Sadyattes fallax (Hennemann, 2025)

Hennemann, Frank H., 2025, A review of Sadyattes Stål, 1875, with the descriptions of six new species from the Philippines (Phasmatodea: Platycraninae: Stephanacridini), Zootaxa 5610 (1), pp. 1-72 : 26-31

Sadyattes fallax (Brunner v. Wattenwyl, 1907) comb. nov.

( Figs. 6 , 21C–D )

Eucarcharus fallax Brunner v. Wattenwyl, 1907: 186. HT, ♀: Coll. Br. V. W., Luzon, Mus. Berlin; der. Br. v. W., Eucarcharus fallax ; 11.332; Holotypus [NHMW, No. 310].

Bruner, 1915: 231.

Brock, 1998: 28. [Type data]

Otte & Brock, 2005: 136.

Brock & Büscher, 2022: 559.

Further material:

Luzon: 1 ♂: Philippinen, South Luzon , II.1986 local collector [ FH, No. 1522-1] ; 1 ♂: Philippinen, South Luzon Island, Prov. Quezon, Infanta Municipality, local collector [ FH, No. 1522-2] ; ♀: Philippinen, Luzon , leg. Jager [ MNHU] ; ♂ Philippinen. Luzon , leg. Jager " Lonchodes feruloides Westwood det.", Nr. 3228 [ MNHU] ; 1 ♀: Vinai Pl. , III.10.1916 (W. Boettcher), Eucarcharus feruloides (Westw.) ♀ Det. Rehn 1934, Hebard Collection [ ANSP] ; 1 ♀: II.14.1916 [ ANSP] ; ♀: Philippines ( Ledyard ), Eucarcharus feruloides (Westw.) ♀ Det. Rehn 1934, A.N. S.P. [ ANSP]. Samar : 2 ♀♀, 1 ♂, 4 ♀♀ (immatures), 1 ♂ (immature n4), 4 immatures: Philippines, Samar Id., Samar Prov., Hinubangan, San Isidro , San Isidro River , 90–200 m, 31.III.–6.IV.1997, leg. et Coll. R. a. Müller [coll. OZ, No. 330] ; 1 ♂: Philippines, Samar Id. , Samar Prov., Hinubangan, Arizona, Mining C., 280 m, 29.III.–6.IV.1997, leg. et Coll. R.A. Müller [coll. OZ, No. 330]. Panay : 1 ♂, 2 ♀♀, 1 ♀ (immature): Philippines, Panay Island, Mt. Madja-as , 1000 m, VIII.2005, R. Cabale legit [ MG, No’s 0222-2225]. No data : 3 ♀♀: no data (presumably Luzon , leg. Jager) [ MNHU]. Doubtful data : 1 ♂: Dapi, Siargao , K. Günther det. [ MHNG] .

