Sadyattes Stål, 1875

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 : 8-13

Sadyattes Stål, 1875

Type species: Sadyattes borrii Stål, 1875: 88 , by original monotypy.

Sadyattes Stål, 1875: 44 , 88.

Kirby, 1904: 358. [in Phryganistriinae]

Redtenbacher, 1908: 444. [in Acrophyllini ]

Bradley & Galil, 1977: 193. [in Phasmatinae : Pharnaciini ] Bragg, 2001: 643.

Zompro, 2004: 321.

Otte & Brock, 2005: 311,

Hennemann & Conle, 2008: 55, 60 [in Stephanacridini ] Seow-Choen, 2017: 150. [in Xeroderinae – in error] Seow-Choen, 2018: 452.

Hennemann, 2020: 175. [in Stephanacridini ]

Seow-Choen, 2021: 772.

Brock & Büscher, 2022: 559.

= Eucarcharus Brunner v. Wattenwyl, 1907: 185 (in part). syn. nov. Günther, 1935b: 138. [Designation of type species]

Bradley & Galil, 1977 193. [in Phasmatinae : Pharnaciini ] Bragg, 2001: 633.

Zompro, 2004: 310.

Otte & Brock, 2005: 136.

Hennemann & Conle, 2008: 132 [in Stephanacridini ]

Bollens, Krijns & Bresseel, 2010a: 10.

Hennemann, 2020: 175. [in Stephanacridini ]

Brock & Büscher, 2022: 559.

Seow-Choen, 2023: 516.

Baculolonga, Bragg, 2001: 377 (in part).

Lonchodes, Westwood, 1859: 45 (in part).

Pharnacia, Kirby, 1904: 359 (in part).

Redtenbacher, 1908: 449 (in part).

Phobaeticus, Brock, 1996: 28 (in part).

Seow-Choen, 2016: 265 (in part).

Seow-Choen, 2018: 421 (in part).

Seow-Choen, 2020: 244 (in part).

Seow-Choen, 2021: 734 (in part).

