Spinocloidea, 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 : 29-33

25. Xylodus Saussure, 1859: 62 . Type-species: Xylodus adumbratus Saussure, 1859: 62 , by original monotypy.

Keys to the genera of Cladomorformia tax. n.

♀♀

1. Body ± round in cross-section and ± stick-like.............................................................. 2

- Body strongly flattened; abdomen prominently dilated laterally; mesosternum tectate and with a granulose longitudinal median keel; NE South America........................................................................ Cranidium

2. Median segment> ¼ the length of metanotum.............................................................. 3

- Median segment <¼ the length of metanotum.............................................................. 4

3. Body length (incl. subgenital plate) <150.0 mm; head sub-cylindrical and notably longer than wide with vertex flattened and often armed..................................................................................... Calynda

- Larger, body length (incl. subgenital plate)> 150.0 mm; head ovoid to sub-globose with vertex convex and unarmed............................................................................................ Spinocloidea gen. n.

4. Typical stick mimics; body without large foliaceous lobes and appendages........................................ 5

- Remarkable crypsis, moss-mimicing; body all over furnished with foliaceous lobes and appendages of variable sizes; Central America & North-western South America........................................................ Trychopeplus

5. Metanotum with posterior margin at best moderataly inflated and with a rounded median protuberance................. 6

- Metanotum armed with a huge anteriad directed median spine; E-Brazil (Antlantic Forest).................... Otocrania

6. Subgenital plate at least reaching to apex of abdomen (if shorter then apical half with a medio-longitudinal furrow ventrally *).................................................................................................. 7

- Subgenital plate small, scoop-shaped and scarcely projecting over posterior margin of abdominal tergum IX; Bolivia................................................................................................... .. Bostriana

7. Gonoplacs ± enlarged and longer than gonapophyses VIII and IX.................................................8

- Gonoplacs shorter than gonapophyses VIII and at best as long as gonapophyses IX................................. 10

8. Mesonotum and mesopleurae unarmed or granulose to sparsely tubercular; gonoplacs at best scarcely projecting over posterior margin of anal segment................................................................................. 9

- Mesonotum and mesopleurae armed with prominent tubercles and blunt spiniform projections; gonoplacs enormously enlarged, paddle-shaped and projecting beyond apex of abdomen by length of anal segment; W-Peru (Coastal Lomas)...... Laciphorus

9. Head ovoid to sub-globose, unarmed or with a pair of spines between the eyes; Western South America east of the Andes............................................................................................. Globocalynda

- Head elongate, sub-cylindrical with vertex flattened and> 1.5x longer than wide, unarmed; Andean regions of Colombia & Ecuador................................................................................... Andeocalynda

10. Abdominal tergum VI with a posterior pair of swellings, protuberances or excrescences............................ 11

- Abdominal tergum VI flat and unarmed................................................................... 12

11. Medioventral carina of meso- and metafemora and tibiae unarmed............................... Hirtuleiodes gen. n.

- Medioventral carina of meso- and metafemora and tibiae armed with prominent and strong spines................. Jeremia

12. Abdominal tergum V with posterior swellings, protuberances or excrescences.................................... 13

- Abdominal tergum V flat and unarmed................................................................... 15

13. Mesothorax elongate,> 4x longer than prothorax combined................................................... 14

- Much stockier insects; mesothorax only 3.5x longer than prothorax......................................... Xylodus

14. Very slender insects; abdominal segment V almost 5x longer than wide; subgenital plate norrowly canaliculate and extending beyond apex of abdomen by considerably more than length of terminal three abdominal terga combined; basitarsi not lobed dorsally.................................................................................... Otocraniella

- Stockier insects; abdominal segment V at best 3x longer than wide; subgenital plate broadened with lateral margins ± undulate and projecting beyond apex of abdomen by no more than length of terminal three abdominal terga taken together; basitarsi with a lobe dorsally............................................................................. Cladomorphus

15. Dorsal carinae of protibiae not notably raised or lamellate..................................................... 16

- Posterodorsal carina of protibiae prominently raised, uniformly lamellate and blade-like............ Ocreatophasma gen. n.

