Medauromorpha regina (Brunner von Wattenwyl, 1907) Bresseel & Constant, 1893

Bresseel, Joachim & Constant, Jérôme, 1893, The new stick insect genus Medauromorpha gen. nov. with one new species from Vietnam and notes on Medauroidea Zompro, 2000 (Phasmida: Phasmatidae: Clitumninae), Belgian Journal of Entomology 45, pp. 1-38 : 1-38

Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov.

Figs 1 A ; 6–7 ; 8 E–H ; 13 A–C ; 14–17 .

Cuniculina regina BRUNNER VON WATTENWYL, 1907: 201 [described].

Baculum spinicornum CHEN & HE, 2000: 396 [described (in Chinese)], Fig. 3 [male habitus and terminalia illustrated]. syn. nov.

Baculum reginum – HAUSLEITHNER, 1988: 195; fig. 2 I [transferred to Baculum ; egg described, illustrated; placed in Baculum hyphereon group]. — BROCK, 1998: 53 [2 female syntypes catalogued; male syntype not traced].

Ramulus reginus – OTTE & BROCK, 2005: 306 [catalogued].

Ramulus spinicornus – OTTE & BROCK, 2005: 307 [transferred to Ramulus , catalogued]. — HENNEMANN et al., 2008: 33 [catalogued from China].

Baculum spinicornum – CHEN & HE, 2008: 269 [described (in Chinese)], Fig. 232 [male habitus and terminalia illustrated].

non Baculum reginum — BROCK, 1998: 53 [typo for B. regulum ].

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TYPE MATERIAL EXAMINED. LT ♂ of Cuniculina regina , by present designation (examined from photographs, Fig. 13 A–C ): Tonkin, Than- Moi, Juni-Juli, H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Cuniculina regina ( NHMW, No. 343).

PLT (2♀♀, examined from photographs – BROCK et al., 2016): 1♀: Tonkin, Montes Mauson, April-Mai, 2-3000 m, H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Cuniculina regina ( NHMW, No. 343); 1♀: Tonkin, Than-Moi, Juni-Juli, H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Cuniculina regina ( NHMW, No. 343) .

ADDITIONAL MATERIAL (17♂♂, 28♀♀, 2♀♀ nymphs, eggs). 4♀♀: Vietnam, Tay Yen Tu Nat. Res. , 21°11’10’’N 106°43’25’’E, 7-11.VII.2013, night collecting, leg. J. Constant & J. Bresseel, I.G.: 32.454 (3♀♀: RBINS, 1♀: VNMN) ; 3♂♂, 5♀♀: Ex breeding Bruno Kneubühler 2014, origin: Vietnam, Tay Yen Tu Nat. Res. , 21°11’10’’N 106°43’25’’E, 7-11.VII.2013, night collecting, leg. J. Constant & J. Bresseel ( RBINS) ; 4♂♂, 3♀♀, 3 eggs: Vietnam Vinh Phuc pr., Me Linh B.S., 21°23’38’’N 105°45’56’’E, 30.VI-1.VII.2015, night collection, Leg. J. Constant & J. Bresseel, I.G.: 33.092 (3♂♂, 2♀♀: RBINS, 1♂, 1♀: VNMN) ; 6♂♂, 10♀♀, 3 eggs: Vietnam, Bac Kan pr., Ba Be N.P., 22°24’19’’N 105°36’55’’E, 2-7.VII.2015, night collecting, Leg. J. Constant & J. Bresseel, I.G.: 33.092 (4♂♂, 8♀♀: RBINS, 2♂♂, 2♀♀: VNMN) ; 4♂♂, 4♀♀, 5 eggs: Vietnam, Cham Chu N.R., 22°12’N 105°06’E, 8-12.VII.2015, night collecting, Leg. J. Constant & J. Bresseel, I.G.: 33.092 (1♂, 3♀♀: RBINS; 1♂, 1♀: VNMN) ; 1♀: Ex breeding Tim Bollens 2016, origin: Vietnam, Cham Chu N.R., 22°12’N 105°06’E, 8-12.VII.2015, night collecting, Leg. J. Constant & J. Bresseel ( RBINS) ; 2♀♀ nymphs: Vietnam, Tuyen Quang prov. 5 km E of Na Hang , 22°20’59’’N, 105°25’36’’E, 290m asl; 4-13 Nov. 2015, Coll D.V. Logunov ( MMUM) .

