Phanocles pleurospinosus, 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 : 152-154

35. Phanocles vosseleri ( Redtenbacher, 1908: 419) [ Bacteria ]. HT, ♀: 52; Guatemala; 52. Phanocles vosseleri Br. [SMNS]. ( Figs. 55C , 60J , 65Q–S , 93G , 97J , 99K )

Remarks: The ♂ and eggs were previously unknown and are described from specimens originating in Belize and Guatemala below.

Distribution: Guatemala: Guatemala [SMNS]; Guatemala, Petén, Tikal [FH]. Belize: Belize, Toledo District, Blue Creek Village [USNM]; Belize, Belmopan, Green Hills [FH, OC].

Keys to the species of Phanocles

♀♀ *

1. Metapleurae unarmed (at best tubercular).................................................................. 2

- Metapleurae armed with prominent conical spines; Costa Rica................................ .. pleurospinosus sp. n.

2. Mesonotum without spines.............................................................................. 3

- Mesonotum armed with several distinct paired spines; Ecuador............................................. decorus

3. Thorax smooth (at best sparsely granulose)..................................................................4

- Thorax distinctly granulose, tubercular or spinulose......................................................... 12

4. Subgenital plate elongated and surpassing tip of abdomen by> 1.5x length of anal segment............................5

- Subgenital plate short and projecting beyond tip of abdomen by hardly more than length of anal segment................ 7

5. Head with a pair of prominent conical horns; meso and metatibiae smooth dorsally.................................. 6

- Head unarmed; meso- and metatibiae prominently and irregularly lobed dorsally; Colombia.................... lobulatus

6. Cephalic horns auriform and compressed laterally; praeopercular organ prominent, a pair of distinct scale-like lobes that project over sternum VII posteriorly; French Guiana & Suriname............................................. maroniensis

- Cephalic horn conical and acutely pointed; praeopercular organ obscure, only a black marking with a small central peg-like protunberance; Bolivia & E-Ecuador....................................................... acutecornutus sp. n.

7. Abdominal tergum VII with lateral margins at best weakly deflexed posteriorly.................................... 8

- Abdominal tergum VII with a distinct, rounded posterolateral lobe; Ecuador............................. falcatus sp. n.

8. Head unarmed or at best with a pair of small spines...........................................................9

- Head armed with a prominent pair of laterally compressed, angularly semi-circular lobes; E-Peru.............. .. virgulatus

9. Head elongate-ovoid and> 1.3x longer than wide........................................................... 10

- Head globose and <1.3x longer than wide.................................................................11

10. Very large (body length incl. subgenital plate> 190.0 mm) and slender species; mesothorax 8.6x longer than pronotum; Colombia.................................................................................... .. remphan

- Smaller (body length incl. subgenital plate <185.0 mm) and stockier species; mesothorax only 7x longer than pronotum; Guatemala & Belize............................................................................. vosseleri

11. Very large (body length> 220.0 mm); subgenital plate with apical portion roundly spatulate; meso-and metatibiae with a prominent rounded dorsal lobe at apex; Panama.................................................. maximus sp. n.

- Smaller (body length <210.0 mm); subgenital plate with apical portion strongly laterally compressed and canaliculated; no obvious apical dorsal lobe on meso- and metatibiae; Ecuador..................................... ecuadoricus sp. n.

12. Meso- and metanotum prominently spinulose; SW-Mexico................................................... 13

- Meso- and metanotum granulose to tubercular............................................................. 14

13. Back of vertex smooth; praeopercular organ formed by a closely spaced pair of spiniform appendices; meso- and metatibiae smooth dorsally................................................................................... horni

- Back of vertex tubercular; praeopercular organ a single peg-like posteromedian protuberance; meso-and metatibiae with three ± distinct dorsal lobes...................................................................... mexicanus sp. n.

14. Subgenital very long, lanceolate and projecting over tip of abdomen by notably more than the combined length of abdominal terga VIII–X, lanceolate............................................................................... 15

- Subgenital plate shorter............................................................................... 16

15. Small (body length incl. subgenital plate <160.0 mm) and very slender species; abdominal tergum VII with lateral margins just slightly deflexed posteriorly; legs slender and unarmed; Mexico...................................... berezini sp. n.

- Large (body length incl. subgenital plate> 200.0 mm) and stocky species; abdominal tergum VII with lateral margins forming a prominent, rounded lobe; Ecuador........................................................... superbus sp. n.

16. Slender to moderately stocky insects with mesonotum> 6x longer than pronotum................................. 17

- Very stocky species with mesonotum only 5.3x longer than pronotum; Ecuador.......................... solidus sp. n.

