Parotocrania panamae, Hennemann & Conle, 2024

Hennemann, Frank H. & Conle, Oskar V., 2024, 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), Zootaxa 5444 (1), pp. 1-454 : 139-146

Parotocrania panamae sp. n.

urn:lsid:zoobank.org:act:48EE3766-2993-4CD0-ACCE-EF9D03CD5F3C

( Figs. 7A , 46A–C, 46F–H , 47C–E , 48A, 48C , 86D–E , 95J , 100C , 110A–C )

HT, ♀: Panamá, Panamá Pr., Parq. Nac. Soberanía, Camino Plantación, II-Sep–1999, D. Quintero A. [MIUP].

PT, ♀ (penultimate instar): Panamá, Panamá Pr., Parq. Nac. Soberanía, Camino Plantación, II-sSp–1999, D. Quintero A. [MIUP].

PT, 1 ♀, 3 ♂♂, 9 eggs: Panama, LP-08, Prov. Coclé, Valle de Antón, Alto La Mesa, Hochplateau , 850–900 m, leg. F. Hennemann 16.– 17.11.2019 (night); Alt.: 850 m, 8°38’09.9”N 80°07’18.9”W, Mountainous rainforest [ FH, No. 1313–1 to 4 & E–10] .

PT, 2 ♂♂: Panama, LP-06, Prov. Coclé, Valle de Antón, along Calle La Medina / La Mesa , nr. Río Guayabo 650–850 m, leg. F. Hennemann 14.– 17.11.2019 (night); Alt.: 650–850 m, 8°37’48.0”N 80°08’17.0”W, Mountainous rainforest [ FH,No’s 1313–5 & 6] .

PT, ♂: Panama, LP-04, Prov. Coclé, Res. Biol. Altos de Campana, Sendero Panamá “ Panama Trail”, 700–900 m, leg. F. Hennemann 13.11.2019 (night); Alt.: 700–900 m, 8°40’57.1”N 79°55’31.3”W, Moist cloud forest [ FH, No. 1313–7] .

PT, 2 ♀♀, 5 ♂♂, 1 egg: Panama: Prov. Coclé, Valle de Antón , 8º37’49.20”N 80º8’21.50”W, leg. O.Conle, F.Hennemann & P.Valero, 17.09.18. [OC–0551–1 to 8] .

PT, ♀: Panama: Prov. Panamá Oeste , Laguna de San Carlos, 8º37’30.07”N 80º3’0.26”W, leg. O.Conle, & P.Valero, 05.07.19. [OC–0551–9] .

Diagnosis. Females of this new species ( Fig. 46A ) are very well characterised by the globose head and pair of huge, white to cream-coloured, peg-like and apically inward angled cephalic horns ( Fig. 48A ), just minutely granulose thoracic segments and wholly unarmed legs. The large cephalic horns at first glance resemble the type-species of the genus Phanocloncrania gen. n., Ph. dorsuria (Stål, 1875) , but these ♀♀ differ by the generic characters such as the lack of a prominent medio-longitudinal keel on the ventral body surface. Males ( Figs. 46B–C ) resemble those of P. regina ( Conle et al., 2011) comb. n. in but are easily recognised and distinguished from this and all other known apterous members of the genus by the globose and tubercular head ( Fig. 48C ), that however lacks large cephalic horns like in P. cassicephalus ( Conle et al., 2011) comb. n. and P. segmentaria ( Redtenbacher, 1908) comb. n., and by having the thoracic segment all over supplied with distinctive white nodes.

Etymology: Named after the distribution in Panama. Neuter.

Description. The colouration is described on the basis of pictures of the live paratypes.

♀♀ ( Fig. 46A , 110A–B ): Medium-sized (body length including subgenital plate 148.0–166.0 mm) and moderately slender for the genus with a very minutely granulose dorsal body surface, a median segment that is almost equal in length to metanotum and huge cephalic horns. General colour uniform green, the ventral body surface brown and the terminal two abdominal terga with a brownish hue. Meso- and metapleurae red posteriorly and granules of the thoracic sterna white ( Fig. 86D ). Cephalic horns pale cream to white and the genae brown at the anterior ventral corner below the eyes ( Fig. 48A ). Clypeus and mandibles dull yellow. Eyes grey. Antennae dull green dorsally and dark grey ventrally. Meso- and metafemora with a ± distinct but irregularly shaped black transverse band sub-basally. Terminal tarsal segment pale cream-coloured.

Head ( Fig. 48A ): Strongly globose, only 1.1x longer than wide and roundly angular in dorsal aspect, the vertex roundly convex and armed with a central pair of huge peg-like, tubercular horns which bear a narrow and acutely, inward-angled point on top and extend by almost the height of the head capsule; surface of vertex otherwise unevenly supplied with node-like tubercles of variable sizes and a pair of somewhat enlarged distant tubercles at posterior margin.A longitudinal, postocular row about 4–5 small granules present. Frons with a shallow median impression in and a pair of obtuse swelling just before the eyes. Eyes small, circular in cross-section and their diameter contained 2.1x in length of genae.Antennae reaching half way along abdominal segment III. Scapus compressed dorsoventrally, carinate laterally with the outer lateral carina roundly deflexed in apical half, the deflexion downward-directed and scapus rectangular in dorsal aspect; 2x longer than wide. Scapus round in cross-section, somewhat narrowing towards the apex and about half the length of pedicellus. III much narrower and somewhat longer than pedicellus.

Thorax: Pronotum slightly longer but narrower than head, basically roundly rectangular in shape with the posterior half weakly dilated. Anterior margin somewhat inflated and followed by a deep transverse furrow which laterally terminates in an angular pit. Transverse median sulcus fairly shallow, weakly arcuate and not reaching lateral margins of segment; surface minutely and sparsely granulose ( Fig. 48A ). Mesothorax uniform in diameter and 7.2x longer than prothorax. Mesonotum minutely granulose and with a longitudinal row of some distantly placed but slightly pronounced granules. Metanotum sculptured like mesonotum, only one-third of whose length and 3x longer than wide. Meso- and metapleurae with a very few node-like granules. Meso- and metasternum unevenly set with white, node-like granules and both with a fine longitudinal lateral carina; mesonotum also with a fine but uneven medio-longitudinal carina.

Abdomen: Median segment scarcely shorter than metanotum, 3x longer than wide with lateral margins notably concave. Segment II about as long as median segment, II–VI gradually increasing in length and VII only about as long as II and narrower than all preceding segments. II–VI roughly uniform in diameter with tergum V about 3x longer than wide. Terga almost smooth except for a single pair of small node-like white granules, sterna II–VII minutely and sparsely granulose and with a fine longitudinal carina close to each lateral margin. Sternum VII with the lateral carinae posteriorly terminating in a somewhat rounded lobe, the praeopercular organ merely represented by a verrucose median protuberance close to posterior margin ( Fig. 46H ). Tergum VIII half the length of VII, weakly widening in posterior half and with the posterolateral angles protruded into a small, digitiform process ( Fig. 46G ); 1.2x longer than wide. IX similar but only two-thirds the length of VIII and somewhat wider than long. Anal segment narrowed at the base and slightly widening towards the posterior margin, which is broadly rounded; the lateral margins notably deflexed and forming and almost semi-circular, downward directed lobe ( Fig. 46F ). Epiproct very small and wholly concealed under anal segment. Cerci small, conical with a rather obtuse apex and slightly projecting over posterior margin of anal segment. Gonapophyses VIII just not reaching to apex of anal segment. Subgenital plate weakly bulgy longitudinally and with an obtuse medio-longitudinal keel in posterior one-third, the same portion gently downcurved ( Fig. 46F ) and the lateral surfaces with a distinct curved longitudinal keel; the apex triangular and projecting beyond tip of abdomen by about the length of anal segment ( Figs. 46G–H ).

Legs: Moderately long, slender and entirely unarmed. Profemora a little longer than mesothorax, mesofemora notably longer than metathorax, metafemora reaching to mid of abdominal segment V and metatibiae slightly projecting beyond tip of abdomen. Mesofemora gently downcurved. Probasitarsi with the dorsal carina weakly rounded, the dorsal carina of the meso- and metabasitarsi uniform in height; all ± as long as following three tarsomeres combined.

♂♂ ( Fig. 46B–C , 110A–C ): Moderately sized (body length 100.0–115.0 mm), moderately stocky for the genus with a globose and tubercular head, nodulose thoracic segments a median segment that is almost equal in length to metanotum. Apterous. Colouration variable and basically occurring in two distinct colour forms ( Figs. 46 B-C). Either brownish green to greenish ochre with the ventral body surface pale orange-brown and the legs green or mid brown with irregular darker speckles, the ventral body surface black and the legs buff. Node-like granules of the body white. Anterior portion of genae pale cream and with a blackish marking below the eye ( Fig. 48C ). Abdominal terga with a variable number of blackish spots and markings, the posterior portion of the poculum pale cream to white ( Fig. 47E ). Meso- and metafemora with a distinct but irregularly shaped black sub-basal transverse band and occasionally irregularly banded or flecked with black in the brown colour form. Antennae with two basal segments coloured like head, the remaining antennomeres ochre to mid drab dorsally and blackish ventrally.