16. Praeopercular organ very indistinct........................................................................17

- Praeopercular organ obvious, represented by a pair of tubercles, protuberances, spiniform to digitiform processes or lobes near posterior margin of abdominal sternum VII................................................................ 18

17. Mesonotum, meso- and metapleurae densely granulose to nodulose...................................... Jeremiodes

- Mesonotum, meso- and metapleurae prominently spinose........................................ Aplopocranidium

18. Mesonotum unarmed or ± evenly granulose to minutely tubercular (rarely spinose)................................. 19

- Mesonotum unevenly tubercular and with a pair or cluster of prominently enlarged conical spines or protuberances pre-medially; Amazon basin................................................................................. Hirtuleius

19. Gonapophyses VIII short and concealed under anal segment, even if subgenital plate is strongly elongated or lanceolate and projects greatly beyond the tip of the abdomen.............................................................. 20

- Gonapophyses VIII always ± elongated and projecting considerably beyond tip of anal segment, at least reaching to tip of subgenital plate if this is not notably elongated............................................................. 21

20. Mesosternum with a prominent medio-longitudinal keel; meso- and metapleurae at best tubercular; Mexico & Northern Central America........................................................................... .. Lanceobostra gen. n.

- Mesosternum weakly tectate longitudinally and usually with two closely spaced, parallel longitudinal rows of tubercleses; meso- and/or metapleurae distinctly spinose; E-Brazil (Atlantic Forest)........................... Globocrania gen. n.

21. Lateral margins of abdominal tergum VII parallel-sided...................................................... 22

- Lateral margins of abdominal tergum VII dilated to form a ± prominent lobe posterolaterally (**); almost entire Neotropical Region comprising S-Mexico, Central America, Lesser Antilles and northern half.......................... .. Phanocles

22. Mesosternum at best weakly tectate medio-longitudinally or with a fine medio-longitudinal ridge..................... 23

- Mesosternum with a very prominent and acute medio-longitudinal keel; head with a huge pair of cephalic horns that project by more than height of head capsule; Costa Rica & W-Panama.................................. Phanoclocrania gen. n.

23. Subgenital plate distinctly keeled medio-longitudinally and laterally compressed in the apical portion (if not reaching tip of abdomen, then apical half with a medio-longitudinal furrow ventrally *); mesopleurae and/or mesosternum occasionally with spines; Central America, Tobago and northern portion of South America (usually north of the Amazon)....... Phanocloidea

- Subgenital plate scoop-shaped, at best very weakly keeled and rather flattened and obtuse in apical portion (***, occasionally with apex tridentate and ± downcurved); no spines on mesopeurae and mesosternum; only Panama and NW South America ( Colombia, Ecuador & N-Peru)............................................................ Parotocrania gen. n.

* Only true for Phanocloidea gracilis ( Burmeister, 1838) comb. n. and Ph. ibaguena ( Giglio-Tos, 1910) comb. n.

** P. aequatorialis ( Redtenbacher, 1908) comb. n., P. chaperi ( Redtenbacher, 1908) comb. n., P. horni ( Redtenbacher, 1908) comb. n., P. remphan ( Westwood, 1859) and P. siginificans ( Redtenbacher, 1908) comb. n. and Ph. vosseleri ( Redtenbacher, 1908) comb. n. have these lobes just weakly developed or almost missing

*** Only exception P. pastazae ( Hebard, 1924) comb. n.

♂♂ *

1. Medioventral carina of profemora distinct and notably displaced towards anteroventral carina; poculum with variable in shape (often with various specialisation) but at best 1.5x higher than body diameter; phallus unarmed....................... 2

- Medioventral carina of profemora obscur e and central on ventral surface of profemur; poculum very Bulgy, deeply tub-shaped and extending ventrally by more than 2x body diametre; phallus spinulose; NE South America................ Cranidium

2. Cerci at best slightly longer than anal segment; vomer well-developed........................................... 3

- Cerci strongly elongated, 90° incurved and 2x longer than anal segment; vomer reduced, spatulate and apically bidentate; Central America............................................................................. Alienobostra

3. Median segment <¼ the length of metanotum...............................................................4

- Median segment> ¼ the length of metanotum.............................................................. 6

4. Lateral margins of abdominal tergum IX straight or at best gently convex, not dilated; anal segment ± parallel-sided; vomer triangular; Central America & Colombia................................................................... 5

- Lateral margins of abdominal tergum IX dilated into a large triangular downward and posteriad directed posterolateral process; anal segment strongly widening towards the posterior and trapezoidal n dorsal aspect; vomer roundly rectangular; Bolivia............................................................................................... Bostriana

5. Body length <110.0 mm; poculum incised and bilobate posteriorly; apex of cerci rounded..................... Calynda

- Body length> 110.0 mm; poculum with posterior margin rounded and entire; apex of cerci flattened dorsoventrally and truncated interiorly...................................................................... Spinocloidea gen. n.