17. Posterior margin of mesonotum flat...................................................................... 18

- Posterior margin of mesonotum swollen to form a tubercular transverse bulge; Costa Rica.................. costaricensis

18. Head bi-cornute..................................................................................... 19

- Head unarmed; Panama......................................................................... subvolans

19. Subgenital plate short and surpassing tip of abdomen by notably less than length of anal segment..................... 20

- Subgenital plate longer................................................................................ 23 20. Head with a distinct pair of spines or horns................................................................ 21

- Head only with a small pair of low conical tubercles; Costa Rica & Panama........................... spectabilis sp. n.

21. Meso- and metanotum nodulose / tubercular; lateral margins of abdominal tergum VII just very weakly dilated posteriorly. 22

- Meso- and metanotum granulose; lateral margins of abdominal tergum VII forming a prominent and obtusely angular lobe; Nicaragua................................................................................... rehni sp. n.

22. Largest tubercles of mesonotum node-like; praeopercular organ obscure; N-Ecuador...................... aequatorialis

- Tubercles of mesonotum more acute; praeopercular organ formed by a pair of low, obtusely rounded lobes; Panama.. ploiaria

23. Cephalic horns ± acute and higher than wide; sculpturing of thorax sparse and/or uneven........................... 24

- Cepahlic horns very obtuse, rounded and wider than high; thorax densely, distinctly and evenly granulose (granules black tipped); Brazil & Venezuela............................................................. satyr

24. Apex of subgenital plate broadened and angulard........................................................... 25

- Subgenital plate tapering with apex narrowly rounded to pointed............................................... 26

25. Stocky species (mesonotum 6.1x longer than pronotum); legs short (metatibiae only reaching posterior of abdominal segment V), stocky, carinated and with meso- and metafemora distinctly downcurved; praeoperculuar organ formed by two huge, elongate and foliaceous lobes; E-Peru.......................................................... brevipes sp. n.

- Slenderer (mesonotum 7.5x longer tan pronotum); legs elongate (metatibiae reaching abdominal segment VIII) and slender, not totably curved; praeopercular organ indistinct; SW-Mexico.............................................. saussurei

26. Praeopercular organ indistinct; granules of thorax not white................................................... 27

- Praeopercular organ formed by a pair of large downward directed lobes; granules of thorax white Colombia..... significans

27. Body length incl. subgenital plate <180.0 mm; lateral margins of abdominal tergum VII at best weakly arounded posteriorly. .................................................................................................. 28

- Larger (body length incl. subgenital plate> 190.0 mm); abdominal tergum VII with a ± distinct lateral lobe; W-Mexico (Veracruz ...................................................................................... burkartii

28. Meso- and metanotum unevenly granulose; paired protuberances of the praeopercular organ distinct and acutely pointed or angular; Lesser Antilles............................................................................... 29

- Meso- and metanotum with scattered nodes; praeopercular organ only represented by a pair of shallow, rounded, rough swellings; Venezuela.............................................................................. chaperi

29. Lateral margins of abdominal tergum VII almost with only a very slight widening posteriorly; lateral margins of abdominal terga VIII–X straight; gonapophyses VIII not or jest very slightly projecting beyond anal segment.................... 30

- Abdominal tergum VII with a distinct rounded posterolateral lobes; lateral margins of abdominal terga VIII–X deflexed and obtusely rounded; gonapophyses VIII projecting considerably beyond anal segment with apex broadened and triangular; Barbados............................................................................. barbadosense sp. n.

30. Praeopercular organ form by two small spiniform projections; Trinidad & Tobago.............................. mutica

- Praeopercular organ formed by two short angular ridges; Dominica, Martinique, St. Lucia, St. Vincent, Grenada & Grenadines.............................................................................. keratosqueleton

[* The ♀♀ of Ph. chiapasense sp. n., Ph. obtusus ( Redtenbacher, 1908) comb. n. and Ph. undulatus Conle, Hennemann & Gutiérrez, 2011 are not known]

♂♂ *

1. Apterous............................................................................................ 2

- Alate.............................................................................................. 15

2. Poculum at best with a small and obtuse central protuberance....................................................3

- Poculum with a large, laterally compressed triangular, tooth-like projection; Mexico.......................... saussurei

3. Median segment <2x longer than metanotum............................................................... 4

- Median segment almost 3x longer than metanotum; Venezuela............................................. obtusus

4. Anal segment not notably broader than terga VIII–IX......................................................... 5

- Anal segment strongly dilated and much broader than preceding terga............................................ 6

5. Median segment slightly shorter than metanotum; anal segment widened medially; SW-Mexico.................... horni

- Median segment almost 2x longer than metanotum; anal segment widening towards posterior and trapezoidal in dorsal aspect; Lesser Antilles........................................................................... keratosqueleton

6. Meso- and metafemora without conspicuous coloration........................................................ 7

- Meso and metafemora pale cream or straw basally and with a distinct black sub-basal transverse band; Costa Rica................................................................................................... costaricensis

7. Abdominal tergum IX at best 1.5x longer than anal segment.................................................... 8

- Abdominal tergum IX distinctly elongated and almost 3x longer than anal segment; Mexico & Guatemala... mexicanus sp. n.

