3.3x longer than wide. All terga with a fine pair of closely placed parallel longitudinal median carinae, which often are raised posteriorly and may terminate in a pair of curved ridges on III–V; a further lateral carina present on VI–VIII. Surface of terga II–VII otherwise sparsely and faintly rugulose and granulose. II–VII parallel-sided and roughly uniform in width. Sterna II–VIII very sparsely nodose and each with a pair of very obtuse and low longitudinal carinae, which are somewhat more raised at posterior margin of each sternum. Tergum VIII almost three-quarters the length of VII, strongly convex and very slightly widening towards the posterior; IX ¾ the length of VIII and indistinctly longer than wide and narrowing towards posterior margin; both trapezoidal in dorsal aspect. Anal segment two-thirds the length of IX, sub-rectangular in dorsal aspect and slightly narrowing towards posterior, weakly tectinate; the posterior margin with a small median indention and the outer portions broadly rounded ( Fig. 19G ); ventral surface of outer portions of posterior margins armed with a few small denticles. Epiproct very small, rounded and fully hidden under anal segment. Cerci small, distinctly oval in cross-section, laterally compressed at base and somewhat tapering towards a fairly obtuse apex; slightly projecting over posterior margin of anal segment. Vomer broadly triangular in shape, somewhat wider than long with the terminal hook narrowed, posterior directed, short but acutely pointed and upcurved ( Fig. 20F ). Poculum moderately convex in basal portion ( Fig. 19F ) with the apical half gradually flattening and roughly reaching to posterior of abdominal tergum IX, the posterior half carinate longitudinally and the posterior margin with a wide, concave median excavation ( Fig. 19H ).

Legs: Relatively longer and slenderer than in ♀♀; profemora a little longer than mesothorax, mesofemora almost three-quarters 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 forming some obtuse, broad tooth-like lobes and the latter with a few obtuse dentations. Anterodorsal carina of protibiae moderately raised and lamellate, usually forming a ± distinct (sometimes bifid) lobe apically; posteroventral carina just very slightly deflexed and slightly wavy. Armature of mid and hind legs generally as in ♀♀ but less pronounced ( Fig. 19M ). Probasitarsus almost as long as following two tarsomeres combined and with a large (sometimes dentate) dorsal lobe; second tarsomere with a much smaller triangular (sometimes bifid) dorsal lobe ( Fig. 19J ). Meso- and metabasitarsus almost as long as following two tarsomeres combined, dorsal carina slightly raised and rounded.

Nymphs: Newly hatched nymphs have a body length of about 20 mm and are mostly dark brown with the lateral portions of the body of a somewhat paler brown or ochre. The meso- and metatibiae bear a conspicuous white transverse median band and the terminal three antennomeres are contrasting white. There are also two distinctive pinkish markings on the frons between the eyes. The head bears the characteristic cephalic tubercles, which are however much more pronounced than in the adults insects and the meso- and metatibiae already bear the characteristic dorsal apical lobe. Later instars are various shades of ochre, grey and brown and in general have the legs more stocky and the armature of the extrimities more distinct than the adult insects. Females have the lateral lobes of abdominal tergum VII much larger than adults.