6. Poculum projecting noticeably over apex of abdomen........................................................ 7

- Poculum not projecting over apex of abdomen............................................................. 10

7. Poculum with a prominent tubular or spatulate apical appendix, that projects beyond tip of abdomen by notably more than length of anal segment (**)............................................................................. 8

- Poculum with apex elongated, spatulate-acuminate and projecting beyond tip of abdomen by less than length of anal segment; Brazil (Atlantic Forest).......................................................................... Otocrania

8. Alate; head convex and ± bi-cornute...................................................................... 9

- Apterous; head flattened and unarmed; Eastern Brazil (Minas Geraïs)................................... Otocraniella

9. Slender insects; mesothorax> 4.3x longer than prothorax.......................................... Cladomorphus

- Stockier; mesothorax 3.75x longer than prothorax...................................................... Xylodus

10. Cerci remarkably enlarged and specialised; thorn pads of anal segment reduced................................... 11

- Cerci oval or cylindrical in cross-section; paired thorn pads of anal segment distinct............................... 13

11. Posterior margin of poculum bidentate; medioventral carina of four posterior tibiae unarmed......................... 12

- Posterrior margin of poculum entire; medioventral carina of four posterior tibiae dentate........................ Jeremia

12. Mesonotum unarmed; cerci obtusely hook-shaped................................................... Jeremiode s

- Mesonotum prominently spinose; cerci laterally compressed and angled inward....................... Aplopocranidium

13. Typical stick mimics; body and legs without large foliaceous lobes and appendages................................ 14

- Remarkable crypsis, moss-mimicing; body and legs furnished with various large foliaceous lobes and appendages; Central America & North-western South America........................................................ Trychopeplus

14. Vomer basically triangular in shape........................................................................15

- Vomer roundly rectangular with a terminal hook at each posterolateral angle............................... Hirtuleius

15. Median segment at best ½ the length of metanotum (***).................................................... 16

- Median segment longer............................................................................... 18

16. Poculum large, bulgy and projecting considerably beyond posterior margin of abdominal tergum IX.................. 17

- Poculum not reaching beyond abdominal tergum IX; Andean regions of Colombia & Ecuador.............. Andeocalynda

17. Head elongated, sub-cylindrical and 1.5x longer than wide; lateral margins of anal segment with a distinct, downward directed digitiform process; West Peru “Coastal Lomas”..................................................... Laciphorus

- Head ovoid, <1.5x longer than wide; no digitiform process on lateral margins of anal segment; Western South America east of the Andes.................................................................................. Globocalynda

18. Poculum unarmed or at best with a small, spiniform central projection that extends by less than half the height of segment. 19

- Poculum with a huge, spiniform, ventro-posterior directed process, that extends by more than ¾ the height of segment; Central America and N-Colombia............................................................. Phanoclocrania gen. n.

19. Slender insects; mesothorax> 6x longer than prothorax; abdominal tergum VI smooth.............................. 20

- Stockier; mesothorax at best 6x longer than prothorax; abdominal tergum VI with a posterior pair of protuberances............................................................................................. Hirtuleiodes gen. n.

20. Cerci ± round in cross-section........................................................................... 21

- Cerci ± paddle-shaped with interior surface impressed and canaliculate longitudinally; exclusively apterous; Mexico & Northern Central America....................................................................... Lanceobostra gen. n.

21. No black medio-longitudinal streak on pronotum; anal segment not considerably widened posteriorly; if widened then poculum bulgy and with posterior margin simple and entire........................................................... 22

- Pronotum with a distinct black medio-longitudinal streak; anal segment strongly deflexed post-medially or posteriorly; poculum scoop-shaped with posterior margin distinctly indented medially and bidentate; E-Brazil (Atlantic Forest).................................................................................................... Globocrania gen. n.

22. Head and body surface smooth or at best with a few minute, dispersed granules on mesothorax........................23

- Head and thorax distinctly granulose or nodulose; Panama, Colombia & Ecuador................... Parotocrania gen. n.

