Phanocles burkartii ( Saussure, 1868 )

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 : 167-170

Phanocles burkartii ( Saussure, 1868)

( Figs. 49A–B , 57F , 61A–C , 88A )

Bacteria burkartii Saussure, 1868: 65 .

Saussure, 1872: 151, pl. 3: 6 (♀).

Redtenbacher, 1908: 421.

Giglio-Tos, 1910: 40.

Passerin d’Entreves, 1981: 57.

Phanocles burkartii, Stål, 1875: 81 .

Kirby, 1904: 353.

Shelford, 1909: 364, pl. 7: 3 (♀).

Zompro, 2001a: 197 (in part—illustrated egg is Phanocles mexicanus sp. n.). Zompro & Brock, 2003: 8.

Otte & Brock, 2005: 262.

López-Mora & Llorente-Bousquets, 2018: 48.

Brock & Büscher, 2022: 513.

Luna, 2022: 93.

= Bacteria sartoriana Kaup & Heyden, 1871: 29 . [Synoynmized by Zompro, 2001a: 197] Kaup, 1871b: 18, pl. 1: 5 (egg).

Zompro, 2001a: 197. [As synonym of P. burkartii ( Saussure, 1868) ]

Zompro, 2001c: 142, fig. 19 (♀ HT).

Zompro, 2004c: 91. [As synonym of P. burkartii ( Saussure, 1868) ]

Otte & Brock, 2005: 262.

Phanocles sartoriana, Kirby, 1904: 354 .

Shelford, 1909: 364.

= Bacteria zehntneri Redtenbacher, 1908: 421 . syn. n.

Shelford, 1909: 364.

Brock, 1993: 21.

Phanocles zehntneri, Zompro & Brock, 2003: 26 .

Otte & Brock, 2005: 67.

Brock, Marshall, Beccaloni & Harman, 2016: 203.

López-Mora & Llorente-Bousquets, 2018: 48.

Luna, 2022: 93.

Further material examined [5 ♂♂, 5 ♀♀]:

MEXICO: 1 ♀: Coll. Br. v. W., St. Martha , Deyrolle; det. Redtenb., B. Burkartii Sauss. ; 73.; 5990 [ NHMW, No. 810] ; 1 ♀: Atoyac, Vera Cruz, Schumann. ; Godman-Salvin Coll. 1908.–168.; B.C.A. Orth. II, Bacteria burkartii Sauss. [ NHMUK] ; 1 ♂: Museum Paris, Mexique ,. Et. de. Vera Cruz, A. Génin 1921 [ MNHN] ; 1 ♀: Mexico, Veracruz, Zargalico, 22/V/92 [ OC] ; 1 ♂: Mexico, Veracruz, Santiago , Tuxtla, Cerro del Vigia, 12.VI.–9.IV.1994 [ OC] ; 1 ♂: Mexico, Veracruz, Santiago , Tuxtla, Cerro del Vigia, 9/VII/95 [coll. OC] ; 1 ♂: Mexico, Veracruz, Santiago , Tuxtla, Cerro del Vigia, 8/VIII/94 [ OC] ; 1 ♂: Mexico, Veracruz, Est. Biol. Los Tuxtla, 14.V.88 [ OC] ; 1 ♀: Mexico, Veracruz, Dos Amates, Catemaco , leg. D. Caroe, 15.VIII.1980 [ FH, No. 1218–1] .

NO DATA: 1 ♀: Bacteria arumatia Stoll Central-Amerika [ ZMUH] .

Diagnosis: This species, the type of Phanocles , is similar to several of the more southward distributed Central American species of the genus but apart from its very northern distribution easily recognised by a number of characteristic features. The morphologically most similar species are P. costaricensis Hennemann, 2002 from Costa Rica and P. rehni sp. n. from Honduras, the latter of which is only known from the ♀. While ♀♀ differ from those of costaricensis by the slightly less globose head and much more prominent cephalic horns (only a pair of shallow, obtuse swellings in costaricensis ), low posterior margin of the mesonotum (forming a tubercular transverse bulge in costaricensis ) and presence of two rounded dorsal lobes on the meso- and metatibiae, ♂♂ can be separated by the less globose head and unarmed head (with a small pair of conical tubercles in costaricensis ), flattened vertex, having the median segment ± equal in length to the metanotum (notably longer than the metanotum in costaricensis ), shorter cerci, less medially indented posterior margin of the anal segment and not annulated meso- and metafemora (pale cream basally with a distinct black transverse sub-basal band in costaricensis ). From rehni the ♀♀ of this species differ by the lack of the prominent peg-like, black posteromedian protrusion of the praeopercular organ and presence of two rounded dorsal lobes on the meso- and metatibiae. The most similar and possibly most closely related Mexican species is P. mexicanus sp. n. from Chiapas, which is only known from the ♀♀ and was confused with P. burkartii by Zompro (2001a; see below). Females of burkartii however may be separated by the much shorter subgenital plate, that projects beyond the apex of the abdomen by no more than the length of the anal segment ( Figs. 61A–C ; projecting by more than the combined length of the two terminal terga in mexicanus ); lack of the distinct black, verrucose posteromedian swelling of the praeopercular organ seen in mexicanus , broadened posterior portion of the anal segment ( Fig. 61B ), which has the outer angles deflexed and obtusely rounded (posterior margin not widened and rather trapezoidal in mexicanus ), on average larger, higher and peg-like cephalic horns and much less pronounced and less numerous tubercles of the vertex, and thorax (vertex all over set with distinct node-like tubercles in mexicanus ).