Diagnosis (♂, ♀). Medium to large (body length ♂♂ 72.3–142.0 mm, ♀♀ incl. subgenital plate 127.8–245.0 mm) almost exceptionally slender and elongate, stick-like Stephanacridini (only ♀ of one species stocky with body notably broadened); females apterous, ♂♂ winged. Body of both sexes smooth and unarmed, often slightly shiny in ♂♂. Sexual dimorphism strongly developed with ♂♂ much smaller and slenderer than ♀♀. Head ovoid to globose, longer than wide with vertex smooth; no ocelli. Gula small, variable in shape and covering less than half of cervical membrane. Antennae of moderate length, filiform and laid back at least reaching to posterior of mesothorax; comparatively longer in ♂♂ and may reach to abdominal segment V; scapus small, slender and not notably dilated and just moderately compressed dorsoventrally. Mesothorax> 3.5x longer than prothorax, usually much longer and slenderer in ♂♂; cylindrical, slender and ± uniform in diameter (only in one species with lateral margins notably deflexed and rounded); mesonotum with a ± distinct transverse ridge or bulge at anterior margin. Meso- and metapleurae simple, unarmed. Meso- and metanotum with a ± distinct medio-longitudinal carina or keel. Median segment in ♀♀ at best three-quarters the length of metanotum, much longer than metanotum in ♂♂. Tegmina in ♂♂ elongate and spatulate in shape with the basal half strongly narrowed and the central protuberance shallow. Alae variable in length and moderately developed, at best and only in one species reaching to abdominal segment V, mostly much shorter; anal fan transparent grey or brown. Abdomen excluding median segment longer than head and thorax taken together. Abdominal segments II-VII slender, much longer than wide and uniform in diameter in ♂♂; in ♀♀ less elongate and slightly sub-uniform in diameter (only in one species with lateral margins of terga weakly expanded and rounded). Sterna II-VII with a ± distinct medio-longitudinal carina or keel (more pronounced in ♂♂). Sternum VII in ♀♀ with a moderate praeopercular organ that is formed by a posteromedian pit and a variably shaped granule or tubercle in front. Terminalia of ♀♀: Anal segment weakly tectate and with a shallow concave to triangular posteromedian notch, the lateral margins with a distinct sometimes semi-circular excavation at base of cerci with the basal portion ± deflexed and occasionally lobe-like. Epiproct small, scale-like and scarcely projecting underneath posterior margin of anal segment; keel medio-longitudinally. Cerci small, round in cross-section and ± tapering towards the tip. Gonapophyses VIII elongated, rather filiform and up-curved but never considerably projecting beyond anal segment. Subgenital plate bulgy in basal portion, variable in length and shape and always projecting beyond tip of abdomen by at least two-thirds the length of anal segment and as much as slightly more than the length of the three terminal terga taken together; the flattened posterior half ranging from broad, spatulate and roundly angular to strongly tapering, pointed and lanceolate. Terminalia of ♂♂: Tergum VIII slightly to distinctly inflated and widened posteriorly, but three terminal segment usually not distinctly broader than preceding segment. Anal segment simple, gently tectate longitudinally with the posterior margin ± emarginated and the outer angles ± expanded and triangular and with a distinct and large thorn-pad ventrally. Epiproct very small and fully concealed under anal segment. Vomer a strongly sclerotised, variably shaped plate that ranges in shape from triangular over heart- or pear-like to having the basal portion sub-circular; always with a single ± arched terminal hook. Phallus enlarged, elongated and often projecting over upper lateral margin of poculum. Cerci fairly small, elongate, round in cross-section, ± curved and with the apex ± club-like. Poculum small and at best scarcely projecting beyond posterior margin of tergum IX; basal portion weakly to moderately bowl-shaped and only in two species with a small, obtuse central protuberance; the posterior margin ± labiate and occasionally with a slight median indention. Legs all long and slender (only ♀♀ of two species have noticeably incrassated meso- and metafemora), even more so in ♂♂; all carinae to a variable degree dentate or serrate and usually no considerably enlarged teeth or lobes present on mid and hind legs except for an enlarged apical tooth on two outer ventral carinae of meso- and metafemora. Medioventral carina of meso- and metafemora dentate to spinose. Armature of limbs generally less developed in ♂♂. Probasitarsus with dorsal carina unarmed; ventral carinae of all protarsi minutely dentate. Dorsal carina of basitarsi in ♀♀ ± raised or crested (occasionally strongly deflexed in probasitarsus); basitarsi always slender in ♂♂; in both sexes notably elongated and at least as long as following three tarsomeres taken together.

Eggs ( Fig. 21 ). Of average size if compared to the eggs of other genera of the tribe and comprising a wide range of different chorion shapes and surface textures; mostly not noticeably longer than wide (2x longer than wide only in one species) and more or less compressed laterally (strongly globose with a smooth surface only in one species). Dorsal surface often with two longitudinal carinae or bulges. Polar area mostly rounded but may be flattened (one species) or with a central protrusion (one species). Surface of chorion often with a mesh-work of ridges, keel or bulges, occasionally pitted to a variable degree. Micropylar plate less than three-fifth the length of chorion and spearhead-shaped to rhomboidal; the lower portion more or less distinctly narrowed; closed internally. Micropylar cup fairly central within the borders of the micropylar plate. Operculum oval and flattened; often arched with the two ends facing the dorsal and ventral egg surfaces distinctly downward-directed and in these species with a conspicuous pit or impression in the portion facing the dorsal egg surface. Capitulum not stalked, variable in size and shape.

Differentiation. Sadyattes is morphologically closest to Nesiophasma , which is distributed throughout most of Wallacea ( Hennemann, 2021: 103), and presumably the sister taxon. Based on the available data there seems to be no distributional overlap between these two genera, which means that they are biogeographically separated by the Wallace’s Line that runs between Borneo and the Philippines in the west and north, and Sulawesi in the east. The islands east of Wallace’s Line are termed Wallacea which is principally regarded a transition zone between the Oriental and Australian regions ( Dickerson, 1928). Although the Philippines are now also considered as part of Wallacea (e. g. Vallejo, 2011), which is in accordance with the interpretation of Wallace’s Line after Huxley (1868) separating the Philippines from Borneo, the distribution of Sadyattes however reflects the faunal boundary originally intended by Wallace that includes the Philippines on the west side of the line ( Wallace, 1876). Such, the Oriental Sadyattes has seven of its fourteen known species distributed in the Philippines, whereas the Wallacean Nesiophasma has no representatives in the Philippines ( Hennemann, 2021: 104).

