Phanocloidea muricata ( Burmeister, 1838 )

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 : 249-255

Phanocloidea muricata ( Burmeister, 1838)

( Figs. 4E , 69B–C , 72J–K , 73E , 74A–C , 75A–C , 87B–C , 91F , 96A , 100L )

Bacteria muricata Burmeister, 1838: 564 .

Westwood, 1859: 179.

Redtenbacher, 1908: 418.

Chopard, 1911: 345.

Moxey, 1972: 176.

Lelong, 1993: 12.

Dyme muricata, Kirby, 1904: 350 .

Phanocloidea muricata, Zompro, 2001: 196 .

Otte & Brock, 2005: 263.

Zompro, 2005: 272.

Delfosse, 2009: 6.

Harman, 2012: 17.

Araujo & Garraffoni, 2012b: 233.

Jourdan, Lelong & Bellanger, 2014: 489. Bellanger, Lelong & Jourdan, 2018: 274.

Conle, Hennemann, Bellanger, Lelong, Jourdan & Valero, 2020: 12.

Brock & Büscher, 2022: 514.

= Phasma (Bacteria) canna Haan, 1842: 101 . syn. n.

Bacteria canna, Westwood, 1859: 21 .

Stheneboea canna, Kirby, 1904: 324 .

Prisomera canna, Bragg, 1996: 110 .

Brock & Büscher, 2022: 529.

Prisomera cannum, Otte & Brock, 2005: 283 .

= Bacteria divergens Redtenbacher , 19978: 418. [Synonymised by Zompro & Brock, 2003: 11]

Lelong, 1993: 12.

Otte & Brock, 2005: 263.

Delfosse, Cliquennois, Depraetere & Robillard, 2019: 199.

Phanocloidea divergens, Zompro, 2001: 196 .

= Dyme dreyfusi Brunner v. wattenwyl, 1907: 324 . syn. n.

Lelong, 1993: 11.

Bacteria dreyfusi, Otte & Brock, 2005: 63 .

Delfosse, Cliquennois, Depraetere & Robillard, 2019: 200.

Brock & Büscher, 2022: 510.

= Bacteria imitans Redtenbacher, 1908: 419 . [Synonymised by Hennemann & Conöle, 2003: 6]

Hennemann, Gehler & Conle, 1995: 437.

Hennemann & Conle, 2003: 6.

Otte & Brock, 2005: 263.

Zompro, 2005: 5.

= Bacteria nodulosa Redtenbacher, 1908: 416 . syn. n.

Shelford, 1909: 362.

Moxey, 1972: 176.

Hennemann, Gehler & Conle, 1995: 437.

Brock, 1998: 46.

Zompro, 2000: 177, figs. (in part—illustrated Venezuelan specimens are Phanocloidea venezuelica sp. n.). Phanocloidea nodulosa, Zompro, 2001: 196 (in part).

Hennemann & Conle, 2003: 6.

Otte & Brock, 2005: 263.

Zompro, 2005: 7.

Delfosse, Cliquennois, Depraetere & Robillard, 2019: 218.

Brock & Büscher, 2022: 514.

= Bostra reducte-dentata Redtenbacher, 1908: 409 (in part—only LT). syn. n.

Bostra reductedentata, Otte & Brock, 2005: 73 .

Brock & Büscher, 2022: 511.

Phanocloidea reductedentata, Hennemann, Conle & Brock, 2023: 182 . [Designation of lectotype]

= Bacteria rubispinosa Audinet-Serville, 1838: 224 . syn. n.

Westwood, 1859: 224.

Phibalosoma rubispinosa, Kirby, 1904: 356 .

Redtenbacher, 1908: 427.

Lelong, 1993: 12.

Cladomorphus rubispinosus , otte & Brock, 2005: 95.

= Bacteria rufopectus Redtenbacher, 1998: 416 . syn. n.

Chopard, 1911: 343.

Lelong, 1993: 11.

Phanocloidea rufopecta, Zompro, 2001: 196 .

Zompro, 2003: 21.

Otte & Brock, 2005: 263.

Brock & Büscher, 2022: 514.

= Bacteria sakai Bates, 1864: 332 , pl. 44: 1 (♀). syn. n.

Redtenbacher, 1908: 418.

Otte & Brock, 2005: 66.

Brock & Büscher, 2022: 511.

Dyme sakai, Kirby, 1904: 350 .

[Not: Phanocloidea nodulosa, Harman, 2012: 16 . → This is Phanocloidea venezuelica sp. n.).