Description. ♂♂: Fairly large (body length 133.0– 170.6 mm) and slender for the genus with a median segment that is very scarcely shorter than the metanotum and an unarmed head; apterous. Body surface wholly smooth. General colour dark buff to drab, the meso- and metathorax and most of median segment dark olive-green. Dorsal surface of head with a bold, washed triangular white marking, whose three angles are positioned at the eyes and near the posteromedian end of vertex; lower portions of genae whitish. Membranes between the abdominal terga blackish brown, the lateral surfaces of abdominal tergum IX with a faint washed pale marking anteriorly and posteriorly. All tibiae with two very faint, washed pale grey bands.

Head: Ovoid, broadest at the eyes and notably narrowing towards the posterior with the vertex flattened and wholly unarmed; 1.25x longer than wide. Frons with two distinct, small impressions between the bases of the antennae and between yes with two very shallow, convex areas. Eyes large, circular in outline and their diameter contained 1.7x in length of genae. Antennae projecting over posterior margin of abdominal segment V. Scapus compressed dorsoventrally (more distinctly towards the base), elliptical in cross-section, rectangular in dorsal aspect and 1.4x longer than wide. Pedicellus cylindrical and about half the length of scapus.

Thorax: Pronotum about as long but notably narrower than head, roundly rectangular in basic shape with the anterior portion somewhat widened and the posterolateral angles rounded; almost 2x longer than wide. Transverse median sulcus shallow, weakly curved and almost expanding over entire width of segment. Mesothorax 10x longer than prothorax and uniform in diameter except for a slight widening at anterior margin and posteriorly. Mesonotum with a very weakly indicated medio-longitudinal line and with a fine, uneven longitudinal carina along lateral margins. Metanotum about 0.4x the length of mesonotum, slightly widened anteriorly and weakly constricted medially. Meso- and metasternum with a rather indistinct and obtuse medio-longitudinal carina.

Abdomen: Median segment scarcely shorter than metanotum, 7.5x longer than width at anterior margin and with the lateral margins notably concave. Segment II shorter than median segment, II–V almost equal in length, VI and VII decreasing in length but all roughly uniform in diameter; II–V on average 4.8x longer than wide, VII only 3.5x longer than wide. Tergum VIII three-fifth the length of VII, trapezoidal in dorsal aspect with posterior margin 1.7x wider than anterior portion. IX narrowed in the posterior half and about as long as VIII; the lateral margins moderately deflexed and rounded (in the anterior half in particular). Anal segment slightly shorter than IX, almost rectangular and about 1.4x longer than wide in dorsal aspect, slightly descendant towards the posterior and the dorsal surface with a fine medio-longitudinal carina; posterior margin somewhat widened, weakly indented medially and the outer angles broadly rounded. Ventral surface of outer posterior angles set with several distinct and acute teeth. Epiproct minute and wholly concealed under anal segment. Vomer very elongate and slender, broad at the base, almost parallel-sided in the median portion and with the apical portion narrowed towards upcurved terminal hook; entire ventral surface with a very distinct and deep longitudinal median fuurow and the lateral margins strongly inflated. Cerci small, much shorter than anal segment, gently arched and slightly club-shaped. Poculum large, bulgy, roundly angular in lateral aspect with the central portion obtusely protruded, the vertical posterior portion carinate, medio-longitudinally and the posterior margin triangular and slightly projecting beyond tergum IX.

Legs: All long but fairly stocky and wholly unarmed except for 1–2 minute sub-apical teeth on the medioventral carina of the meso- and metafemora. Profemora almost as long as pro- and mesothorax combined, mesofemora about as long as mesothorax, metafemora projecting somewhat over posterior margin of abdominal segment V and metatibiae projecting beyond tip of abdomen by almost the length of the five terminal abdominal segments taken together. Basitarsi all slender, compressed laterally with the dorsal carina lamellate but uniform in height; all longer than remaining tarsomeres combined.

