Phanocles vosseleri ( Redtenbacher, 1908 )

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 : 222-229

Phanocles vosseleri ( Redtenbacher, 1908)

( Figs. 55C , 60J , 65Q–S , 93G , 97J , 99K , 128C )

Bacteria vosseleri Redtenbacher, 1908: 419 .

Shelford, 1909: 364.

Hennemann, Gehler & Conle, 1995: 437.

Phanocles vosseleri, Hennemann & Conle, 2003: 6 .

Otte & Brock, 2005: 262.

Brock & Büscher, 2022: 514, fig.

Further material examined [4 ♂♂, 11 ♀♀, eggs]:

GUATEMALA: 1 ♀: Guatemala, 86 [ SMNS] ; 2 ♂♂, 5 ♀♀, eggs: ex Zucht: M. Duytschaever 2017, Herkunft: Guatemala, Petén, Tikal, F 3– Generation [ FH. No‘s 0704–3 to 9, E2] ; 4 ♀♀: ex Zucht: M. Duytschaever 2018, Herkunft: Guatemala, Petén, Tikal, F 4– Generation [ FH. No‘s 0704–10 to 13] .

BELIZE: 1 ♂: Belize: Toledo Dist., Blue Creek Village , 9 June 1981, W. E. Steiner; Earthwatch Belize Expedition 1981, D. H. Messersmith, W. E. Steiner, at al. [ USNM] ; 1 ♂, 1 ♀, 23 eggs: ex Zucht: B. Kneubühler 2009, F1– Generation, Herkunft: Belize, Greenhills, Cayo, leg. J. Meermann 2008 [ FH, No’s 0704–1, 2 & E] .

Diagnosis. Females of this species may be distinguished from all other Central American members of the genus by the distinctively elongate-ovoid and minutely bi-cornute head, that has the vertex just very weakly convex and is 1.5x longer than wide, as well as the acute medio-longitudinal keel of the mesonotum and otherwise almost smooth meso- and metathorax (distinctly granulose and/or tubercular in all other known species). The only other species with a rather elongate-ovoid head and a smooth (but not medio-longitudinally keeled) mesonotum is Ph. maximus sp. n. from Panama, but this huge species readily differs by the much larger size and slenderer shape, lack of a cephalic pair of spines and characters of the terminalia, such as a subgenital plate that projects beyond the apex of the abdomen by notably more than the length of the anal segment (much less than length of anal segment in vosseleri ). Moreover, the praeopercular organ on abdominal sternum VII has a short and obtuse posteromedian bulge that is not seen in any of the other known Central American ♀♀ of the genus. The apterous ♂♂ ( Fig. 55C ) resemble those of the type-species Ph. burkartii Saussure, 1868 and Ph. chiapasense sp. n. both from Mexico. From those of burkartii they differ by the smaller size, lack of the whitish dorsal portion of the vertex and somewhat more ovoid head ( Fig. 60J ), rather flattened and posteriorly narrowed anal segment that has the posterior margin deeply notched and bi-lobed ( Fig. 65R ; tectate longitudinally, posteriorly widened and merely with a very shallow median emargination in burkartii ), broader and more triangular vomer ( Fig. 97J ) and less bulgy, flat-angled poculum that has the central protrusion notably less pronounced ( Fig. 65Q ; rectangular in lateral aspect in burkartii ). From those of the latter species they may be separated by the stockier shape and relatively shorter body segments (mesothorax 7.5x longer than prothorax vs. 9.5x in chiapasense ), more ovoid head and elongated head ( Fig. 60J ), distinct medio-longitudinal carina on the meso- and metanotum, longer and slenderer cerci and a vomer that has the lateral margins less inflated, the central portion less impressed and exhibits several transverse furrows throughout almost the entire ventral surface ( Fig. 97J ).

Description. The colouration is described from colour photographs of live wild and captive reared insects ( Fig. 128C ).