Further material examined [35 ♀♀, 49 ♂♂, 40 nymphs]:

BRAZIL: 1 ♀ (in alcohol): Brasilien , Amazonien , Igarape , Cachoeira, Rio Cuieras, 20. IV. 1961 [ ZSMC] ; 1 ♀: Unt. Amazonas Tabajos , Mte. Christo; Brasilien, J. Arp ded., Eing. Nr. 145, 1933; PHA 88, Zoologisches Museum Hamburg [ ZMUH] ; 1 ♂: Unt. Amazonas , Rio Tabajos, Marituba; Brasilien, J. Arp ded., Eing. Nr. 145, 1933; PHA 92, Zoologisches Museum Hamburg [ ZMUH] ; 1 ♀: Unt. Amazonas , Obidos; Brasilien, J. Arp ded., Eing. Nr. 145, 1933; PHA 89, Zoologisches Museum Hamburg [ ZMUH] ; 1 ♂: Unt. Amazonas , Taperinha; Brasilien, J. Arp ded., Eing. Nr. 145, 1933; PHA 90, Zoologisches Museum Hamburg [ ZMUH] ; 1 ♀: Brasil , Obidos (Traira), Est. Pará, Dirings, Out 1961 [ MZUSP] ; 1 ♀: Brasil , Camargo , Mun. de Itaituba ( Rio Tapajoz ), Est. Pará, Dirings; Abr 1964 [ MZUSP] ; 1 ♂: Brasil , Itaituba ( Rio Tapajos ) Estado Pará, Dirings; Abr 1961 [ MZUSP] ; 1 ♀: Santaremsinho , Itaituba—Pará, Rio Tapajoz, III/64, Acrydoidea [ MZUSP] ; 1 ♂: Canindé , Rio Garupí, Poará, IV.1963, Malkin & Pinheiros col. [ MZUSP] ; 1 ♀: Mangabeìra , Mocajuba , Para, Brasil, XI–1952, Orlando Rego [ UFPR] .

FRENCH GUIANA: 1 ♂, 1 ♀: Museum Paris, Guyane Française, P. Grandcolas rec., Montagne de Kaw , Piste PK 36, 23.IX.1989 [ MNHN] ; 1 ♀: Museum Paris, Guyane francs., Dr. Borgrarid 1913 [ MNHN] ; 1 ♀: Museum Paris, Cayenne, Roux 255–36 [ MNHN] ; 3 ♀♀: Guyane Franc., St-Laurent du Maroni, Collection Le Moult ; Octobre, Mus. Paris, Coll. Lucien Chopard [ MNHN] ; 1 ♀ (subadult): Museum Paris, Cayenne, H. Deyrolle 1891 [ MNHN] ; 1 ♀, 1 ♂: Guyane, Route du Nouveau Chantier, St. Laurent , 15.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN] ; 3 ♀♀, 1 ♂, 1 nymph: Saûl, Guyane, Roubaud [ MNHN]; 8 nymphs: Saûl, Guyane, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN] ; 4 ♀♀, 17 ♂♂: Guyane, Petit Saut, 20.–31.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN]; 8 nymphs: Regina , en forêt, 9.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN]; 3 nymphs: St. Laurent du Maroni , 13.–19.VIII.92 [ MNHN]; 1 nymph: Guyane, St. Laurent du Maroni, 13.–18.VII.92, Roubaud rec. [ MNHN] ; 1 ♂: Piste du Plateau des Meno , St. Jean du Maroni, 13.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN] ; 1 ♂, 4 nymphs: Montagne de Kaw , 16.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN] , 2 ♀♀: Piste de Kaw, PK 36, 5.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN] ; 10 ♂♂ (nymphs): Piste de Kaw , 5.VIII.92, Roubaud, Rarchaert, Morin, Auvray rec. [ MNHN]; 4 nymphs: without data [ MNHN] ; 1 ♂: 620 85, Cayenne, Prudhomme [ MHNG] ; 1 ♀: Guyane Franc. , St-Laurent du Maroni, Collection Le Moult,; Décembre; Bacteria muricata Burm . ♀ [ ANSP] ; 1 ♂: Guyane Franc. , St-Laurent du Maroni, Collection Le Moult; Décembre; Bacteria rufopectus Redt . ♂. [ ANSP] ; 3 ♀♀, 2 ♂♂, eggs: Französisch Guyana , Montagnes de Roura, Eskol Route, 50 m, 4°42´N, 52°17´W, leg. Hennemann & Conle, 19. & 29. IX. 2001 [ FH 0473–1 to 3 , 19 & 20] ; 1 ♂: Französisch Guyana , Umgebung Matoury, 30 m, 4°51´N, 52°21´W, leg. Hennemann & Conle, 21 & 22. IX. 2001 [ FH 0473–11 ] ; 2 ♀♀, 4 ♂♂, 6 ♀♀, 1 ♂ (nymphs): Französisch Guyana , Montagne Grand Matoury, 10 m, 4°53´N, 52°20´W, leg. Hennemann & Conle, 20., 24. & 30. IX. 2001 [ FH 0473–4 , 7 , 15 , 17 , 18 , 21 , 24 to 30 ] ; 1 ♀, 1 ♂ (in Copula), 1 ♂: Französisch Guyana , Umgebung Cacao, Dégrad Menado Route, 50 m, 4°35´N, 52°24´W, leg. Hennemann & Conle, 23. IX. 2001 [ FH 0473–5 & 10] ; 3 ♀♀ (nymphs), 3 ♂♂: Französisch Guyana , Montagnes de Roura, Piste de Kaw, 200 m, 4°38´´N, 52°16´W, leg. Hennemann & Conle, 26. IX. 2001 [ FH 0473–6 , 8 , 12 , 14 , 16 & 23] ; 3 ♂♂: Französisch Guyana , Montagnes de Kaw, Piste de Kaw, 250 m, 4°34´N, 52°11´W, leg. Hennemann & Conle, 20., 27. & 29. IX. 2001 [ FH 0473–9 , 13 & 22] ; 2 ♀♀, 4 ♂♂, eggs: ex Zucht : F. Hennemann, urspr.: Französisch Guayana, F 1– Generation 2002 [ FH 0473–31 to 36 & ED] .

