Hesperophasmatini

Frank H. Hennemann, Oskar V. Conle & Daniel E. Perez-Gelabert, 2016, Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “ Anareolatae ”: Phasmatidae: Cladomorphinae), Zootaxa 4128 (1), pp. 1-211 : 21-28

4.2.4. Hesperophasmatini Bradley & Galil, 1977

( Figs. 20–35 , 47–48 )

Type-genus: Hesperophasma Rehn, 1901: 271 .

Hesperophasmatini Bradley & Galil, 1977: 188 (in part). Cladoxerinae Karny, 1923: 237 (in part).

Diapheromerini “ Clonistria -group” Zompro, 2001: 228 (in part). Haplopodini Günther, 1953: 557 (in part).

Phibalosomini (Sectio V: Phibalosomata), Redtenbacher, 1908: 399 (in part). Phibalosominae, Shelford, 1908: 343, 355 (in part).

Phibalosomatinae, Moxey, 1971: 44 (in part).

Differentiation ( Table 2 ): Close relation to Pterinoxylini n. trib. and Haplopodini is obvious and a detailed but as yet provisional differentiation from these two tribes is presented in Table 2 . These three tribes are likely to represent a monophyletic clade that differs from Cladomorphinae sensu stricto by a number of characters (→ see above). The differentiation can at this point only be provisional and deserves considerably more research, since the authors have at hand several as yet undescribed genera and species that would need to be incorporated for providing a sufficient and satisfying differential diagnosis (→ see comments below). Eggs of the currently known genera readily differ from both these tribes by the distinct hairy structures of the capsule and operculum ( Figs. 24–27 ), but eggs of some of the as yet undescribed taxa have almost entirely smooth capsules. A character that is unique within this clade and the entire Cladomorphinae is the secondary beak-like ovipositor in ♀♀ of Agamemnon Moxey, 1971 and Rhynchacris Redtenbacher, 1906 , which is formed by an elongated subgenital plate and epiproct.

The most obvious characters that distinguish Hesperophasmatini from Pterinoxylini n. trib. are: the much smaller size (body lengths: ♀♀ 33.0–94.0 mm, ♂♂ 24.5–90.0 mm); more robust and stocky body (♀♀ in particular); rather rectangular cross-section of the profemora (distinctly triangular with the anterodorsal strongly raised in Pterinoxylini n. trib.); not broadly expanded and lamellate dorsal carinae of the protibiae and not displaced medioventral carina of the meso- and metafemora of both sexes. Furthermore, Hesperophasmatini lack developed tegmina or alae in ♀♀ and at best have very small and scale-like rudiments of alae in Hypocyrtus Redtenbacher, 1908 . Males are either apterous, brachypterous or have fully developed alae. The presence of sensory areas on the probasisternum and/or profurcasternum ( Figs. 28–30 ) is shared with Pterinoxylini n. trib., which is likely to be a synapomorphy of Pterinoxylini n. trib. + Hesperophasmatini . This would support a sistergroup relationship between these two tribes and distinguishes both from their possible sister-group, the Haplopodini ( Fig. 409 ). The lack of a gula In Hesperophasmatini might be an autapomorphy but this deserves evaluation since this character is shared with Haplopodini.

From Haplopodini members of this tribe at once differ by: the generally more robust body and shorter legs (♀♀ in particular); presence of rough sensory areas on the probasisternum and/or profurcasternum (exception Lamponius Stål, 1875 ); generally more elongate antennomeres, and more rectangular cross-section of the profemora (± triangular in Haplopodini). Males of the currently known genera furthermore differ by being mostly brown or grey, while those of Haplopodini are often very colourful insects.

Comments: No detailed and definite diagnosis of Hesperophasmatini is presented here, since it appears as yet premature. The authors have at hand a large number of new species and even new genera mostly from Hispaniola, whose description will be the subject of forthcoming publications. Several of these new taxa exhibit characters, which deserve comprehensive examination and evaluation and will change our present view of current Hesperophasmatini in various aspects. Hence, any diagnosis would now only be preliminary and cause more confusion instead of supporting the actual systematic position and clarifying the relationships of Hesperophasmatini within the subfamily Cladomorphinae . Some morphological features of the seven currently recognized genera however warrant a more detailed discussion, which is presented below.