Differentiation. Females of this species show the morphologically closest affinity to the Luzonian S. feruloides ( Westwood, 1859) but may be separated by the stockier overall form (mesothorax just 6.8x longer than prothorax vs. 8x in feruloides ) and more robust limbs, which lack the distinctive dorso-apical tooth on the meso- and metatibiae seen in feruloides , have the cristate dorso-apical lobe of the meso- and metatibiae smaller and the basitarsi less distinctly crested dorsally. Moreover, distinction is rendered possible by the broadly triangular posteromedian notch of the anal segment ( Fig. 6I ; shallow and concave in feruloides ), the somewhat shorter and tapering subgenital plate, that protrudes beyond the tip of the abdomen by less than the combined length of the two terminal terga and has the apex obtusely triangular ( Figs. 6H–J ; spatulate with the lateral margins roughly parallel-sided and the apex broad and roundly angular in feruloides ) as well as the less distinct median node of the praeopercular organ. Males resemble those of the Luzonian S. matipuno sp. nov., Mindanaoan S. leytensis ( Zompro, 1997) and S. maganda sp. nov. from the island of Mindoro, with which they share the general colour pattern and contrastive yellowish head ( Figs. 6D–E ) but differ by the notably slenderer overall shape, more delicate limbs and trapezoidal anal segment, that strongly widens towards the posterior and has the outer angles somewhat protruded and triangular (rather parallel-sided or narrowing posteriorly in matipuno and maganda ). From matipuno they may also be separated by the more globose head, which bears a distinct black postocular streak (a faint green streak in matipuno ), green mesonotum that lacks the black lateral streaks seen in matipuno , shorter alae that do not reach the posterior margin of abdominal segment III and the distinctly annulated meso- and metatibiae. From the ♂♂ of leytensis they may be also be differentiated by the notably larger size (body length <95.0 mm in leytense), relatively larger head and more contrastive and defined postocular streak, shorter alae (reaching about half the way along abdominal segment IV in leytense), that have the costal region not distinctly striped longitudinally, lack of the black apex of the meso- and metafemora seen in leytense, as well as the smaller and narrower, rather triangular vomer (heart-shaped in leytense). Finally, from maganda these ♂♂ are also distinguishable by the relatively larger and much more globose head and presence of a black postocular streak ( Figs. 6E–F ; ovoid and lacking a postocular streak in maganda ), not longitudinally striped costal region of the alae, lack of the black lateral markings and blotches of abdominal terga II-VIII seen in maganda , lack of the black apex of the femora and much smaller and triangular vomer (broad with base almost circular in maganda ). Although the colour pattern is essentially similar, the colouration in fallax is notably less striking and contrastive if compared to maganda . From the known eggs of Sadyattes , those of fallax ( Figs. 21C–D ) most strongly resemble the ones of mindanaense sp. nov. but differ by the more fine-meshed network of ridges and bulges of the chorion, lack of the posteroventral indention seen in the eggs of mindanaense as well as the somewhat smaller and rather pear-shaped micropylar plate, which has the polar extension notably shorter (rather spearhead-shaped in mindanaense ).

Description. Since this species has so far only been known from the ♀ and the very brief and insufficient original description was based on the strongly shrivelled and discoloured holotype only, a detailed redescription of the ♀ as well as a first description of the ♂ are here provided. Since all ♀♀ examined were provisionally stored in spirits, the colouration has certainly faded and care should be taken because the colour of live specimens may be different, for instance might also comprise green tones that are discoloured to yellow in the dried examples at hand. Measurements in table 3 below.

♀ ( Figs. 6A, F–J, N ). Medium-sized (body length incl. subgenital plate 146.5–172.0 mm), form average for the genus with a short subgenital plate, that projects beyond the tip of the abdomen by less than length of anal segment ( Figs. 6H–J ). Colour various tones of ochre, buff and brown and to a variable degree with darker brown and pale greyish to whitish speckles; the meso- and metafemora and tibiae to a variable intensity with three irregular, pale annulae. The holotype rather uniformly ochre with the head, prothorax and posterior portions of the mesonotum and median segment sepia and with a fine but uncontinuous dark brown medio-longitudinal line on the meso- and metanotum and median segment. Moreover, the holotype lacks the annulae of the legs seen in all the other specimens. Genae white in ventral portion and with a faintly indicated brown postocular streak ( Fig. 6F ).

Head ( Fig. 6F ): Sub-oval but somewhat compressed dorso-ventrally and broadest just behind the eyes with the posterior portion narrowed; the vertex roundly convex with a shallow median impression in front, the coronal line fairly distinctly, the lateral furrow slightly indented. Frons with two closely spaced, small and C-shaped impressions between bases of antennae. Eyes rather small and just weakly projecting, round in outline and their diameter corresponding to 0.4x length of gena. Antennae reaching about three-quarters the way along mesonotum; scapus small, rectangular and about 1.5x longer than wide; pedicellus much shorter and round in cross-section.

Thorax: Pronotum noticeably shorter and narrower than head, basically rectangular with the lateral margins gently concave and about 1.7x longer than wide; the anterior margin concave, weakly bi-gibbose medially and followed by a furrow; the transverse median sulcus rather shallow, gently arched and almost expanding over entire width of notum. Mesothorax almost uniform in width and just slightly widened posteriorly, 6.3x longer than prothorax. Mesonotum with a very fine medio-longitudinal carina and at anterior margin with a narrow transverse ridge, which has the median portion somewhat anterior-directed. Metanotum almost parallels-sided, 2.7x longer than wide and somewhat less than half as long as mesonotum; the medio-longitudinal carina scarce. Meso- and metasternum with a weak and shallow medio-longitudinal keel.