23. Apterous, brachypterous or alate; bases of profemora never red; meso- and metathorax never conspicuously colourful; abdominal tergum IX parallel-sided; poculum strongly bulgy, fornicate and angular in lateral aspect, projecting beyong abdominal tergum IX and with a ± distinct obtuse to spiniform central protuberance or projection (no specialisations of posterior margin); almost entire Neotropical Region comprising S-Mexico, Central America, Lesser Antilles and northern half of South America.................................................................................................. Phanocles

- Almost exclusively apterous (****); bases of profemora often red; meso- and metathorax may be multi-coloroured; abdominal tergum IX ± dilated laterally (often forming a rounded or triangular lobe); poculum at best moderately bulgy and scoop-shaped, usually not projecting beyond abdominal tergum IX and often with specialisations of the posterior margin but never with a central protuberance (*****); Central America, Tobago and northern portion of South America (usually north of the Amazon).......................................................................................... Phanocloidea

* Males of Ocreatophasma gen. n. are not known.

** Exception Cladomorphus guianensis Conle et al., 2020 → appendix of poculum lacking

*** Exception Globocalynda colombiae ( Hebard, 1919) comb. n. → median segment two-thirds the length of metanotum

**** Exceptions are Ph. semiptera sp. n. and Ph. schulthessi ( Redtenbacher, 1908)

***** Only species with a central protuberance is Ph. incolumis (Brunner v. Wattenwyl, 1907 ) comb. n., but the poculum stays considerably before the posterior margin of abdominal tergum IX and the latter has strongly expanded almost semi-circular lateral margins

5.1. Genus Alienobostra Zompro, 2001

( Figs. 4C , 6C–D , 8 , 85E , 95H , 98L–M , 101A , 105 )

Type-species: Calynda brocki Hausleithner, 1987: 178 , figs. 1–2, 5–6 (= Bostra remiformis Rehn, 1904: 58 ), by original designation.

Alienobostra Zompro, 2001: 200 , figs. 10–11, 80–81, 126–127. Zompro & Brock, 2003: 13, 27, figs. 33–34. Otte & Brock, 2005: 43.

Brock & Büscher, 2022: 509.

Bostra, Rehn, 1904: 58 (in part).

Rehn, 1905: 72.

Redtenbacher, 1908: 406 (in part).

Bostranova Villet, 2023: 150 (in part).

Calynda, Hausleithner, 1987: 178 , figs 1–2, 5–6 (in part).