Head ( Fig. 48C ): Sub-globose, 1.2x longer than wide, broadest just behind the eyes with the genae scarcely narrowing towards the posterior, the vertex strongly convex, rounded and in the middle with a pair of obtusely conical swellings; surface otherwise set with some distinct, node-like granules. Frons with a shallow median impression between bases of antennae. Eyes prominent, projecting hemispherical, circular in outline and their diameter contained 1.5x in length of genae. Antennae reaching half way along abdominal segment IV; otherwise mostly as in ♀♀ but the outer margin not notably deflexed anteriorly.

Thorax: Pronotum basically as in ♀♀ with the transverse median sulcus somewhat more pronounced ( Fig. 48C ). Mesothorax uniform in diameter and 8.1x longer than prothorax, metanotum less than 0.4x the length of mesonotum. Meso- and metanotum irregularly but rather sparsely supplied with a variable number of node-like white granules (these less pronounced on metanotum although). A few of these nodes also present on meso- and metapleurae and sterna; a medio-longitudinal carina on meso- and metasternum weakly indicated.

Abdomen: Median segment about as long as metanotum and somewhat widened posteriorly; surface with a few small, node-like granules. Segment II about as long as median segment, segments II–V roughly uniform in length, VI and VII slightly decreasing in length; II–VI about uniform in width and VII very weakly widening towards the posterior; V 5.3x longer than wide and VII only 3.5x longer than wide. Each tergum and sternum with one or two small pairs of white nodes. Tergum VIII slightly widening towards the posterior and a little more than half the length of VII, IX notably longer weakly dilated anteriorly and strongly convex longitudinally with the lateral margins gradually deflexed towards the posterior and almost straight; both with obtuse longitudinal lateral bulge. Anal segment only about half the length of IX, notably wider than long, constricted at anterior margin, the lateral margins gently convex and the swollen posterior margin weakly sinuate with the outer angles obtusely rounded ( Fig. 47D ); ventral surface of posterior margin wholly set with small denticles ( Fig. 95J ). Epiproct minute, transverse triangular and at best scarcely projecting over posterior margin of anal segment. Vomer roundly triangular with a rather short but acute somewhat dextral directed terminal hook, the outer margins somewhat inflated ( Fig. 95J ). Cerci slender, round in cross-section, straight and with a slightly club-like and in-ward curved apex. Poculum moderately bulgy, rounded with a small and obtusely peg-like central protrusion and the posterior margin weakly labiate and obtusely triangular ( Fig. 95J ); reaching to posterior margin of tergum IX ( Fig. 47C ).

Legs: All moderately long and slender and mostly entirely unarmed. Occasionally with one or two blunt teeth or irregularly shaped, slender lobes on the two outer ventral carinae of the meso- and metafemora. Profemora about as long as head, pro- and mesothorax combined, mesofemora notably longer than metathorax, metafemora projecting over apex of abdominal segment V and metatibiae projecting greatly beyond tip of abdomen. All basitarsi notably longer than following three tarsomeres taken together.

Variability. Males occur in a green ( Figs. 46B , 110C ) and a brown colour form ( Figs. 46C , 110A–B ) that are described in more detail below. Variability is seen in the meso- and metafemora of brown specimens, which are either wholly unarmed or have one or two teeth or lobes on the two outer ventral carinae. The size and shape of these appendages shows considerable variability. Slight variability is also seen in the shape of the cephalic horns in ♀♀. The small number of available specimens however makes it impossible to identify the complete range of possible intraspecific variability.

Egg ( Fig. 100C ). Moderately sized and of very unusual shape. Capsule somewhat compressed laterally, oval in cross-section, notably higher than wide and scarcely higher than long (at anterior margin), in lateral aspect strongly narrowed towards the polar area and the polar area flatted, giving the capsule a roundly trapezoidal shape; lateral margins almost parallel to each other. Entire surface of capsule surface coriaceous to weakly rugose. Micropylar plate oval, scarcely more than half the length of capsule, positioned centrally on dorsal surface. Both ends rounded but posterior with a small median gap in which the small, knob-like micropylar cup is placed. Median line indistinct and almost reaching to polar end of capsule. Operculum oval, flat and with a very large hollow structure, that in lateral aspect is roughly equal in size to the capsule and has the upper margin irregularly crenulate and incurved; this structure formed by a membranous excrescence of the outer margin; surface slightly uneven. Capsule and micropylar plate plain black, opercular excrescence reddish brown.

Measurements [mm]: Overall length 5.2, capsule length 2.8, width 2.1, height 3.0, length of micropylar plate 1.7.

Table 28: Measurements of Parotocrania panamae sp. n.

5.20. Genus Phanocles Stål, 1875

( Figs. 4A , 6B , 8B, 8D , 49–67 , 87 M-N, 88, 92–93, 97, 99, 100A–B, 104A, 109C, 111–119, 120A–B)

Type-species: Bacteria burkartii Saussure, 1868: 65 , by subsequent designation of Kirby, 1904: 353.

Phanocles Stål, 1875b: 28 , 81 (in part).

Kirby, 1904: 353.

Redtenbacher, 1908: 412. [Listed as a synonym of Bacteria Latreille et al., 1827 in error]

Shelford, 1909, 363, pl. 7: 3b.

Bradley & Galil, 1977: 190.

Bragg, 2001: 626.

Zompro, 2001a: 197, figs. 3–5, 72–75. [ Phanocles re-established]

Zompro, 2001b: 17, figs. 1–2.

Hennemann, 2002: 8, figs. 1–6.

Hennemann & Conle, 2003: 6.

Zompro, 2004a: 318.

Zompro, 2004b: 91.

Otte & Brock, 2005: 262 (in part).

Eilmus, 2008: 78.

Conle, Hennemann & Gutiérrez, 2011: 59 (in part).

Jourdan, Lelong & Bellanger, 2014: 489.

López-Mora & Llorente-Bousquets, 2018: 48 (in part).

Conle, Hennemann, Bellanger, Lelong, Jourdan & Valero, 2020: 6.

Brock & Büscher, 2022: 513 (in part).

Luna, 2022: 93.

Bacteria, Bates, 1865: 330 , 331, pl. 44: 13a (in part).

Westwood, 1859: 27, 79, pl. 22: 3 & pl. 13: 4 (in part).

Saussure, 1868: 65.

Redtenbacher, 1908: 412 (in part).

Shelford, 1909: 362 (in part).

Chopard, 1911: 345 (in part).

Hebard, 1924: 150, pl. 7: 1–3.

Otte & Brock, 2005: 262 (in part).

Brock & Büscher, 2022: 510 (in part).

Luna, 2022: 91 (in part).

Bostra, Rehn, 1904: 57 .

Redtenbacher, 1908: 46 (in part).

Shelford, 1909: 359 (in part).

Hebard, 1919: 159, 161, pl. 22: 5–6 & 19: 10–11.

Otte & Brock, 2005: 262 (in part).

Brock & Büscher, 2022: 511 (in part).

Luna, 2022: 91 (in part).

Bostranova Villet, 2023: 150 (in part).

Clonistria, Redtenbacher, 1908: 403 (in part).

Brock & Büscher, 2022: 512 (in part).

Dyme, Brunner v. Wattenwyl, 1907: 324 , 326 (in part).

= Paraphanocles Zompro, 2001a: 198 , figs. 6–7, 76–77, 124–125. (Type-species: Mantis keratosqueleton Olivier, 1792 , by original designation). syn. n.

Brock & Büscher, 2022: 513.

Phanocloidea Zompro, 2001a: 196 (in part).

Otte & Brock, 2005: 262 (in part).

