Phanocles mexicanus, 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 : 192-195

Phanocles mexicanus sp. n.

urn:lsid:zoobank.org:act:5148C229-BE73-46E3-9B1C-52006C933CE1

( Figs. 52B–C , 59A–C , 60D , 64 O-Q, 67G–H, 88D)

Phanocles burkartii, Zompro, 2001a: 198 , pl. 1: 2 [♀), pl. 5: 72–73 (egg). [Misidentification]

HT, ♀: Museum Paris, Mexique, Amedegnato/ Poulain rec.; Chis., Huixtla/Motozintla, km. 30, 700m, 27 X 86 [ MNHN] .

PT, ♀: Südl. Mexico, Tapachula, A. Kulow leg. 1898 [ ZMUH] .

PT, ♀: Mexico, Soconusco (Chiapas), 1200 m üb. Meer, Herm. Hans Kolow, 6.7.1900; Phanocles burkartii (Saussure) 1868 , det. O. Zompro XI.1997, hierzu ein Ei [ ZMUH] .

PT, ♀: Mexico, Soconusco (Chiapas), 1200 m üb. Meer, Herm. Hans Kolow, 6.7.1900 ; Phanocles burkartii (Saussure) 1868 , det. O. Zompro XI.1997 [ ZMUH] .

PT, ♀: Museum Paris, Guatemala, Dép. De Chimaltenango, Env. de Chimaltenango (800 a 1600 m. Dalt.), R. Guérin 1908; Bacteria voit de satyr [MNHN].

PT, ♂: 3/28, Guatemala, Mr. H. de Sauss. [ MHNG] .

PT, ♂: GUAT. Dept. Chemaltenango, Yepocapa, 1300±m, vii.10.1949, 14, L.C. Stuart; Bostra sp. prob. not mirata Redt., Det. T.H. Hubbell 1954 ; UMMZ–191538 [UMMZ].

PT, ♂: Costa Rica, Underwood?; Bostra n. sp. near B. incompta Rehn diff subg pl, ceci etc. [ANSP].

Diagnosis. Females if this species resembles the Mexican P. horni ( Redtenbacher, 1908) from Oaxaca and Guerrero and P. burkartii ( Saussure, 1868) from Veracruz. From horni , with they they share the long subgenital plate, they can easily be distinguished by the tubercular posterior portion of the vertex (smooth in horni ) and monochromatic head (most of dorsal portion whitish in horni ), much more numerous but on average smaller tubercles of the thoracic segments (less numerous and more spiniform in horni ), distinctly granulose to tubercular meso- and metasternum, differently shaped praeopercular organ, more prominent posterolateral lobes of abdominal tergum VII, presence of two dorsal lobes on the meso- and metatibiae and dorsally crested basitarsi (slender in horni ). From burkartii , with which this species has been confused previously ( Zompro, 2001a), they may readily be separated by the much longer subgenital plate (projecting beyond apex of the abdomen by no more than the length of the anal segment in burkartii ), presence of the distinct black, verrucose posteromedian swelling of the praeopercular organ on abdominal sternum VII (no such swelling in burkartii ), narrowed and trapezoidal posterior margin of the anal segment (broadened with the outer angles deflexed and obtusely rounded in burkartii ), smaller and rather conical pair of horns of the vertex and much more prominently tubercular vertex and thorax (vertex just weakly and sparsely granulose in burkartii ). Males are characteristic for the very short anal segment and elongated abdominal tergum IX, with the anal segment being less than half the length of tergum IX, and having the legs distinctly annulated. They strongly resemble ♂♂ of Ph. vosseleri ( Redtenbacher, 1908) but readily differ by the two features mentioned.

Etymology. Named after the distribution in Mexico. Masculine.

Description. ♀♀ ( Fig. 52B–C ): Medium sized to large species (body length including subgenital plate 168.0– 196.0 mm), shape average with a moderately globose, acutely bi-cornute head, a prominently tubercular to spinulose head and thorax, lobate legs and a fairly long, naviculate and obtuse to pointed subgenital plate, that projects over the tip of the abdomen by more than the two terminal abdominal terga taken together. General colouration ranging from grey over greyish ochre to drab (the holotype with a very faint olive wash) with all the thoracic armature dull yellow with the tips ochre; the larger tubercles with the tip reddish brown. Dorsal surface of the subgenital plate often dark brown to almost black. Eyes pale yellowish brown and flecked with sepia. Antennae drab and slightly darker brown ventrally, gradually darkening towards the apex (occasionally with a slight reddish hue).