♀♀: Moderately sized (body length incl. subgenital plate 134.0– 184.5 mm) and averagely robust species with an elongate and oval, minutely bi-spinose head, almost entirely smooth body and a short subgenital plate, that scarcely projects beyond the apex of the abdomen. Body surface very slightly glossy. General colouration various shades of grey and brown and to a variable degree irregularly flecked with pale drab or dark brown, sometimes also with a slight olive or greenish wash. The holotype is dark greyish brown with irregular buff to ochre mottling. Occasionally specimens may have a white or cream-coloured or reddish ( Fig. 128C ) medio-longitudinal stripe running along portions or the entire dorsal body surface or the medio-longitudinal of the mesonotum orange. Pronotum often with a fine black medio-longitudinal line. Eyes yellow and irregularly flecked with mid brown. All femora and tibiae to a variable degree furnished with faint irregular dark brown transverse bands. Antennae except for scapus and pedicellus drab or greyish brown dorsally, dark greyish brown ventrally and becoming ochreous to buff towards the apex.

Head: Elongate-ovoid, broadest just behind the eyes, weakly narrowing towards the posterior and with the vertex just very gently rounded; about 1.5x longer than wide. Frons with a shallow C-shaped impression in posterior portion. Vertex smooth and with an impressed coronal line in posterior portion, behind the eyes with a pair of small but acutely pointed, slightly anteriad directed spines. Eyes strongly projecting, circular in outline and their diameter contained about 2.3x in length of genae. Antennae reaching to posterior margin of abdominal segment III. Scapus distinctly compressed dorsoventrally, very slightly narrowing towards the base and with the lateral margins straight; 1.6x longer than wide. Pedicellus round in cross-section, slightly club-shaped and a little less than half the length of scapus.

Thorax: Pronotum somewhat shorter and narrower than head, 1.6x longer than wide with a slight narrowing pre-medially. Anterior margin with a pair of small, conical median tubercles, surface otherwise smooth. Transverse median sulcus moderately distinct, gently curved and almost reaching lateral margins of segment. Mesothorax 7x longer than prothorax, weakly constricted anteriorly and the posterior portion gently expanded. Metanotum about 0.4x the length of mesonotum with lateral margins gently concave and roughly 3x longer than wide. Meso- and metanotum with a fine medio-longitudinal keel (more pronounced on mesonotum) and a minutely granulose longitudinal carina close to lateral margins; metanotum otherwise smooth but mesonotum with some scattered minute granules in anterior portion and along medio-longitudinal keel. Mesopleurae with a minutely granulose longitudinal marginal carina, this much less distinct on metapleurae. Meso- and metasternum sparsely and minutely granulose, the mesosternum weakly tectate longitudinally.

Abdomen: Median segment scarcely shorter than metanotum, noticeably constricted medially and with a fine medio-longitudinal carina; about 2.9x longer than width at anterior margin. Segment II as long as median segment and 2x longer than wide. II–IV slightly increasing in length, V as long as IV, longest segments and 2.1x longer than wide. VI–VII decreasing length with VII only three-fifth the length of IV and V. Terga IV–VI with a small, scale-like posteromedian projection, otherwise all terga smooth except for a fairly distinct longitudinal lateral keel close and parallel to lateral margins. Tergum VII basically narrower than preceding, the lateral margins slightly expanded posteriorly to form a small, rounded lateral lobe ( Fig. 93G ). Sterna smooth except for a fine longitudinal carina close to lateral margins. Praeopercular organ on sternum VII formed by an elongate a short and obtuse posteromedian bulge and a slight, rounded posterior expansion of the two lateral carinae ( Fig. 93G ). Terga VIII–X uniform in width and slightly narrower than VII, VIII almost two-thirds the length of VII, about 1.7x longer than wide and parallel-sided, IX a little more than half the length of VIII and roughly quadrate in dorsal aspect. Anal segment slightly longer than IX, finely carinate medio-longitudinally with the posterior portion widely rounded and the posterior margin with a shallow median indention. Cerci small and tapering towards a narrow apex. Gonapophyses VIII elongate, upcurved in the apical portion and reaching to posterior margin of anal segment. Subgenital plate canaliculate and obtusely keeled longitudinally with the apex broadly rounded and projecting over apex of abdomen by less than length of anal segment; the basal half with a very obtuse lateral longitudinal bulge.