GUYANA: 1 ♀, 2 ♂♂: Demerara, II –III.04, R. Haensch; Bacteria rufopectus Redt. , det. H. Dohrn [ ZMPA] ; 1 ♀: Niederl. Guyana, Distr. Para, V. 08, C. Heller leg. Vend 4.VIII.1908; PHA 93, Zoologisches Museum Hamburg [ ZMUH] .

Diagnosis. Females of this species ( Fig. 69B ) resemble P. globocephala Conle et al., 2011 , P. turgida ( Westwood, 1859) comb. n. and P. venezuelica sp. n. in general appearance but readily differ from all of these by the distinctly tubercular to heavily spinose mesonotum ( Figs. 72 J-K), dorsal lobe of the basitarsi and on average larger size (body length including subgenital plate> 160.0 mm). Males are very similar to those of the Venezuelan P. semiptera sp. n. in many aspects, but differ at first glance by the lack of wings and slenderer shape ( Fig. 96C ). From the similar looking ♂ of P. venezuelica sp. n. they can be separated by the longer metanotum, that is more than half the length of the median segment ( Fig. 73K ; only about one-third the length of median segment in venezuelica ), larger lateral lobes of abdominal tergum IX ( Fig. 75A ), emarginate and bidentate posterior margin of the poculum ( Fig 75C ; obtusely triangular in venezuelica ), slenderer, elongate and arcuate vomer ( Fig. 96A ) and different colouration.

Description. ♀♀ ( Fig. 69B ): Large to very large (body length including subgenital plate 160.0–210.0 mm) and slender species with a median segment that is slightly longer than the metanotum, moderately globose and unarmed head, strongly tubercular to spinose mesonotum and a subgenital plate that just slightly projects over the tip of the abdomen. Surface of head very sparsely and of thorax densely but very minutely granulose. Colouration variable and ranging from light or dark green over straw, ochre and drab to various tones of grey and mid to dark brown; occasionally with numerous darker speckles or several bold white markings on the thorax and abdomen. Mesothoracic armature dark red to brown, the granulations of the head and thorax usually cream to white. Antennae greenish brown in green and brown in straw or brown specimens, ventral surface of all antennomeres except for scapus und pedicellus black.

Head ( Figs. 72J–K ): Sub-globose, about 1.25x longer than wide, broadest at eyes and notably narrowing towards the posterior; vertex weakly convex and smooth except for a few scattered minute granules. Frons with a small but distinct trapezoidal impression between the bases of the antennae. Between the eyes with shallow and smooth swellings. Eyes almost circular, moderately projecting from head capsule and their diameter contained about 1.7x in length of genae. Antennae almost reaching to posterior margin of abdominal segment III. Scapus almost 2x longer than wide, compressed dorsoventrally and rectangular in dorsal aspect. Pedicellus slightly oval in cross-section, weakly narrowing towards the apex and constricted at the base with the median portion somewhat inflated, about about 0.6x the length of scapus. III notably narrower than pedicellus.

Thorax: Pronotum about as long as head but notably narrower, rectangular in outline and 1.7x longer than wide; posterior margin rounded; the anterolateral angles each with a shallow triangular impression. Transverse median sulcus moderately distinct, weakly curved and almost reaching to lateral margins of segment; surface otherwise minutely granulose and with the longitudinal median line slightly impressed ( Figs. 72J–K ). Mesothorax cylindrical and uniform in diameter with only the extreme posterior portion gently widened and 8.3x longer than pronotum. Mesonotum armed with a variable number (7–40 in total) of obtuse conical to spiniform tubercles or spines of variable sizes, the largest of which may be somewhat arched and anteriad directed ( Figs. 72J–K ); these unevenly dispersed throughout entire surface. Mesopleurae with a longitudinal row of 9–14 pointed tubercles or small spines. Metanotum about one-third the length of mesonotum, parallel-sided and about 4x longer than wide; unarmed. Metapleurae often with a longitudinal row of pointed tubercles. Meso- and metasternum minutely but densely granulose, the mesosternum weakly tectate longitudinally ( Fig. 87B ).

