Phanocles ploiaria ( Westwood, 1859 ) 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 : 197-201

publication ID

https://doi.org/ 10.11646/zootaxa.5444.1.1

publication LSID

lsid:zoobank.org:pub:5DE4A9DD-99F7-4E23-AD50-58DC491BB75E

persistent identifier

https://treatment.plazi.org/id/03FD87D9-FF64-D8AD-FF55-F6B32AEEE53C

treatment provided by

Plazi

scientific name

Phanocles ploiaria ( Westwood, 1859 )
status

comb. nov.

Phanocles ploiaria ( Westwood, 1859) comb. n.

( Figs. 4A View FIGURE 4 , 8B, 8D View FIGURE 8 , 49E View FIGURE 49 , 55E View FIGURE 55 , 58C View FIGURE 58 , 62K–M View FIGURE 62 , 67N–O View FIGURE 67 , 88E–F View FIGURE 88 , 93H View FIGURE 93 , 97H View FIGURE 97 , 99G View FIGURE 99 , 117C View FIGURE 117 , 118A–B View FIGURE 118 )

Phibalosoma ploiaria Westwood, 1859: 79 View in CoL , pl. 13: 4 (♂).

Cladoxerus ploiaria , 1904: 358.

Bacteria ploiaria, Redtenbacher, 1908: 416 View in CoL .

Shelford, 1909: 363, pl. 7: 3 (♂).

Hebard, 1923: 360.

Robinson, 1969: 291, fig. 10. [Defensive behaviour]

Berger, 2004: 18. [Ecology]

Otte & Brock, 2005: 66.

Simoens & Rabaey, 2009: 5, figs.

Harman, 2013: 13.

Brock, Marshall, Beccaloni & Harman, 2016: 189.

Bock & Büscher, 2022: 511.

[Not: Bacteria subvolans Redtenbacher, 1908: 416 → erroneous synonym of Hebard, 1923: 360]

Further material examined [65 ♂♂, 54 ♀♀, eggs]:

