Studies on neotropical Phasmatodea XVII: Revision of Phantasca Redtenbacher, 1906, with the descriptions of six new species (Phasmatodea: Diapheromeridae: Diapheromerinae) Author Hennemann, Frank H. Author Conle, Oskar V. Author Bellanger, Yannick Author Lelong, Philippe Author Jourdan, Toni text European Journal of Taxonomy 2018 2018-05-15 435 1 62 journal article 30011 10.5852/ejt.2018.435 ae4cf61a-06d2-4fd9-a12c-3457bf050e88 1255145 861CF951-45BE-458F-B0F7-79530DEE06CE Phantasca quadrilobata ( Chopard, 1911 ) comb. nov. Figs 82–103 Bacteria 4-lobata Chopard, 1911 : 343 . Bacteria quadrilobata – Otte & Brock 2005 : 66 . — Jourdan, Lelong & Bellanger 2014 : 489 , fig.1 ( ♀ ). Phantasca phantasma – Chopard 1911 : 338 . Phantasca sp. II – Jourdan, Lelong & Bellanger 2014 : 489 . Material examined Lectotype ( here designated ) FRENCH GUIANA : ♀ , “Museum Paris, Collection, Lucien Chopard, 191; Type ; Bacteria 4-lobata Chopd. ♀; Cayenne ” (MNHN). Figs 82–89. Phantasca quadrilobata ( Chopard, 1911 ) , ♀ . 82 . Habitus, dorsal view. 83 . Habitus, ventral view. 84 . Head and pronotum, dorsal view. 85 . Head and pronotum, lateral view. 86 . Apex of abdomen, dorsal view. 87 . Apex of abdomen, lateral view. 88 . Apex of abdomen, ventral view. 89 . Basal portion of left mesotibia, dorsal view. Paralectotype FRENCH GUIANA : ♀ , “ Guyane Française, Cayenne , Coll Le Moult ; Fevrier; Museum Paris, collection Lucien Chopard, 191; Bacteria 4-lobata Chopd.♀” ( MNHN ). Other material FRENCH GUIANA : 1 ♂ , “Forêt de Balaté ( St Laurent Du Maroni ), km 28, 16.X.1969 ” ( MNHN ) ; 1 ♂ , “ Guyane Françse, Nouveau Chantier, collection Le Moult ; Museum Paris, Collection Lucien Chopard; Phantasca phantasma Westw. ♂ ” ( MNHN ) ; 1 ♀ , “ Guayana franc., Rio Maroni , ex Le Moult ” ( MNHN ) ; 2 ♀♀ , “Saut Boko, Crique Elepousing, 13-III-1976 ; Museum Paris , Guyane ( Oyapock ), M. Descamps Rec. ” ( MNHN ) ; 1 ♂ : Guyane , Montagne de Kaw , 5–12 VIII 92 , Roubaud, Auvray, Rarchaert rec” ( MNHN ) ; 1 ♀ , “ Guyane Française, leg. GEP ♀ , E. Delfosse Det. 07/05” ( MNHN ) ; 2 ♀♀ , 42 eggs , “ Guyane , Saül , 8–23.X.2013 , GUY13-080, leg. Y. Bellanger , P. Lelong & T. Jourdan ” ( ASPER ) ; 1 ♀ , “ Guyane , Saül , 8–23.X.2013 , GUY13-040, leg. Y. Bellanger , P. Lelong & T. Jourdan ” ( ASPER-PL ) ; 1 ♂ , “ Guyane , Saül , 8–23.X.2013 , GUY13-039, leg. Y. Bellanger , P. Lelong & T. Jourdan ” ( ASPER-PL ) ; 1 ♂ , “GUY15-049, Guyane , route de Roura , N 4°42.905'W 52°18.252', 25.VII.2015 , rec T. Jourdan ” (ASPER- PL) ; 1 ♂ , “GUY15-084, Guyane , route de Mana , 21.VIII.2015 , rec T. Jourdan ” ( ASPER-PL ) ; 1 ♀ , Figs 90–97. Phantasca quadrilobata ( Chopard, 1911 ) , ♂ . 