Abdomen: Median segment slightly longer than metanotum, almost 5x longer than wide with lateral margins gently concave. Segment II about three-quarters the length of median segment. II–V gradually increasing in length, VI as long as V and VII shorter than all preceding segments; II–III weakly widening and VI–VII narrowing, on average all 2.8x longer than wide. Terga smooth, but occasionally a more or less prominent posteromedian tubercle present on V. Sternum VII with a prominent praeopercular organ at posterior margin, that is formed by two fairly large, obtusely rounded, transverse lobes ( Fig. 91F ). Tergum VIII half the length of VII and widening towards the posterior; IX notably shorter, strongly convex and slightly wider than long. All terga with a fine, granulose longitudinal carinae close to lateral margins. Anal segment slightly longer than IX, with a fine but acute medio-longitudinal carina, about as long as wide and with the posterior margin somewhat widened; the posterior margin with a shallow concave median excavation and the outer angles deflexed and obtusely rounded ( Fig. 74B ). Epiproct very small fully concealed by anal segment. Gonapophyses VIII elongate, slender, gently upcurved and projecting by no more than length of anal segment ( Fig. 74A ). Subgenital plate keeled longitudinally, parallel-sided with the apex rounded to obtusely triangular ( Fig. 74C ); length variable and projecting over tip of abdomen by at least half the length of anal segment but no more than combined length of the terminal two terga.

Legs: All very long and slender, profemora about as long as mesothorax, mesofemora longer than metanotum and median segment combined and hind legs slightly projecting beyond apex of abdomen. All unarmed except for the occasional occurrence of a more or less distinct blunt tooth or triangular lobe about one-quarter the way from the base of the meso- and metafemora. Medioventral carina of meso- and metafemora rather distinct; unarmed. Dorsal carinae of meso- and metatibiae weakly rounded apically. Probasitarsus as long as remaining tarsomeres combined, meso- and metabasitarsus slightly shorter than remaining tarsomeres except claw combined; dorsal carina forming a ± distinct, rounded to obtusely triangular lobe.

♂♂ ( Fig. 96C ): Medium-sized to large (body length 114.0– 131.5 mm) and very slender with a median segment that is notably longer than the metanotum, large triangular lateral lobes of abdominal tergum IX and characteristic colouration; apterous. Body surface smooth. General colour of the body and head medium to dark creamish brown, the abdomen usually of a somewhat lighter tone than head and thorax and often with a slight orange wash. Head uniformly dark orange ( Fig. 73E ). Mesonotum with two elongate, longitudinal, parallel black streaks, that run almost the complete length of the segment except for the very anterior and posterior portions; the narrow space between these markings reddish brown and the lateral surfaces dark green ( Fig. 73K ). Meso- and metapleurae and most of median segment dark green. Meso- and metasternum yellow and with a broad medio-longitudinal red line ( Fig. 87C ). All abdominal terga with a pale cream-coloured to whitish longitudinal line along lateral margins. Basal half of profemora red; apical half brown with a pale transverse band near the apex. Meso- and metafemora greenish straw basally and becoming dark brown towards the apex; with a faint pale transverse median band and a more distinct band near the apex. Tibiae mid brown with two very faint and slightly lighter brown annulae. Antennae brown with ventral surfaces of all antennomeres except for scapus and pedicellus black.

Head ( Fig. 73E ): Sub-globose, indistinctly longer than wide, broadest at eyes and strongly narrowed towards the posterior. Between the eyes with a pair of shallow swellings; surface entirely smooth and otherwise as in ♀♀. Eyes large, circular, strongly convex and projecting hemispherical from head capsule; their diameter contained 1.4x in length of genae. Antennae projecting over posterior margin of abdominal segment V, otherwise like in ♀♀ but scapus almost 1.5x longer than wide.

Thorax: Pronotum slightly longer but narrower than head; basically, as in ♀♀ but transverse median sulcus somewhat more impressed medially ( Fig. 73E ). Mesothorax narrower and 9.5x longer than prothorax, uniform in diameter except for being slightly widened at the posterior. Metanotum only one-quarter the length of mesonotum and about 4x longer than wide. Meso- and metasternum with a distinct and acute medio-longitudinal keel ( Fig. 87C ); this becoming gradually less pronounced towards the posterior margin of metasternum. Very minute and rudimentary tegmina and alae present (length <0.5 mm; Fig. 73K ).

Abdomen: Median segment almost 1.5x longer than metanotum, about 7x longer than wide and parallel-sided. Segment II only about two-thirds the length of median segment and longer than all following segments; almost 6.5x longer than wide. III–VII gradually decreasing in length with VII only about 3.6x longer than wide; all roughly uniform in diameter but very weakly constricted medially. VII with posterior portion somewhat widened. Sterna II–VII with the median line weakly indicated. Tergum VIII about 0.6x length of VII and trapezoidal with posterior margin 1.6x wider than anterior margin. IX notably longer than VIII strongly convex longitudinally, with lateral margins prominently deflexed and dilated towards the posterior to form a large triangular lobe, that ventrally projects by more than height of tergum VIII and projects over ventral surface of poculum ( Fig. 75A ). Anal segment a little more than half the length of IX and slightly wider than long, narrowed anteriorly with the lateral margins weakly rounded and the posterior margin deeply and roundly emarginated ( Fig. 75B ); the lateral angles obtusely rounded and ventrally set with small black denticles ( Fig. 96A ). Vomer very slender and gradually narrowing towards a pointed and somewhat sinistral directed terminal point, strongly arched in lateral aspect and the ventral surface with a moderately impressed medio-longitudinal furrow ( Fig. 96A ). Cerci shorter than anal segment, roughly cylindrical, slender and almost straight. Poculum moderately convex, cup-like ( Fig. 75A ) and reaching to posterior margin of tergum IX; lateral margins each with a prominent, rounded indention with the upper margin in this portion labiate and downcurved; the posterior margin thickened, labiate concave and with the outer angles downcurved and glossy ( Fig. 75C , 96A ).