Bradley & Galil (1977: 188) established Hesperophasmatini to replace Haplopodini Günther, 1953. Zompro (2004: 135) removed the genera Aploploides Rehn & Hebard, 1938 , Haplopus Burmeister, 1838 and Diapherodes Gray, 1835 and erroneously placed them in the tribe Cranidiini (→ 4.2.3). The genera Hesperophasma Rehn, 1901 , Lamponius Stål, 1875 , Rhynchacris Redtenbacher, 1908 and Taraxippus Moxey, 1971 were retained in Hesperophasmatini by Zompro (2004: 139), who however omitted Agamemnon Moxey, 1971 and erroneously synonymised Hypocyrtus Redtenabcher, 1908 with Lamponius (see below).

Examination has shown the monotypical genus Laciphorus Redtenbacher, 1908 (Type-species: Laciphorus lobulatus Redtenbacher, 1908: 351 ) from Peru is not a member of Hesperophasmatini but belongs in Diapheromerinae : Diapheromerini , hence is here removed from Hesperophasmatini (→ 8.3). A redescription of Laciphorus and a discussion of its systematic position within Diapheromerini by the authors is in progress (Hennemann & Conle, in preparation). The monotypical Tersomia Kirby, 1904 (Type-species: Tersomia brasiliensis Kirby, 1904 ) from Brazil is obviously misplaced and here removed from Hesperophasmatini . Females, the only sex known, are large and very slender, typically stick-like insects with a long, spatulate subgenital plate, a very short median segment, very long and slender, unarmed legs and a pair of horns between the eyes. These features as well as the remarkably short antennae, which consist of no more than 23 segments, and morphology of the genitalia place Tersomia in Heteronemiidae : Heteronemiinae : Paraleptyniini (→ 8.3).

Although lacking an area apicalis, Zompro (2004: 139) transferred Hesperophasmatini from the anareolate Phasmatidae : Cladomorphinae to the areolate Pseudophasmatidae : Xerosomatinae . The author stated they simply represented derived Xerosomatinae , in which the area apicalis is reduced, and postulated Hesperophasmatini were a subordinate taxon of Xerosomatinae . However, the very few features on which Zompro (2004: 139) based his action appear arbitrary (e.g. “eggs bullet-shaped”, “interodorsal carina of the protibiae lamelliform” and “mid- and hind legs agree completely”). Indeed, some Hesperophasmatini resemble certain representatives of Xerosomatinae : Xerosomatini at first glance, but presuming a close relation between these two tribes mostly based on convergences such as a similar general resemblance, basally curved and compressed profemora (a character widely distributed throughout the entire Phasmatodea and also typical for all members of the family Phasmatidae and subfamily Cladomorphinae respectively) and trapezoidal meso- and metafemora is the result of superficial examination. The fact that Xerosomatini (e.g. Creoxylus Audinet-Serville, 1838 or Xylospinodes Zompro, 2004 ) of similar habitus occur in geographically close regions or even in the same habitats as certain Hesperophasmatini (e.g. Rhynchacris in Central America) is not at all surprising. It merely shows the “similar resemblance” discussed by Zompro (2004: 139) are convergent developments caused by similar or identical evolutional pressure in the common habitats of these taxa. Detailed examination and comparison of several representatives from both groups undertaken in the course of this work have revealed numerous morphological characters of the insects and eggs that clearly separate Hesperophasmatini not only from Xerosomatinae but from the entire family Pseudophasmatidae . This is clearly supported by phylogenetic studies based on external morphology (Bradler, 2009: 99) and molecular data ( Whiting et al., 2003; Buckley et al., 2009; Buckley et al., 2010). Bradler (2009: 99) furthermore stated that he could not find any synapomorphies that Lamponius Stål, 1875 (the only genus of Hesperophasmatini this author examined) had in common with genera of Xerosomatinae and that none of the apomorphies of the family Pseudophasmatidae were present in this Hesperophasmatini genus. Consequently, Zompro's placement of Hesperophasmatini as a subordinate taxon of Pseudophasmatidae : Xerosomatinae cannot be supported and the tribe is here re-transferred back to Cladomorphinae , where it was originally placed by Günther (1953: 557). In fact, close relation to Pterinoxylini n. trib. and Haplopodini is obvious and the most important characters that support the position within Cladomorphinae are discussed in more detail below.