90 . Habitus, dorsal view. 91 . Habitus, lateral view. 92 . Head and pronotum, dorsal view. 93 . Head and pronotum, lateral view. 94 . Left ala. 95 . Apex of abdomen, dorsal view. 96 . Apex of abdomen, lateral view. 97 . Apex of abdomen, ventral view. “GUY15-014, Guyane , St Elie , N5°17.725 ' W53°03.093' , 05.VIII.2015 , rec. T. Jourdan ” ( ASPER-PL ); 1 ♂ , “GUY15-094, Guyane , route de Saut Maripa , N 3°52.078'W 51°51.277', 14.VIII.2015 , rec. T. Jourdan ” ( ASPER-PL ) ; 1 ♂ , “GUY15-099, Guyane , route de Roura , N 4°41.757'W 52°18.339', 09.VIII.2015 , rec. T. Jourdan ” ( ASPER-PL ) ; 1 ♂ , “ GUYKAW16-044 , 12.XI.2016 , Guyane fr. - Route de Kaw , Camp Caïman, PK22 depuis Roura ” ( ASPER-YB ) ; 1 ♂ , “ GUYKAW16-021 , 11.XI.2016 , Guyane fr. - Route de Kaw, Camp Caïman, PK28 depuis Roura , N4°34'11,7" ; W52°12'41,8" , alt. 300 m .” ( ASPER-YB ) ; 1 ♂ , “ GUYKAW16-022 , 11.XI.2016 , Guyane fr. - Route de Kaw, Camp Caïman, PK28 depuis Roura , N4°34'11,7" ; W52°12'41,8" , alt. 300 m .” ( ASPER-PL ) ; 1 ♀ , “ GUYKAW16-051 , 09.XI.2016 , Guyane fr. - Route de Kaw, Camp Caïman, PK27 depuis Roura , N4°34'13,1" ; W52°12'53,9" , alt. 300 m .” ( ASPER- PL ) ; 2 ♀♀ , “ GUYKAW16-052 & - 053 , 09.XI.2016 , Guyane fr. - Route de Kaw, Camp Caïman, PK27 depuis Roura , N4°34'13,1" ; W52°12'53,9" , alt. 300 m .” ( ASPER-YB ) ; 1 ♂ , “ Französisch Guyana , Commune de Roura, Montagne des Chevaux, RN2 PK22 , 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Brûlé 0 8.02.2014 , Gen 0010” ( OC 0334-1 ) ; 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Figs 98–103. Phantasca quadrilobata ( Chopard, 1911 ) , eggs. 98 . Type B, dorsal view. 99 . Type B, lateral view. 100 . Type B, anterior view. 101 . Type A, dorsal view. 102 . Type A, lateral view. 103 . Type A, anterior view. Brûlé 13.09.2014 ” (OC 0334-3); 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Brûlé 14.09.2013 , Lichtfang ” (OC 0334-4); 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Brûlé 20.01.2013 ” (OC 0334-5); 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Brûlé 06.09.2014 ” (OC 0334-6); 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Brûlé 24.04.2011 ” (OC 0334-11); 1 ♂ , “Französisch Guyana , Commune de Matoury , Réserve du Mont Grand Matoury , 4°44'56"N – 52°26'28"W , 75 m , forest edge, leg. Stéphane Brûlé 28.09.2014 ” (OC 0334-2); 1 ♂ , “Französisch Guyana , Commune de Matoury, La Désirée, 4°51'0"N – 52°19'60"W, SEAG , leg. Stéphane Brûlé 0 4.01.2014, Lichtfang ” (OC 0334-7); 1 ♂ , “Französisch Guyana , Commune de Régina, Nouragues, Saut Pararé, 4°02'N – 52°41'W , SEAG , leg. Stéphane Brûlé 27.03.2010 ” (OC 0334-8); 1 ♂ , “Französisch Guyana , Commune de Régina, Nouragues, Saut Pararé, 4°02'N – 52°41'W , SEAG , leg. Stéphane Brûlé 03.09.2010 ” (OC 0334-10); 1 ♂ , “Französisch Guyana , Commune de Saül, Aussichtspunkt Belvedere de Saül, 3°37'22"N – 53°12'57"W , 326 m , SEAG , leg. Stéphane Brûlé 19.10.2010 ” (OC 0333-9); 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , leg. Stéphane Brûlé 24.04.2011 ” (OC 0334-11); 1 ♂ , “Französisch Guyana , Commune de Roura , Réserve Naturelle Trésor , Route de Kaw RN6 PK18, 4°36'39"N – 52°16'46"W , alt. 252 m , SEAG , leg. Stéphane Brûlé 05.10.2009 ” (OC 0334-12); 1 ♂ , “Französisch Guyana , Commune de Régina , Piste forestière Tibourou , SEAG , Lichtfang , leg. Stéphane Brûlé 04.09.2008 ” (OC 0334-13); 1 ♂ , 3 ♀♀ , “Französisch Guyana , Ex Zucht O. Conle 2015, Gen F 2, Commune de Saül , 10.2013, leg. Yannick Bellanger & ASPER ” ( OC 0334- 14 –0334- 17 ); 1 ♂ , “Französisch Guyana , Commune de Roura , Montagne des Chevaux , RN2 PK22, 4°44'56"N – 52°26'28"W , alt. 75 m , SEAG , Lichtfang , leg. Stéphane Brûlé 20.08.2016 ” (OC 0334-18); 1 ♂ , “Französisch Guyana , Commune de Matoury , La Désirée , Forêt de colline littorale, N 4°50'42,34” O 52°20'54,12”, S.E.A.G, Lichtfang , leg. Stéphane Brûlé 27.09.2014 ” (OC 0334-19); 1 ♂ , 3 ♀♀ , 55 eggs , “ex Zucht F. Hennemann 2015, Herkunft : French Guiana , Saül , leg. Bellanger , Lelong & Jourdan 8.