Abdomen: Median segment rectangular, 2.3x longer than wide and scarcely more than half the length of metanotum. Segment II about 1.25x longer than median segment, II–IV increasing in length, V as long as IV and VVII decreasing in length with VII shorter than all preceding; IV and V about 2.5x longer than wide. II–VII just very slightly sub-uniform in diameter. Tergum V with a fairly distinct gibbose transverse posterior swelling ( Fig. 6N ). Sterna II–VII smooth and each at best only with a slightly indicated medio-longitudinal carina in anterior portion. Praeopercular organ on VII formed by a short, granular longitudinal ridge some distance off the weakly medially notched posterior margin; the whole area unevenly verrucose and including the ridge black in colour ( Fig. 6J ). Tergum VIII about half as long as VII and slightly widened posteriorly, IV roughly half as long as VIII and scarcely wider than long. Anal segment with a large, almost semi-circular posterolateral emargination ( Fig. 6H ); the anterior portion of lateral margins somewhat deflexed and the posterior margin widely bi-lobed with a small triangular median indention ( Fig. 6I ); dorsal surface with a shallow medio-longitudinal carina. Epiproct small, wider than long with a distinct medio-longitudinal keel and a posteromedian notch; filling the median gap of posterior margin of anal segment but not protruding beyond it ( Fig. 6I ). Cerci fairly large, conical and tapering towards a narrow tip; roughly reaching to posterior margin of anal segment. Gonapophyses VIII up-curved, filiform and notably projecting under anal segment ( Fig. 6I ). Subgenital plate scoop-shaped, obtusely keeled medio-longitudinally and with a prominent ledge-like keel along lateral margins in the basal one-third; apex obtuse, rounded and with the tip somewhat protruded, projecting beyond tip of abdomen by less than length of anal segment ( Figs. 6H–J ).

Legs: Moderately long and fairly stocky with essentially all carinae minutely denticulated; the meso- and metafemora very slightly incrassate sub-basally with the two outer ventral carinae somewhat deflexed. Profemora somewhat longer than mesothorax, mesofemora about four-fifths the length of mesothorax, metafemora almost reaching to posterior margin of abdominal segment IV and metatibiae reaching about halfway along abdominal segment VII. Dentations on anterodorsal carina of profemora prominent and serrate, those of the posteroventral carina slightly smaller and rather tooth-like; posterodorsal carina supplied only with very small denticles. The apical tooth of two outer ventral carinae of the mesofemora strongly enlarged and triangular, often of contrastive black colour. Medioventral carina of meso- and metafemora supplied with about 15 small spines; that of the meso- and metatibiae raised and forming a distinct, rounded lobe sub-basally. Meso- and metabasitarsus about as long as following three tarsal joints taken together with all three carinae minutely but densely denticulate; probasitarsus notably longer than following three tarsomeres combined and wholly unarmed; all with dorsal carina slightly raised.

♂ ( Figs. 6B–E, G, K–M ). Of average size (body length 116.3–118.5) and typical form for the genus, with a sub-spherical, yellow head with a dark postocular streak, a black transverse band on the abdominal terga II-VI and short alae, that on average only reach only halfway along abdominal segment III (length 24.5–26.6 mm). Colour basically olive with the abdomen ochre to buff. Head straw-coloured to yellow with a rather slender and washed brown postocular streak that vanishes towards the posterior of gena ( Fig. 6E ). Antennae brown with the basal eight or so joints black ventrally, the basal three joints blackish brown also dorsally. Prothorax greyish buff. Mesonotum occasionally with a slight brownish was dorsally. Tegmina and costal region alae greyish green to cream with the anterior margin marked by a broad creamy yellow band, which is interiorly bordered by a narrow brown longitudinal stripe; longitudinal veins green; anal fan hyaline with a slight greyish hue towards the outer margin and with all veins dark olive. Abdominal terga and sterna II-VI with a black transverse marking at posterior margin, which is broadest on II and becomes gradually narrower towards VI. Tergum VIII with a washed cream-coloured posterolateral marking, IX with a similar but more elongate marking. Profemora ochre with the compressed and curved base dark green, meso- and metafemora olive with the apex slightly brownish ( Fig. 6G ). Protibiae orangey ochre, meso- and metatibiae greyish brown with three distinct, broad yellow annulae.