Legs: All moderately long, slender and distinctly carinated. Profemora a little shorter than mesothorax, mesofemora reaching about half the way along abdominal segment II, metafemora projecting over posterior margin of abdominal segment IV and metatarsi slightly projecting over apex of abdomen. Two outer ventral carinae of meso- and metafemora ± deflexed and rounded sub-basally; the same carinae forming a fairly distinct triangular apical tooth. Posteroventral carina of meso- and metatibiae gently deflexed and rounded apically. Legs otherwise unarmed. All basitarsi with a fairly prominent, roundly triangular dorsal crest; probasitarsus distinctly, meso- and metabasitarsi just slightly longer than corresponding following three tarsomeres combined.

♂♂ ( Fig. 55C ): Moderately sized (body length 111.0–113.0 mm) and slender, apterous insects with an oval, unarmed head. Body surface entirely smooth but very faintly tectate medio-longitudinally and slightly glossy. General colour of body greyish ochre to mid brown, great parts of meso- and metathorax with a bluish or bluish green wash and posterior margins of abdominal terga II–VII with a dark brown to black transverse. Prothoracic epimerum black. Abdominal tergum IX with an irregularly shaped white marking anterolaterally and two elongate black markings antero-medially. Head with an indistinct dark brown postocular line ( Fig. 60J ). Eyes dark yellow and irregularly flecked with reddish brown. Legs greyish mid brown with very faint pale annulations, the meso- and metafemora pale red towards the base. Apex of all femora black. Antennae except for scapus and pedicellus reddish mid to dark brown and blackish ventrally.

Head ( Fig. 60J ): Ovoid with the vertex very gently convex and the genae distinctly narrowing towards the posterior; broadest at the eyes and about 1.5x longer than wide, proportionally somewhat shorter than in ♀♀. Between the bases of the antennae with a small, circular impression and between the eyes with a gently convex area. Eyes large, circular in outline and projecting hemispherical from head capsule; their diameter contained only 1.4x in length of genae. Antennae reaching to abdominal segment VII. Scapus and pedicellus as in ♀♀, but pedicellus not club-shaped but rather cylindrical.

Thorax: Pronotum basically as in ♀♀, slightly narrower and a little shorter than head, rectangular in dorsal aspect with the lateral margins slightly concave pre-medially ( Fig. 60J ); 1.6x longer than wide. Surface with an impressed longitudinal median line and a distinct, gently curved transverse median sulcus, that spans the whole width of segment and laterally terminates in a shallow pit. Mesothorax very long, about 7.5x longer than prothorax, uniform in diameter except for a slight widening posteriorly and a little narrower than prothorax. Meso- and metanotum very faintly tectate medio-longitudinally and with a longitudinal lateral row of minute granules close to lateral margins. Meso- and metasternum with a fairly distinct medio-longitudinal keel.

Abdomen: Median segment 0.8x the length of metanotum, very gently constricted medially and about 5.3x longer than width at anterior margin; surface weakly tectate medio-longitudinally. Segments II–VII uniform in diameter. II four-fifths the length of median segment and 3.6x longer than wide. II–V roughly uniform in length and a little longer than II or VI; on average 4x longer than wide. VI and VII decreasing in length with VII only two-thirds the length of III–V. All sterna very weakly tectate medio-longitudinally. Tergum VIII widest segment, about three-fifths the length of VII and trapezoidal in dorsal aspect with posterior margin 1.4x wider than anterior margin. IX elongate, 1.3x longer but narrower than VIII, slightly constricted medially and with posterior margin somewhat narrower than anterior margin; lateral margins somewhat deflexed posteriorly and straight ( Fig. 65R ). Lateral surfaces of VIII and IX each with an obtuse but distinct longitudinal bulge, that is notably arched on IX. Anal segment short, scarcely more than half the length of IX, 1.5x longer than wide, slightly gradually narrowing towards the apex; the posterior margin distinctly bi-lobed with a deep median notch ( Fig. 65R ). Ventral surface of outer angles densely set with minute dark brown teeth ( Fig. 97J ). Cerci long, slender, gently incurved and with the apex slightly club-like; about three-quarters the length of anal segment. Vomer elongate-triangular in shape and gradually tapered towards a short but acute terminal hook ( Fig. 97J ); the outer margins somewhat inflated, the central area somewhat impressed and with several irregular transverse grooves. Poculum small, moderately convex, reaching only about two-thirds the way along tergum IX ( Fig. 65S ) and with a small and obtuse, angular basal protrusion ( Fig. 65Q ).