–23.X.2013 ” (FH 0866-1–4, E). Diagnosis The colouration of males is similar to that of P. guianensis sp. nov. and P. phantasma , but the distinctive transverse pale cream, yellowish or almost white marking between the eyes well characterises this species ( Fig. 92 ). The genital morphology indicates a close relationship to the latter species,, with which it shares the short and upcurving cerci. However, the cerci in P. quadrilobata are arcuate to almost sickle-shaped and gradually tapered towards an acutely pointed apex (blunt in P. phantasma ), the anal segment has the lateral margins distinctly convex and the posterior margin only, with a small triangular excavation and the poculum is considerably smaller, roundly bowl-shaped, with the lateral margins indented (labiate and angled downward in P. phantasma ). From P. guianensis sp. nov. , they readily differ by the short cerci, which hardly reach to the posterior margin of the anal segment (very long and greatly projecting beyond apex of abdomen in P. guianensis sp. nov. ) as well as the shape of the anal segment (parallel-sided in P. guianensis sp. nov. ). Females are similar to those of P. guianensis sp. nov. but differ by the somewhat smaller size, more globose head, which bears a faint cream marking on the frons, presence of two transversely carinate swellings or humps on abdominal tergum VI, smaller epiproct, slightly shorter cerci, distinctly more slender legs and the absense of the red bases of the profemora seen in P. guianensis sp. nov. Description Male ( Figs 90–91 ) BODY. Size fairly large (body length 54.5–66.5 mm , Table 11 ); form very slender and typical for the genus. COLOUR. Body green, yellowish brown or pale brown, with very characteristic dark brown to black markings. Head greyish pale to mid brown, with a bold black transverse marking on frons and a roughly V-shaped black marking on vertex. Between these black markings a distinctive pale cream, yellowish or almost white transverse area between the eyes. Lower portions of cheeks black. Scapus pale brown, with a faint dark brown spot extero-medially. Pedicellus of same colour as scapus and the rest of the antennae mid brown, with a slight reddish hue. Pronotum, with several dark longitudinal lines, including a fairly bold median stripe. Mesonotum, with a very slender dark line near lateral margins and four weakly defined dark brown markings in posterior portion. Metanotum mid to dark brown. Meso- and metasternum, with several irregular dark markings and speckles. Tegmina and costal region of alae ochre to greyish mid brown and all major veins dull yellow, the latter with a faint sepia longitudinal stripe near anterior margin. Anal region transparent with all