Legs: All very long and slender and completely destitute of spines or teeth; profemora slightly longer than mesothorax, mesofemora reaching posterior margin of abdominal segment II and hind legs greatly projecting beyond apex of abdomen. Basitarsus long, slender with the dorsal carina weakly lamellate and somewhat longer than remaining tarsomeres combined.

Variability. Both sexes show a considerable range of morphological variability in general size (see table 58 for measurements). While the colouration is very constant in ♂♂, ♀♀ range from light or dark green over pale straw and drab to various tones of grey and mid to dark brown. Occasionally , there are ♀♀ that have numerous darker speckles or several bold white markings on the thorax and abdomen. Specimens collected by the authors at Cacao are considerably larger than from other localities (body lengths: ♂ 131.5 mm, ♀ 198.4 mm), with ♀♀ showing a more pronounced mesothoracic armature ( Fig. 72K ). The comparatively smallest specimens are those from Cayenne , Matoury and Montagne de Matoury. The original type specimens of B. muricta in the collection of MNHU from the Pará State of Brazil are even smaller (♂ 115.5 mm, ♀ 161.0 mm). Another feature in ♀♀, that underlies considerable variability is the armature of the mesonotum, with the total number of tubercles/spines ranging from 7–40 in the specimens examined. The mesonotum ranges from being sparsely (e. g. ♀ from Dermerara, Guyana in the collection of ZMPA) to densely covered with spiniform tubercles or obtuse spines (e. g. in the paralectotype of Burmeister’s muricata , Fig. 72J ) or at the other extreme can be all over armed with prominent, acutely pointed spines of variable sizes, the largest of which may be somewhat arcuate and directed towards the head (e. g. a captive reared F 1 ♀ in coll. FH, No. 0473–31; Fig. 72K ). The basal lobe of the two outer ventral carinae of the mesofemora of ♀♀ may be distinct (at least on posteroventral carina), very indistinct or completely absent. The ♀ from Monte Cristo at the Rio Tapajos, Pará, Brasil in the ZMUH collection is remarkable for the very prominent dorsal lobe of the basitarsi. Nymphs usually have the lobes of the mid and hind legs and basitarsi more prominently developed than adults .

Egg ( Fig. 100L ). Capsule elongate, 1.7x longer than wide, compressed laterally with lateral surfaces slightly sub-parallel to each other, oval in cross-section and slightly narrowed towards the polar end; angular in lateral aspect with polar-area angular and flattened, the dorsal surface more convex than ventral surface. Complete surface covered with an irregular network of ridges; the areas in between either somewhat indented minutely granulose and slightly glossy or covered with a raised scale-like plate. A strongly raised collar of these plates surrounding the micropylar plate and polar-area. Polar area with a flat oval region and a tubercle in the centre. Micropylar plate elongate, almost parallel-sided and covering about three-quarters the length of dorsal capsule surface; plate slightly indented with surface mostly smooth and minutely granulose. Anterior end of plate rounded and posterior end with a median notch. Micropylar cup an elongate swelling and placed in posteromedian gap of plate. Median line represented by a short but distinct bulge posterior of micropylar plate. Internal micropylar plate shiny white and with a small posteromedian incision. Operculum slightly oval, outer margin with a very prominent, high, hollow and crest-like structure that is formed by membranous, lamellate extensions of the outer margin; these extending by about two-thirds the length of capsule. All raised portions of capsule dark cream-coloured to ochre, the indented areas reddish mid brown. Polar area with an oval blackish central marking. Micropylar plate dark blackish brown. Raised opercular structures dark yellow to straw-coloured.

Measurements [mm]: Overall length 5.9–6.5, length 3.9–4.2, width 2.0–2.2, height 2.6–2.9, length of micropylar plate 2.9–3.2.