Description. ♀, ♂ ( Figs. 49–56 ): Medium to very large (body length ♂♂ 90.0–154.0 mm, ♀♀ incl. subgenital plate 120.0–285.0 mm), moderately slender to fairly stocky Cladomorphini with a median segment that is equal in length or longer than the metanotum. ♂♂ apterous, brachypterous or alate. Body surface smooth in ♂♂, at least thorax almost always granulose or tubercular to a variable degree in ♀♀ (mesonotum with a few conical spines in one species). Colour of ♀♀ various tones of straw, grey or brown to almost black, more rarely green. Colouration of ♂♂ usually more complex and often with meso- and metathorax different from rest of body; head with lower portion of genae white to light cream-coloured and with a ± defined dark postocular streak; mostly lateral margins of abdominal tergum IX and/or dorsal surface of anal segment whitish to light cream-coloured. Head of both sexes very variable in shape and armature; in ♀♀ ranging from ovoid to almost spherical with vertex ± convex; in ♂♂ alike but occasionally sub-cylindrical with vertex flattened. The vertex may be unarmed but mostly bears a pair of shallow humps, tubercles, spines or prominent auriform horns in ♀♀ ( Figs. 57–58 , 59A–C, 59E ); more frequently unarmed or bi-cornute in ♂♂ ( Figs. 59F–J , 60A–J ). Eyes small, circular and strongly projecting, relatively larger and more prominent in ♂♂. Antennae very long and filiform, ± equal (♀♀) or considerably longer than head and complete thorax combined (♂♂). Scapus compressed dorsoventrally, roundly rectangular in dorsal aspect and longer than wide. Pedicellus notably shorter than scapus and ± round in cross-section; antennomere III longer than pedicellus. Pronotum ± as long but narrower than head (♀♀ in particular), roughly rectangular and distinctly longer than wide. Mesothorax elongate, at least 4.5x longer than prothorax in ♀♀ and 6x longer in ♂♂; roughly equal in width to prothorax and occasionally somewhat inflated pre-medially in ♀♀ and slightly narrower than prothorax in ♂♂. Posterior margin of mesonotum sometimes somewhat raised bulgy in ♀♀. Metathorax less than two-thirds the length of mesothorax, smooth in ♂♂ and sculptured like mesothorax in ♀♀. Meso- and metapleurae smooth in ♂♂, smooth, granulose or tubercular in ♀♀ (metapleurae armed with strong spines in one species). Mesosternum obtusely tectate longitudinally in ♀♀ and ± distinctly keeled medio-longitudinally in ♂♂; surface smooth in ♂♂, unarmed or to a variable degree set with scattered granules or tubercles in ♀♀ (only in one case with a few short spines). Metasternum mostly simple but occasionally with a very shallow medio-longitudinal carina in ♀♀. If wings present in ♂♂ the tegmina slender and spatulate with a shallow central hump and roughly reaching to posterior margin of metanotum; alae variable in length rather narrow in shape but reaching no further than half the way along abdominal segment IV. Abdomen excluding median segment longer than head and complete thorax combined. Median segment equal in length or longer than metanotum, much longer than metanotum in alate ♂♂. Abdominal segment II slightly shorter than median segment and III. II–VI longer than wide and roughly uniform in diameter, VII shorter than all preceding and V–VI longest segments. Terga V and/or VI often with a posteromedian tubercle or crest (♀♀ in particular). Tergum VII in ♀♀ with lateral margins ± deflexed and forming a rounded or sub-truncated posterolateral lobe. Sternum VII of ♀♀ with a prominent praeopercular organ that is formed by a pair of spiniform or digitiform protuberances or appendages or ± large, rounded to sub-truncated lobes at posterior margin ( Figs. 92–93 ). Terminalia of ♀♀ ( Figs. 61–64 ): Terga VIII–X considerably shorter and narrower than preceding and ± uniform in width. Anal segment subquadrate in dorsal aspect and flattened towards the apex, with a ± decided medio-longitudinal carina and the posterior margin broad with a minute median excavation. Epiproct very small and mostly or wholly concealed under anal segment. Cerci small, ± round in cross-section and much shorter than anal segment. Gonapophyses VIII strongly elongated, filiform, slightly upcurved at the apex and surpassing posterior margin of anal segment (in species with a short subgenital plate ± reaching to whose tip, in species with a rather long and lanceolate subgenital plate not reaching tip). Subgenital plate keeled longitudinally and variable in length but always projecting over tip of anal segment; usually projecting no more than combined length of terga VIII–X; shape ranging from obtusely spatulate to canaliculate and fairly lanceolate. Gonapophyses IX roughly as long as gonoplacs, the gonoplacs ± broadened. Terminalia of ♂♂ ( Figs. 65–66 , 67A–K ): Terga VIII–X noticeably broader than preceding segments and club-like. Tergum VIII ± trapezoidal and broadened towards the posterior. IX narrowed towards the posterior and ± constricted medially; lateral margins straight to very moderately rounded and deflexed. Anal segment variable in shape and length, longer than wide, rather flattened and usually somewhat declining towards the posterior margin; the latter rounded and entire or more frequently with a ± distinct posteromedian indention or excavation and the posterolateral angles rounded and occasionally ± protruded; ventral surface of posterolateral angles set with a variable number of small denticles (= thorn-pads). Epiproct very small and mostly or wholly concealed under anal segment. Vomer well-developed and sclerotised, variable in shape triangular to digitiform in shape, occasionally strongly arched and with a single, ± slender and acute terminal hook ( Figs. 97A– O ). Cerci slender, ± round in cross-section, slightly curved and projecting over posterior margin of anal segment; length variable. Poculum bulgy, strongly convex, cup-shaped, ± fornicate and with a ± prominent, obtuse basal protuberance (only one species with a large tooth-like appendage); roughly reaching to posterior margin of tergum IX. Legs long and moderately slender, profemora ± as long as (♀♀) or longer than mesothorax (♂♂), mesofemora longer than metathorax, and metatibiae just not reaching (♀♀) or notably projecting over apex of abdomen (♂♂); tibiae longer than corresponding femora.Profemora triangular in cross-section with the anterodorsal carina raised and lamellate (♀♀ in particular); medioventral carina distinct, lamellate and distinctly displaced towards anteroventral carina ( Fig. 8B ). Meso- and metafemora and all tibiae trapezoidal in cross-section with the medioventral carina centrally on ventral surface. Meso- and metafemora slender and mostly unarmed in ♂♂ (rarely with one or two sub-basal spines on two outer ventral carinae of mesofemora or with some carinae gently undulate); in ♀♀ meso- and metafemora often with a sub-basal teeth, expansions or lobe(s) on the two outer ventral carinae, and occasionally lobes are also present in the basal half of the posterodorsal carina. Posterodorsal carina of tibiae in ♀♀ usually with a ± distinct, rounded apical lobe and occasionally with further lobes in the remaining portion; a further ± distinct apical lobe is present on the posteroventral carina. Basitarsi slender or with the dorsal carina very gently raised and rounded in ♂♂, in ♀♀ with a ± prominent, rounded or triangular crest or lobe ( Fig. 8D ). Basitarsi slightly shorter (♀♀) or longer (♂♂) than remaining tarsomeres combined.

Eggs ( Figs. 99A–P , 100A–B ). Size variable, shape angularly ovoid with the polar-area ± flattened and the dorsal egg surface strongly convex in the anterior portion; oval in cross-section. Entire surface of capsule and micropylar plate ± distinctly punctured. Micropylar plate elongate-oval, at least half as long as capsule; sculptured like capsule but with the pits smaller and the entire surface gently convex longitudinally. Median line indistinct, short and not reaching to posterior pole of capsule. Operculum oval to almost circular and with a hollow capitulum either represented by a rounded net-like structure with circular pits or formed by lamellate extensions of the outer margin; these longitudinally connected by thin membranes. Height of capitulum very variable but at best tw-thirds the length of capsule. Capsule ± plain grey, drab, ochre or greyish brown, posterior portion of micropylar plate often dark brown to black. Capitulum dull yellow, orange or reddish brown.

Differentiation. This very speciose and widely distributed genus is morphologically most similar to Lanceobostra gen. n. and Phanocloidea Zompro, 2001 . From Lanceobostra gen. n. both sexes can be distinguished by the relatively longer median segment that is at least equal in length to the metanotum or longer (shorter than metanotum in Lanceobostra ). Females can also be separated from those of Lanceobostra gen. n. by having the mesosternum just obtusely tectate longitudinally (with a distinct and acute medio-longitudinal keel in Lanceobostra ), more globose to almost spherical head and longer, filiform gonapophyses VIII, that always project beyond the posterior margin of the anal segment. Males may be distinguished by having cerci that are ± round in cross-section (spatulate and with an oval impression interiorly in Lanceobostra ), generally more globose head and on average bulgier poculum that rarely bears a distinct spiniform central projection at the angle. Moreover, ♂♂ of Lanceobostra gen. n. are exclusively apterous, while in Phanocles several species have wings. From the very similar Phanocloidea ♀♀ may be distinguished by the longer gonapophyses VIII, which project over the posterior margin of the anal segment, longitudinally tectate mesosternum and almost always by the presence of a lateral lobe of abdominal tergum VII (missing only in a few species → see comments below). Males lack the red bases of the profemora that are frequently seen in Phanocloidea and have a much bulgier poculum, which is ± fornicate, often bears a basal spine or protuberance and always has the posterior margin unspecialised, rounded to sub-truncated and entire (roundly bowl-shaped and posterior margin often with two points or a lateral impression in Phanocloidea ). Furthermore, the anal segment is longer at an average and notably longer than wide (often subquadrate and scarcely longer than wide in Phanocloidea ) and ♂♂ of Phanocles are not as strikingly colourful as many of Phanocloidea are. The characteristic eggs of Phanocles readily differ from both genera by the distinctly punctured capsule surface.