NOTES. (1) One additional female of M. regina labelled “ Tonkin, Than-Moi, Juni-Juli, H. Fruhstorfer” in the RBINS collections, was also examined. It bears a “ syntype ” label typically added by some RBINS staff and is placed in the collections together with two syntypes of Myronides magnificus Brunner von Wattenwyl, 1907 . However, the specimen does not match in any way the description given by BRUNNER VON WATTENWYL (1907). The female in the collection at RBINS has a body length of 141.1 mm while Brunner von Wattenwyl’s specimen of M. magnificus measures 128 mm. Its typical status was already questioned by VANSCHUYTBROECK & COOLS (1981). However, it was erroneously mentioned in the type series of M. magnificus given by BROCK (1998).

We here exclude this specimen from the type-series of M. magnificus .

(2) Baculum spinicornum Chen & He, 2000 was described from a single male preserved in the collections of CAU, with the following data: China, Guangxi Province, Longzhou County, Longgang, 240m, 19.v.1982, leg. Li Fasheng. It is here synonymized under M. regina based on the examination of the illustrations of the habitus and terminalia given in CHEN & HE (2000; 2008).

(3) BRUNNER VON WATTENWYL (1907) described Cuniculina regina from two females and one male. In his catalogue of the phasmid types in the collections of NHMW, BROCK (1998) stated that he had not been able to trace the male ST of C. regina . However, he erroneously listed two female ST for Cuniculina regula Brunner von Wattenwyl, 1907 . The species regula was actually described on a single female HT (BRUNNER VON WATTENWYL, 1907), and one of the two “ST” recognized by BROCK (1998) is actually a male specimen. This male is conspecific with the two female C. regina ST and represents the missing male ST of the latter species, as confirmed by the examination of the labels attached to the specimen ( Fig. 13 C ). We designate this male as LT of C. regina in order to stabilize the nomenclature in the group. The status of the female HT of C. regula is treated infra.

27 (4) HAUSLEITHNER (1988) described the egg of M. regina based on a broken egg from a Than-Moi specimen, and placed the species in his “ Baculum hyphereon ” (erroneously spelled “hypereon”) species group. The species group was defined by the same author in a previous paper (HAUSLEITHNER, 1986), from the shape of the eggs, and contained four species from Sri-Lanka. The eggs of M. regina is much more elongated and shows a network-like structure, minute hooks and mushroom-like structures which are absent in B. hyphereon group eggs. On this basis, M. regina is excluded from the group.

DESCRIPTION.

MALE ( Figs 6 ; 14 A–G ; 15 A ; 16 A–H ).

Measurements: see Table 3.

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Body: consistent in colour following the localities. Mostly brown with irregular small black spots. Pro-, meso-, metastenum and first four abdominal segments with black longitudinal line laterally. Black line thickened at the apex of each segment.

Head: longer than wide, slightly tapering posteriorly with few minute granules. Vertex slightly elongated, raised apically and split by a longitudinal median furrow creating two “bumps”. Dorsal portion of head flattened. Between eyes two definite, thick black spines. Shape of spines variable. Eyes circular and strongly projecting hemispherically. Antennae about as long as profemora with about 22 segments. Scapus strongly flattened dorsoventrally and slightly rounded laterally. Pedicellus slightly flattened. Apical antennomere slightly shorter than two preceding ones combined.

Thorax: pronotum trapezoidal and rugose, shorter than head and widening posteriorly. Anterior margin strongly incurved and raised, followed by a transverse impression, curving and getting deeper towards anterolateral angles. Median longitudinal groove starting behind anterior transverse depression, not reaching posterior margin. Centrally with a short transverse impression not reaching lateral margins of pronotum. Posterior margin convex. Mesonotum more than six times longer than pronotum, slightly widening towards the posterior. Smooth with a mediolongitudinal line. Metanotum as mesonotum, about four times longer than pronotum. Meso- and metasterna smooth.

Legs: profemora slightly shorter than head and thorax combined; compressed and curved basally. Dorsal carina faint, more definite towards apex. Anteroventral carinae raised and laterally flattened. Mesofemora shorter than head, pro- and mesonotum combined. Carinae indistinct and simple. Metafemora distinctly shorter than profemora. Carinae indistinct, more distinct towards the posterior. Protibiae considerably longer than profemora, triangular in cross section and with few minute teeth on the posteroventral carinae. Probasitarsus much longer than following tarsomeres combined; tarsomeres with small posteromedian lobe and distinctly setose ventrally. Fourth tarsomere of all legs very short. Claws very small in relation to body size. Mesotibiae longer than mesofemora, all carinae armed with minute saw-like teeth; more densely armed towards apex, tarsomeres and claws as in forelegs. Metatibiae as mesotibiae.