Head: Moderately globose, about 1.2x longer than wide with the vertex weakly convex and with a pair of variably sized horns behind the eyes; these ranging from obtuse conical swellings to acute horns that project by almost half the height of head capsule, if developed as horns the anterior surface is obtusely keeled medially. Vertex unevenly supplied with nodes and tubercles of variable sizes, the largest of them being rather spiniform; usually one or two pair of moderately sized tubercles also present between the two horns. Genae nodulose with the nodes behind the eyes largest and those in the lower portion very small. Frons with a shallow indention between bases of antennae and a pair of weakly developed humps posteriorly. Eyes moderately sized, almost circular in outline, projecting almost hemispherical and their diameter contained about 2x in length of genae. Antennae slightly projecting over posterior margin of median segment. Scapus compressed dorsoventrally, roughly rectangular in dorsal aspect and 1.5x longer than wide. Pedicellus round in cross-section, somewhat inflated and globular and scarcely more than half the length of scapus.

Thorax: Pronotum slightly shorter and noticeably narrower than head, 1.7x longer than wide with a fairly distinct narrowing pre-medially; the posterior margin rounded. Surface with two longitudinal, sub-parallel rows of somewhat unevenly sized tubercles, that converge in the anterior half and slightly diverge in the posterior half of segment; otherwise supplied with a few rather small, scattered tubercles and notes. Anterior margin somewhat raised and set with a few tubercles, the median two of which are notably larger than all others. Transverse median sulcus distinctly impressed, almost straight and short, covering scarcely more than half of segments width. Mesothorax 7.4x longer than prothorax and uniform in width with only a slight posterior widening; metanotum somewhat less than one-third the length of mesonotum, rectangular in outline and roughly 3x longer than wide. Meso- and metanotum both with a shallow and obtuse medio-longitudinal keel and wholly and densely set with nodes and tubercles of variable sizes, the largest of which are acutely pointed and rather spiniform; those along the lateral margins roughly arranged in two sub-parallel longitudinal lines. Meso- and metapleurae also densely tubercular and with a longitudinal row of somewhat enlarged spiniform tubercles. Meso- and metasternum weakly tectate longitudinally and densely set with rather small, but unevenly sized tubercles that are clustered along the mid-line.

Abdomen: Median segment slightly shorter than metanotum, about 2.5x longer than width at posterior margin with the lateral margins gently concave; surface unevenly nodulose and tubercular but sculpturing somewhat less pronounced than that of metanotum. Segment II about as long as median segment, parallel-sided and about 2.2x longer than wide. II–IV increasing in length, IV–VI roughly equal in length and VII shorter than all preceding segments, being only about four-fifths the length of II; all roughly uniform in diameter and IV–V about 3.3x longer than wide. All terga supplied with a few scattered nodes that become gradually less in number and less pronounced towards the apex of abdomen. Tergum VII with lateral margins ± distinctly deflexed and rounded in the posterior half; the deflexion extending laterally at best by about half the width of segment. Sterna II–VII sparsely and minutely granulose and with a distinct longitudinal lateral carina close to lateral margins, that is most pronounced in VII. Praeopercular organ formed by a black, verrucose posteromedian swelling on sternum VII. Tergum VIII slightly more than half the length and somewhat narrower than VII, about 1.5x longer than wide, slightly trapezoidal in dorsal aspect and strongly convex longitudinally. IX about two-thirds the length of VIII, rectangular dorsal aspect and about 1.3x longer than wide. VIII and IX with a fairly distinct longitudinal lateral carina. Anal segment a little shorter than IX, parallel-sided with the posterior portion somewhat narrowed, carinate medio-longitudinally, the posterior margin almost straight with just a slight median indention and the outer angles rounded. Cerci small and tapered towards a blunt apex. Gonapophyses VIII not considerably elongated, weakly upcurved and only reaching to posterior margin anal segment. Subgenital plate variable in length and shape and projecting beyond apex of abdomen by at least the combined length of the two two two terminal segments; almost parallel-sided and keeled medio-longitudinally in the apical two-thirds and with the apex ranging from obtusely to narrowly triangular; in the basal one-third with a prominent, obtuse longitudinal lateral bulge.