Comments. Stål (1875b: 28) originally established Phanocles for the two Mexican species Bacteria burkartii Saussure, 1868 and Bacteria aetolus Westwood, 1859 , of which Kirby (1904: 353) designated the first as the type-species. Redtenbacher (1908: 412) was the first who misinterpreted Phanocles and erroneously synonymised it with Bacteria , as did all subsequent authors. Zompro (2001a: 197) re-established Phanocles and was the first who recognised some of the characteristic features, which distinguish Phanocles from its close relatives. However, the new diagnosis provided by Zompro (2001a: 196) is fairly insufficient and inaccurate in several aspects. This obviously resulted from a lack of material, as Zompro (2001a) listed only two species as to belong to Phanocles . Hence it is not surprising, that Zompro (2001a) failed in recognizing the complete range of variability of the genus and erroneously stated Phanocles to be restricted to Central America, which is definitely not the case. Instead, this speciose genus has a very wide distribution throughout almost the entire Neotropical Region and extends from Mexico to as far south as Bolivia.

Morphologically this genus is very diverse and definite individual characters that distinguish all known species from all representatives of closely related genera are at the present state very difficult to delimit to full satisfaction. Some species violate one of the main characteristics here revealed for the genus but agree in almost all other morphological aspects, thus in aspect of all characteristic taken together clearly key out as Phanocles . For example, the ♀♀ of some species have the posterolateral lobe of abdominal tergum VII very weakly developed and almost unknowable, e. g. P. aequatorialis ( Redtenbacher, 1908) , P. berezini sp. n., P. horni ( Redtenbacher, 1908) , P. maximus sp. n. or P. significans ( Redtenbacher, 1908) . Characters such as the elongated gonapopohyses, that project considerably beyond the tip of the abdomen, very prominent praeopercular organ and longitudinally tectate mesosternum of ♀♀, bulgy and fornicate poculum and long anal segment of ♂♂ and punctured egg capsule however place these species in Phanocles . Examples for species whose ♀♀ exhibit unique traits within the genus are P. decorus Zompro, 2001 which is typical for having the mesonotum armed with several prominent conical spines, P. pleuracanthus sp. n. which has the metapleurae armed with a row of strong, conical spines ( Fig. 59D ) and P. saussurei (Redtenbacher, 19089 whose ♂♂ bear a large conical central projection on the poculum, that resembles the genus Phanoclocrania gen. n. ( Fig. 67D ). All three characters are not present in any other known species of Phanocles . A species that violates several of the characteristic traits of the genus at the same time and thus takes on a fairly isolated position within the genus is the Mexican P. horni ( Redtenbacher, 1908) . Females of this species is characteristic for the prominently spinose meso- and metathorax and median segment, have the posterolateral lobe of abdominal tergum VII very obscure, but violate the characteristics of Phanocles in having an almost smooth and just very faintly tectate mesosternum ( Fig. 87N ) as well as short gonapophyses VIII, that do not project over the posterior margin of the anal segment ( Fig. 64M ). In addition to an almost entirely white head ( Fig. 60E ) ♂♂ have an anal segment that is enlarged and strikingly wider than all preceding segments ( Fig. 67B ). Another contrasting trait is represented by the egg-morphology ( Fig. 100B ), the eggs differing from all other known eggs of the genus by the almost ovoid and just minutely coriaceous but not punctured capsule, that has the polar area rounded and is blackish in colour with the dorsal surface pale grey, and huge opercular excrescence, that is much higher than in all other known eggs of Phanocles . Although there is a great range of morphological variability within the genus and some species violate individual characters of the genus, it appears impossible to define species-groups or present a proper intrageneric systematization at the present state.

The strong sexual dimorphism and intersexual incongruence of most morphological traits considerably hinder the matching of ♂♂ and ♀♀ of individual species without having at hand a series or at least both sexes from the same locality. For instance species with horned ♀♀ may have ♂♂ with or without cephalic armature, species whose ♀♀ possess well-developed leg ornaments may have ♂♂ with armed or wholly unarmed limbs, or species with a more or less equal length relation between the metanotum and median segment in the ♀♀ may have apterous or alate ♂♂. Even if cephalic armature is present in both sexes of an individual species, the spines or horns can be very different in size and shape, which is also true for teeth, lobes or appendages of the legs and basitarsi within certain species. The authors have examined several further and potentially still undescribed species, but all are only known from single and sometimes incomplete specimens, and some are likely to represent the opposite sexes of described species. Thus, and taking into account the great difficulty in the matching of sexes, these are not described herein and omitted in the present study.

Distribution ( Fig. 104A ). The very wide distribution of this speciose genus comprises almost the entire Neotropical Region except for the Greater Antilles. It extends from southern Mexico and the Lesser Antilles in the north to Peru and Bolivia in the south, which according to Morrone (2006: 471, fig. 2) includes the Caribbean, Amazonian and Chacoan Subregions.

Species included:

Figures

FIGURE 4. Terminalia of ♂♂: A. Cladomorphus phyllinus Gray, 1835 [coll. FH]; B. Phanocles mutica (Brunner v. Wattenwyl, 1907) [coll. FH]; C. Alienobostra remiformis (Rehn, 1904) [coll. FH]; D. Xylodus adumbratus Saussure, 1859 [ZMUH]; E. Bacteria ferula (Berthold, 1827) [Diapheromerini: “Bacteroformia”; the white arrow indicates how stigmae VIII are shifted away from the anterior margin of abdominal segment VIII.

FIGURE 6. Terminalia during copulation: A. Paraotocrania panamae gen. n., sp. n. paratypes (Panama, Prov. Coclé, Alle de Antón, XI.2019); B. Phanocles horni (Redtenbacher, 1908) cultured specimens; C. Alienobostra remiformis (Rehn, 1904) ventral view showing how the elongated cerci of the ♂ are used as claspers to grasp the abdomen of the ♀ (Costa Rica, Prov. Alajuela, El Rodeo Experimental Forest, XI.2012); D. same specimens as in figure D in lateral view.

FIGURE 7. Morphology of extremities: A. Basal portion of profemur of ♀ Aplopocranidium waehneri (Günther, 1940) in antero-lateral view [MNHN]; B. Basal portion of profemur of ♀ Phanocles ploiaria (Westwood, 1859) in antero-lateral view [coll. FH]; C. Basal portion of profemur of ♀ HT of Hirtuleiodes peruanus gen. n., sp. n. in ventral view [MNHN]; D. Protarsus and apical portion of protibia of ♀ Phanocles ploiaria (Westwood, 1859) in antero-lateral view [coll. FH].Arrow points towards apical end of femur.

FIGURE 8. Alienobostra remiformis (Rehn, 1904): A. ♀ (Costa Rica, Surubres nr. San Mateo), habitus dorsal view [ANSP]; B. ♀ (captive reared from Costa Rica, Prov. Puntarenas, Santa Elena), habitus dorsolatera view [coll. FH, No. 0049–34]; C. ♂ (captive reared from Costa Rica, Prov. Puntarenas, Santa Elena), habitus dorsolateral view [coll. FH, No. 0049–59]; D. ♀ (Costa Rica, Surubres nr. San Mateo), lateral view of terminalia [ANSP]; E. ♀ (Costa Rica, Surubres nr. San Mateo), dorsal view of terminalia [ANSP]; F. ♀ (Costa Rica, Surubres nr. San Mateo), ventral view of terminalia [ANSP]; G. ♀ (Costa Rica, San José), lateral view of terminalia [ZMUH]; H. ♀ (Costa Rica, San José), dorsal view of terminalia [ZMUH]; J. ♂ (captive reared from Costa Rica, Prov. Puntarenas, Santa Elena), lateral view of terminalia [coll. FH]; K. ♂ (captive reared from Costa Rica, Prov. Puntarenas, Santa Elena), dorsal view of terminalia [coll. FH]; L. ♂ (captive reared from Costa Rica, Prov. Puntarenas, Santa Elena), ventral view of terminalia [coll. FH]; M. ♀ with small cephalic tubercles (Costa Rica, Surubres nr. San Mateo), head and prothorax in lateral view [ANSP]; N. ♀ with well-developed cephalic lobes (Costa Rica, Surubres nr. San Mateo), head and prothorax in lateral view [ANSP]; O. ♂ (captive reared from Costa Rica, Prov. Puntarenas, Santa Elena), head and prothorax in lateral view [coll. FH].