Comments. Burmeister (1838: 564) originally described Bacteria muricata based a ♂ and ♀ from the State of Pará, Brazil in the collection of MNHU, but provided only a very brief characterisation. For confirming the new synonymies here established and stability of Burmeister’s species the ♂ is here selected as the lectotype. Due to its strong intraspecific variability and wide geographic range Ph. muricata has subsequently been described nine more times under nine distinct species names and research on closely related taxa has revealed seven new synonyms. When Redtenbacher (1908: 418) described Bacteria divergens , which has been synonymized with muricata by Zompro & Brock (2003: 11) he already presumed it to be synonymous with Bacteria sakai Bates, 1865 . Examination of the type specimens of sakai in OUMNH confirmed Redtenbacher´s statement and has proven this to be a junior synonym of muricata as well (syn. n.). A lectotype is selected for Bacteria sakai to validate the new synonymy. Phasma (Bacteria) canna Haan, 1842 was described from a ♂ and ♀ from “Promont. Bonae Spei” (= Cape of Good Hope, South Africa) in the RMNH collection. Examination of the two type specimens has however shown the type-locality to be wrong and the species to be a synonym of Burmeister’s muricata (syn. n.). The ♂ is designated as the lectotype to validate this new synonymy. Although the type specimen of Bacteria rubispinosa Serville, 1838 is not traced the type locality “Cayenne” and description leave no doubt it is a synonym of muricata (syn. n.), which is very common throughout French Guiana and also found in the surroundings von Cayenne. Bostra reducte-dentata Redtenbacher, 1908 , Bacteria nodulosa Redtenbacher, 1908 , Bacteria rufopectus Redtenbacher, 1908 and Dyme dreyfusi Brunner v. Wattenwyl, 1907 were all described from the ♂♂ only and examination of their type specimens have clearly shown all them to represent Burmeister’s species. Thus, all four species are here synonymised (syn. n.).

Ph. muricata is very common and readily encountered in various different habitats throughout most of French Guiana. It has a wide distributional range and is in addition to French Guiana also found in the northern regions of Brazil (state of Pará) and Surinam. Observations in its habitats, carried out by the authors in 2001 in French Guiana, have shown Ph. muricata to be highly polyphagous because both nymphs and adults were observed feeding on a variety of different plants. In captivity in French Guiana this species readily accepted Cecropia peltata ( Moraceae ).

Culture stock of P. muricata (Burmeister) originating in French Guiana has been imported to Europe on several occasions, but so far none of the cultures was maintained for more than a few generations. Nevertheless, it was included on the Phasmid Study Group culture-list as culture No. 170. Lelong (1995) provided a culture report with illustrations of both sexes, the nymphs and eggs and listed firethorn ( Pyracantha spp , Rosaceae ), rose ( Rosa spp. , Rosaceae ) and bramble ( Rubus fruticosus , Rosaceae ) as alternative food plants in captivity. Lelong (1995: 18) reported the mortality of the newly hatched nymphs to be very high and stated the species to be difficult to culture successfully. Nymphs which hatched from eggs laid by wild specimens collected by the authors in French Guiana in 2001 were raised to adult using bramble ( Rubus fruticosus , Rosaceae ) and rose ( Rosa spp. , Rosaceae ) as alternative food plants. The nymphs which hatched from the eggs laid by the F1–generation ♀♀ were however very fragile and difficult to maintain.

Table 58: Measurements of Phanocloidea muricata ( Burmeister, 1838)

* Including subgenital plate

** The ♀ from Montagne de Kaw , collected by P. Grandcolas in 1989 in MNHN (only body length measured) .

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 69. Phanocloidea Zompro, 2001: A. P. incolumis (Brunner v. Wattenwyl, 1907) ♀ (Belize, San Ignacio), habitus dorsal view [NHMUK]; B. P. muricata (Burmeister, 1838) ♀ (French Guiana, Montagne de Roura), habitus dorsolateral view [coll. FH, No. 0473–2]; C. P. muricata (Burmeister, 1838) ♂ (French Guiana, nr. Cacao), habitus dorsolateral view [coll. FH, No. 0473–10]; D. P. incompta (Rehn, 1904) ♀ (Costa Rica, Prov. Limón, Matina), habitus dorsolateral view [NHMW]; E. P. baculus (Olivier, 1792) ♀ HT, lateral view [RMNH].

FIGURE 72. Phanocloidea Zompro, 2001: A. P. globocephala (Conle, Hennemann & Gutiérrez, 2011) ♂ (Colombia, Dept. Caldas), habitus lateral view [CEBUC, © Yeisson Gutiérrez]; B. P. gracilis (Burmeister, 1838) ♀ (Venezuela, Carabobo, San Esteban), habitus dorsolateral view [ANSP]; C. P. gracilis (Burmeister, 1838) ♂ HT, habitus dorsolateral view [MNHU]; D. P. gracilis (Burmeister, 1838) ♀, lateral view of head and prothorax [ANSP]; E. P. submutica (Redtenbacher, 1908) ♀, lateral view of head and prothorax [ANSP]; F. P. turgida (Westwood, 1859) ♀ with obtuse cephalic swellings, dorsolateral view of head and prothorax [coll. FH]; G. P. turgida (Westwood, 1859) ♀ with acutely pointed cephalic horns, dorsolateral view of head, prothorax and anterior portion of mesothorax [ZMUH]; H. P. incolumis (Brunner v. Wattenwyl, 1907) ♀, dorsolateral view of head, prothorax and anterior portion of mesothorax [coll. AJH]; J. P. muricata (Burmeister, 1838) ♀ PLT, dorsolateral view of head, prothorax and anterior portion of mesothorax [MNHU]; K. P. muricata (Burmeister, 1838) ♀, dorsolateral view of head, prothorax and anterior portion of mesothorax [coll. FH, No. 0473–31]; L. P. venezuelica sp. n. ♀ PT, dorsolateral view of head, prothorax and anterior portion of mesothorax [coll. FH, No. 0400–7].