Description. ♀, ♂ ( Fig. 8 ): Moderately sized to large (body length ♂♂ 73.0–115.0 mm, ♀♀ including subgenital plate 128.0–205.0 mm) and slender Cladomorphini with a median segment that is slightly shorter than the metanotum, strongly elongated and inward angled cerci and a reduced vomer in ♂♂ and a long, lanceolate subgenital plate and indistinct praeopercular organ in ♀♀. ♂♂ apterous. Body surface of both sexes slightly shiny; entirely smooth in ♂♂, in ♀♀ smooth except for granules or tubercles on meso- and metathorax. Colour of ♀♀ ranging from green over yellow and straw to dark brown ( Figs. 105A–D ), ♂♂ either green, grey or brown with the lower portion of the genae white. Head elongate, ovoid and notably narrowed towards the posterior with the vertex flattened; unarmed in ♂♂ ( Fig. 8O ), in ♀♀ vertex either unarmed or with a pair of ± distinct protuberances or spines in front ( Fig. 8N ). Eyes rather small and their diameter contained at least 2x in length of genae. Antennae longer than head and complete thorax in ♀♀ and roughly ¾ the length of body in ♂♂. Scapus moderately compressed dorsoventrally, roundly rectangular in dorsal aspect and longer than wide. Pedicellus roughly half the length of scapus and round in cross-section. III considerably longer than pedicellus. Pronotum ± as long but narrower than head, roughly rectangular and longer than wide. Mesothorax uniform in diameter and at least 6.5x (♀♀) or 7.5x (♂♂) longer than prothorax. Mesonotum in ♀♀ usually with a variable number of granules or tubercles along lateral margins and meso- and metapleurae with a distinct longitudinal lateral carina or ridge that is almost parallel to the lateral margins and to a variable degree set with granules or spiniform tubercles (less decided on metapleurae). Meso- and metasternum in ♀♀ with a fine medio-longitudinal carina (more pronounced on mesosternum) and mesosternum sometimes with a few scattered granules; in ♂♂ just very faintly tectate longitudinally. Abdomen excluding median segment longer than combined length of head and complete thorax. Median segment about three-quarters the length of metanotum ( Fig. 8A ). Abdominal segment II shorter than median segment and slightly shorter than following segments. II–VII all considerably longer than wide, parallel-sided and of uniform diameter, II–V gradually increasing in length with V–VI longest segments. VII shorter than preceding, the lateral margins parallel-sided in ♀♀. Abdominal sterna II– VII smooth but occasionally with a few minute scattered granules in ♀♀. Praeopercular organ of ♀♀ very indistinct and merely represented by minute median tubercle at posterior margin of sternum VII. Terminalia of ♀♀ ( Figs. 8D–H ): Terga VIII–X considerably shorter and slightly narrower than all preceding segments, roughly of uniform width. Anal segment carinate longitudinally with posterior margin obtusely angular and with a very small and shallow median indention. Epiproct small, basically triangular in shape and hardly projecting over posterior margin of anal segment. Cerci small, conical and tapered towards a pointed tip, weakly incurved. Gonapophyses VIII short and not projecting over apex of anal segment, just slightly longer than gonapophyses IX. Gonoplacs digitiform and shorter than gonapophyses IX. Subgenital plate keeled longitudinally, lanceolate and greatly projecting over apex of abdomen; at least 1.5x longer than terga VIII–X combined and apex ± acuminate. Terminalia of ♂♂ ( Figs. 4C , 8J–L ): Tergum VIII and IX roughly equal in length, wider than all preceding segments and both trapezoidal with VIII slightly broadened towards posterior and IX narrowed posteriorly; both with an obtuse longitudinal lateral ridge. Lateral margins of IX strongly deflexed, rounded, downcurved and with posterolateral angles strongly approaching each other ventrally ( Figs. 4C , 8L ). Anal segment constricted basally, broadened in the apical half and gradually flattened towards the apex; the posterior margin straight but with the lateral angles protruded into an obtuse, digitiform process ( Fig. 8K ), that is set with several small black teeth ventrally ( Figs. 4C , 8L ). Epiproct very small, triangular and fully concealed by anal segment. Vomer strongly reduced and merely represented by a small, spatulate and apically bifid, sclerotized plate ( Figs. 4C , 95H ). Poculum small, moderately convex, angular with an obtuse central protuberance and roughly reaching two-thirds along tergum IX. Cerci prominently enlarged, much longer than anal segment, the apical half roughly 90° angled inward, the apical half compressed dorsoventrally, carinate, broadened and paddle-shaped ( Figs. 8J–L ). Legs all long and slender, profemora roughly as long (♀♀) or longer than mesothorax (♂♂), mesofemora longer than metathorax, and metatibiae not reaching (♀♀) or distinctly projecting over apex of abdomen (♂♂); tibiae longer than corresponding femora. Profemora with anterodorsal carina strongly raised and the medioventral carina distinct, lamellate and considerably displaced towards anteroventral carina (♀♀ in particular). Meso- and metafemora with the medioventral carina centric and armed with 2–4 minute teeth apically; legs otherwise unarmed in ♂♂. Meso- and metafemora in ♀♀ occasionally with a ± distinct sub-basal tooth or lobe on the two outer ventral carinae ( Fig. 8A ) and more rarely with posterodorsal carina forming an obtusely triangular tooth apically. Basitarsi slender in ♂♂, either slender or with dorsal carina weakly convex in ♀♀; longer than following three tarsomeres combined.

Egg ( Figs. 98 L-M). Small (capsule length <3.0 mm), roundly angular in lateral aspect with the anterodorsal portion convex and roundly angular, the lower half gradually narrowed and the polar area flattened; oval in cross-section. Capsule surface minutely granulose, slightly shiny and covered with a blunt network of indistinctly raised areas. Micropylar plate positioned in centre of dorsal egg-surface, ± two-thirds the length of capsule and roughly 3x longer than wide with the posterior portion gently widened and rounded; surrounded by a gently raised fringe. Micropylar cup represented by a blunt, rounded tubercle near polar end of plate. Median line short but distinct. Operculum oval and with a hollow capitulum formed by lamellate extensions of the outer margin; these longitudinally connected by thin membranes. Height of capitulum less than half of capsule length. Colour dark grey or brown with the raised areas of the capsule pale cream or grey. Micropylar plate dark brown to black. Capitulum straw or yellowish.