Hesperophasmatini is a principally Antillean taxon, with only two genera represented in Central America ( Hypocyrtus Redtenbacher, 1908 and Rhynchacris Redtenbacher, 1908 ). The vast number of taxa is distributed througout the West Indies and as yet only a fraction of the tribe's true diversity is known. The authors have at hand numerous new species and genera mostly from Hispaniola, which multiply the number of currently known taxa. No representatives of Hesperophasmatini are so far known from the Bahamas.

The northernmost distributed representative of this tribe, the Central American Hypocyrtus deserves special mention, since the genital morphology of ♂♂ differs considerably from that of all other members of Hesperophasmatini . While there is a well-developed and sclerotized vomer ( Figs. 31–33 ) in ♂♂ of all other currently known genera, it is strongly reduced or missing and concealed by the paraprocts (= sternum XI) in Hypocyrtus ( Hennemann & Conle, 2012: 66, see Fig. 34 ). Furthermore, the anal segment is much more tectiform than in the other genera and has the interior portions of the posterior margin facing each other and armed with prominent in-curving spines ( Fig. 35 ). In all other genera, which do have a well-developed vomer, the posteromedian portion is merely incised and interiorly set with a variable number of small denticles. These striking genital differences are particularly remarkable, since the ♀♀ genitalia perfectly match with those of other Hesperophasmatini as does the insect and egg-morphology. Despite these striking differences of the external ♂♂ genital morphology and other obvious distinguishing features Zompro (2004: 140) erroneously synonymised Hypocyrtus with the exclusively Caribbean Lamponius . This misinterpretation was corrected by Eilmus (2009: 30), who re-established Hypocyrtus . This was confirmed by Hennemann & Conle (2012), who provided a detailed revision of the genus at the species-level. The Central American Rhynchacris and the Antillean Agamemnon are the only two genera whose ♀♀ exhibit a beak-like secondary ovipositor, that is formed by the elongated subgenital plate and an elongated epiproct. The eggs of both genera are more elongate and bullet-shaped than those of other genera and instead of simply being dropped to the ground are laid into a substrate ( Figs. 25–26 ).

Distribution: West Indies (excluding Bahamas) and Central America, ranging from Central Mexico as far south as the northern portions of Colombia.

Below is a more detailed discussion of certain important characters, which distinguish and clearly exclude Hesperophasmatini from Pseudophasmatidae : Xerosomatinae :

1. Area apicalis: Zompro (2004: 139) stated that Hesperophasmatini strikingly “resemble” species of Xerosomatinae : Xerosomatini , and that the lack of an area apicalis on the tibiae was the only significant difference between these two tribes. The author postulated the area apicalis was merely reduced and hence interpreted the lack of this structure as an autapomorphy of Hesperophasmatini . However, as not only Xerosomatinae , but the entire family Pseudophasmatidae have a distinct area apicalis on all three tibiae and there are no transitions between these two extremes, Hesperophasmatini is, based solely on this feature, most unlikely to belong in Pseudophasmatidae .