FIGURE 73. Phanocloidea Zompro, 2001: A. P. pallidenotata (Redtenbacher, 1908) ♀, lateral view of head, prothorax and anterior portion of mesothorax [coll. TR]; B. P. pallidenotata (Redtenbacher, 1908) ♀, dorsal view of head, prothorax and anterior portion of mesothorax [coll. TR]; C. P. schulthessi (Redtenbacher, 1908) ♀, lateral view of head, prothorax and anterior portion of mesothorax [ZMUH]; D. P. schulthessi (Redtenbacher, 1908) ♂, lateral view of head, prothorax and anterior portion of mesothorax [ANSP]; E. P. muricata (Burmeister, 1838) ♂, lateral view of head, prothorax and anterior portion of mesothorax [coll. FH]; F. P. turgida (Westwood, 1859) ♂, lateral view of head, prothorax and anterior portion of mesothorax [coll. FH]; G. P. venezuelica sp. n. ♂ PT, lateral view of head, prothorax and anterior portion of mesothorax [coll. FH]; H. P. sanguinea sp. n. ♂ HT, lateral view of head and prothorax [MNHU]; J. P. venezuelica sp. n. ♂ PT, lateral view of metathoraxwith median segment showing the vestigial tegmina and alae [coll. FH]; K. P. muricata (Burmeister, 1838) ♂, lateral view of metathoraxwith median segment showing the rudimentary tegmina and alae [coll. FH]; L. P. incolumis (Brunner v. Wattenwyl, 1907) ♀, lateral view of terminalia [NHMUK]; M. P. incolumis (Brunner v. Wattenwyl, 1907) ♀, dorsal view of terminalia [NHMUK]; N. P. schulthessi (Redtenbacher, 1908) ♀, lateral view of terminalia [NHMUK]; O. P. schulthessi (Redtenbacher, 1908) ♀, dorsal view of terminalia [NHMUK].

FIGURE 74. Phanocloidea Zompro, 2001 terminalia of ♀♀: A. P. muricata (Burmeister, 1838), lateral view [coll. FH]; B. P. muricata (Burmeister, 1838), dorsal view [coll. FH]; C. P. muricata (Burmeister, 1838), ventral view [coll. FH]; D. P. turgida (Westwood, 1859), lateral view [coll. FH]; E. P. turgida (Westwood, 1859), dorsal view [coll. FH]; F. P. turgida (Westwood, 1859), ventral view [coll. FH]; G. P. turgida (Westwood, 1859) specimen with elongate subgenital plate, ventral view [ZMUH]; H. P. pallidenotata (Redtenbacher, 1908), lateral view [coll. TR] (Arrow indicating the unusual posterolateral lobe of abdominal tergum VII); J. P. pallidenotata (Redtenbacher, 1908), dorsal view [coll. TR]; K. P. pallidenotata (Redtenbacher, 1908), ventrall view [coll. TR] (Arrow indicating the unusual posterolateral lobe of abdominal tergum VII); L. P. venezuelica sp. n. ♀ PT, lateral view [coll. FH]; M. P. venezuelica sp. n. ♀ PT, dorsal view [coll. FH]; N. P. venezuelica sp. n. ♀ PT, ventral view [coll. FH]; O. P. submutica (Redtenbacher, 1908), lateral view [ANSP]; P. P. submutica (Redtenbacher, 1908), dorsal view [ANSP]; Q. P. submutica (Redtenbacher, 1908), ventral view [ANSP]; R. P. incompta (Rehn, 1904), lateral view [NHMW]; S. P. gracilis (Burmeister, 1838), lateral view [ANSP] (Arrow indicating the unusually short subgenital plate); T. P. gracilis (Burmeister, 1838), dorsal view [ANSP]; U. P. gracilis (Burmeister, 1838), ventral view [ANSP] (Arrow indicating the unusually short subgenital plate).

FIGURE 75. Phanocloidea Zompro, 2001 terminalia of ♂♂: A. P. muricata (Burmeister, 1838), lateral view [coll. FH]; B. P. muricata (Burmeister, 1838), dorsal view [coll. FH]; C. P. muricata (Burmeister, 1838), ventral view [coll. FH]; D. P. venezuelica sp. n. ♀ PT, lateral view [coll. FH] (Arrow indicating the characteristically specialised posterior margin of the poculum also seen in M); E. P. venezuelica sp. n. ♀ PT, dorsal view [coll. FH]; F. P. venezuelica sp. n. ♀ PT, ventral view [coll. FH]; G. P. submutica (Redtenbacher, 1908), lateral view [coll. FH]; H. P. submutica (Redtenbacher, 1908), dorsal view [coll. FH]; J. P. submutica (Redtenbacher, 1908), ventral view [coll. FH]; K. P. pallidenotata (Redtenbacher, 1908), lateral view [MNHN] (Arrow indicating the characteristically specialised posterior margin of the poculum, also seen in J);; L. P. pallidenotata (Redtenbacher, 1908), dorsal view [MNHN]; M. P. pallidenotata (Redtenbacher, 1908), ventral view [MNHN]; N. P. semiptera sp. n. ♂ HT, lateral view [USNM]; O. P. semiptera sp. n. ♂ HT, dorsal view [USNM]; P. P. semiptera sp. n. ♂ HT, ventral view [USNM]; Q. P. globocephala (Conle, Hennemann & Gutiérrez, 2011), lateral view [CEBUC, © Yeisson Gutiérrez]; R. P. globocephala (Conle, Hennemann & Gutiérrez, 2011), dorsal view [CEBUC, © Yeisson Gutiérrez] (Arrow in A, C and N indicating the large triangular posterolater projectiongh of abdominal tergum IX).