Abdomen: median segment short and simple. Abdominal segments II-IV gradually increasing in length; IV-VI about the same length. Tergum VII about as long as tergum II; II-VII smooth with a definite mediolongitudinal carina; VIII shorter than II, slightly widening posteriorly; III-VIII with a small lobe anterolaterally covering the spiracles; spiracles only visible in ventral apect. Tergum IX shorter than VIII and tectiform; X apically notched, about as long as VII and with median longitudinal carina. Inner portion of semi-tergite swollen apically and armed with several short denticles. Posterolateral angles rounded. Poculum rounded, reaching apex of tergum IX; apex of poculum narrowing and rounded. Cerci short, laterally flattened and slightly setose, reaching apex of tergum X and with apex rounded. Vomer well developed, visible as an elongated sclerite, narrowing towards apex; apex acute and darkened ( Fig. 6 E–H ). Abdominal sterna II-VII rugose.

FEMALE ( Figs 7 ; 14 H–Q ; 15 ; 16 I–L ; 17 ).

Measurements: see Table 3.

Body: females vary in colour; main colour is various shades of brown. Sometimes completely brown, often with darker markings scattered over body and legs, and a conspicuous black mediolongitudinal marking on posterior part of prosternum and on base of subgenital plate. Some specimens with darker longitudinal markings and some with paler streaks or with almost white patches.

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Head: longer than wide, slightly tapering posteriorly and granulose. Vertex slightly elongated, slightly raised apically, and split by median line creating two apical bumps. Dorsal portion of head flattened with area just behind eyes slightly raised; raised area with transverse ridge, and a spine at each side. Eyes circular and strongly projecting hemispherically. Antennae reaching less than half length of profemora, with 24-28 segments. Scapus strongly flattened dorsoventrally and rounded laterally. Pedicellus shorter, narrower and flattened. Apical antennomere about as long as three to four preceding ones combined.

Thorax: pronotum trapezoidal, widest part subapically; shorter than head and slightly granulose. Anterior margin incurved and raised, followed by a shallow median longitudinal impression which does not reach posterior margin. Central transverse impression not reaching lateral margins of pronotum. Mesonotum about five times the length of pronotum, granulose and “wrinkled” with a fine median longitudinal line. Metanotum with a fine mediolongitudinal line and slightly granulose, more than three times longer than median segment.

Legs: profemora about as long as meso-, metanotum and median segment combined; compressed and curved basally. Anterodorsal carina with small serrations, more prominent and numerous anteriorly. Posterodorsal carina indistinct anteriorly, becoming more distinct towards the posterior. Anteroventral carina present, slightly raised and laterally compressed. Posteroventral carina with some small saw-like teeth anteriorly. Mesofemora with posterodorsal carina armed with three large lobes. First lobe positioned anteriorly and resulting into two spines. Median lobe as anterior lobe and posterior lobe positioned more vertically on carina, resulting in a single curved tooth. Anterodorsal carina also with three distinct lobes, but smaller than the ones on posterodorsal carina. Two lobes positioned in the basal half and bearing two saw-like teeth; third lobe positioned medially, also bearing two saw-like teeth. Mesofemora a small spine subapically. Ventral carinae unarmed. Metafemora with dorsal carinae armed with some minute teeth. Ventral carinae unarmed. Protibiae slightly longer than thorax and median segment combined, almost triangular in cross section. Anterodorsal carinae with few minute, saw-like teeth at regular intervals. Posterodorsal carina with fewer saw-like teeth. Ventral carinae distinct. Medioventral carina strongly raised and laterally flattened. Outer ventral carina with few minute saw-like teeth. Inner ventral carina unarmed. Mesotibiae slightly longer than mesofemora. Posterodorsal carina with a conspicuous lobe in anterior part and bearing two spines; some saw-like teeth subapically. Anterodorsal carina with few small saw-like teeth subapically. Ventral carinae distinct. Medioventral carinae raised and laterally flattened with one conspicuous lobe anteriorly, resulting into one to two teeth. Other portions of ventral carinae, armed with minute saw-like teeth. Metatibiae slightly longer than metafemora, almost triangular in cross section. Dorsal carinae with few small saw-like teeth, more numerous posteriorly. Ventral carinae distinct. Medioventral carinae raised and laterally flattened with one lobe-like tooth anteriorly. Other portions, especially outer carinae, armed with minute saw-like teeth, more numerous posteriorly. Tarsi setose, with basitarsi as long or longer than following tarsomeres combined. Arolium and claw relatively small in relation to body size.