2. Genitalia (♀♀): The genitalia of ♀♀ of Hesperophasmatini differ fundamentally from those of Xerosomatinae by having a distinctly keeled, scoop-like subgenital plate, which is tapered towards the apex and ± distinctly projects over the apex of the abdomen. Two genera ( Agamemnon and Rhynchacris ) even have the epiproct conspicuously elongated to form a beak-like secondary ovipositor together with the elongated subgenital plate. Very different specializations of the ♀ ovipositor are observed in Xerosomatinae . The subgenital plate is either strongly reduced, very small and scale-shaped (e.g. Acanthoclonia Stål, 1875 , Metriophasma Uvarov, 1940 ), or slender, ± tube-like and down-curving with the lower gonapophyses enlarged and sclerotized (e.g. certain species of Creoxylus ) forming a conspicuous derived appendicular ovipositor. The gonoplacs are lacking in Hesperophasmatini , while they are present and often conspicuously enlarged in representatives of Xerosomatinae (Bradler, 2009: 99) . All mentioned features do not at all support the treatment of Hesperophasmatini by Zompro (2004: 139). Instead, the elongated subgenital plate and lack of gonoplacs are shared with Pterinoxylini n. trib. and Haplopodini and hence clearly prove the very close relation to these two tribes, which together form a supposedly monophyletic clade within Cladomorphinae ( Fig. 409 ).

3. Genitalia (♂♂): Also the genital morphology of ♂♂ of Hesperophasmatini differs fundamentally from that of Xerosomatini . In Xerosomatini the anal segment is more or less distinctly tectiform and has the posteromedian portion sub-fissately divided with the bounding lips facing each other, labiately thickened and on their interior surfaces with several small in-curving teeth. The vomer is strongly reduced and missing (see Bradler, 2009: 99, figs. 22b–d). In contrast, Hesperophasmatini have the anal segment flattened and the thorn-pads at the posterior margin directed downward. The vomer is well developed, sclerotized and appearing as a roundly triangular plate usually with a single, prominent hook-like apex ( Figs. 31–33 , also see Bradler, 2009: 99, fig. 22a). An exception is represented by Hypocyrtus whose genital morphology resembles members of Xerosomatini by the tectiform anal segment and reduced vomer (see above, Figs. 34–35 ). However this genus perfectly matches with Hesperophasmatini in every other aspect. These striking morphological differences of the genitalia also do not support the systematic placement of Hesperophasmatini suggested by Zompro (2004: 139), but instead show remarkable affinity to Pterinoxylini n. trib. and Haplopodini and confirm the close relation to these three tribes here suggested ( Fig. 409 ).

4. Sensory areas: Most of the currently known genera of Hesperophasmatini (exceptions are Lamponius and Taraxippus ) possess a large and well developed sensory area on the profurcasternum, an important feature completely neglected and overlooked by Zompro (2004). Sometimes, two further much smaller sensory areas are present near the exavations of the procoxae on the probasisternum (e.g. Rhynchacris , Fig. 29 ). The sensory area of the profurcasternum is often very large, slightly convex and may cover a considerable part of the segments surface ( Figs. 28, 30 ). It is formed by a cluster of small granules or wart-like humps, each of which bears 1–5 hollow, tubelike spines or papillate evaginations. The sensory areas on the probasisternum of e.g. Rhynchacris are much smaller, consist of a single hump-like swelling and usually have the papillate evaginations relatively longer. In Lamponius these swellings of the probasisternum are present but lack the evaginations seen in Rhynchacris . The function of these sensory areas is as yet unknown, but most certainly they serve as tactile or olfactory organs, since the live insects are frequently observed rubbing their prosternum on various surfaces, such as twigs or foliage of the host plant. In addition to several other common features the presence of sensory areas on the profurcasternum and probasisternum support close relation to Pterinoxylini n. trib., which corresponds to the results presented by Bradler (2009) and Buckley et al. (2009) (→ 4.2.5). In Pterinoxylini n. trib. however the sensory area of the profurcasternum is rather small, elliptical and strongly convex and the evaginations are considerably shorter than in members of Hesperophasmatini , being roughly cone-shaped with a deep central pit. Similar sensory areas may be present on the profurcasternum of certain members of Xerosomatini (e.g. some species of Acanthoclonia or Creoxylus ), while there are never sensory areas on the probasisternum. The sensory areas on the profurcasternum of certain Xerosomatini-members are however rather different in structure and in contrast to Hesperophasmatini cover almost the complete surface of the segment, being formed by a cluster of densely arranged small, wart-like swellings, which lack any kind of evaginations or papillate structures. The different structure of the sensory areas do also not support the position of Hesperophasmatini supposed by Zompro (2004: 139). An ultrastructural study of these sensory areas is necessary for any broader discussion on their function or importance for phylogeny (Bradler, 2009: 32).