Head ( Figs. 6D–E ): Strongly globose, sub-spherical with vertex convex; broadest at the eyes and noticeably narrowing towards posterior; coronal line and lateral furrow fine and weakly indented. Vertex with two shallow indentions in front, the frons with a small, but deep transverse pit between bases of antennae. Eyes very large, projecting sub-spherically and their diameter corresponding to 0.66x length of gena. Antennae reaching about halfway along abdominal segment III; scapus roundly rectangular and 1.6x longer than wide, pedicellus roughly two-thirds the length of scapus and round in cross-section.

Thorax: Pronotum basically as in ♀ but the lateral margins less concave and the anterior margin with the two median swellings much smaller ( Figs. 6D–E ). Mesothorax slender, elongate and just weakly widened posteriorly; 7.5x longer than prothorax. Mesonotum with a very fine medio-longitudinal line that is only clearly recognized in the anterior portion; the anterior margin with a low and almost straight transverse ridge ( Figs. 6D–E ). Meso- and metasternum with a fairly distinct medio-longitudinal keel, which is most prominent in the anterior section of mesosternum. Tegmina elongate, spatulate in shape with the basal two-thirds gradually narrowing and the apical one-third rounded; central hump very shallow. Alae somewhat variable in length, reaching no more than two-thirds the way along abdominal segment III (occasionally just reaching to posterior margin of II).

Abdomen: Segments II–V slightly increasing, VI and VII decreasing in length with VII shorter than all preceding, all except for VII uniform in diameter; V about 6.3x longer than wide, VII somewhat constricted medially and about two-thirds the length of V. Terga V and VI with a slight posteromedian swelling. Sterna with a distinct medio-longitudinal keel. Tergum VIII distinctly trapezoidal in outline with posterior margin about 1.3x wider than anterior margin, about 0.6x length of VII and broadest of all segments. IX constricted medially with posterior portion noticeably narrower than anterior portion and scarcely longer than VIII; dorsal surface strongly convex longitudinally. Anal segment much shorter than IX, the anterior portion broad with lateral surface convex and the posterior portion narrowed and strongly tectate longitudinally; posterior margin separated by a distinct transverse furrow, the outer angles rounded and inflated ( Figs. 6K–L ) and the ventral surfaces minutely denticulate and facing each other at an angle of no more than 40°. Vomer small, roundly triangular in outline, longer than width across base, the lateral portions inflated and the central portion impressed longitudinally; terminal hook rather strong and up-curved ( Fig. 6M ). Cerci round in cross-section with apex very slightly club-shaped, gently up-curved and projecting notably beyond anal segment. Phallus projecting notably over posterior and left lateral margin of poculum, rather broad and pointed apically ( Figs. 6K–M ). The poculum of moderate size, bowl-shaped with base bulgy and rounded ( Fig. 6K ), the posterior margin somewhat labiate, narrowed and scarcely reaching to posterior of tergum IX ( Fig. 6M ).

Legs: Long, slender and with all carinae minutely and fairly regularly dentate; the medioventral carina of meso- and metafemora narrow and only supplied with about ten (mesofemora) or fifteen (metafemora) very minute spines. Profemora notably longer than head, pro- and mesothorax taken together, mesofemora about as long as pro- and mesothorax combines, metafemora reaching roughly halfway along abdominal segment VI and metatibiae projecting beyond tip of abdomen by combined length of six terminal abdominal segments. Tarsi elongate, meso- and metabasitarsi as long as remaining joints taken together, probasitarsus slightly longer; all destitute of teeth.

Egg ( Figs. 21 C– D). Two eggs were extracted from the abdomen of the holotype in NHMW and can serve for a brief characterisation. Care should be taken however, because these eggs may not be fully developed. This in particular concerns to the sculpturing of the chorion as well as the colouration .