8. Poculum without a central protrusion.......................................................................9

- Poculum with a central protrusion or protuberance.......................................................... 10

9. Head without distinct dark markings; anal segment notched posteromedially; vomer asymmetrical with a curled dextral hook and a short digitiform sinistral point; Trinidad & Tobago.................................................. muticus

- Head with a black postoclar streak and an angularly C–shaped black marking on frons; anal segment with posterior margin angular; vomer slender and gradually narrowing towards apex; Peru..................................... .. virgulatus

10. Median segment shorter than metanotum.................................................................. 11

- Median segment longer than metanotum.................................................................. 12

11. Very delicate insects, mesonotum 9.5x longer than pronotum; meso- and metanotum smooth; Mexico (Chiapas)................................................................................................ chiapasense sp. n.

- Stockier, mesonotum only 7.5x longer than pronotum; meso- and metanotum with a fine medio-longitudinal carina; Guatemala & Belize...................................................................................... vosseleri

12. Vomer roughly symmetrical and gradually narrowed towards the apex.......................................... 13

- Vomer asymmetrical and with an additional small sinistral point near base; Barbados................. barbadosense sp. n.

13. Anal segment with posterior margin roundly angular or notched medially........................................ 14

- Anal segment tapering and narrowly rounded posteriorly; Colombia...................................... significans

14. Large (body length> 130.0 mm); median segment scarcely longer than metanotum; vomer almost symmetrical and gradually narrowing towards the apex; Mexico................................................................ burkartii

- Smaller (body length <120.0 mm); median segment almost 1.7x longer than metanotum; vomer asymmetrical and with an additional small sinistral point near base; Barbados........................................... barbadosense sp. n.

15. Head with a pair of tubercles, spines or prominent horns...................................................... 16

- Head unarmed; Colombia........................................................................ .. remphan

16. Alae at least reaching to abdominal segment III............................................................. 17

- Brachypterous, alae just slightly projecting over posterior margin of median segment; Panama................ .. subvolans

17. Head armed with a large pair of laterally compresed, carinate and auriform horns.................................. 18

- Head at best with a pair of tubercles or small conical spines................................................... 20

18. Mid and hind legs simple, slender; basitrasi not crested dorsally................................................19

- Mid and hind legs with various undulations; basitarsi with a triangular dorsal lobe; Colombia out a central protuberance, posterior margin indented medially and bidentate; cerci small and much shorter than anal segment; Bolivia & Ecuador.......................................................................................... acutecornutus sp. n.

- Head with a black postocular streak; poculum with a central protuberance and posterior margin triangular; cerci elongate, slender and as long as anal segment; French Guiana & Suriname....................................... maroniensis

20. Poculum with a conical central protuberance; Ecuador....................................................... 21

- Poculum roundly angular in lateral aspect and lacking a central protuberance......................................22

21. Anal segment tapering towards a narrowly rounded apex; vomer very elongate, slender and strongly arched... falcatus sp. n.

- Anal segment with lateral margins convex and posterior margin broadly rounded; vomer triangular in outline.................................................................................................. ecuadoricus sp. n.

22. Brown or grey insects; anal segment notably shorter than tergum IX............................................ 23

- Meso- and metathorax bright green; anal segment equal in length to tergum IX and almost 2x longer Than wide; Costa Rica & Panama................................................................................. spectabilis sp. n.

23. Head flattened and vertex only with a pair of shallow and obtuse swellings....................................... 24

- Head globose and vertex with a pair of acute but short conical spines; anal segment whitis dorsally; Panama........ ploiaria

24. Small insects (body length <100.0 mm); mesonotum with a few scattered node-like granules; meso-and metatibae simple; basitarsi slender; Ecuador..................................................................... aequatorialis

- Large insects (body length> 125.0 mm); mesonotum smooth; meso- and metatibiae with a rounded dorsal lobe at apex; basitarsi crested; Panama.................................................................... maximus sp. n.

[* The ♂♂ of Ph. berezini sp. n., Ph. brevipes sp. n., Ph. chaperi (Redtenacher, 1908) comb. n., Ph. cuzcoense sp. n., Ph. decorus Zompro, 2001 , Ph. lobulatus Conle, Hennemann & Gutiérrez, 2011 , Ph. pleurospinosus sp. n., Ph. rehni sp. n., Ph. satyr ( Redtenbacher, 1908) , Ph. solidus sp. n. and Ph. superbus sp. n. are not known]

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 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 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]