Legs: All long, slender and wholly unarmed. Profemora about equal in length to head, pro- and mesothorax combined, mesofemora somewhat longer than mesothorax, metafemora reaching about half the way along abdominal segment VI and metatibiae projecting considerably beyond apex of the abdomen. All basitarsi slender and considerably longer than remaining corresponding tarsomeres taken together.

Egg ( Fig. 99K ). Fairly small but typical for the genus. Shape angularly ovoid with the polar-area flattened and the dorsal egg surface strongly convex in the anterior portion; oval in cross-section. Capsule surface deeply and densely punctured, the pits small but deep and sharply defined. Micropylar plate elongate, the anterior half slender and the posterior half roundly widened and about four-fifths the length of capsule; structured like capsule and with the central portion slightly convex. Micropylar cup small and with a short longitudinal keel dorsally. Median line distinct and almost reaching to polar-area. Operculum oval and with a roundly convex hollow excrescence that is formed by the outer margin and which has about seven angular lateral impressions and a round central impression on top; height of excrescence about one-sixth of capsule length. Colour of capsule plain grey, the polar-area with a dark grey spot and a variably shaped dark grey marking on each lateral surface. Opercular collar dark grey, the opercular excrescence dark orange.

Table 52: Measurements of Phanocles vosseleri ( Redtenbacher, 1908)

Measurements [mm]: Length incl. operculum 3.7, length 3.1, width 2.0, height 2.5, length of micropylar plate 2.0.

Comments. The previously unknown ♂ and egg are here described and illustrated. Since the original diagnosis of the ♀ provided by Redtenbacher (1908: 419) is very brief and insufficient, a more detailed description appears warranted and is presented as well. Hennemann & Conle (2003: 6) correctly transferred this species from the genus Bacteria Berthold, 1827 to Phanocles Stål, 1875 .

Culture stock was imported to Europe on two occasions. A first stock from Green Hills Butterfly Ranch, Belmopan, Belize collected by Jan Meerman in 2008 and first reared in captivity by Bruno Kneubühler (Luzern, Switzerland) and a second stock from Tikal, Guatemala collected in 2013 and first reared in captivity by Mieke Duytschaever ( Belgium) in 2014 ( Fig. 128C ). The species has however not proven particularly easy to maintain in culture and hence both populations were lost after just a few generations. Although the natural host plants are not known it accepted wild bramble ( Rubus fagifolius , Rosaceae ) and Cupania belizensis ( Sapindaceae ) in captivity at Green Hills Butterfly Ranch, Belmopan, Belize (personal communication with Jan Meerman). In captivity in Europe bramble ( Rubus spp. , Rosaceae ) was frequently accepted as an alternative food plant. At average temperatures of 20–23°C eggs hatched at high ratios of over 75% after an incubation of about four months.At the same temperatures ♂♂ reached maturity after ten weeks, while ♀♀ took some twelve weeks to become adult. Females are fairly prolific and produce an average 4– 5 eggs per day, which are simply flicked away by an abrupt movement of the abdomen.

5.21. Genus Phanoclocrania gen. n.

urn:lsid:zoobank.org:act:13ED6216-5A42-45AE-9B9C-E4918193C466

( Figs. 68 , 86K–L , 95J–K , 100F , 104B )

Type-species: Bostra dorsuaria Stål, 1875: 79 , by present designation.

Bacteria, Redtenbacher, 1908: 412 (in part).

Bostra Stål, 1875: 79 (in part).

Redtenbacher, 1908: 406 (in part).

Zompro, 2001a: 200 (in part).

Phanoclocrania in litt, Conle, Hennemann & Gutiérrez, 2011: 60, 312.