Variability. The few ♀♀ examined show slight variability in the colouration, shape and size of the cephalic horns, number and size of the thoracic tubercles, shape of the two dorsal lobes of the meso- and metatibiae and shape of the dorsal crest of the metatibiae. The horns of the vertex and tubercles of the mesonotum in particular are most strongly developed in the ♀ from the Sierra de Santa Marta in the collection of NHMW ( Fig. 49A ) and least developed in the ♀ within the first author’s collection (coll. FH; Fig. 49B ), This latter specimen also has the sub-basal lobe or deflexion of the posteroventral carina of the meso- and metafemora less acute than all other specimens and more obtuse and just weakly rounded. The two dorsal lobes of the meso- and metatibiae are most pronounced in the specimen from Zargalico in the second authors collection (coll. OC). According to Saussure (1870: 151) the holotype has a body length of only 190.0 mm, which is notably shorter than the three other examined specimens (see table 34 below), except for the ZMUH specimen, which measures 198.0 mm. This latter specimen has the subgenital shorter than all other examined ♀♀ and just very scarcely projecting beyond the apex of the abdomen. Other than variation in size and slightly variable colouration noteworthy variability is not seen in ♂♂ .

Comments. Saussure (1870: 151, pl. 3: 6) provided a fairly detailed description of the ♀ along with an illustration. An illustration of the ♀ in the collection of NHMUK was presented by Shelford (1909: 364, pl. 7: 3b), which in combination with Saussure’s description is sufficiently detailed to resign from providing another description of the ♀ of P. burkartii at this place. Zompro (2001a: 198) was correct in stating that both, the drawings by Saussure and Shelford erroneously show the species without posterolateral lobes of abdominal tergum despite these are clearly mentioned in the description provided by Saussure (1870: 152) “ ...le [segment] 6e ayant ses bords latéroux gradullement dilatés en forme de lobes arrondis... ” (Saussure did not count the median segment as the first abdominal segment, thus referred to VI instead of VII) and present in the specimen in NHMW. Unfortunately, Saussure’s holotype has not been traced in the collection of MHNG ( Zompro & Brock, 2003: 8). Thus, it might become necessary to designate a neotype, once the loss of the holotype can be fully confirmed. The previously unknown ♂ is here described and illustrated for the first time.

The egg briefly characterised and sketched by Zompro (2001a: 198, plate 5: 72–73) was extracted from the ovipositor of a ♀ from Soconusco, Chiapas, Mexico in the collection of ZMUH. Examination of the specimen has shown this was misidentified and is not P. burkartii . The concerned specimen is now a paratype of P. mexicanus sp. n., why the illustration and diagnosis of the egg refer to this species instead. Kaup (1871b: 18, pl. 1: 5) provided a short description and illustration of the egg of his synoymous B. sartoriana . Although the illustrated egg was extracted from the abdomen of the holotype and is not fully developed, it shows a fairly typical egg for the genus with the typically punctured surface however not yet developed.

Distribution. There has been some confusion concerning to the distribution of this species. Based on the ♀ in the collection of NHMW, Redtenbacher (1908: 421) recorded P. burkartii from Colombia, because he misinterpreted the label data “Santa Martha”. Comparison with other specimens of P. burkartii and evaluation of all localities has shown that this relates to Sierra de Santa Marta in Veracruz, Mexico, but not Santa Marta in the Departemento Magdalena of Colombia. Shelford (1909: 364) copied this erroneous record and also recorded P. burkartii from the Valle de Chiriquí in W-Panama, based on an immature ♀ in the collection of NHMUK. Examination of the concerned specimen however, has shown this record to be based on a misidentification and the Panamanian specimen to be a distinct species of unknown identity. Also, the record “ Guatemala ” for the untraced immature ♀ paralectotype of the synoynmous B. zehntneri Redtenbacher, 1908 (syn. n.) in MHNG is doubtful and most certainly based on a not conspecific specimen. Since all definite records of P. burkartii restrict to the state of Veracruz along the central Caribbean coast of Mexico, this species is estimated to be restricted to the biogeographic region termed the Mexican Gulf Province ( Morrone, 2006: fig. 2).