Whereas ♀♀ of both genera are virtually identical in means of morphology and very difficult to separate from another ( Hennemann, 2021: 104), ♂♂ of Sadyattes readily differ from the consistently apterous ♂♂ of Nesiophasma by the presence of tegmina and alae and therefore much longer median segment, that is considerably longer than the metanotum. In ♀♀ on Nesiophasma the anal segment shows a deep and narrow posteromedian incision or slit, whereas in Sadyattes there merely is a shallow concave to triangular excavation. The mostly distinct lateral excavation of the anal segment at the base of the cerci on the other hand is shared between both genera. Females have the gonapophyses VIII always hidden under and never projecting beyond the tip of the anal segment like in most Nesiophasma -species, but there is one species of Nesiophasma , namely N. plateni ( Dohrn, 1910) that also has short gonapophyses VIII ( Hennemann, 2021: 117, Figs. 45P-R) and therefore renders this possible distinguishing character not fully reliable. This is also the case with the variable subgenital plate, which is fairly short and broadly spatulate in several species, a condition never seen in Nesiophasma , but other species have it long and lanceolate like in Nesiophasma -species. The known eggs of Sadyattes are remarkably polymorphic and show a wide range of different shapes and chorion surface textures that also render a reliable delimitation and distinction from eggs of Nesiophasma impossible. All eggs, except for those of the Bornean S. decoris ( Seow-Choen, 2016) comb. nov. which is here transferred from the Pharnaciini genus Phobaeticus Brunner v. Wattenwyl, 1907, have the chorion more bulgy and less elongate than in Nesiophasma being not considerably longer than wide. The eggs of several Philippine species have a conspicuous pit or impression in the portion of the operculum that faces towards the dorsal egg surface and moreover have the operculum notably arched with the two ends facing the dorsal and ventral egg surfaces distinctly downward-directed. Both features are not seen in eggs of Nesiophasma . Altogether, reliable morphological differences between Sadyattes and Nesiophasma , other than the presence of absence of wings in the ♂♂ and minor differences mentioned above, are very hard to define why it is hoped that molecular data can assist with the delimitation of these two genera in due course.

Remarks. Stål (1874: 44, 88) originally described Sadyattes based on a unique ♂ without locality in the collection of RBINS, namely S. borrii , and the true identity of the genus has long been questionable although subsequent authors linked it with taxa that now belong in Clitumninae : Pharnaciini . Hennemann & Conle (2008: 55, 60) examined the holotype specimen in detail and transferred Sadyattes from Pharnaciini to the tribe Stephanacridini Günther, 1953 , mainly based on morphological characteristics of the ♂ terminalia, like the presence of a well-developed vomer and simple, non-split anal segment. Still however, the identity of the genus has remained unsolved because the ♀ and eggs were not known. It was until now that the availability of new material of the type-species S. borrii Stål, 1875 has rendered a clarification of the taxonomic position and delimitation of Sadyattes within Stephanacridini possible and proven this species to be distributed on Java and Sumatra. It also became obvious that Stål’s borrii is congeneric to the type-species of Eucarcharus Brunner v. Wattenwyl, 1907, the Philippine E. feruloides ( Westwood, 1859) , why Eucarcharus needs to be synonymised under Sadyattes (syn. nov.). This has already been suggested previously by Hennemann (2021: 104). Moreover, further investigation of the two type specimens of Westwood’s feruloides , one from the Philippines (lectotype) and one from Java (paralectotype), has shown that these represent two distinct species, which has already been suggested by Seow-Choen ( Seow-Choen, 2023: 516). In fact, the Javanese paralectotype is a specimen of borrii , the type-species of Sadyattes . Comprehensive investigation of species with questionable generic affiliations within Hermarchus Stål, 1875 (tribe Stephanacridini ) and Phobaeticus Brunner v. Wattenwyl, 1907 ( Clitumninae : Pharnaciini ) has uncovered three further species that actually belong in Sadyattes and are here transferred (comb. nov.).