Description. ♀, ♂ ( Fig. 68 ): Large to very large (body length: ♂♂ 100.0– 159.5 mm, ♀♀ incl. subgenital plate 172.0–227.0 mm), fairly elongate and slender Cladomorphini , with large cephalic horns (may be missing in ♂♂ although), an acute medio-longitudinal keel (♀♀) or carina (♂♂) on meso- and metasternum and a median segment that is about ¾ the length of the metanotum. ♂♂ apterous. Colour various shades of ochre, buff and brown, more rarely green; the pronotum mostly with a black longitudinal median line or streak ( Fig. 68F ), which is often continued on rest of dorsal body surface. Body surface smooth. Head strongly globose, hardly longer than wide with vertex moderately convex and armed with a huge pair of laterally compressed and apically pointed horns, which usually extend by ± the height of head capsule in ♀♀ ( Figs. 68D–E ); these cephalic horns occasionally reduced or missing in ♂♂ ( Fig. 68F ). Antennae reaching to posterior margin of median segment (♀♀) or to abdominal segment III (♂♂). Scapus compressed dorsoventrally and roughly rectangular in dorsal aspect, about 2.2x longer than wide. Pedicellus almost round in cross-section and roughly half the length of scapus. III slightly longer than pedicellus. Pronotum about as long but much narrower than head; roundly rectangular and notably longer than wide. Mesothorax elongate, uniform in diameter and at least 6x (♀♀) or 10x (♂♂) longer than prothorax. Mesosternum acutely keeled medio-longitudinally ( Figs. 86K–L ). Abdominal segment II roughly as long as median segment. II–VII parallel-sided and roughly equal in length, about 3.2x (♀♀) to 8x (♂♂) longer than wide. Tergum VII of females with lateral margins parallel-sided. All sterna with an obtuse medio-longitudinal carina (more pronounced in ♀♀). Sternum VII of ♀♀ with a distinct praeopercular organ that is formed by a pair of large, slender and acutely pointed lobes at posterior margin ( Figs. 68 G-H). Terminalia of ♀♀ ( Figs. 68G–H ): Terga VIII–X tectate medio-longitudinally ( Fig. 68H ), roughly uniform in width, just indistinctly narrower but much shorter than preceding segments. IX shorter than VIII. Anal segment with a widely triangular posteromedian excavation and the outer angles deflexed and broadly rounded. Epiproct very small and almost fully hidden under anal segment. Cerci small, short, somewhat compressed laterally, conical and with a fairly obtuse apex ( Fig. 68H ). Gonapophyses VIII moderately elongated, filiform, slightly upcurved, broadened sub-apically and projecting over apex of abdomen by less than length of anal segment ( Fig. 68G ). Subgenital plate naviculate, keeled longitudinally and gradually tapered towards a fairly acute apex; projecting over abdomen by less than length of anal segment. Terminalia of ♂♂ ( Figs. 68J–K , 95J–K ): Terga VIII and IX somewhat wider than all preceding segments, VIII slightly gradually widening towards the posterior. IX somewhat swollen anteriorly, constricted medially and with posterior half considerably narrowed; the lateral margins moderately to strongly deflexed and either rounded or protruded into an obtusely triangular lobe posteriorly ( Fig. 68J ). Anal segment longer than wide, distinctly narrower than IX, parallel-sided in dorsal aspect, declining in lateral aspect and the posterior margin bi-labiate with a distinct posteromedian indention ( Fig. 68K ). Posterolateral angles swollen and set with various small teeth ventrally; occasionally protruded into a downward-directed obtusely digitiform process. Epiproct small. Cerci slender, elongate, ± incurved and variable in length ( Figs. 95J–K ). Vomer either basically triangular in with a single terminal hook, or of complex shape with apex basically trifid, rather spatulate with the basal two-thirds roughly parallel-sided and the lateral margins each terminating in an obtuse tooth and the terminal hook slender. Poculum large, strongly bulgy, cup-like and convex with a huge, posteriad directed, spiniform basal projection ( Fig. 68J ). Legs long and slender and all distinctly carinate; meso- and metafemora and corresponding tibiae weakly downcurved in ♀♀. Medioventral carina of profemora lamellate and conspicuously displaced towards anteroventral carina, on meso- and metafemora roughly centric on ventral surface of femur and unarmed. Meso- and metafemora often with a ± prominent, tooth-like to foliaceous appendage about one-third off the base on two outer ventral carinae (more pronounced in ♀♀). All basitarsi of both sexes with the dorsal carina ± raised and rounded (more distinct in ♀♀); ± as long as remaining tarsomeres combined.