PANAMA: 2 ♂♂: Panamá, Panamá Pr., Parq. Nac. Soberanía , Camino Plantación, II-sep–1999, D. Quintero [ MIUP] ; 1 ♂: Panamá, Panamá Pr., Calzeda Largo , Chilibre, 21–feb-2002, S. Herkham [ MIUP] ; 2 ♀♀, 10 eggs: Panama, LP-02, Prov. Panamá, P.N. Soberanía, Camino de Plantación “Plantation Road” ca. 40 m, leg. F. Hennemann, 14.– 17.11.2019 (night); Alt.: ca. 40 m, 9°04’32.5”N 79°39’32.3”W, Moist lowland forest [ FH, No’s 0705–8, 9 & E3] GoogleMaps ; 1 ♀: Panama, LP-08, Prov. Coclé, Valle de Antón, Alto La Mesa , Hochplateau , 850–900 m, leg. F. Hennemann 16.– 17.11.2019 (night); Alt.: 850 m, 8°38’09.9”N 80°07’18.9”W, Mountainous rainforest [ FH, No. 0705–10] GoogleMaps ; 2 ♂♂, 1 ♀, 10 eggs: Panama, LP-06, Prov. Coclé, Valle de Antón, along Calle La Medina / La Mesa , nr. Río Guayabo 650–850 m, leg. F. Hennemann 14.– 17.11.2019 (night); Alt.: 650–850 m, 8°37’48.0”N 80°08’17.0”W, Mountainous rainforest [ FH, No’s 0705–11 to 13, E4] GoogleMaps ; 1 ♂: Barro Colorado Isl. , Panama, June 25, 1933, Hood, Hood & Hook; B. ploiaria ♂ (Westw.), Det. Hebard 1935 [ ANSP] ; 1 ♂: Muséum Paris, Barro Colorado , Dr. Gregoria, 1954 [ MNHN] ; 1 ♀, eggs: ex Zucht: F. Hennemann 2009, Herkunft: Panama, Gamboa , leg. T. & P. James, PSG No. 280 [ FH, No. 0705–1, E1, E2] ; 2 ♂♂: ex Zucht: Rob Krijns 2009, Herkunft : Panama, Gamboa , leg. T. & P. James, PSG No. 280 [ FH, No. 0705–2 & 3] ; 1 ♂: ex Zucht: K. Rabaey & R. Simoens 2010, Herkunft: Panama, Gamboa , leg. T. & P. James, PSG No. 280 [ FH, No. 0705–4] ; 1 ♂, 2 ♀♀, eggs: ex Zucht: F. Hennemann 2011, Herkunft: Panama, Gamboa , leg. T. & P. James, PSG No. 280 [ FH, No’s 0705–5 to 7] ; 18 ♂♂, 21 ♀♀, eggs: ex Zucht F. Hennemann 2020–2021, Herkunft: Panama, Prov. Coclé, Valle de Antón , Los Manadrinos Lodge , leg. S. Eilmus VII.2019 [ FH, No’s 0705–14 to 52 & E5] ; 5 ♂♂, 4 ♀♀, eggs: Panama: Prov. Coclé, Valle de Antón , 8º37’49.20”N 80º8’21.50”W, leg. O. Conle, F. Hennemann & P. Valero 17.09.2018 [ OC –0553–1 to 10] GoogleMaps ; 9 ♂♂, 9 ♀♀, eggs: Ex Zucht O. Conle, 2019, F1, Panama: Prov. Coclé, Valle de Antón , 8º37’49.20”N 80º8’21.50”W, leg. O. Conle, F. Hennemann & P. Valero 17.09.2018 [ OC –0553–11 to 29)] GoogleMaps ; 7 ♂♂, 8 ♀♀: Ex Zucht O. Conle, 2020, F2, Panama: Prov. Coclé, Valle de Antón , 8º37’49.20”N 80º8’21.50”W, leg. O. Conle, F. Hennemann & P. Valero 17.09.2018 [ OC –0553–30 to 44] GoogleMaps ; 10 ♂♂, 3 ♀♀, eggs: Ex Zucht O. Conle, 2021, F3, Panama: Prov. Coclé, Valle de Antón , 8º37’49.20”N 80º8’21.50”W, leg. O. Conle, F. Hennemann & P. Valero 17.09.2018 [ OC –0553–45 to 57] GoogleMaps ; 5 ♂♂, 2 ♀♀, eggs: Panama: Prov. Coclé, Valle de Antón , 8º37’49.20”N 80º8’21.50”W, leg. O. Conle & P. Valero 2– 3.07.2019 [ OC –0553–58 to 65] GoogleMaps .

Diagnosis. Females of this species resemble those of other Central American species of the genus, including Ph. costaricensis Hennemann, 2002 , Ph. spectabilis sp. n. and Ph. subvolans ( Redtenbacher, 1908) . From the first species they differ by the slenderer and elongate shape with the body segments proportionally longer, more prominent cephalad horns (only represented as obtusely rounded humps in costaricensis ), slightly less prominently tubercular meso- and metathorax, flat posterior margin of the mesonotum (with a transverse tubercular bulge in costaricensis ), much smaller posterolateral lobes of abdominal tergum VII and on average shorter subgenital plate, that projects beyond the apex of the abdomen by notably less than the length of the anal segment ( Figs. 62K–L View FIGURE 62 ). From the latter two species the ♀♀ of ploiaria can easily be separated by the pair of fairly distinct but short and obtusely conical cephalic horns ( Fig. 58C View FIGURE 58 ); from spectabilis sp. n. they additionally differ by the much slenderer and elongate shape and proportionally longer body segments and from subvolans they can also be differentiated by the rather rectangular anal segment ( Fig. 62L View FIGURE 62 ) and broader, much shorter subgenital plate (laterally compressed and projecting beyond apex of abdomen by almost the length of the two terminal terga taken together in subvolans ). Among the Central American representatives of the genus, the winged ♂♂ of this species are particularly similar to those of Ph. spectabilis sp. n. and Ph. subvolans (Redtenbacher) . From those of the first species they may be separated by the dark olive or brown mesothorax (green in spectabilis ), more pronounced and obtusely conical pair of cephalad tubercles, posteromedially less decidedly notched and more flattened anal segment, much shorter and broader vomer ( Fig. 97H View FIGURE 97 ; very elongate, slender and almost parallel-sided in spectabilis ) and noticeably smaller and less bulgy poculum. From the ♂♂ of Ph. subvolans they readily differ by the much longer alae that almost reach to the posterior margin of abdominal segment III ( Fig. 55E View FIGURE 55 ; just slightly projecting over posterior margin of median segment in subvolans ), much less tectate and posteriorly flattened anal segment and more distinct cephalad tubercles.