Differentiation. Mostly based on the anatomy of the terminalia of ♂♂ this exclusively Central American genus takes on a somewhat isolated position within Cladomorphini . Males are at once separated from those of all other genera of Cladomorphini by the strongly enlarged cerci, that are considerably longer than the anal segment and ca. 90° angled inward with the apex compressed dorsoventrally and paddle-shaped ( Figs. 4C , 8J–L ), as well as the very small, secondarily reduced, spatulate and apically bidentate vomer ( Figs. 4C , 95H ). In almost all other morphological aspects they strongly resemble the ♂♂ of Lanceobostra gen. n. The laterally deflexed abdominal tergum IX resembles only certain members of that genus (e. g. L. guatemalensis ( Redtenbacher, 1908) comb. n.) but also a Central American representative of Phanocloidea Zompro, 2001 ( P. incolumis ( Redtenbacher, 1908) comb. n.). Females of Alienobostra strikingly resemble those of Lanceobostra gen. n., but differ by the weakly developed praeopercular organ ( Fig. 9F ), less distinct medio-longitudinal keel of the meso- and metasternum ( Fig. 85E ) and very short gonapophyses VIII, which are scarcely longer than gonapophyses IX.

Comments. Although Zompro (2001a: 200) recognised most of the distinguishing features of ♂♂ this author did not provide any characters that would distinguish ♀♀ from related genera. Moreover, the original description is questionable and inaccurate in some aspects, why a new diagnosis of the genus is presented here. There has been considerable confusion concerning the identities of species that belong in Alienobostra , caused by untraced or immature type specimens. As a result of detailed investigation of all seven taxa concerned, two new generic combinations and five new synonymies were detected and are established herein.

Bradler (2009: 101) suggested Alienobostra to be the possible sister-taxon of Sermyleformia and mentioned the strongly elongated, incurved cerci as a possible synapomorphy of Alienobostra + Sermyleformia. This position of Alienobostra however is not supported by other morphological characters of the insects and eggs, because the genus agrees with Cladomorphini in almost all other aspects and does not possess any of the apomorphies mentioned for Sermyleformia ( Bradler, 2009). The cerci of ♂♂ lack the typical intero-basal protuberance seen in Sermyleformia, the vomer is notably reduced but is present (completely lacking in Eusermyleformia), the anal segment has ventral thorn pads at the posterolateral angles and ♀♀ have a long and lanceolate subgenital plate. Molecular data ( Forni et al., 2022; Bank & Bradler, 2022) also confirm Alienobostra as a subordinate taxon of Cladomorphini ( Fig. 133 ).

Distribution ( Fig 101A ). Central America ( Costa Rica, Nicaragua, Guatemala & S-Mexico). Records of this distinctive genus are restricted to the two biogeographical provinces Chiapas and Eastern Central America, both part of the Mesoamerican Dominion of the Caribbean Subregion (see Morrone, 2006: 471, fig. 2).

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 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 9. Andeocalynda Hennemann & Conle, 2021: A. A. tenuis Hennemann & Conle, 2020 ♀ HT, habitus dorsolateral view [NHMUK]; B. A. tenuis Hennemann & Conle, 2020 ♀ HT, habitus dorsal view [NHMUK]; C. A. decorata Hennemann & Conle, 2020 ♀ HT, habitus dorsal view [NHMUK]; D. A. decorata Hennemann & Conle, 2020 ♂ PT, habitus dorsal view [NHMUK]; E. A. tenuis Hennemann & Conle, 2020 ♂ PT, habitus dorsolateral view [NHMUK].