Abdomen: median segment transverse. Abdominal segments II-V gradually increasing in length, granulose and with fine mediolongitudinal line. Tergum VI about as long as IV; VII slightly shorter than VI and with a wart-like structure posteriorly; VIII two thirds as long as VII. Terga III–VIII with a small lobe anterolaterally, covering the spiracles; spiracles only visible in ventral aspect. Tergum IX about half as long as III and with a posteromedian definite hump, not projecting over apex; X tectiform with a raised mediolongitudinal carina, and facing obliquely downwards. Epiproct short, triangular and tectiform, with a median longitudinal carina. Sterna II–VI with a variable number of blunt spines; posteriorly with two symmetric, laterally compressed, ridges. Sternum VII with a distinct praeopercular organ

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32 present as a large black hump slightly notched posteriorly. Subgenital plate elongated, laterally compressed and not reaching apex of abdomen; smooth, with a small posteromedian carina and with apex rounded. Anteriorly with a distinct black marking. Cerci small, laterally compressed, not reaching apex of tergum X ( Fig. 7 E–G ).

NYMPH ( Fig. 14 R–T ).

Newly hatched nymphs slender and rather large, 20 to 25 mm long; mostly pale brown with a greenish tinge on head. Head bigger than pronotum and with a broad black postocular line continuing on sides of pronotum and basal part of mesonotum. Antennae very short with apical antennomere oval, knob-like and black. Legs long, lobes not developed. Legs mottled dark brown and white.

EGG ( Fig. 8 E–H ).

Measurements [in mm]: length: 16.5; length without opercular rim: 15.9; width: 1.5; height: 1.4.

33 Capsule light to dark brown, strongly elongated and gently curved in lateral aspect. Slightly convex ventrally and slightly concave dorsally. Broadest centrally, slightly narrowing towards anterior and posterior ends. Capsule surface with a network of smooth ridges fusing around the micropylar plate, creating a smooth surface around the plate. Ridges with minute mushroom and hook-like structures. Polar area very distinct, cylindrical with apex shallowly excavated; shaped much like operculum and about 1 mm long; surface wrinkled and armed with several hook-like tubercles and a fringe-like structure apically; broadening towards apex. Operculum with thickened opercular rim; smooth basally, later with minute mushroom- and hook-like structures; apex deeply excavated. Micropylar plate small and almost round, positioned close to polar area. Micropylar plate surface smooth. Micropylar cup black and distinct, followed by an elongated median line.

NOTE. Eggs from Tay Yen Tu N.R. have a drop-shaped micropylar plate, tapering towards the operculum.

INTRASPECIFIC VARIABILITY. The redescription of this species is based on specimens from Ba Be N.P. because these agree completely with BRUNNER VON WATTENWYL’ s (1907) specimens. Following the localities, several small differences can be seen between the populations. Variation between specimens of the same location is also observed.

Horns of males: ( Fig. 16 A–E ) the horns on the males’ head can vary to a certain extent. Specimens from Tay Yen Tu N.R. have horns longer than scapus, incurving, thickened at base and gradually tapering towards apex. One specimen from Me Linh B.S. has the spines short,

34 not longer than scapus, and straight. Another specimen has the spines longer than scapus and slightly incurving. Specimens from Cham Chu N.R. have the spines slender, narrowing, strongly tapering and incurving, and not completely black, but brown on the posterior side. Some specimens from Ba Be N.P. have conspicuously thickened spines; others are like those from Cham Chu N.R.

Male terminalia: ( Fig. 16 F–H ) the anal segment in male varies only slightly, being sometimes more elongated and less rounded.

Female legs: ( Fig. 16 I–L ) in females, lobes on the metafemora are absent in specimens from Cham Chu N.R. and Ba Be N.P. Specimens from Me Linh B.S. can have no lobes or a small lobe basally on the anteroventral carina of the metafemora. Specimens from Tay Yen Tu N.R. can have a developed lobe on the anterodorsal and posterodorsal carina on the basal portion of the metafemora, or, more often, two developed lobes on the posterodorsal carina and one on the anteroventral carina. The lobe on the anteroventral carina is between the other two lobes and always the biggest of the three.