FIGURE 46. Parotocrania gen. n.: A. P. panamae gen. n., sp. n. ♀ PT, habitus dorsolateral view [coll. FH, No. 1190–1]; B. P. panamae gen. n., sp. n. ♂ PT green colour form, habitus dorsolateral view [coll. FH, No. 1190–6]; C. P. panamae gen. n., sp. n. ♂ PT brown colour form, habitus dorsolateral view [coll. FH, No. 1190–2]; D. P. acutilobata gen. n., sp. n. ♀ HT, habitus dorsolateral view [NHMUK]; E. P. acutilobata gen. n., sp. n. ♂ PT, habitus dorsolateral view [MNCN]; F. P. panamae gen. n., sp. n. ♀ PT, terminalia in lateral view [coll. FH]; G. P. panamae gen. n., sp. n. ♀ PT, terminalia in dorsal view [coll. FH]; H. P. panamae gen. n., sp. n. ♀ PT, terminalia in ventral view [coll. FH]; J. P. acutilobata gen. n., sp. n. ♀ HT, terminalia in lateral view [NHMUK]; K. P. acutilobata gen. n., sp. n. ♀ HT, terminalia in ventral view [NHMUK]; L. P. dilatipes (Conle, Hennemann & Gutiérrez, 2011) ♀, terminalia in lateral view [MNHN]; M. P. dilatipes (Conle, Hennemann & Gutiérrez, 2011) ♀, terminalia in dorsal view [MNHN]; N. P. dilatipes (Conle, Hennemann & Gutiérrez, 2011) ♀, terminalia in ventral view [MNHN]; O. P. acutilobata gen. n., sp. n. ♀ HT, intero-lateral view of basal portion of left mesofemur [NHMUK]; P. P. dilatipes (Conle, Hennemann & Gutiérrez, 2011) ♀, intero-lateral view of right miD–leg [MNHN].

FIGURE 47. Parotocrania gen. n.: A. P. dilatipes (Conle, Hennemann & Gutiérrez, 2011) ♀ (Ecuador, Prov. Napo, Tena, MIsahualli), habitus dorsolateral view [MNHN]; B. P. curvata gen. n., sp. n. ♂ PT, habitus dorsal view [coll. YB, © Yannick Bellanger]; C. P. panamae gen. n., sp. n. ♂ PT, terminalia in lateral view [coll. FH]; D. P. panamae gen. n., sp. n. ♂ PT, terminalia in dorsal view [coll. FH]; E. P. panamae gen. n., sp. n. ♂ PT, terminalia in ventral view [coll. FH]; F. P. acutilobata gen. n., sp. n. ♂ PT, terminalia in lateral view [MNCN]; G. P. acutilobata gen. n., sp. n. ♂ PT, terminalia in ventral view [MNCN]; H. P. curvata gen. n., sp. n. ♂ HT, terminalia in lateral view [ZSMC]; J. P. segmentaria (Redtenbacher, 1908) ♂ HT, terminalia in lateral view [NHMW]; K. P. segmentaria (Redtenbacher, 1908) ♂ HT, terminalia in dorsal view [NHMW]; L. P. segmentaria (Redtenbacher, 1908) ♂ HT, terminalia in ventral view [NHMW].

FIGURE 48. Parotocrania gen. n.: A. P. panamae gen. n., sp. n. ♀ PT, head and prothorax in lateral view [coll. FH]; B. P. acutilobata gen. n., sp. n. ♀ HT, head and prothorax in dorsolateral view [NHMUK]; C. P. panamae gen. n., sp. n. ♂ PT, head and prothorax in lateral view [coll. FH]; D. P. curvata gen. n., sp. n. ♂ HT, head and prothorax in dorsolateral view [ZSMC]; E. P. dilatipes (Conle, Hennemann & Gutiérrez, 2011) ♀, head and prothorax in lateral view [MNHN]; F. P. curvata gen. n., sp. n. ♂ HT, habitus dorsolateral view [ZSMC].

FIGURE 49. Phanocles Stål, 1875 habitus (in scale): A. P. burkartii (Saussure, 1868) ♀ (SE-Mexico), dorsal view [NHMW]; B. P. burkartii (Saussure, 1868) ♀ (Mexico, Veracruz, Dos Amates, Catemaco), dorsolateral view [coll. FH, No. 1218–1]; C. P. costaricensis Hennemann, 2002 ♀ (captive reared F1 from Costa Rica, Proc. Alajuela, Bosque del Niño N.P.), dorsolateral view [coll. FH, No. 0472–4]; D. P. spectabilis sp. n. ♀ PT (captive reared from Costa Rica, Prov. Puntarenas, La Gamba), dorsolateral view [coll. FH, No. 0563–5]; E. P. ploiaria (Westwood, 1859) ♀ (Panamá, Prov. Coclé, Valle deAntón 650–850 m), dorsolateral view [coll. FH, No. 0705–11].

FIGURE 50. Phanocles Stål, 1875 habitus (in scale): A. P. maximus sp. n. ♀ HT (Panamá, Prov Panamá, Chagres N.P., Cerro Azul), dorsolateral view [MIUP]; B. P. subvolans (Redtenbacher, 1908) ♀ (Costa Rica) dorsolateral view [ANSP]; C. P. saussurei (Redtenbacher, 1908) ♀ (Mexico, Veracruz, Puerto Escondido), dorsolateral view [coll. FH No. 0650–1]; D. P. rehni sp. n. ♀ HT (Honduras, Dept.Atlantida, Lancetilla), dorsolateral view [ANSP]; E. P. cuzcoense sp. n. ♀ HT (Peru, Dept. Cuzco, Quince Mil), dorsolateral view [ANSP].

FIGURE 51. Phanocles Stål, 1875 habitus (in scale): A. P. ecuadoricus sp. n. ♀ HT (Ecuador, Prov. Imbabura, Paramba), dorsolateral view [ZMUH]; B. P. falcatus sp. n. ♀ PT (Ecuador, Prov. Napo, Misahualli), dorsolateral view [coll. FH, No. 0919–1]; C. P. virgulatus (Redtenbacher, 1908) ♀ (Peru, Dept. Huanuco, Rio Lullapichis, Panguana) dorsolateral view [coll. FH, No. 0538–3]; D. P. brevipes sp. n. ♀ HT (Peru, Prov. Satip, Junín, Satipo), dorsal view [ANSP]; E. P. satyr (Redtenbacher, 1908) ♀ (S-Venezuela, San Esteban), dorsolateral view [ANSP].

FIGURE 52. Phanocles Stål, 1875 habitus (A–C in scale): A. P. horni (Redtenbacher, 1908) ♀ (captive reared from SW-Mexico, Oaxaca/Chiapas, Est. D. Tehuantepec), dorsolateral view [coll. FH, No. 0700–28]; B. P. mexicanus sp. n. ♀ HT (Mexico, Chiapas, Huixtla/Motozintla 700 m), dorsal view [MNHN]; C. P. mexicanus sp. n. ♀ PT (Guatemala, Dept. Chimaltenango), dorsal view [MNHN]; D. P. superbus sp. n. ♀ HT (Ecuador, Prov. Pichincha, Tandapi 1700 m), dorsal view [MNHN].

FIGURE 53. Phanocles Stål, 1875 habitus: A. P. pleurospinosus sp. n. ♀ HT (Costa Rica, Prov. Limón, Pococí, Bosque Lluvioso), lateral view [MNCR-A]; B. P. pleurospinosus sp. n. ♀ HT, dorsal view [MNCR-A]; C. P. muticus (Brunner v. Wattenwyl, 1907) ♀ (captive reared from Trinidad), dorsolateral view [coll. FH, No. 0837–13]; D. P. barbadosense sp. n. ♀ HT (Barbados), dorsal view [NHMW]; E. P. barbadosense sp. n. ♀ HT (Barbados), dorsolateral view [NHMW, © 2019 Natural History Museum Vienna, NOaS Image Collection / H. Bruckner]; F. P. acutecornutus sp. n. ♀ HT (Ecuador, Prov. Napo, Yasuni), dorsal view [MNHN]; G. P. acutecornutus sp. n. ♀ HT (Ecuador, Prov. Napo, Yasuni), dorsolateral view [MNHN].

FIGURE 54. Phanocles Stål, 1875 habitus: A. P. maroniensis (Chopard, 1911) ♀ HT (French Guiana, St. Jean de Maroni, Nouveau Chantier), dorsolateral view [MNHN]; B. P. aequatorialis (Redtenbacher, 1908) ♀ (captive reared from Ecuador, Prov. Esmeraldas, Durango), dorsolateral view [coll. FH, No. 0659–3]; C. P. decorus Zompro, 2001 ♀ HT (Ecuador), dorsolateral view [MNHU]; D. P. berezini sp. n. ♀ PT (captive reared from Mexico, Oaxaca, Xadani village nr. Huantulco), dorsolateral view [coll. FH, No. 0718–8]; E. P. solidus sp. n. ♀ HT (Ecuador, Prov. El Oro, Arenillas), dorsolateral view [ANSP]; F. P. aequatorialis (Redtenbacher, 1908) ♂ (captive reared from Ecuador, Prov. Esmeraldas, Durango), dorsolateral view [coll. FH, No. 0659–1]; G. P. acutecornutus sp. n. ♂ PT (Bolivia, Prov. Chapare, Villa Tunari), dorsal view [ZSMC].