Description. ♀ ( Fig. 49E View FIGURE 49 ): The colouration is described from live captive reared specimens and colour photographs of live wild specimens from different localities.

Variable in size, rather small to moderately large (body length including subgenital plate 148.0–205.0 mm) and slender for the genus with a globose, obtusely bi-cornute head ( Fig. 58C View FIGURE 58 ), an unevenly tubercular thorax and a fairly short subgenital plate, that projects beyond the tip of the abdomen by no more than the length of the anal segment ( Figs. 62K–L View FIGURE 62 ). General colour variable and ranging from pale buff or greyish ochre over various shades of brown to almost black; most of abdominal tergum IV pale cream to straw. Occasionally with a slight olive wash and greenish marbling in paler specimens. Granules and tubercles of the thorax dark ochre or olive and tipped with black. Eyes yellowish green and irregularly flecked with dark brown. Antennae drab dorsally and somewhat darker ventrally.

Head ( Fig. 58C View FIGURE 58 ): Moderately globose with the vertex roundly convex, broadest at the eyes and the genae gently narrowing towards the posterior; about 1.2x longer than wide. Just behind the eyes with a pair of moderately prominent and obtusely conical, somewhat forward directed horns. Vertex and genae minutely and sparsely granulose. Frons with a distinct C–shaped impression behind the bases of the antennae and two low swellings between the eyes. Eyes rather weakly convex, roughly circular in outline and their diameter contained almost 1.8x in length of genae. Antennae reaching about half the way along median segment. Scapus compressed dorsoventrally, roundly rectangular in dorsal aspect but slightly narrowing towards the base with the interior margin gently rounded and about 2x longer than wide. Pedicellus round in cross-section, constricting towards the apex and roughly half the length of scapus.

Thorax: Pronotum about as long and scarcely narrower than head, 1.5x longer than wide and roundly rectangular in dorsal aspect with the lateral margins gently concave pre-medially. Anterior margin somewhat raised and with a median pair of small tubercles. A shallow pair of median tubercles just in front of the median sulcus and two slightly diverging longitudinal rows of small tubercles in posterior half; the surface otherwise rather unevenly supplied with granules and small node-like tubercles ( Fig. 58C View FIGURE 58 ). Transverse median sulcus moderately impressed, arched and expanding by little more than half the width of segment. Mesothorax 5.5x longer than prothorax, slender and uniform in diameter with just a slight expansion posteriorly. Metanotum one-third the length of mesonotum, 2.2x longer than wide and roughly rectangular in outline. Meso- and metanotum all over densely but unevenly supplied with tubercles and nodes of variable sizes which are only fewer along the medio-dorsal line.; meso- and metapleurae irregularly tubercular and nodose. Meso- and metasternum with somewhat elongate, tubercles in the median portion, that are roughly arranged in two closely spaced longitudinal rows on mesosternum and both with a distinct granulose carina close to lateral margins; the median tubercles often dull orange in colour ( Fig. 88E View FIGURE 88 )