It seems worth mentioning that members of Sadyattes are remarkably similar in overall appearance to species of Phobaeticus Brunner v. Wattenwyl, 1901, a genus of Clitumninae : Pharnaciini , that has frequently been shown to be not closely related (e. g. Hennemann & Conle, 2008; Bradler, 2009; Buckley et al., 2009; Robertson et al., 2018, Bank & Bradler, 2022; Forni et al., 2023). Representatives of both genera have widely overlapping distributions, are large to very large slender stick insects with wingless ♀♀ and much smaller, more colourful winged ♂♂ and often have a more or less elongated subgenital plate in the ♀♀. It may be for these reasons why, several of the species here transferred to Sadyattes have previously been misplaced in Phobaeticus . One species originally described in the genus Phobaeticus and the world’s longest known insect, the Bornean Phobaeticus chani Bragg, 2008 , has even been erroneously attributed to Sadyattes subsequently but was re-transferred to Phobaeticus ( Hennemann, 2021: 104) . As argued by Hennemann (2021: 104) Phobaeticus chani shows all the distinctive morphological traits that separate members of Clituminiae: Pharnaciini from the tribe Stephanacridini , this the split anal segment in ♂♂ that consists of two hemi-terga, lack of a vomer, short gonapophyses in ♀♀ and the lamellate, displaced medioventral carina of the profemora.

Distribution ( Fig. 1 ). Java, Sumatra (including the surrounding islands of Banka, Enggano and Nias), Peninsular Malaysia, Nicobar Islands, Borneo and Philippines.

Species included:

1. Sadyattes annulatus ( Redtenbacher, 1908: 451) [ Pharnacia ]. ( Fig. 2 )

Distribution: Borneo.

2. Sadyattes banwaon sp. nov. ( Fig. 3 )

Distribution: Philippines (Mindanao).

3. Sadyattes borrii Stål, 1875: 88 . ( Figs. 4–5 )

= Nesiophasma zanus Hennemann, 1999: 213 , Figs. 1–2 . syn. nov.

Distribution: Java and Sumatra.

4. Sadyattes decoris ( Seow-Choen 2016: 267, figs.) [ Phobaeticus ]. comb. nov.

Distribution: Borneo.

5. Sadyattes enganensis ( Redtenbacher, 1908: 451) [ Pharnacia ].

Distribution: Enggano Id. and Nias Id. East of Sumatra and Peninsular Malaysia.

6. Sadyattes fallax (Brunner v. Wattenwyl, 1907: 186) [ Eucarcharus ]. comb. nov. ( Fig. 6 )

Distribution: Philippines (Luzon, Samar and Panay).

7. Sadyattes feruloides ( Westwood, 1859: 45, pl. 6: 5) [ Lonchodes ]. comb. nov. ( Fig. 7 )

Distribution: Philippines (Luzon).

8. Sadyattes incertus (Brunner v. Wattenwyl, 1907: 185) [ Phobaeticus ]. comb. nov. ( Figs. 8–9 ) = Nearchus grubaueri Redtenbacher, 1908: 448 . syn. nov.

Distribution: Sumatra, Banka Id., Peninsular Malaysia and Nicobar Islands. 9. Sadyattes leytensis ( Zompro, 1997: 38, figs.) [ Hermarchus ]. comb. nov. ( Figs. 10–11 )

Distribution: Philippines (Leyte, Mindanao and Panay).

10. Sadyattes maganda sp. nov. ( Figs. 12–13 )

Distribution: Philippines (Mindoro).

11. Sadyattes matipuno sp. nov. ( Figs. 14–15 )

Distribution: Philippines (Luzon).

12. Sadyattes mindanaense sp. nov. ( Figs. 16–18 )

Distribution: Philippines (Mindanao).

13. Sadyattes nigricornis ( Redtenbacher, 1908: 451) [ Pharnacia ].

Distribution: Borneo.

14. Sadyattes panayense sp. nov. ( Fig. 19 )

Distribution: Philippines (Panay)

15. Sadyattes tubaense sp. nov. ( Fig. 20 )

Distribution: Philippines (Luzon)

Keys to the species of Sadyattes

♀♀ *

1. Body slender and stick-like; lateral margins of mesonotum slender.............................................. 2

- Body stocky; lateral margins of mesonotum and abdominal terga II-VII dilated and rounded; colour uniformly bright green; Philippines (Leyte, Panay & Mindanao); Fig. 10 ...................................................... leytensis

2. Protarsus light cream-coloured to whitish; abdominal terga VIII-X with a distinct lateral lobe; not Borneo or Philippines... 3