Legs: All moderately long and slender. Profemora notably shorter than mesothorax, mesofemora about as long as metathorax, metafemora almost reaching to posterior margin of abdominal segment IV and metatarsi reaching to abdominal segment VII. Two outer ventral carinae of meso- and metafemora with a ± distinct, rounded to obtusely angular deflexion sub-basally and the medioventral carina with 4–5 small spines sub-apically. Meso- and metatibiae with posterodorsal carina forming two lobes that are very variable in size and range from an indistinct deflexion to a large, semi-circular, foliaceous lobe; one about one-third off the base and a smaller one about one-third before the apex. All basitarsi with a prominent, roundly triangular dorsal crest, that is most prominent and angular on probasitarsus; all slightly longer than corresponding three following tarsomeres taken together.

♂♂: The following description is mostly based upon the ♂ in the collection UMMZ because this is the most complete and least discoloured specimen.

Fairly small (body length estimated ca. 77.0–88.0 mm), moderately robust for the genus and apterous with a notably elongated abdominal tergum IX and abbreviated anal segment. Meso- and metathorax very minutely coriaceous and sparsely granulose, body surface otherwise smooth and slightly glossy. General colour of body ochre brown, most of meso- and metathorax with on olive wash. Head dark ochreous dorsally with a washed dark brown postocular stripe and the lower portion of the genae cream. Eyes reddish mid brown. Legs mid brown, all femora except for profemora and all tibiae with three pale annulae; the apical portion of all femora dark brown. Antennae reddish mid brown dorsally and rather dark brown ventrally.

Head: Ovoid, broadest at the eyes, gently narrowing towards the posterior and roughly 1.2x longer than wide; vertex very weakly rounded and smooth. Posterior margin with two small humps medially and between the eyes with two shallow, rounded swellings that are followed by a pair of obtuse and low granules. A transverse bulge between the bases of the antennae that is followed by a roundly triangular impression. Eyes large, circular in outline, projecting hemispherical and their diameter contained only 1.4x in length of genae. Scapus somewhat compressed dorsoventrally and oval in diameter, the lateral margins converging towards the base and in dorsal aspect and about 1.3x longer than wide. Pedicellus slightly globose, round in cross-section and only about half as long as scapus. III notably longer than pedicellus.

Thorax: Pronotum about as long and indistinctly narrower than head, the lateral margins weakly rounded with the median portion broadest and a slight narrowing pre-medially; posterior margin somewhat narrower than anterior margin; about 1.6x longer than maximum width. Surface with a slightly impressed medio-longitudinal line and a short but distinctly impressed, gently arched transverse median sulcus; anterior margin with a median pair of granules. Mesothorax 8x longer than prothorax and uniform in diameter with a slight widening at anterior margin and the posterior portion. Mesonotum very minutely coriaceous and very sparsely and weakly granulose, with a weakly indicated medio-longitudinal keel and along lateral surfaces with an obtuse and granulose longitudinal bulge parallel to lateral margin. Mesopleurae with a longitudinal row of minute granules, the mesosternum weakly tectate longitudinally and supplied with a few small, scattered granules. Metanotum basically like mesonotum and hardly one third of whose length of mesothorax; metasternum smooth.

Abdomen: Median segment 0.8x the length of metanotum, gently narrowed medially and with the posterior margin broader than anterior margin; about 3.5x longer than wide. Abdominal segment II scarcely longer than median segment and about 3.3x longer than wide. II–VI uniform in diameter, II–V roughly uniform in length, VI somewhat shorter than preceding and VII hardly more than two-thirds the length of VI and gently widening in posterior one third; II–V on average 4x longer than wide. Terga II–V each with a slight posteromedian swelling, II–VII with a very fine longitudinal lateral carina. Sterna smooth. Tergum VIII about two thirds the length of VII, wider than all preceding segments and trapezoidal in dorsal aspect with posterior margin 1.3x wider than anterior margin; lateral surfaces with an obtuse longitudinal bulge. IX 1.4x longer than VIII, notably constricted medially with the anterior portion wider than posterior margin and about 3x longer than width in median portion; the lateral margins almost straight and just weakly deflexed and angular posteriorly. Anal segment small, notably less than half the length of tergum IX, triangular in lateral aspect, roundly rectangular in dorsal aspect with a median indention posteriorly and the outer portions of posterior margin obtusely rounded and somewhat inflated; dorsal surface with an obtuse and low medio-longitudinal keel. Ventral surface of the rounded portions of posterior margin unevenly supplied with dark reddish brown denticles. Cerci of moderate length, gradually tapering towards a weakly club-like apex and gently incurved; projecting slightly beyond apex of anal segment. Vomer fairly broad, short and roundly triangular in shape with the lateral margins prominently inflated and bulgy and the central portion strongly impressed; upcurved in lateral aspect with a moderately long and pointed terminal hook. Poculum bulgy, strongly convex, angular in lateral aspect with a rather small and obtuse, laterally compressed protuberance at the angle and the vertical posterior portion carinate medio-longitudinally; the posterior margin weakly indented medially and roughly reaching half the way along tergum IX