Abdomen: Median segment about 1.6x longer than metanotum, notably constricted medially and 3.5x longer than width of anterior margin; surface sparsely and irregularly tubercular. Following segments just sparsely set with scattered granules that become less in number towards the apex of abdomen and II–VI with a distinct, granulose longitudinal carina close to lateral margins. Segment II slightly shorter than median segment and 2.3x longer than wide. II–V slightly increasing and VI–VII decreasing in length; V longest and about 3x longer than wide. II–V almost uniform in diameter, VI and VII somewhat narrowing. Tergum VII with lateral margins almost entirely straight and just very weakly deflexed and rounded posteriorly ( Fig. 62K View FIGURE 62 ). Sterna II–VII sparsely and minutely granulose and with a fine longitudinal lateral carina. Praeopercular organ ( Figs. 4A View FIGURE 4 , 93H View FIGURE 93 ) formed by a rounded, almost semi-circular elevation of the two lateral carinae near posterior margin of sternum VII. Terga VIII–X uniform in width and as wide as VII; VIII slightly more than half the length of VII, 2x longer than wide; IX somewhat more than half the length of VIII, rectangular in dorsal aspect and scarcely wider than long. VIII and IX both with a weakly indicated medio-longitudinal line, the lateral margins straight and not notably deflexed. Anal segment as long as IX, roughly rectangular and carinate medio-longitudinally, the posterior margin with a very shallow median emargination and the outer angles broadly rounded ( Fig. 62L View FIGURE 62 ). Epiproct very small, triangular and almost fully concealed under anal segment. Cerci very small, oval in cross-section, obtuse and slightly gradually tapering towards the apex. Gonapophyses VIII moderately elongated, upcurved in the apical portion and roughly reaching to tip of subgenital plate; apex fairly pointed ( Figs. 4A View FIGURE 4 , 62 View FIGURE 62 KL). Subgenital plate obtusely keeled longitudinally with the apex bluntly triangular and projecting beyond tip of abdomen by no more than length of anal segment ( Figs. 62K–L View FIGURE 62 ); basal one-third with a longitudinal bulge laterally ( Fig. 62M View FIGURE 62 ).

Legs: All of moderate length, profemora slightly shorter than mesothorax, mesofemora notably longer metathorax, metafemora reaching about two-thirds the way along abdominal segment IV and metatibiae roughly reaching to posterior margin of abdominal segment VI. Two lower outer carinae of meso- and metafemora ± expanded and forming a variably sized, obtusely triangular or sometimes weakly bifid lobe sub-basally ( Figs. 67 View FIGURE 67 N- O; much less pronounced on metafemora). Posterodorsal carina of meso- and metatibiae with a rounded lobe at the apex. Dorsal carina of all basitarsi with a large, roundly triangular lobe. Probasitarsus almost as long as remaining tarsomeres taken together except for the claw ( Fig. 8D View FIGURE 8 ); meso- and metabasitarsus a little longer than combined length of following three tarsomeres.

Variability. This species shows considerable variability in size depending on the locality. Specimens collected and reared from lowland habitats in the Canal Zone (Parque Nacional Soberanía) are notably smaller than specimens collected and reared from mountainous habitats in Valle de Antón at 650–900 metres. The size ranges (body length including subgenital plate) are 148.0–171.0 mm for the lowland specimens and 167.0–205.0 mm for the highland specimens. The same is observed in ♂♂ but the size differences are less striking between the two populations with lowland specimens usually below and highland specimens over 100.0 mm in body length. No other frequent morphological differences are however to be observed between these two populations. Females reared from Valle de Antón show interesting colour variability ranging in general colour from pale buff or greyish ochre to almost black with lighter specimens usually more distinctly flecked with darker tones and sometimes with olive or greenish tones. Occasionally, specimens may have a dark brown medio-longitudinal streak indicated along the dorsal body surface and more rarely may have the largest meso- and metathoracic tubercles distinctively marked by white dots. Specimens from both populations almost always have most of abdominal tergum and sternum IV pale cream to straw and occasionally a pale central marking also occurs on abdominal tergum III. No such chromatic variability is seen in ♂♂ which are more consistent and just vary in being of a more light or darker general colour, which ranges from pale olive drab to dark brown.