major veins brown. Abdominal tergites II–VI each, with a very fine dark longitudinal median line and four dark spots posteriorly, VII, with two large dark markings post-medially. Two bold black, slightly diverging stripes on anal segment and sternum VII, usually with a pair of longitudinal black markings on posterior half. Poculum, with a transverse dark band medially. Front legs pale to mid brown, the profemora green basally. Mid and hind legs mid to dull green and with fairly contrasting black annulae. Knees cream coloured. HEAD. Ovoid, with the vertex very gently convex and smooth; broadest at the eyes ( Figs 92–93 ). Eyes large, projecting hemispherically and their length contained about 1.5× in that of cheeks.Antennae longer than body. Scapus rectangular but somewhat narrowed basally and 1.6× as long as wide, pedicellus subglobose and antennomere III a little longer than scapus and pedicellus combined. THORAX. Pronotum slightly shorter but considerably narrower than head and very gently narrowed postmedially ( Fig. 92 ). Transverse median sulcus moderately distinct, straight and expanding almost over entire width of segment. Mesothorax almost 3.2× as long as head and pronotum combined. Tegmina slender, spatulate, strongly constricted in the basal half and with the apical portion moderately broadened and angular. Alae roughly reaching to posterior margin of abdominal segment IV. ABDOMEN. Median segment about 2.8× as long as metanotum. Segments II–VII gradually decreasing in length. II about 6× and VII hardly 3× as long as wide; the latter somewhat deflexed posteriorly. Tergum VIII shortest and broadest of all segments, strongly swollen and hardly longer than wide; the lateral margins very indistinctly separated from sternum VIII, with the fissure only visible in the posterior portion ( Fig. 96 ). IX a little longer than VIII, narrowed anteriorly and gradually widening towards the posterior; the lateral margins deflexed, curved inward ventrally and almost completely fused with one another (an indistinct longitudinal fissure may be seen in the posterior portion) leaving only an almost circular, distinctly marginated hole or pit roughly in the middle ( Fig. 97 ). Anal segment as broad as posterior portion of VIII, a little longer than wide and slightly cucullate; the posterior margin, with a narrow but fairly distinct triangular median excavation and the outer angles obtusely triangular ( Fig. 95 ). The region of the median excavation, with about 3–4 blunt black teeth ventrally. Vomer very small, triangular in shape and with an acute apical hook ( Fig. 97 ). Cerci short, arcuate to almost sickleshaped, upcurving and gradually constricted towards an acutely pointed tip; hardly projecting beyond anal segment ( Figs 96–97 ). Sternum VIII strongly swollen and sub-globose ( Fig. 96 ). Poculum small, bowl-shaped, with the basal portion strongly convex, the upper lateral margins distinctly excavated and the posterior margin upcurving, entire and broadly rounded ( Fig. 96 ). Table 11. Measurements of Phantasca quadrilobata (Chopard, 1911) comb. nov. (in mm). LT = Lectotype; PLT = Paralectotype.