Medium-sized, chorion longer than high, flattened laterally and distinctly oval in cross-section; polar-area rounded; on the dorsal surface these ridges indicate a pair of arched, longitudinal bulges. Anterior margin bulgy. Surface wholly covered by an irregular meshwork of shallow keels that leave impressed areas inbetween them; surface of the indented areas very minutely granular. Micropylar plate very small with the broaded anterior portion oval and somewhat narrowed anteriorly and the lower portion strongly narrowed with the raised outer margin fused. Internal surface concave and with a small bowl-shaped micropylar cup at lower end of widened anterior area. Operculum elliptical and noticeably arched with the dorsal and ventral portions downward-directed.Anterior portion with a small oval pit. Capitulum an irregular, mushroom-like protrusion that consists of about seven irregular ridges that fuse in centre. Colour yellowish ochre with the raised portions of the chorion straw-coloured. Micropylar plare, operculum and capitulum orangey brown.

Measurements [mm]: Length incl. operculum 3.9, length 3.4, width 2.3, height 2.8, length of micropylar plate 1.3.

Remarks. This species was originally described upon a unique ♀ from the island of Luzon, Philippines in the collection of NHMW. Since, the data label states “Luzon, Mus. Berlin” it is most likely that the specimen is part of the series of specimens labelled “Luzon, leg. Jager” in the collection of MNHU. All have provisionally been preserved in spirits and the right mid leg glued to the holotype in NHMW is apparently a hind leg, that does not belong to the specimen. Frequently, specimens of this species have been misidentified as S. feruloides ( Westwood, 1859) . In fact, fallax is the more commonly encountered species throughout Luzon and exhibits a remarkable distributional pattern within the Philippines, since it is also found in the Eastern Visayas (Samar) and Western Visayas (Panay). On the island of Luzon, fallax appears to be restricted to the southern half of the island, although many more records will be needed for a delimitation of its distribution. The collection of MHNG houses a ♂ that is stated to be from Dapa on the island of Siargao northeast of Mindanao. This distribution however is doubtful because it violates biogeographic data, that regard Siargao as part of the Mindanao Region, which essentially has no common species with the Greater Luzon Region or the Visayas. Therefore, this record is here excluded from the known distribution of fallax and deserves further evaluation.

Distribution. Philippines. Luzon: “Luzon” [MNHU]; Province Quezon (Infanta Municipality [FH]). Samar: Province Samar (Hinubangan, San Isidro, San Isidro River, 90–200 m [OZ]). Panay: Province Antique (Mt. Madja-as, 1000 m [MG]).

Figures

FIGURE 6. Sadyattes fallax (Brunner v. Wattenwyl, 1907) comb. nov.: A. Habitus of ♀ in dorsolateral view [MNHU]; B: Habitus of ♂, dorsal view [FH 1522-2]; C. Habitus of ♂, dorsolateral view [FH 1522-2]; D. Head and prothorax of ♂, lateral view [FH 1522-2]; E. Head and prothorax of ♂, dorsal view [FH 1522-2]; F. Head and prothorax of ♀, lateral view [MNHU]; G. Knee-region of left midleg, anterolateral view [FH 1522-1]; H. Terminalia of ♀ in lateral view [MNHU]; I. Terminalia of ♀ in dorsal view [MNHU]; J. Terminalia of ♀ in ventral view [MNHU]; K. Terminalia of ♂ in lateral view [FH 1522-1]; L. Terminalia of ♂ in dorsal view [FH 1522-1]; M. Terminalia of ♂ in ventral view [FH 1522-1]; N. Terminal six abdominal segments of ♀ in dorsolateral view with arrow indicating the posteromedian protuberance of tergum V [MNHU]. Pha = phallus.

FIGURE 21. Eggs of Sadyattes spp., dorsal view (left) and lateral view with micropylar plate at left hand side (right): A–B. S. mindanaense sp. nov. (paratype); C–D. S. fallax (Brunner v. Wattenwyl, 1907) comb. nov.; E–F. S. feruloides (Westwood, 1859) comb. nov.; G–H. S. leytensis (Zompro, 1997) comb. nov.; I–J. Sadyattes matipuno sp. nov. (paratype); K–L. S. decoris (Seow-Choen, 2016) comb. nov. [© Francis Seow-Choen]; M–N. S. incertus (Redtenbacher, 1906) comb. nov. from ovipositor of holotype of synonym Nearchus grubaueri Redtenbacher, 1908 [© NHMW, Harald Bruckner]; O–P. S. enganensis (Redtenbacher, 1908) [© Francis Seow-Choen].