FIGURE 55. Phanocles Stål, 1875: A. P. ecuadoricus sp. n. ♂ PT (Ecuador, Prov. Esmeraldas, Rio Verde), dorsolateral view [MNCN]; B. P. falcatus sp. n. ♂ PT (Ecuador, Prov. Napo, Misahualli), dorsolateral view [coll. FH, No. 0919–3]; C. P. maximus sp. n. ♀ PT (captive reared from Panamá, Prov. Coclé, Valle deAntón), dorsolateral view [coll. FH, No. 1360–3]; D. P. spectabilis sp. n. ♂ PT (captive reared from Costa Rica, Prov. Puntarenas, La Gamba), dorsolateral view [coll. FH, No. 0563–18]; E. P. ploiaria (Westwood, 1859) ♀ (Panamá, Prov. Coclé, Valle de Antón 650–850 m), dorsolateral view [coll. FH, No. 0705–12]; F. P. obtusus (Redtenbacher, 1908) ♂ (Venezuela, Aragua N.P., Rancho Grande), dorsolateral view [ANSP].

FIGURE 56. Phanocles Stål, 1875: A. P. costaricensis Hennemann, 2002 ♂ (captive reared F1 from Costa Rica, Proc.Alajuela, Bosque del Niño N.P.), dorsolateral view [coll. FH, No. 0472–15]; B. P. virgulatus (Redtenbacher, 1908) ♂ (Peru, Dept. Huanuco, Rio Lullapichis, Panguana) dorsolateral view [coll. FH, No. 0538–4]; C. P. vosseleri (Redtenbacher, 1908) ♂ (captive reared from Belize, Cayo, Greenhills) dorsolateral view [coll. FH, No. 0704–2]; D. P. horni (Redtenbacher, 1908) ♂ (captive reared from SW-Mexico, Oaxaca/Chiapas, Est. D. Tehuantepec), dorsolateral view [coll. FH, No. 0700–42]; E. P. chiapasense sp. n. ♂ HT (Mexico, Chiapas), dorsolateral view [ZMUH]; F. P. saussurei (Redtenbacher, 1908) ♂ (Mexico, Veracruz, Puerto Escondido), dorsolateral view [coll. FH No. 0650–2]; G. P. muticus (Brunner v. Wattenwyl, 1907) ♂ (captive reared from Trinidad), dorsolateral view [coll. FH, No. 0837–35]; H. P. barbadosense sp. n. ♂ PT (Barbados), dorsal view [NHMW, © 2019 Natural History Museum Vienna, NOaS Image Collection / H. Bruckner].

FIGURE 57. Phanocles Stål, 1875 head and prothorax: A. P. acutecornutus sp. n. ♀ HT, dorsolateral view [MNHN]; B. P. berezini sp. n. ♀ PT, lateral view [coll. FH]; C. P. brevipes sp. n. ♀ HT, dorsolateral view (head is turned upside down) [ANSP]; D. P. barbadosense sp. n. ♀ PT, lateral view [ANSP]; E. P. muticus (Brunner v. Wattenwyl, 1907) ♀, dorsolateral view [coll. FH]; F. P. burkartii (Saussure, 1868) ♀, dorsolateral view [coll. FH]; G. P. saussurei (Redtenbacher, 1908) ♀, lateral view [coll. FH]; H. P. falcatus sp. n. ♀ PT, dorsolateral view [MNHN]; J. P. falcatus sp. n. ♀ PT, dorsolateral view [MNHN]; K. P. superbus sp. n. ♀ HT, dorsolateral view [MNHN].

FIGURE 58. Phanocles Stål, 1875 head and prothorax: A. P. aequatorialis (Redtenbacher, 1908) ♀, lateral view [coll. FH]; B. P. costaricensis Hennemann, 2002 ♀, lateral view [coll. FH]; C. P. ploiaria (Westwood, 1859) ♀, lateral view [coll. FH]; D. P. maroniensis (Chopard, 1911) ♀ HT, lateral view [MNHN]; E. P. satyr (Redtenbacher, 1908) ♀ dorsolateral view [ANSP]; F. P. ecuadoricus sp. n. ♀ PT, lateral view [coll. FH]; G. P. maximus sp. n. ♀ PT, dorsolateral view [coll. FH]; H. P. spectabilis sp. n. ♀ PT, dorsolateral view [coll. FH]; J. P. spectabilis sp. n. ♀ PT, dorsolateral view [coll. FH]; K. P. subvolans (Redtenbacher, 1908) ♀, dorsolateral view [ANSP]; L. P. horni (Redtenbacher, 1908) ♀, dorsolateral view [coll. FH].

FIGURE 59. Phanocles Stål, 1875: A. P. mexicanus sp. n. ♀ HT (with small cephalic tubercles), lateral view of head and prothorax [MNHN]; B. P. mexicanus sp. n. ♀ PT (with prominent cephalic horns), lateral view of head and prothorax [MNHN]; C. P. mexicanus sp. n. ♀ HT, dorsal view of head, pro and mesonotum [MNHN]; D. P. pleurospinosus sp. n. ♀ HT, dorsolateral view of mesothorax with median segment showing the typical spines of the metapleurae [MNCR-A]; E. P. solidus sp. n. ♀ HT, lateral view of head and prothorax [ANSP]; F. P. maroniensis (Chopard, 1911) ♂, lateral view of head and prothorax [MNHN]; G. P. virgulatus (Redtenbacher, 1908) ♂, dorsolateral view of head and prothorax [coll. FH]; H. P. ecuadoricus sp. n. ♂ PT, dorsolateral view [MNCN]; J. P. falcatus sp. n. ♂ PT, dorsolateral view of head and prothorax [coll. FH].

FIGURE 60. Phanocles Stål, 1875 head and prothorax: A. P. aequatorialis (Redtenbacher, 1908) ♂, lateral view [coll. FH]; B. P. costaricensis Hennemann, 2002 ♂, lateral view [coll. FH]; C. P. muticus (Brunner v. Wattenwyl, 1907) ♂, lateral view [coll. FH]; D. P. mexicanus sp. n. ♂ PT, lateral view [UMMZ, © Erika Tucker]; E. P. horni (Redtenbacher, 1908) ♂, lateral view [coll. FH]; F. P. maximus sp. n. ♂ PT, dorsolateral view [coll. FH]; G. P. spectabilis sp. n. ♂ PT, lateral view [coll. FH]; H. P. saussurei (Redtenbacher, 1908) ♂, lateral view [coll. FH]; J. P. vosseleri (Redtenbacher, 1908) ♂, lateral view [coll. FH].

FIGURE 61. Phanocles Stål, 1875 terminalia of ♀♀: A. P. burkartii (Saussure, 1868), lateral view [coll. FH]; B. P. burkartii (Saussure, 1868), dorsal view [coll.FH]; C. P.burkartii (Saussure, 1868), ventral view [coll. FH]; D. P.costaricensis Hennemann, 2002, lateral view [coll. FH]; E. P. costaricensis Hennemann, 2002, dorsal view [coll. FH]; F. P. costaricensis Hennemann, 2002, ventral view [coll. FH]; G. P. ecuadoricus sp. n. ♀ HT, lateral view [ZMUH]; H. P. ecuadoricus sp. n. ♀ HT, dorsal view [ZMUH]; J. P. ecuadoricus sp. n. ♀ HT, ventral view [ZMUH]; K. P. aequatorialis (Redtenbacher, 1908), lateral view [coll. FH]; L. P. aequatorialis (Redtenbacher, 1908), dorsal view [coll. FH]; M. P. aequatorialis (Redtenbacher, 1908), ventral view [coll. FH]; N. P. maroniensis (Chopard, 1911) ♀ HT, lateral view [coll. FH]; O. P. maroniensis (Chopard, 1911) ♀ HT, dorsal view [coll. FH]; P. P. maroniensis (Chopard, 1911) ♀ HT, ventral view [coll. FH]; Q. P. saussurei (Redtenbacher, 1908), lateral view [coll. FH]; R. P. saussurei (Redtenbacher, 1908), dorsal view [coll. FH]; S. P. saussurei (Redtenbacher, 1908), ventral view [coll. FH].

FIGURE 62. Phanocles Stål, 1875 terminalia of ♀♀: A. P. maximus sp. n. ♀ PT, lateral view [coll. FH]; B. P. maximus sp. n. ♀ PT, dorsal view [coll. FH]; C. P. maximus sp. n. ♀ PT, ventral view [coll. FH]; D. P. rehni sp. n. ♀ HT, lateral view [ANSP]; E. P. rehni sp. n. ♀ HT, dorsal view [ANSP]; F. P. rehni sp. n. ♀ HT, ventral view [ANSP]; G. P. subvolans (Redtenbacher, 1908), lateral view [ANSP]; H. P. subvolans (Redtenbacher, 1908), dorsal view [ANSP]; J. P. subvolans (Redtenbacher, 1908), ventral view [ANSP]; K. P. ploiaria (Westwood, 1859), lateral view [coll. FH]; L. P. ploiaria (Westwood, 1859), dorsal view [coll. FH]; M. P. ploiaria (Westwood, 1859), ventral view [coll. FH]; N. P. spectabilis sp. n. ♀ PT, lateral view [coll. FH]; O. P. spectabilis sp. n. ♀ PT, dorsal view [coll. FH]; P. P. spectabilis sp. n. ♀ PT, ventral view [coll. FH]; Q. P. virgulatus (Redtenbacher, 1908), lateral view [coll. FH]; R. P. virgulatus (Redtenbacher, 1908), dorsal view [coll. FH]; S. P. virgulatus (Redtenbacher, 1908), ventral view [coll. FH].