♀, LT (MNHN)	♀, PLT (MNHN)	♂♂	♀♀
Body	109.5	105.0	54.5–66.5	102.5–118.0
Pronotum	3.9	3.8	1.3–2.1	4.0–4.5
Mesonotum	26.0	25.0	11.2–14.0	24.7–29.0
Metanotum	7.5	7.2	2.8–2.9	2.7–3.2
Median segment	9.0	8.4	7.4–7.6	12.5–13.9
Tegmina	-	-	2.3–3.2	-
Alae	-	-	19.6–29.3	-
Profemora	27.0	24.2	18.6–20.3	25.6–29.0
Mesofemora	18.6	17.0	12.4–14.0	17.6–22.0
Metafemora	24.7	22.9	16.2–18.0	21.3–27.5
Protibiae	30.7	28.1	20.5–23.8	31.0–33.5
Mesotibiae	20.0	18.2	13.2–15.2	18.7–23.0
Metatibiae	28.1	27.9	19.5–21.8	25.0–32.2
Antennae	> 60.0	59.5–68.0	57.0–73.8

LEGS. All very long and slender, profemora somewhat longer than head, pro- and mesothorax combined and metatibiae projecting much beyond apex of abdomen. All basitarsi considerably longer than combined length of remaining tarsomeres. Female ( Figs 82–83 ) BODY. Size large (body length 102.5–118.0 mm, Table 11 ); form slender, with characteristic humps on abdominal tergum VI. Body smooth. COLOUR. Variable; ranging from dull green over various shades of brown and grey, either plain or, with a variable degree of white or dark speckles and spots. Occassionally, with bold white markings on head and some or all body segments. Sometimes brown specimens may have green legs. Head usually, with a more or less defined dark V-shaped marking on vertex and a more pale (sometimes white) area between the eyes. Legs usually, with fairly distinct but irregular dark annulae. Basal portion of profemora always mid to dull green. HEAD. Ovoid, broadest at the eyes, the vertex very convex and with a scarce longitudinal median sulcus; cheeks slightly narrowing towards the posterior ( Figs 84–85 ). Eyes fairly small, circular and their length contained about 1.8× in that of cheeks. Antennae reaching about halfway along abdominal segment IV. Scapus and pedicellus as in males. THORAX. Pronotum rectangular, somewhat shorter but considerably narrower than head; slightly constricted pre-medially ( Fig. 84 ). Transverse median sulcus moderately distinct, gently curved and just not expanding over entire width of segment. Longitudinal median sulcus distinct. Mesothorax about 3× as long as head and pronotum combined, of uniform width. ABDOMEN. Median segment about 1.2× as long as metanotum and very gently constricted medially. Abdominal segments II–V slightly increasing, VI–X decreasing in length; V on average some 2.5× as long as wide. II–V of uniform width, VII–X somewhat narrower than preceding. VI swollen and broadest of all segments, tergum, with posterior portion raised and protruded into two bluntly conical or transversely carinate swellings; these fairly variable in size and shape. Praeopercular organ formed by a small, obtuse median swelling at posterior margin of sternum VII ( Fig. 88 ). Anal segment almost equal in length to IX. Tergites VIII–X of uniform width and somewhat narrower than preceding. Anal segment about equal in length to IX, slightly deflexed basally and somewhat narrowed in the posterior portion; the posterior margin roundly angular and with a very shallow and small median indention ( Fig. 86 ). Epiproct very small, roundly triangular. Cerci small, conical, with the apex gently incurving and slightly projecting beyond posterior margin of anal segment ( Fig. 87 ). Gonapophyses VIII elongated, filiform, upcurving at the apex and roughly reaching to apex of subgenital plate ( Fig. 87 ). The latter boat-shaped, keeled longitudinally, with the lateral margins strongly angled downward apically; apex blunt and just not reaching the posterior margin of the anal segment ( Fig. 88 ). LEGS. All fairly long and slender. Anterodorsal carina of profemora moderately raised sub-basally. Medioventral carina of meso- and metatibiae deflexed to form a distinctive, rounded lobe sub-basally ( Fig. 89 ). All basitarsi longer than remaining tarsomeres combined. Egg ( Figs 98–103 ) There are two different types of eggs in this species, whose differences in colour and surface sculpturing of the capsule are summarized separately below. Medium-sized, ovoid, distinctly oval in cross-section, about 1.2× as long as high and about 1.8× as long as wide. Opercular collar distinct. Micropylar plate elongate, slender and very gently widened