Table 34: Measurements of Phanocles burkartii ( Saussure, 1868)

* According to Saussure (1870: 151); body length 192.0 mm according to Saussure (1868: 65)

** Metathorax (including median segment)

Figures

FIGURE 49. Phanocles Stål, 1875 habitus (in scale): A. P. burkartii (Saussure, 1868) ♀ (SE-Mexico), dorsal view [NHMW]; B. P. burkartii (Saussure, 1868) ♀ (Mexico, Veracruz, Dos Amates, Catemaco), dorsolateral view [coll. FH, No. 1218–1]; C. P. costaricensis Hennemann, 2002 ♀ (captive reared F1 from Costa Rica, Proc. Alajuela, Bosque del Niño N.P.), dorsolateral view [coll. FH, No. 0472–4]; D. P. spectabilis sp. n. ♀ PT (captive reared from Costa Rica, Prov. Puntarenas, La Gamba), dorsolateral view [coll. FH, No. 0563–5]; E. P. ploiaria (Westwood, 1859) ♀ (Panamá, Prov. Coclé, Valle deAntón 650–850 m), dorsolateral view [coll. FH, No. 0705–11].

FIGURE 57. Phanocles Stål, 1875 head and prothorax: A. P. acutecornutus sp. n. ♀ HT, dorsolateral view [MNHN]; B. P. berezini sp. n. ♀ PT, lateral view [coll. FH]; C. P. brevipes sp. n. ♀ HT, dorsolateral view (head is turned upside down) [ANSP]; D. P. barbadosense sp. n. ♀ PT, lateral view [ANSP]; E. P. muticus (Brunner v. Wattenwyl, 1907) ♀, dorsolateral view [coll. FH]; F. P. burkartii (Saussure, 1868) ♀, dorsolateral view [coll. FH]; G. P. saussurei (Redtenbacher, 1908) ♀, lateral view [coll. FH]; H. P. falcatus sp. n. ♀ PT, dorsolateral view [MNHN]; J. P. falcatus sp. n. ♀ PT, dorsolateral view [MNHN]; K. P. superbus sp. n. ♀ HT, dorsolateral view [MNHN].

FIGURE 61. Phanocles Stål, 1875 terminalia of ♀♀: A. P. burkartii (Saussure, 1868), lateral view [coll. FH]; B. P. burkartii (Saussure, 1868), dorsal view [coll.FH]; C. P.burkartii (Saussure, 1868), ventral view [coll. FH]; D. P.costaricensis Hennemann, 2002, lateral view [coll. FH]; E. P. costaricensis Hennemann, 2002, dorsal view [coll. FH]; F. P. costaricensis Hennemann, 2002, ventral view [coll. FH]; G. P. ecuadoricus sp. n. ♀ HT, lateral view [ZMUH]; H. P. ecuadoricus sp. n. ♀ HT, dorsal view [ZMUH]; J. P. ecuadoricus sp. n. ♀ HT, ventral view [ZMUH]; K. P. aequatorialis (Redtenbacher, 1908), lateral view [coll. FH]; L. P. aequatorialis (Redtenbacher, 1908), dorsal view [coll. FH]; M. P. aequatorialis (Redtenbacher, 1908), ventral view [coll. FH]; N. P. maroniensis (Chopard, 1911) ♀ HT, lateral view [coll. FH]; O. P. maroniensis (Chopard, 1911) ♀ HT, dorsal view [coll. FH]; P. P. maroniensis (Chopard, 1911) ♀ HT, ventral view [coll. FH]; Q. P. saussurei (Redtenbacher, 1908), lateral view [coll. FH]; R. P. saussurei (Redtenbacher, 1908), dorsal view [coll. FH]; S. P. saussurei (Redtenbacher, 1908), ventral view [coll. FH].

FIGURE 88. Mesosternum: A. Phanocles burkartii (Saussure, 1868) ♀ [coll. FH]; B. Phanocles maximus sp. n. ♀ PT [coll. FH]; C. Phanocles maximus sp. n. ♂ PT [coll. FH]; D. Phanocles mexicanus sp. n. ♀ HT [MNHN]; E. Phanocles ploiaria (Westwood, 1859) ♀ [coll. FH]; F. Phanocles ploiaria (Westwood, 1859) ♂ [coll. FH]; G. Phanocles subvolans (Westwood, 1859) ♀ [ANSP]; H. Phanocles ecuadoricus sp. n. ♀ HT [ZMUH]; J. Phanocles maroniensis (Chopard, 1911) ♂ [MNHN]; K. Phanocles satyr (Redtenbacher, 1908) ♀ [ANSP]; L. Phanocles saussurei (Redtenbacher, 1908) ♀ [coll. FH]; M. Phanocles superbus sp. n. ♀ HT [MNHN]; N. Phanocles spectabilis sp. n. ♂ HT [coll. FH]; O. Phanocles falcatus sp. n. ♂ HT [ANSP]. Not in scale.