Variability: Both sexes, but ♀♀ in particular, show considerable intraspecific variability relating to several morphological characters. These include overall size, colour, armature of the head, pronotum and mesothorax, length of the alae as well as size and shape of the teeth and lobes of the extremities. In ♀♀ the shape and size of the pair of posterior projections on abdominal terga II–IV as well as the length and shape of the subgenital plate are fairly variable. While the latter has the posterior margin always obtusely tri-dentate, the projections of abdominal terga II–IV may either be almost obsolete or protruded into bi- or trifid spiniform projections, that are always most pronounced on III. The most noteworthy variability however is seen in the shape and armature of the mesonotum. While this is almost parallel–sided and irregularly tuberculose to rugulose in specimens from Costa Rica and Panama, several of the Colombian examples have the mesonotum slightly swollen pre-medially and set with numerous fairly prominent, obtusely spiniform tubercles (most pronounced in the ♀ from Andagoya in the NHMUK collection ( Fig. 17E ) and ♀ from Valle de Cauca in MEFLG). The ♀ from Andagoya is also stockier than all other specimens at hand and has the mesothorax only 3.7x longer than the prothorax (4.0–4.3x longer in specimens from Panama and Costa Rica). Similar variability can be observed in the mesothoracic armature of ♂♂, which ranges from obtuse tubercles to slender and acute spines. In ♀♀ the alae show considerably variability in length, mostly reaching no more than half way along abdominal segment II but in the specimen from Guatemala in the collection of USNM reaching as far back as to the posterior margin of that segment. While ♂♂ from Panama and Costa Rica are various shades of grey and brown, greenish colourations of the thorax and fore legs in particular are sometimes seen in specimens from Colombia. Furthermore it is noteworthy, that in general Colombian specimens of both sexes have the two outer ventral carinae of the meso– and metafemora more distinctly denticulate than examples from Panama, Costa Rica or Belize.

Eggs: ( Figs. 22G–H ): The following description is based on two eggs laid by a ♀ from Balboa, Panama in the collection of NHMUK, an egg that was extracted from the ovipositor of the ♀ from Reventazon, Costa Rica in the ZMUH collection and eggs of the culture stock from Greenhills, Belize,

Very large, alveolar, capsule more than 3x longer than wide, almost cylindrical in cross-section; very slightly curved in lateral aspect and dorsal surface somewhat more convex than ventral and lateral surfaces. Surface of capsule very minutely but densely granulose, distinctly pitted and with several irregular, longitudinal but fairly low and indistinct ridges. 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, spear-shaped in outline. Micropylar cup small and at posterior margin of plate. Outer margin slightly raised and darker brown than capsule. Operculum almost circular and with a minute central tubercle. Outer margin with a prominent, hollow crest-like protuberance which has the anterior margin with five conspicuous crown-like teeth; height almost one-thirds of capsule length. General colour plain reddish mid brown (examples from Panama and Costa Rica) or dark ochraceous brown (examples from Belize). Some variability is seen in the length-diameter ratio, with some eggs being notably more elongate in general appearance than others. Measurements [mm]: Overall length 12.4–12.7, capsule length 9.3–9.6, width 2.8–3.2, height 3.1–3.0, length of micropylar plate 2.7–2.9, height of operculum 3.0–3.2.

Comments: Redtenbacher (1908: 428) decribed P. spinulosus based on a ♂ in the collection of NHMW and a ♂ nymph in the NHMUK collection both from Valle del Chiriquí in NE-Panama and provided an illustration of the NHMW specimen (Plate 20, Figure 3 ) which has been designated as the lectotype by Conle, Hennemann & Gutiérrez (2011: 64). The ♂ penultimate instar nymph (paralectotype) in NHMUK is not the same species but represents P. perarmatus ( Redtenbacher, 1908) . This is evident by the large pair of cephalic lobes and strongly spinose mesonotum. Rehn (1957) presented a very detailed description of the ♀ based on three examples from the Canal Zone of Panama in ANSP and provided notes on the stridulation observed in ♀♀. The ♀ and egg from Reventazon, Costa Rica in ZMUH referred to as P. difformipes Serville, 1838 by Zompro (1997: 180) are clearly misidentified and represent P. spinulosus ( Fig. 17D ). The ♀ from Balboa (now part of Panama City), which laid the two eggs in NHMUK that served for the description here presented, is not traced. Clark–Sellick (1998: 221, fig. 30h, i) briefly described and figured the internal micropylar plate of the egg.

After a storm in 2016 Jan Meerman ( Belize) collected a ♀ on the area of his butterfly farm “Green Hills” near Belmopan, Belize. The specimen was found on a fallen over tree of Coccoloba belizensis (Polygonaceae) and kept alive for some time in captivity. Since leaves of this tree were frequently eaten it is very like to be a host plant of P. spinulosus in Belize. Eggs were sent to Bruno Kneubühler (Zürich, Switzerland), who first successfully reared this species in captivity. It has proven not difficult to culture and accepts Salal ( Gaultheria shallon , Ericaceae ) and bramble ( Rubus spp. , Rosaceae ) as alternative food plants. Eggs take some 4–6 months to hatch at average temperatures of 20°C and nymphs are fairly slow growing.