Legs: All of moderate long and wholly unarmed. Profemora almost as long as head, pro- and mesothorax combined, mesofemora equal in length to mesothorax, metafemora reaching to posterior margin of abdominal segment V and metatibiae projecting considerably beyond apex of abdomen. Mesobasitarsi as long as, the pro- and metabasitarsi longer than the remaining corresponding tarsomeres taken together.

Variability. Females show considerable variability in size, the size of the cephalic pair of horns, degree of the thoracic armature, length and shape of the subgenital plate as well as the size and shape of the leg furniture. The holotype has the cephalic horns just weakly developed and merely represented as two rather obtuse, conical swellings. The paratype from Papachula in the collection of ZMUH has them notably more pronounced but represents and intermediate if compared to the three other paratypes from Soconusco (collection of ZMUH) and Chimaltenango (collection of MNHN), which all have them forming two large and conical, acutely pointed spines that project by almost half the height of the head capsule. One of the specimens in ZMUH even has them all over supplied with minute tubercles while they are somewhat peg-like with the base gently constricted in the other ZMUH specimen. The armature of the vertex and thoracic terga is strongly variable in means of density but also in the shape and size of the tubercles and spines. Noteworthy variability is also seen in the length and shape of the subgenital plate, which projects beyond the apex of the abdomen by at least the length of the two terminal abdominal terga taken together but may also project by more the combined length of terga VIII–X (specimens from Soconusco in ZMUH). These latter specimens also have the apex of the subgenital much more narrowed and acutely pointed than all other examined specimens. The paratype from Chimaltenango is remarkable for having the two dorsal lobes of the meso- and metatibiae much more strongly developed than all examined specimens and almost semi-circular in outline. These are least developed and almost absent on the metatibiae in the holotype, whereas the three specimens in ZMUH represent intermediate steps, the example from Tapachula having the sub-basal lobe weakly bifid .

Comments. Although meant to support future studies on the group and nomenclatural stability, the lectotype designation for Clonistria guatemalensis Redtenbacher, 1908 by Zompro & Brock (2003: 13) was very unlucky and in several aspects hard to understand. The original type series consists of three specimens all from Guatemala but which are two distinct species. The adult ♂ in the collecdtion of MHNG (now a paralectotype) perfectly matches with the original description and measurements provided by Redtenbacher (1908: 405) and is obviously the specimen on which Redtenbacher based his description. Hence, this specimen should actually have been selected as the lectotype. The other two immature specimens, a ♀ in MNHU and a ♂ in MHNG (the latter was not even listed by Redtenbacher (1908: 405)) are specifically and generically different from the ♂ in MHNG (see comments on Lanceobostra guatemalensis ( Redtenbacher, 1908) comb. n.). Now, the adult MNHG ♂ becomes a paratype of this new species. The specimen is strongly damaged and incomplete, lacking the entire antennae except for the right scapus, both pro- and metatarsi, most of the mesotarsi as well as abdominal segments IV–V. Hence, the description above is mostly based on the more complete ♂ in the collection of UMMZ. A further ♂ in the collection of ANSP labelled “ Costa Rica, Underwood?” apparently represents this new species but as already suggested by the question mark on the data label, the locality is most certainly wrong. It is discoloured to plain buff by provisional storage in spirits.

The terminalia of the ♀ and egg illustrated by Zompro (2001a) and misidentified as P. burkartii ( Saussure, 1868) were based on of the two ♀ paratypes from Soconusco, Chiapas in the collection of ZMUH. Thus, both the illustration and this record of P. burkartii actually refer to P. mexicanus sp. n. Unfortunately, the egg illustrated by Zompro (2001a, plate 5: 72–73) could not be examined because it appears to have not been returned from a loan by that author and was retained in the personal collection of O. Zompro (coll. OZ). The three ♀♀ in ZMUH have all been preserved in spirits for a considerable time and were dried subsequently, hence are discoloured to plain drab and have their abdomens shrivelled.

Table 41: Measurements of Phanocles mexicanus sp. n.

* Abdominal segments IV and V are missing (estimated to be ca. 85.0 mm)

Figures

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

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

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

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