5. Egg-morphology: Zompro (2004: 139) stated the eggs of Hesperophasmatini and Pseudophasmatidae : Xerosomatinae were similar in the aspect that they are “bullet-shaped”, and the micropylar plate is small, placed roughly in the “centre of the capsule” [certainly meaning centre of the dorsal egg surface] and projects from the capsule surface. These observations are surprising since eggs of Xerosomatini are never “bullet-shaped” as stated by Zompro (2004: 139) but ± ovoid. Eggs of Hesperophasmatini are variable in shape, ranging from distinctly elongate and bullet-like with the polar-area more or less decidedly tapered over ovoid to almost spherical. In Xerosomatinae the capsule surface is merely covered by ± distinctly raised, net-like structures, while the capsule surface and operculum are to a variable degree covered with conspicuous hairy structures in all currently known genera of Hesperophasmatini ( Figs. 24–27 ). Xerosomatinae have a small, roughly circular to elliptical micropylar plate, which distinctly projects from the capsule surface and is open internally with a rather narrow posteromedial gap and a distinct median line ( Fig. 51 ). In contrast, the micropylar plate of Hesperophasmatini never projects considerably from the capsule surface, is relatively larger and variable in shape, being slightly transverse and shield-shaped to distinctly longer than wide with the anterior end narrowed and the posterior portion broadened. Internally the plate is open with a wide, triangular notch but without a median line ( Figs. 47–48 ). Consequently, also the egg-morphology does not support the relation between Hesperophasmatini and Xerosomatinae postulated by Zompro (2004: 139). Instead, the lack of a median line is shared with Pterinoxylini n. trib. and Haplopodini and also confirms the close relation to these two tribes revealed in the present study and also suggested by the results of Bradler (2009) and Buckley et al. (2009).

Genera included:

1. Agamemnon Moxey, 1971: 71 . Type-species: Agamemnon iphimedeia Moxey, 1971: 75 , by original designation of Moxey, 1971: 71.

2. Hesperophasma Rehn, 1901: 271 . Type-species: Phantasis saussurei Bolivar, 1888: 137 , by subsequent designation of Kirby, 1904a: 343. [Replacement name for Phantasis Saussure, 1870 ]

= Phantasis Saussure, 1870: 188 . Type-species: Phantasis saussurei Bolivar, 1888: 137 , by subsequent designation of Kirby, 1904a: 343.

3. Hypocyrtus Redtenbacher, 1908: 355 . Type-species: Hypocyrtus substrumosus Redtenbacher, 1908: 357 (= Hypocyrtus postpositus Redtenbacher, 1908 ), by subsequent designation of Zompro, 2000: 95.

4. Lamponius Stål, 1875: 19 , 74. Type-species: Pygirhynchus guerini Saussure, 1868: 64 , by monotypy. = Antillophilus Carl, 1913: 38 . Type-species: Antillophilus brevitarsus Carl, 1913: 38 (= Lamponius guerini ( Saussure, 1868) , by monotypy.

5. Rhynchacris Redtenbacher, 1908: 354 . Type-species: Rhynchacris ornata Redtenbacher, 1908: 354 , by monotypy. = Pseudoceroys Hebard, 1922: 354 . Type-species: Pseudoceroys harroweri Hebard, 1922: 355 , pl. 15: 1–2 by original designation. [Synonymised by Hennemann & Conle, 2012: 81]

6. Tainophasma Conle, Hennemann & Perez-Gelabert, 2014: 29 . Type-species: Tainophasma monticola Conle, Hennemann & Perez, Gelabert, 2014: 31 , by original designation.