FIGURE 87. Mesosternum: A. Phanocloidea incolumis (Brunner v. Wattenwyl, 1907) ♀ [NHMUK]; B. Phanocloidea muricata (Burmeister, 1838) ♀ [coll. FH]; C. Phanocloidea muricata (Burmeister, 1838) ♂ [coll. FH]; D. Phanocloidea venezuelica sp. n. ♀ PT [coll. FH]; E. Phanocloidea venezuelica sp. n. ♂ PT [coll. FH]; F. Phanocloidea turgida (Westwood, 1859) ♀ [coll. FH]; G. Phanocloidea turgida (Westwood, 1859) ♂ [coll. FH]; H. Phanocloidea tobagoense (Langlois & Bellanger, 2012) ♀ HT [MNHN]; J. Phanocloidea submutica (Redtenbacher, 1908) ♀ [ANSP]; K. Phanocloidea schulthessi (Redtenbacher, 1908) ♀ [ANSP]; L. Phanocloidea schulthessi (Redtenbacher, 1908) ♂ [ANSP]; M. Phanocles solidus sp. n. ♀ HT [ANSP]; N. Phanocles horni (Redtenbacher, 1908) ♀ [coll. FH]. Not in scale.

FIGURE 91. Praeopercular organ of ♀♀: A. Lanceobostra aetolus (Westwood, 1859) [ANSP]; B. Lanceobostra guatemalensis (Redtenbacher, 1908) HT of Bostra intermedia Redtenbacher, 1908 [MHNG]; C. Lanceobostra oaxacae sp. n. ♀ HT [ANSP]; D. Phanoclocrania dorsuaria (Stål, 1875) HT of Bacteria monticola Carl, 1913 [MHNG]; E. Phanocloidea gracilis (Burmeister, 1838) [ANSP]; F. Phanocloidea muricata (Burmeister, 1838) [coll. FH]; G. Phanocloidea venezuelica sp. n. PT [coll. FH]; H. Phanocloidea turgida (Westwood, 1859) [coll. FH]; J. Phanocloidea schulthessi (Redtenbacher, 1908) [ANSP]. Not in scale.

FIGURE 96. Vomer of ♂♂ (ventral view): A. Phanocloidea muricata (Burmeister, 1838) [coll. FH]; B. Phanocloidea venezuelica sp. n. PT [coll. FH]; C. Phanocloidea turgida (Westwood, 1859) [ZMUH]; D. Phanocloidea incolumis (Brunner v. Wattenwyl, 1907) [coll. AJH]; E. Phanocloidea pallidenotata (Redtenbacher, 1908) [MNHN]; F. Phanocloidea tobagoense (Langlois & Bellanger, 2012) AT [MNHN]; G. Phanocloidea gracilis (Burmeister, 1838) [ANSP]; H. Phanocloidea schulthessi (Redtenbacher, 1908) [MNHN]; J. Phanocloidea procera (Conle et al., 2020) PT [MNHN]; K. Phanocloidea globocephala Conle, Hennemann & Gutierréz, 2011 [CEBUC]; L. Trychopeplus laciniatus (Westwood, 1874) [coll. OC]. Not in scale.

FIGURE 100. Eggs: A. Phanocles berezini sp. n. [coll. FH]; B. Phanocles horni (Redtenbacher, 1908) [coll. FH]; C. Parotocrania panamae gen. n., sp. n. PT [coll. FH]; D. Trychopeplus laciniatus (Westwood, 1874) [coll. OC]; E. Cranidium gibbosum (Burmeister, 1838) [coll. FH]; F. Phanoclocrania gen. n. dorsuaria (Stål, 1875) [coll. FH]; G. Phanocloidea procera (Conle et al., 2021) [coll. FH]; H. Phanocloidea incolumis (Brunner v. Wattenwyl, 1907) [coll. FH]; J. Phanocloidea schulthessi (Redtenbacher, 1908) [coll. FH]; K. Phanocloidea pallidenotata (Redtenbacher, 1908) [coll. TR]; L. Phanocloidea muricata (Burmeister, 1838) [coll. FH]; M. Phanocloidea venezuelica sp. n. PT [coll. FH]; N. Phanocloidea tobagoense (Langlois & Bellanger, 2012) [coll. FH]; O. Phanocloidea turgida (Westwood, 1859) [coll. FH]; P. Phanocloidea incompta (Rehn, 1904) [coll. FH]. [Scale = 1 mm]