FIGURE 63. Phanocles Stål, 1875 terminalia of ♀♀: A. P. cuzcoense sp. n. ♀ HT, lateral view [ANSP]; B. P. cuzcoense sp. n. ♀ HT, dorsal view [ANSP]; C. P. cuzcoense sp. n. ♀ HT, ventral view [ANSP]; D. P. brevipes sp. n. ♀ HT, lateral view [ANSP]; E. P. brevipes sp. n. ♀ HT, dorsal view [ANSP]; F. P. brevipes sp. n. ♀ HT, ventral view [ANSP]; G. P. pleurospinosus sp. n. ♀ HT, lateral view [MNCR-A]; H. P. pleurospinosus sp. n. ♀ HT, dorsal view [MNCR-A]; J. P. pleurospinosus sp. n. ♀ HT, ventral view [MNCR-A]; K. P. falcatus sp. n. ♀ PT, lateral view [coll. FH]; L. P. falcatus sp. n. ♀ PT, dorsal view [coll. FH]; M. P. falcatus sp. n. ♀ PT, ventral view [coll. FH]; N. P. solidus sp. n. ♀ HT, lateral view [ANSP]; O. P. solidus sp. n. ♀ HT, dorsal view [ANSP]; P. P. solidus sp. n. ♀ HT, ventral view [ANSP]; Q. P. satyr (Redtenbacher, 1908), lateral view [ANSP].

FIGURE 64. Phanocles Stål, 1875 terminalia of ♀♀: A. P. superbus sp. n. ♀ HT, lateral view [MNHN]; B. P. superbus sp. n. ♀ HT, dorsal view [MNHN]; C. P. superbus sp. n. ♀ HT, ventral view [MNHN]; D. P. acutecornutus sp. n. ♀ PT, lateral view [ZSMC]; E. P. berezini sp. n. ♀ PT, lateral view [coll. FH]; F. P. berezini sp. n. ♀ PT, dorsal view [coll. FH]; G. P. berezini sp. n. ♀ PT, ventral view [coll. FH]; H. P. muticus (Brunner v. Wattenwyl, 1907), lateral view [coll. FH]; J. P. muticus (Brunner v. Wattenwyl, 1907), dorsal view [coll. FH]; K. P. muticus (Brunner v. Wattenwyl, 1907), ventral view [coll. FH]; L. P. horni (Redtenbacher, 1908), lateral view [coll. FH]; M. P. horni (Redtenbacher, 1908), dorsal view [coll. FH]; N. P. horni(Redtenbacher, 1908), ventral view [coll. FH]; O. P. mexicanus sp. n. ♀ PT, lateral view [MNHN]; P. P. mexicanus sp. n. ♀ PT, dorsal view [MNHN]; Q. P. mexicanus sp. n. ♀ HT, ventral view [MNHN]; R. P. barbadosense sp. n. ♀ PT, lateral view [ANSP]; S. P. barbadosense sp. n. ♀ PT, ventral view [ANSP].

FIGURE 65. Phanocles Stål, 1875 terminalia of ♂♂: A. P. costaricensis Hennemann, 2002, lateral view [coll. FH]; B. P. costaricensis Hennemann, 2002, dorsal view [coll. FH]; C. P. costaricensis Hennemann, 2002, ventral view [coll. FH]; D. P. falcatus sp. n. ♂ PT, lateral view [coll. FH]; E. P. falcatus sp. n. ♂ PT, dorsal view [coll. FH]; F. P. falcatus sp. n. ♂ PT, ventral view [coll. FH]; G. P. virgulatus (Redtenbacher, 1908), lateral view [coll. FH]; H. P. virgulatus (Redtenbacher, 1908), dorsal view [coll. FH]; J. P. virgulatus (Redtenbacher, 1908), ventral view [coll. FH]; K. P. maximus sp. n. ♂ PT, lateral view [ANSP]; L. P. maximus sp. n. ♂ PT, dorsal view [coll. FH]; M. P. maximus sp. n. ♂ PT, ventral view [coll. FH]; N. P. spectabilis sp. n. ♂ PT, lateral view [coll. FH]; O. P. spectabilis sp. n. ♂ PT, dorsal view [ANSP]; P. P. spectabilis sp. n. ♂ PT, ventral view [ANSP]; Q. P. vosseleri (Redtenbacher, 1908), lateral view [coll. FH]; R. P. vosseleri (Redtenbacher, 1908), dorsal view [coll. FH]; S. P. vosseleri (Redtenbacher, 1908), ventral view [coll. FH].

FIGURE 66.Phanocles Stål, 1875 terminaliaof ♂♂: A. P.maroniensis (Chopard, 1911), lateral view [MNHN]; B. P.maroniensis (Chopard, 1911), dorsal view [MNHN]; C. P. maroniensis (Chopard, 1911), ventral view [MNHN]; D. P. barbadosense sp. n. ♂ PT, lateral view [NHMW, © 2019 Natural History Museum Vienna, NOaS Image Collection / H. Bruckner]; E. P. barbadosense sp. n. ♂ PT, dorsal view [NHMW, © 2019 Natural History Museum Vienna, NOaS Image Collection / H. Bruckner]; F. P. barbadosense sp. n. ♂ PT, ventral view [NHMW, © 2019 Natural History Museum Vienna, NOaS Image Collection / H. Bruckner]; G. P. keratosqueleton (Olivier, 1792), ventral view (specimen from St. Vincent) [FSCA]; H. P. aequatorialis (Redtenbacher, 1908), lateral view [coll. FH]; J. P. acutecornutus sp. n. ♂ PT, lateral view [ZSMC]; K. P. acutecornutus sp. n. ♂ PT, dorsal view [ZSMC]; L. P. acutecornutus sp. n. ♂ PT, ventral view [ZSMC]; M. P. muticus (Brunner v. Wattenwyl, 1907), lateral view [coll. FH]; N. P. muticus (Brunner v. Wattenwyl, 1907), dorsal view [coll. FH]; O. P. muticus (Brunner v. Wattenwyl, 1907), ventral view [coll. FH]; P. P. ecuadoricus sp. n. ♂ PT, lateral view [MNHN]; Q. P. ecuadoricus sp. n. ♂ PT, dorsal view [MNHN].

FIGURE 67. Phanocles Stål, 1875: A. P. horni (Redtenbacher, 1908), lateral view of terminalia [coll. FH]; B.. horni (Redtenbacher, 1908), dorsal view of terminalia [coll. FH]; C. P. horni (Redtenbacher, 1908), ventral view of terminalia [coll. FH]; D. P. saussurei (Redtenbacher, 1908), lateral view of terminalia [coll. FH]; E. P. saussurei (Redtenbacher, 1908), dorsal view of terminalia [coll. FH]; F. P. saussurei (Redtenbacher, 1908), ventral view of terminalia [coll. FH]; G. P. mexicanus sp. n. ♂ PT, lateral view of terminalia [UMMZ, © Erika Tucker]; H. P. mexicanus sp. n. ♂ PT, dorsal view of terminalia [UMMZ, © Erika Tucker]; J. P. chiapasense sp. n. ♂ HT, lateral view of terminalia [ZMUH]; K. P. chiapasense sp. n. ♂ HT, dorsal view of terminalia [ZMUH]; L. P. maroniensis (Chopard, 1911) ♀ HT, right mesofemur in intero-lateral view [MNHN]; M. P. solidus sp. n. ♀ HT, right mesofemur in intero-lateral view [ANSP]; N. P. ploiaria (Westwood, 1859) ♀, right mesofemur in interolateral view [coll. FH]; O. P. ploiaria (Westwood, 1859) ♀, right mesofemur in intero-lateral view [coll. FH]; P. P. brevipes sp. n. ♀ HT, right metatibia in intero-lateral view [ANSP]; Q. P. brevipes sp. n. ♀ HT, left mid leg in intero-lateral view [ANSP].

FIGURE 86. Mesosternum: A. Globocrania culmus (Bates, 1865) ♀ [MNHN]; B. Globocrania emesa (Westwood, 1859) ♀ HT of Bostra imperialis Redtenbacher, 1908 [MNHU]; C. Otocrania aurita (Burmeister, 1838) ♂ [NHMW]; D. Parotocrania panamae gen. n., sp. n. ♀ PT [coll. FH]; E. Parotocrania panamae gen. n., sp. n. ♂ PT [coll. FH]; F. Parotocrania acutilobata gen. n., sp. n. ♀ HT [NHMUK]; G. Parotocrania dilatipes (Conle, Hennemann & Guiérrez, 2011) ♀ [MNHN]; H. Spinocloidea panamaense gen. n., sp. n. ♀ PT [coll. FH]; J. Spinocloidea panamaense gen. n., sp. n. ♂ PT [coll. FH]; K. Phanoclocrania dorsuaria (Stål, 1875) ♀ [ANSP]; L. Phanoclocrania dorsuaria (Stål, 1875) ♂ [MNCN]; M. Phanocloidea magistralis (Redtenbacher, 1908) ♀ HT [NHMUK]. Not in scale.