Egg ( Fig. 100F ). The following description is based on two examples of the type-species Ph. dorsuaria (Stål, 1875) comb. n. from Guacimo, Rio Frío, Province Límon, Costa Rica in the first author’s collection (coll. FH, No. 01242–E). Fairly large with a very high opercular extension. Capsule angular and slightly compressed laterally with the lateral surfaces flattened; capsule surface unevenly coriaceous and slightly shiny. Polar area angular, flattened. Entire ventral surface and polar area surrounded by a distinctly raised somewhat crenulate ridge, which encircles an elongate and parallel-sided, smooth area that resembles a micropylar plate. This area of slightly darker colour than rest of capsule. Micropylar plate four-fifth as long as capsule, somewhat pear-shaped with a slight constriction just above mid and roughly 2.3x longer than wide. Outer margin with a lamellate fringe, that has the outer margin minutely serrate. Lower central portion with a distinct, conical black swelling, the micropylar cup distinct and placed at posterior end of plate. Median line very short. Operculum almost circular and furnished with a large hollow structure formed by lamellate extensions of the outer margin; these longitudinally connected by thin membranes. Height of opercular extension almost equal to length of capsule. Colour of capsule ranging from dark grey over buff to dark brown, the fringe of the micropylar plate and ventro-posterior ridge drab. Capitulum dark orange. Measurements [mm]: overall length 5.0, capsule length 2.7, width 2.1, height 2.9, length of micropylar plate 2.2.

Etymology. The name is a combination of the generic name Phanocles and the Greek κραΝΊΟΝ or cranium (= skull) used as an epithet to indicate the close relation to that genus and refer to the impressive cephalic armature of members of this new genus. Feminine.

Differentiation. This new genus is similar to Phanocloidea Zompro, 2001 and Parotocrania gen. n. but is readily recognised by the very prominent, spiniform basal projection of the poculum of ♂♂ ( Fig. 68J ). From both genera it can frequently be separated by features such as the medio-longitudinally keeled ventral body surface (at best mesosternum faintly tectate in Phanocloidea and Parotocrania ), as well as the dorsally lobed basitarsi of ♂♂. From Parotocrania gen. n. ♂♂ of this new genus may also be separated by the mostly smooth surface of the head and thorax (granulose to nodulose head and occasionally also with single granules and nodes on abdomen in Parotocrania ) and ♀♀ differ by the much more distinct medio-longitudinal keel of the subgenital plate.

In several aspects Phanoclocrania gen. n. is also similar to Otocrania Redtenbacher, 1908 and Globocrania gen. n. both of which are peculiar to the Atlantic Forest along the Brazilian east coast. The distinct medio-longitudinal keel of the meso- and metasternum, dorsally lobed basitarsi and black longitudinal median line or streak of the pronotum of both sexes as well as the huge pair of cephalad horns in ♀♀ (occasionally also present in ♂♂) are shared with both genera. From Otocrania this new genus however readily differs by the much less globose and convex head of both sexes ( Figs. 68D–F ), lack of a prominent posterior spine on the metanotum, shorter median segment that is notably shorter than the metanotum, distinct praeopercular organ ( Figs. 68 G-H), much shorter subgenital plate and gonapophyses VIII ( Fig. 68G ) of ♀♀ as well as the lack of tegmina and alae ( Fig. 68A ) and short poculum of ♂♂, which bears a very prominent, spiniform basal protuberance ( Fig. 68J ) but is not extended and acuminate apically. From Globocrania gen. n. it can be separated by the more distinct and acute medio-longitudinal keel of the meso- and metasternum of both sexes, smooth meso- and metapleurae and more prominent praeopercular organ of ♀♀ as well as the very prominent, transverse basal protuberance of the poculum of ♂♂.