posteriorly; on average some 5× as long as wide and about ¾ the length of capsule. Outer margin slightly raised and the interior portion somewhat bulgy longitudinally. Micropylar cup small, bowl-shaped. Median line very short. Operculum flat and oval, with outer portion distinctly marginated; capitulum represented by a raised rim and some irregularly shaped acute ridges in the centre. Type A ( Figs 101–103 ): General colour plain dark brown, with a pale cream to grey rim just below opercular collar; entire surface strongly shiny. Micropylar plate, with inner portion dark blackish brown and outer portion reddish mid brown. Operculum dark brown, with a slight reddish wash and the capitular structures dull yellowish to orange. Capsule surface almost smooth and strongly shiny; minute pits can only be seen at strong magnification. Type B ( Figs 98–100 ): Plain dull grey, with a bold washed black central marking on lateral surfaces. Micropylar plate black, with the outer portion and median line dull ochre. Operculum dull grey and the capitular structures pale brown, with a slight yellowish hue. Complete capsule surface and micropylar plate minutely and densely pitted. Measurements (in mm). Length (incl. operculum) 2.69, length 2.45, width 1.34, height 2.06, length of micropylar plate 1.85, diameter of operculum 0.77 × 1.11. Remarks Chopard (1911: 343) described Bacteria 4-lobata based on two female syntypes from the collection of L. Le Moult in MNHN. The broader female, which has the front legs folded, is here selected as the lectotype to guarantee stability of the name. Examination of the male from Nouveau Chantier, in the collection of Le Moult in MNHN referred to as P. phantasma ( Westwood, 1859 ) by Chopard (1911: 338) , has proven to be P. quadrilobata ( Chopard, 1911 ) . Both sexes show considerable variability in colouration. Females mostly show various shades of brown or grey but sometimes have a slight greenish hue or are almost entirely dark green or occasionally have large white markings on the head and almost all body segments. The two distinctive protuberances on abdominal tergum VI show considerable variability in size and shape, ranging from bluntly conical swellings to transversely carinate scales. Also, the sub-basal ventral lobe of the meso- and metatibiae is fairly variable in shape. While males are mostly buffy or ochre to mid brown, specimens may be green during the first days after their final ecdysis. Quite some variability is also seen in the shape of the black markings on the head, pronotum and terminal abdominal tergites. The eggs occur in two different types , which differ in colouration and sculpturing of the egg capsule. Breeding in captivity was attempted from stock collected in Saül in 2013 but has proven difficult. The culture was lost after the second generation but allowed the authors to be sure of the matching between males and females. Hatching rates in the first generation were fairly low, with only about 50%. At average temperatures of 20°C males reached maturity after about five months while females took a little longer. Rose ( Rosa spp., Rosaceae ), St. John’s Wort ( Hypericum patulum ‘Hidcote’, Hypericaceae ) and eucalyptus ( Eucalyptus spp., Myrtaceae ) were readily accepted as alternative food plants. In the wild this species is known to feed on Gonzalagunia spp. ( Rubiaceae ), but presumably several other plants serve as a natural diet, since most species appear to be fairly polyphagous. Distribution French Guiana : Cayenne (MNHN); Nouveau Chantier (MNHN); St Laurent Du Maroni , Forêt de Balaté (MNHN); St Laurent Du Maroni , Rio Maroni (MNHN); Saut Boko, Crique Elepousing (MNHN); Commune de Saül (ASPER); Commune de Roura , Montagne de Kaw (MNHN); Commune de Roura , Route de Kaw (ASPER); Commune de Roura , Montagne des Chevaux (OC); Commune de Roura, Réserve Naturelle Trésor, Route de Kaw (OC); Commune de Roura , route de Roura (ASPER); Commune de Matoury , Réserve du Mont Grand Matoury (OC); Commune de Saül , Belvedere (OC); Commune de Régina , Piste forestière Tibourou (OC); Commune de Régina , Nouragues , Saut Pararé (OC); St. Elie (ASPER); Commune de Matoury , La Désirée , Forêt de colline littorale (OC).