FIGURE 87. Mesosternum: A. Phanocloidea incolumis (Brunner v. Wattenwyl, 1907) ♀ [NHMUK]; B. Phanocloidea muricata (Burmeister, 1838) ♀ [coll. FH]; C. Phanocloidea muricata (Burmeister, 1838) ♂ [coll. FH]; D. Phanocloidea venezuelica sp. n. ♀ PT [coll. FH]; E. Phanocloidea venezuelica sp. n. ♂ PT [coll. FH]; F. Phanocloidea turgida (Westwood, 1859) ♀ [coll. FH]; G. Phanocloidea turgida (Westwood, 1859) ♂ [coll. FH]; H. Phanocloidea tobagoense (Langlois & Bellanger, 2012) ♀ HT [MNHN]; J. Phanocloidea submutica (Redtenbacher, 1908) ♀ [ANSP]; K. Phanocloidea schulthessi (Redtenbacher, 1908) ♀ [ANSP]; L. Phanocloidea schulthessi (Redtenbacher, 1908) ♂ [ANSP]; M. Phanocles solidus sp. n. ♀ HT [ANSP]; N. Phanocles horni (Redtenbacher, 1908) ♀ [coll. FH]. Not in scale.

FIGURE 92. Praeopercular organ of ♀♀: A. Phanocles berezini sp. n. PT [coll. FH]; B. Phanocles horni (Redtenbacher, 1908) [coll. FH]; C. Phanocles chaperi (Redtenbacher, 1908) HT [MNHN]; D. Phanocles keratosqueleton (Olivier, 1792) St. Vincent [coll. ASPER, © Philippe Lelong]; E. Phanocles keratosqueleton (Olivier, 1792) Dominica [coll. ASPER, © Philippe Lelong]; F. Phanocles mutica (Brunner v. Wattenwyl, 1907) Trinidad [coll. ASPER, © Philippe Lelong]; G. Phanocles ecuadoricus sp. n. PT [ZMUH]; H. Phanocles maximus sp. n. [coll. FH]; J. Phanocles spectabilis sp. n. PT [coll. FH]. Not in scale.

FIGURE 93. Praeopercular organ of ♀♀: A. Phanocles significans (Redtenbacher, 1908) [ANSP]; B. Phanocles decorus Zompro, 2001 [MNHU]; C. Phanocles falcatus sp. n. PT [ANSP]; D. Phanocles maroniensis (Chopard, 1911) HT [MNHN]; E. Phanocles brevipes sp. n. HT [ANSP]; F. Phanocles virgulatus (Redtenbacher, 1908) [coll. FH]; G. Phanocles vosseleri (Redtenbacher, 1908) [coll. FH]; H. Phanocles ploiaria (Westwood, 1859) ♀ [coll. FH]. Not in scale.

FIGURE 95. Vomer of ♂♂ (ventral view): A. Calynda coronata Carl, 1913 [ANSP]; B. Spinocloidea panamaense gen. n., sp. n. PT [coll. FH]; C. Lanceobostra aetolus (Westwood, 1859) [ANSP]; D. Lanceobostra ornata sp. n. HT [ANSP]; E. Lanceobostra jaliscensis Rehn, 1904 [ANSP]; F. Lanceobostra guatemalensis (Redtenbacher, 1908) [ANSP]; G. Bostriana boliviana (Piza, 1939) [coll. OC]; H. Alienobostra remiformis (Rehn, 1904) LT of Bostra jugalis (Rehn, 1905) [ANSP]; J. Phanoclocrania speciosa Conle, Hennemann & Gutiérrez, 2011 [CEBUC, © Yeisson Gutierréz]; K. Phanoclocrania dorsuaria (Stål, 1875) [ZMUH]; L. Parotocrania panamae gen. n., sp. n. PT [coll. OC]; M. Parotocrania regina (Conle, Hennemann & Gutiérrez, 2011 [CEBUC, © Yeisson Gutierréz]; N. Globocrania emesa (Westwood, 1859) HT of Pseudobacteria longipes Piza, 1938 [MZUSP]; O. Globocrania culmus (Bates, 1865) [CEIOC]. Not in scale.

FIGURE 97. Vomer of ♂♂: A. Phanocles falcatus sp. n. HT lateral view [NHMUK]; B. Phanocles falcatus sp. n. PT [MNHN]; C. Phanocles maroniensis (Chopard, 1911) [MNHN]; D. Phanocles costaricensis Hennemann, 2002 [coll. FH]; E. Phanocles maximus sp. n. PT Laguna San Carlos, Panamá [coll. FH]; F. Phanocles significans (Redtenbacher, 1908) [ANSP]; G. Phanocles virgulatus (Redtenbacher, 1908) [coll. FH]; H. Phanocles ploiaria (Westwood, 1859) [coll. FH]; J. Phanocles vosseleri (Redtenbacher, 1908) [coll. FH]; K. Phanocles spectabilis sp. n. PT [ANSP]; L. Phanocles horni (Redtenbacher, 1908) [coll. FH]; M. Phanocles acutecornutus sp. n. PT [ZSMC]; N. Phanocles mutica (Brunner v. Wattenwyl, 1907) [coll. ASPER, © Philippe Lelong]; O. Phanocles keratosqueleton (Olivier, 1792) St. Vincent [coll. ASPER, © Philippe Lelong]. Not in scale.

FIGURE 99. Eggs (in scale): A. Phanocles aequatorialis (Redtenbacher, 1908) [coll. FH]; B. Phanocles costaricensis Hennemann, 2002; C. Phanocles cuzcoense sp. n. ex ovipositor HT [ANSP]; D. Phanocles ecuadoricus sp. n. PT [coll. FH]; E. Phanocles maximus sp. n. [coll. FH]; F. Phanocles mutica (Brunner v. Wattenwyl, 1907) [coll. FH]; G. Phanocles ploiaria (Westwood, 1859) Panamá, Prov. Coclé, Valle de Antón [coll. FH]; H. Phanocles rehni sp. n. ex ovipositor HT [ANSP]; J. Phanocles spectabilis sp. n. PT [coll. FH]; K. Phanocles vosseleri (Redtenbacher, 1908) [coll. FH]; L. Phanocles virgulatus (Redtenbacher, 1908) [coll. FH]; M. Phanocles keratosqueleton (Olivier, 1792) Martinique [coll. FH]; N. Phanocles keratosqueleton (Olivier, 1792) St. Vincent [coll. FH]; O. Phanocles chaperi (Redtenbacher, 1908) [coll. FH]; P. Phanocles falcatus sp. n. PT [coll. FH]. [Scale = 1 mm]

FIGURE 100. Eggs: A. Phanocles berezini sp. n. [coll. FH]; B. Phanocles horni (Redtenbacher, 1908) [coll. FH]; C. Parotocrania panamae gen. n., sp. n. PT [coll. FH]; D. Trychopeplus laciniatus (Westwood, 1874) [coll. OC]; E. Cranidium gibbosum (Burmeister, 1838) [coll. FH]; F. Phanoclocrania gen. n. dorsuaria (Stål, 1875) [coll. FH]; G. Phanocloidea procera (Conle et al., 2021) [coll. FH]; H. Phanocloidea incolumis (Brunner v. Wattenwyl, 1907) [coll. FH]; J. Phanocloidea schulthessi (Redtenbacher, 1908) [coll. FH]; K. Phanocloidea pallidenotata (Redtenbacher, 1908) [coll. TR]; L. Phanocloidea muricata (Burmeister, 1838) [coll. FH]; M. Phanocloidea venezuelica sp. n. PT [coll. FH]; N. Phanocloidea tobagoense (Langlois & Bellanger, 2012) [coll. FH]; O. Phanocloidea turgida (Westwood, 1859) [coll. FH]; P. Phanocloidea incompta (Rehn, 1904) [coll. FH]. [Scale = 1 mm]

FIGURE 104. Maps showing the known distributions of the genera A. Phanocles Stål, 1875, B. Phanoclocrania gen. n., C. Phanocloidea Zompro, 2001, D. Spinocloidea gen. n., E. Trychopeplus Shelford, 1909 and F. Xylodus Saussure, 1859.

FIGURE 110. Live insects: A. Parotocrania panamae gen. n., sp. n. mating couple at Valle de Antón, Panamá (♂ brown form); B. Parotocrania panamae gen. n., sp. n. mating couple at Valle de Antón, Panamá (♂ brown form); C. Parotocrania panamae gen. n., sp. n. ♂ (green form) at Valle de Antón, Panamá; D. Parotocrania curvata gen. n., sp. n. immature ♀ at Tarapoto, Peru (© Johann Chretien); E. Jeremiodes pachycercus (Redtenbacher, 1908) ♂ at Tarapoto, Peru (© Jean- Francois Brousseau).