Size: varies considerably. In Tay Yen Tu N.R., only females were collected and breeding the species in captivity (B. Kneubühler) gave us access to additional material including males. One captive reared female measures 155.2 mm whereas the wild caught females and all other captive bred specimens measure between 119.5-139.6 mm. The two female PLT studied by BRUNNER VON WATTENWYL (1907) differ also considerably in length (see measurements table 3).

Shape of eggs: ( Fig. 8 E–H ) eggs from Cham Chu N.R, Ba Be N.P. and Me linh B.S. have the micropylar plate almost round. Eggs from Tay Yen Tu N.R. have a drop-shaped micropylar plate, tapering towards the operculum. Eggs from Tay Yen Tu are also the largest ones (measurements above).

(* according to BRUNNER VON WATTENWYL, 1907).

DISTRIBUTION. Fig. 1 A .

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Figures

Fig. 1. Medauromorpha gen. nov. spp., distribution maps. A, M. baviensis sp. nov. and M. regina (Brunner von Wattenwyl, 1907) comb. nov. B, M. foedata (Brunner von Wattenwyl, 1907) comb. nov.

Fig. 6. Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov., ♂, Me Linh B.S. A, habitus, dorsal view. B, habitus, lateral view. C, habitus, ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia, ventral view. G, terminalia, posteroventral view. H, terminalia, lateral view. I, head and thorax, lateral view.

Fig. 7. Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov., ♀, Me Linh B.S. A, habitus, dorsal view. B, habitus, lateral view. C, habitus, ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia, ventral view. G, terminalia, lateral view. H, head and thorax, lateral view.

Fig. 8. Medauromorpha gen. nov. spp., eggs (above scale bar, lateral view; below scale bar, dorsal view). A, M. baviensis sp. nov. — B–D, M. foedata (Brunner von Wattenwyl, 1907) comb. nov. B, Cuc Phuong N.P. C, Tam Dao N.P. D, Tay Yen Tu N.R. — E–H, M. regina (Brunner von Wattenwyl, 1907) comb. nov. E, Ba Be N.P. (operculum missing). F, Cham Chu N.R. G, Me Linh B.S. H, Tay Yen Tu N.R. (operculum missing in 3 above pictures).

Fig. 13. Type specimens in NHMW (©Natural History Museum Vienna, Orthoptera Image Collection, published with permission). A–C, Cuniculina regina Brunner von Wattenwyl, 1907, LT ♂. A, lateral view. B, dorsal view. C, labels. D–E, Cuniculina regula Brunner von Wattenwyl, 1907, HT ♀. D, lateral view. E, labels.

Fig. 14. Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov., captive reared (B. Kneubühler) from Tay Yen Tu N.R. (photographs by B. Kneubühler). A–G, ♂. A, lateral view. B, dorsal view. C, ventral view with typical black marking. D, head and pronotum, lateral view. E, terminalia, dorsal view. F, terminalia, lateral view. G, terminalia, ventral view. — H–Q, ♀. H, lateral view. I, dorsal view. J, dorsal view K, lateral view. L, ventral view with typical black marking. M, lobes on metafemora. N, head and pronotum, lateral view. O, terminalia, dorsal view. P, terminalia, lateral view. Q. terminalia, ventral view. — R–T, ♀ nymph. R, dorsal view. S, dorsal view. T, head and thorax, dorsal view.

Fig. 15. Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov., in nature. A, mating pair, Ba Be N.P., 2.VII.2015. B, ♀, Cham Chu N.R., 8.VII.2015.

Fig. 16. Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov., intraspecific variability. A–E, ♂, head, lateral view. A, Ba Be N.P. B, Cham Chu N.R. C, Me Linh B.S. D, Me Linh B.S. E, Tay Yen Tu N.R. — F–H, ♂, terminalia, lateral view. F, Ba Be N.P. G, Cham Chu N.R. H, Tay Yen Tu N.R. — I–L, ♀, left metafemur. I, Ba Be N.P. J, Cham Chu N.R. K, Me Linh B.S. L, Tay Yen Tu N.R.

Fig. 17. Medauromorpha regina (Brunner von Wattenwyl, 1907) comb. nov., in nature, ♀, Tay Yen Tu N.R., 10.VII.2013.

Fig. 3. Medauromorpha baviensis sp. nov., paratype ♀. A, habitus, dorsal view. B, habitus, lateral view. C, habitus, ventral view. D, head and thorax, dorsal view. E, terminalia, dorsal view. F, terminalia, ventral view. G, terminalia, lateral view. H, head and thorax, lateral view.