Distribution ( Fig. 33 ): Panama: (Prov. Colón, Canal Zone, Parque Nacional Soberanía: Barro Colorado Island; Balboas; Bruja Point; Gamboa; Prov. Colón: Valle de Antón; Prov. Colón: Distrito Donoso, Cerro Botija; Prov. Panamá: Islas Perlas, Isla San José; Prov. Panamá: Arraijun; Prov. Los Santos: Reserva Forestal Tronosá, El Cortezo; Prov. Chiriquí: Valle del Chiriquí). Costa Rica: (Prov. Limón: Veragua Rainforest, Restaurant, 400–440 m; Guápiles; Prov. Puntarenas: Osa Peninsula, Corcovado National Park, Est. Sirena 0–100 m; Térraba; Prov. Guanacastae: Est. Pistilla 700 m; Prov. Cartago: Tucurrique & Turrialba; Prov. Heredia: Sarapiqui, Estacion El Ceibo 500–600 m; Prov. Puntarenas: Río Térraba; Cachi & Madero Negro, Reventazón). Honduras: Dept. Atlántida (Guaimas district, Tela); Dept. Colón (Río Paulaya, Barranco). E-Guatemala: Polochic River (Chacoj). Belize: (Cayo, Green Hills nr. Belmopan). Colombia: (Dept. Chocó: Rio Condoto, Andagoya; Dept. Caldas: Anserma; Dept. Antioquia: Porce, 1500 m; Dept. Quindío: San Luis, Vereda La Tebaida, Intradomita). Ecuador (Prov. Esmeraldas, Montalvo, La Mayronga).

Figures

FIGURE 3: Pterinoxylus cocoense n. sp. ♂ PT [MNCR-A, collection no. #107146] and egg. A. PT, dorsolateral view; B. PT, dorsal view; C. PT, ventral view; D. Apex of abdomen of PT in lateral view; E. Apex of abdomen of PT in dorsal view; F. Apex of abdomen of PT in ventral view; G. Egg in dorsolateral view, PT (operculum lacking).

FIGURE 17: Pterinoxylus spinulosus Redtenbacher, 1908 ♀♀. A. Specimen from Barro Colorado Island, Panama, dorsal view (specimen figured by Rehn, 1957) [ANSP]; B. Same specimen, dorsolateral view [ANSP]; C. Specimen from Barro Colorado Island, Panama, dorsolateral view [ANSP]; D. Specimen from Reventazon, Costa Rica, dorsal view [ZMUH]; E. Specimens from Andagoya, Colombia in dorsolateral view, showing very strong thoracic armature [NHMUK].

FIGURE 18: Pterinoxylus spinulosus Redtenbacher, 1908 ♂♂. A. LT from Chiriquí, Panama, dorsolateral view [NHMW, No. 825]; B. Specimen from Barro Colorado Island, Panama, dorsal view (Photo by Paul D. Brock) [USNM]; C. Specimen from San Luis, Colombia, dorsolateral view [CEUA].