Egg ( Fig. 99G View FIGURE 99 ). Moderately sized for the genus and of fairly typical shape. Capsule angular and about 1.3 longer than high with the polar-area slightly impressed and the dorsal egg surface prominently convex and rectangular in the anterior portion; oval in cross-section. Whole capsule surface deeply and densely punctured, the pits fairly large and sharply defined but inhomogeneous in size and shape. Micropylar plate elongate, very slightly widening towards the posterior, about four-fifths the length of capsule and roughly 3.5x longer than width in the median portion; sculptured like capsule but the pits comparatively smaller and with the central portion obtusely bulgy longitudinally. Micropylar cup a small node with a short bulge anteriorly. Median line very short but reaching to outer margin of impressed portion of polar area. Operculum oval and with a roundly convex, hollow excrescence that is formed by the outer margin; this with a circular row of radial impressions in dorsal half, which in centre unite to form an irregularly defined crater-like central impression; height about one-fifth of capsule length. Colour of capsule plain grey with a variably shaped dark grey marking on each lateral surface and the polar-area with a dark grey central spot; opercular collar dark greyish brown. Most of micropylar plate blackish, the outer margin pale grey or cream. Operculum excrescence dull orange to reddish mid brown.

Measurements [mm]: Length including operculum 3.8–4.0, length 3.1–3.3, width 2.0–2.1, height 2.5–2.8, length of micropylar plate 2.4–2.7.

Remarks: The previously unknown eggs are here formally described for the first time. A brief description of the ♀ was provided by Hebard (1923: 360), but the concerned specimen is the previously unknown ♀ of Ph. subvolans ( Westwood, 1859) , which has erroneously been synonymised with Ph. ploiaria by Shelford (1909: 263) but is reinstated herein (stat. rev.). Brief characterisations of both sexes and the eggs were presented by Simoens & Rabaey (2009: 6), who provided information on the captive breeding of Ph. ploiaria . A formal description of the ♀ is therefore presented above. Berger (2004: 18) recorded this species from Barro Colorado Island and provided information on its dietary habits and stated that 34% of individuals were found on plants of the family Rhamnaceae (e. g. Gouania lupuloides ) and exclusively along forest edges. The observation that in captivity also representatives of the plant families Bignoniaceae , Melastomaceae , Sapindaceae , Sterculiaceae and Tiliaceae were accepted ( Berger, 2004: 22) suggest Ph. ploiaria to be rather polyphagous. This is supported by the wide range of alternative food plants that are accepted in breeding conditions in Europe (see below).

Captive breeding is fairly easy in large, well-ventilated cages but high humidity. At average temperatures of 22–25°C eggs hatch after 4–5 months and nymphs reach maturity after 6–7 months. Females lay an average of 5– 6 eggs per day and simply flick these away by an abrupt movement of the abdomen. These sturdy insects can reach almost one year in age meaning that a female can easily produce more than 1000 eggs in a lifetime. Males are notably more short-lived and hardly reach six months of age. Mating usually takes place at night and as typical for Phanocles -species both sexes but also the nymphs show very hectic behaviour when disturbed and handled. Females in particular usually drop to the ground and hectically tumble around to finally walk away. Alternative fooD–plants that are frequently accepted in captivity in Europe include bramble ( Rubus spp. , Rosaceae ), raspberry ( Rubus idaeus , Rosaceae ), wild roses ( Rosa spp. , Rosacea), hazel ( Corylus avellana , Betulaceae ), oak ( Quercus robur, Fagacceae ) and beech ( Fagus sylvatica , Fagaceae ).

Table 43: Measurements of Phanocles ploiaria ( Westwood, 1859) comb. n.

FH

Fort Hays

ANSP

Academy of Natural Sciences of Philadelphia

MNHN

Museum National d'Histoire Naturelle

OC

Oberlin College

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Phasmida

Family

Diapheromeridae

Genus

Phanocles

Loc

Phanocles ploiaria ( Westwood, 1859 )

Hennemann, Frank H. & Conle, Oskar V. 2024
2024
Loc

Bacteria ploiaria

Redtenbacher, J. 1908: 416
1908
Loc

Phibalosoma ploiaria

Westwood, J. O. 1859: 79
1859
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