FIGURE 85. Mesosternum: A. Bostriana boliviana (Piza, 1939) ♀ [coll. OC]; B. Globocalynda ruficollis sp. n. ♀ PT [ANSP]; C. Globocalynda unilobata (Brunner v. Wattenwyl, 1907) ♀ Prov. Sara, Bolivia [MNHU]; D. Calynda bicuspis Stål, 1875 ♀ Volcán Baru, Chiriqui, Panamá [coll. FH]; E. Alienobostra remiformis (Rehn, 1904) ♀ Surubres nr. San Mateo, Costa Rica [ANSP]; F. Lanceobostra aetolus (Westwood, 1859) ♀ Venvidio, Sinaloa, Mexico [ANSP]; G. Lanceobostra deplanata (Redtenbacher, 1908) ♀ HT [NHMUK]; H. Lanceobostra freygessneri (Redtenbacher, 1908) ♀ HT [MHNG]; J. Lanceobostra similis (Redtenbacher, 1908) ♀ HT [NHMUK]; K. Ocreatophasma elegans gen. n., sp. n. ♀ HT [MNHN]; L. Ocreatophasma modestum gen. n., sp. n. ♀ HT [MNHN]; M. Ocreatophasma fragile gen. n., sp. n. ♀ HT [MNHN]. 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 98. Eggs: A. Andeocalynda decorata Hennemann & Conle, 2020 [coll. FH]; B. Andeocalynda tenuis Hennemann & Conle, 2020 [coll. FH]; C. Calynda bicuspis Stål, 1875 [coll. FH]; D. Calynda coronata Carl, 1913 [coll. FH]; E. Spinocloidea panamaense gen. n., sp. n. PT [coll. FH]; F. Bostriana boliviana (Piza, 1939) [coll. FH]; G. Globocalynda unilobata (Brunner v. Wattenwyl, 1907) ex abdomen PLT of Globocalynda simplex (Brunner v. Wattenwyl, 1907) [NHMW, © 2019 Natural History Museum Vienna, NOaS Image Collection / H. Bruckner]; H. Cladomorphus phyllinus Gray, 1835 [coll. FH]; J. Cladomorphus tuberculatus (Piza, 1936) [coll. FH]; K. Jeremia gymnota Günther, 1930 [ZSMC]; L. Alienobostra remiformis (Rehn, 1904) Costa Rica, Monteverde [coll. FH]; M. Alienobostra remiformis (Rehn, 1904) Costa Rica, San Mateo [ANSP]; N. Lanceobostra aetolus (Westwood, 1859) [ANSP]; O. Globocrania culmus (Bates, 1865) [MZUSP, © Pedro Ivo Chiquetto-Machado]; P. Globocrania tabida (Redtenbacher, 1908) Brazil, Est. Rio de Janeiro, Itatiaia N.P. [coll. FH]. [Scale = 1 mm]

FIGURE 101. Maps showing the known distributionsof the genera A. Alienobostra Zompro, 2001, B. Andeocalynda Hennemann & Conle, 2020, C. Aplopocranidium Zompro, 2004, D. Bostriana Hennemann, Conle & Valero, 2021, E. Calynda Stål, 1875 and F. Cladomorphus Gray, 1835.

FIGURE 105. Live insects of Alienobostra remiformis (Rehn, 1904): A. Mating couple at El Rodeo Experimental Forest 900 m, Costa Rica; B. Captive reared ♀, brown form with white medio-longitudinal streak on body (stock from Costa Rica, Monteverde); C. Mating couple at Monteverde 1530 m, Costa Rica (© Kenji Nishida); D. Captive reared ♀, green form with two white medio-longitudinal streaks on body (stock from Costa Rica, Monteverde).

FIGURE 133. Phylogenetic trees showing the most recent result concerning the subfamily Diapheromerinae based on molecular data. Both trees are simplified with root lengths and support values ignored and showing only the section that includes the taxa of the subfamily Diapheromerinae. Names of taxa are updated and corrected according to the taxonomic changes conducted in the present study. Trees were generated with iTool v6 (https://itol.embl.de) based on the original files in newick format. Cladomorformia tax. n. is highlighted in green with the arrow indicating the corresponding node. The genus Calynda is marked by an orange starbecause it is hereincluded inCladomorformia based on morphological traits,which is in contradiction to the here shown molecular results. A. according to Bank & Bradler (2022, additional file 2, File S1), modified (Remarks: *Paraphanocles sp. (Trinidad) is apparently misidentified and not a member of Paraphanocles (= Phanocles); **Paraclynda picta (Bruinner v. Wattenwyl, 1907) and Paracalynda utilaensis Zompro, 2001 are undoubtedly congeneric, thus the result postulated in the tree is implausible); B. according to Forni et al. (2022, Supplementary figure 1), modified (Remark: ***Alienobostra remiformis (Rehn, 1904) appears twice in the tree with the position as sister to Phantasca quadrilobata (Chopard, 1911) implausible; “Bacteriformia” is split into three different lineages, which is in contradiction to the tree of Bank & Bradler (2022) shown in A, which in this aspect is considered more relieable and plausible).