FIGURE 19: Pterinoxylus spinulosus Redtenbacher, 1908. A. Apex of abdomen of ♀ from Parque Nacional Corcovado, Costa Rica, in lateral view [MNCR-A, collection no. 598337]; B. Apex of abdomen of same specimen in dorsal view [MNCR-A, collection no. 598337]; C. Apex of abdomen of ♀ from Reventazon, Costa Rica on dorsal view [ZMUH]; D. Apex of abdomen of same specimen in ventral view [ZMUH]; E. Apex of abdomen of ♀ from Polochic River, Guatemala, in ventral view (Photo by Paul D. Brock) [USNM]; F. Apex of abdomen of ♂ from M.N. Guayabo, Costa Rica, in lateral view [MNCR-A, collection no. #74105]; G. Apex of abdomen of same specimen in dorsal view [MNCR–A, collection no. #74105]; H. Apex of abdomen of same specimen in ventral view [MNCR-A, collection no. #74105]; I. Head, pro- and mesonotum of ♂ from Barro Colorado Island, Panama, in dorsolateral view [MNHN]; J. Head, pro- and mesonotum of ♂ from M.N. Guayabo, Costa Rica, in dorsal view [MNCR-A, collection no. #74105]; K. Left protibia of ♀ from Greenhills, Belize in dorsal view, arrows indicate exterior and apical direction [coll. OC]; L. Right mid leg of ♀ from Greenhills, Belize in anterolateral view [coll. OC]; M. Right mid leg of ♂ from Greenhills, Belize in anterolateral view [coll. OC].

FIGURE 20: Pterinoxylus spp. venation of alae. A. P. speciosus n. sp. ♀ HT, dorsal view (mirrored horizontally) [MNCR-A, collection no. 042809]; B. P. spinulosus Redtenbacher, 1908 ♀, ventral view with tympanal area coloured yellow [coll. OC]; C. P. speciosus n. sp. ♂ PT, dorsal view (mirrored horizontally) [MNCR-A, collection no. 52571]. Explanation of abbreviations: 1AA-7AA = first–seventh anterior anal; 1PA-7PA = first–seventh posterior anal, C = costa, Cu = cubitus, Cu1 cubitus 1, Cu2 = cubitus 2, E = empusal, M = media, MA = media anterior, Mp1 = media posterior 1, Mp2 = media posterior 2, R = radial, R1 = radius 1, Rs = radial sector, Sc = subcostal, TA = tympanal area.

FIGURE 21: Pterinoxylus spp. terminal two abdominal segments of ♂♂ in ventral aspect, showing vomer and poculum. A. P. cocoense n. sp.: PT [MNCR–A, collection no. #107146]; B. P. crassus (Kirby, 1889): Specimen from Martinique [coll. FH, No. 0588–21]; C. P. eucnemis (Burmeister, 1838): Specimen from French Guiana [coll. OC]; D. P. perarmatus (Redtenbacher, 1908): Specimen from Osa Peninsula, Costa Rica [coll. FH, No. 0841–6]; E. P. speciosus n. sp.: PT [MNCR–A, collection no. 52571]; F. P. spinulosus Redtenbacher, 1908: Specimen from Greenhills, Belize [coll. OC]. Scale = 1 mm.

FIGURE 22: Pterinoxylus spp. eggs. A. P. crassus Kirby, 1889, dorsal view; B. P. crassus Kirby, 1889, lateral view; C. P. eucnemis (Burmeister, 1838), dorsal view; D. P. eucnemis (Burmeister, 1838), lateral view; E. P. perarmatus (Redtenbacher, 1908), dorsal view; F. P. perarmatus (Redtenbacher, 1908), lateral view; G. P. spinulosus Redtenbacher, 1908 (Belize), dorsal view; H. P. spinulosus Redtenbacher, 1908 (Belize), lateral view. Explanation of abbreviations: Cer = cercus, Pa = paraproct, SG = subgenital plate (poculum), T9 = abdominal tergum 9, T10 = abdominal tergum 10 (anal segment), DF = thorn pads (“Dornenfeld”), Vo = vomer.

FIGURE 26: P. spinulosus Redtenbacher, 1908; captive reared specimens from Belize (© Bruno Kneubühler). Mating couple with ♂ showing the typical resting attitudes. The inset shows a ♂ in startle display with wings opened.

FIGURE 27: P. spinulosus Redtenbacher, 1908; captive reared from Belize. A. ♀ in resting position; B. Same specimen in startle display showing the convex contour of the alae; C. Same specimen in startle display showing the convex contour of the alae.

FIGURE 33: Map showing the known records for Pterinoxylus spinulosus Redtenbacher, 1908.