The eggs of Phanoclocrania gen. n. ( Fig. 100F ) strongly resemble those of Phanocloidea and Globocrania gen. n. in having a fairly angular capsule, strongly inflated or sculptured outer margin of the micropylar plate and a huge, hollow peripheral opercular extension, that extends by more than half of the capsule length. While they differ from the eggs of both genera by the smoother, merely coriaceous and slightly glossy capsule surface, they may additionally be distinguished from the eggs of Globocrania by the larger and more or less parallel-sided micropylar plate (pear-shaped with the anterior end gradually narrowing in Globocrania ).

For a more detailed comparison and distinction of Phanoclocrania gen. n. from Otocrania and Globocrania gen. n. see table 8 above and for a distinction from Phanocloidea and Parotocrania gen. n. see table 53 below.

A similarly prominent spiniform basal projection of the poculum as in this new genus is otherwise only seen in ♂♂ of the Mexican Phanocles saussurei ( Redtenbacher, 1908) comb. n., but this species has a simple vomer with a single terminal hook and lacks the distinct medioventral keel of the meso- and metasternum and lobed dorsal carina of the basitarsi, that are typical for Phanoclocrania gen. n.

Comments. Examination of both sexes of B. dorsuaria has shown this remarkable species to be misplaced in the preoccupied Bostra Stål, 1875 (= Phanocloidea Zompro, 2001 ), because it violates the generic description in several morphological aspects (see differentiation above). Unfortunately, there has been some confusion with the authority of this new genus, that was caused by the preliminary use of the name by Conle et al. (2011: 59). Brock & Büscher (2022: 514) therefore listed the genus as “ Phanoclocrania Hennemann & Conle, 2022 ” but unfortunately there has been massive delay of the present work, which now formally describes Phanoclocrania as a new genus. Hence, the authority should be referenced to this current publication. The eggs are here described and illustrated for the first time

Distribution ( Fig 104B ). Southern Central America ( Panama & Costa Rica) and Colombia.

Table 53: Comparison of Phanocloidea Zompro, 2001 , Phanoclocrania gen. n. and Otocrania Redtenbacher, 1908

......Continued on the next page

Table 53: (continued)

* Exceptions are Phanocloidea gracilis ( Burmeister, 1838) comb. n., P. incompta ( Rehn, 1904) comb. n., P. lanceolata

Conle, Hennemann & Gutiérrez, 2011 and P. schulthessi ( Redtenbacher, 1908)

Species included:

Figures

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 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 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 68. Phanoclocrania gen. n. dorsuaria Stål, 1875: A. ♂ (Costa Rica, Prov. Limón, Las Mercedes), habitus dorsolateral view [ZMUH]; B. ♀ with large sub-basal ventral lobes on meso- and metafemora (Costa Rica, Prov. Heredia, Sarapiqui), habitus dorsolateral view [coll. FH, No. 1242–1]; C. ♀ with unarmed meso- and metafemora(Costa Rica, Prov. Heredia, Sarapiqui), habitus dorsolateral view [coll. FH, No. 1242–2]; D. ♀, head and prothorax in lateral view [coll. FH]; E. ♂ with large cephalic horns (arrow), head and prothorax in lateral view [ANSP]; F. ♂ with unusually reduced cephalic armature (arrow), head and prothorax in lateral view [ZMUH]; G. ♀, lateral view of terminalia [MNCR-A, © Kenji Nishida]; H. ♀, ventral view of terminalia [MNCN]; J. ♂, lateral view of terminalia [MNCN] (arrow indicating the large, dstinctive central projection of the poculum); K. ♂, dorsal view of terminalia [ANSP]; L. ♀, intero-lateral view of basal half of left mesofemur [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 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 128. Live insects. A. Phanocloidea pallidenotata (Redtenbacher, 1908) ♀ at Montagne de Cheavaux, French Guiana (© Stepháne Brulé); B. Phanocloidea pallidenotata (Redtenbacher, 1908) ♂ at Montagne de Cheavaux, French Guiana (© Stepháne Brulé); C. Phanocles vosseleri (Redtenbacher, 1908) ♀ at Tikal, Guatemala (© Mieke Duytschaever); D. Mating couple of Phanocloidea incompta (Rehn, 1904) at Volcann Tenorio, Costa Rica (© Gernot Kunz).