7. Taraxippus Moxey, 1971: 67 . Type-species: Taraxippus paliurus Moxey, 1971: 70 , by original designation of Moxey, 1971: 67.

Figures

FIGURES 409. Possible relationships within the subfamily Cladomorphinae sensu lato. The tree shown above is merely meant to visualise the possible relationships here suggested. It does not restrict to defining apomorphies for each clade or taxon, but also uses key-features that are helpful for distinction. Hence, it should not be interpreted as a strict phylogenetic cladogramm.

FIGURES 20 – 23. Tribe Hesperophasmatini Bradley & Galil, 1977. 20. Lamponius guerini (Saussure, 1868) ♀: captive reared from Guadeloupe, Bassé Terre [coll. FH, No. 0084 - 39]; 21. Lamponius portoricensis Rehn, 1903 ♂: captive reared from Puerto Rico, Luquillo Experimental Forest [coll. FH, No. 0467 - 6]; 22. Hesperophasma planulum (Westwood, 1859) ♂: Dominican Republic, Province Barahona [USNM]; 23. Hesperophasma planulum (Westwood, 1859) ♀: Dominican Republic [MNCN].

FIGURES 24 – 30. Tribe Hesperophasmatini Bradley & Galil, 1977. 24. Lamponius guerini (Saussure, 1868) egg (dorsolateral view): captive reared from Guadeloupe, Bassé Terre [coll. FH, No. 0084 - E]; 25. Agamemnon cornutus (Burmeister, 1838) egg (dorsal view): captive reared from Virgin Islands, Tortola [coll. FH, No. 0589 - E]; 26. Rhynchacris ornata Redtenbacher, 1908 egg (dorsal view): captive reared from S-Costa Rica, Prov. Limón [coll. FH, No. 0134 - E]; 27. Hypocyrtus scythrus (Westwood, 1859) egg (dorsolateral view): captive reared from Mexico, Veracruz, Tuxtlas [coll. FH, No. 0638 - E]; 28. Rhynchacris ornata Redtenbacher, 1908 ♀: sensory-area of profurcasternum [coll. FH, No. 0134 - 12]; 29. R. ornata ♀: left sensory area of prosternum [coll. FH, No. 0134 - 12]; 30. Hypocyrtus scythrus (Westwood, 1859) ♀: sensory-area of profurcasternum [coll. FH, No. 0638 - 1].

FIGURES 31 – 36. Tribes Hesperophasmatini Bradley & Galil, 1977 and Pterinoxylini n. trib., genitalia of ♂♂. 31. Lamponius portoricensis Rehn, 1903: ventral view of anal segment [coll. FH, No. 0467 - 6]; 32. Agamemnon cornutus (Burmeister, 1838): ventral view of anal segment [coll. FH, No. 0589 - 6]; 33. Rhynchacris ornata Redtenbacher, 1908: ventral view of anal segment [coll. FH, No. 0134 - 32]; 34. Hypocyrtus scythrus (Westwood, 1859): ventral view of anal segment [coll. FH, No. 0638 - 7]; 35. Hypocyrtus scythrus (Westwood, 1859): caudal view of anal segment [coll. FH, No. 0638 - 7]; 36. Pterinoxylus crassus Kirby, 1889: ventral view of anal segment [coll. FH, 0588 - 7].

FIGURES 45 – 51. Internal micropylar plates of Cladomorphinae and Xerosomatinae. 45. Tribe Haplopodini: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825); 46. Tribe Pterinoxylini n. trib.: Pterinoxylus crassus Kirby, 1889; 47. Tribe Hesperophasmatini: Lamponius guerini (Saussure, 1868); 48. Tribe Hesperophasmatini: Rhynchacris ornata Redtenbacher, 1908; 49. Tribe Cranidiini: Cranidium gibbosum (Burmeister, 1838); 50. Tribe Cladomorphini: Cladomorpus rubus (Saussure, 1861); 51. Pseudophasmatidae: Xerosomatinae: Xerosomatini: Creoxylus spinosus (Fabricius, 1793).