- Protarsus not conspicuously contrastive in colour; no distinct lateral lobes on abdominal terga VIII-X.................. 4

3. Abdominal segments III-VI almost uniform in diameter; apex of subgenital plate obtuse ( Figs. 4F–G ); Java & Sumatra ..................................................................................................... borrii

- Abdominal segments III-VI notably inflated and swollen; apex of subgenital plate acute; Peninsular Malaysia, Enggano Id. & Nias Id. (east of Sumatra)....................................................................... enganensis

4. Subgenital plate short, projecting beyond abdomen by no more than combined length of two terminal abdominal terga, apex broad and obtuse; Philippines............................................................................ 5

- Subgenital plate long and more or less lanceolate with apex acute, projecting beyond abdomen by more than combined length of two terminal abdominal terga.......................................................................... 7

5. Mesothorax <7x longer than prothorax; no enlarged dorsa-apical tooth on meso- and metafemora; subgenital plate tapering towards apex......................................................................................... 6

- Slender species, mesothorax 8x longer than prothorax; meso- and metafemora with a distinct triangular ventral tooth apically ( Fig. 7D ); subgenital plate broadly spatulate with lateral margins parallel-sided and apex broad and roundly angular (Figs.- 7B–C); Luzon................................................................................. feruloides

6. Posteromedian indention of anal segment broadly triangular ( Fig. 6I ); ventral dentations of meso- and metafemora small; Luzon, Samar & Panay.............................................................................. fallax

- Median indention of anal segment shallow and rather concave ( Figs. 17C–D ); ventral dentations of meso- and metafemora distinct and more spinose in the basal half ( Fig. 17I ); Mindanao................................ mindanaense sp. nov.

7. Slender species; mesothorax roughly uniform in diameter; meso- and metafemora slender.............................8

- Stocky species ( Figs. 14A–C ); mesothorax notably constricted anteriorly ( Fig. 14E ); meso- and metafemora strongly incrassated ( Fig. 14F ); East Luzon.................................................................... matipuno sp. nov.

8. Subgenital plate projecting beyond abdomen by no more than combined length of abdominal terga VIII-X; head sub-globose with vertex distinctly convex; Philippines & Borneo.......................................................... 9

- Subgenital plate long, lancet-like and projecting beyond abdomen by more than combined length of abdominal terga VIII-X ( Figs. 8F–H ); head elongated, narrowed posteriorly with vertex flattened ( Figs. 8 C-D); Peninsular Malaysia, Bangka Island & Nicobar Islands.................................................................................. incertus

9. Two outer ventral carinae of tibiae narrow; posterodorsal carina of profemora with definite but small teeth; Philippines... 10

- Two outer ventral carinae of tibiae notably widened; posterodorsal carina of profemora with very minute denticles; Borneo................................................................................................ decoris

10. Body length incl. subgenital plate> 140.0 mm (usually notably larger); epiproct small and full or mostly Concealed under anal segment; meso- and metatibiae not crested apically; not Mindanao............................................. 11

- Smaller species (body length incl. subgenital plate 139.0 mm); epiproct large, shield-shaped and projecting notably beyond anal segment ( Fig. 3H ); posterodorsal carina of meso- and metatibiae forming a distinct rounded apical crest ( Fig. 3A ); Mindanao...................................................................................... banwaon sp. nov.

10. Head unicoloured ( Figs. 19 D-E); dentations on two outer ventral carinae of meso- and metafemora spinose and no conspicuously enlarged apical tooth present ( Fig. 19F ); Panay................................................ panayense sp. nov.

- Head with dark postocular markings (indicating a streak; Fig. 12D ); dentations on two outer ventral carinae of meso- and metafemora represented by distinct and black serrate teeth, the apical tooth distinctly enlarged ( Figs. 12E–F ); Mindoro........................................................................................... maganda sp. nov.

* ♀♀ of S. annulatus ( Redtenbacher, 1908) and S. tubaense sp. nov. are not known.

♂♂ *

1. Head with a distinct dark postocular streak; only Philippines................................................... 2

- No postocular streak on genae........................................................................... 4

2. Mesonotum unicoloured; abdominal terga II–VI with a black transverse marking at posterior margin................... 3 Mesonotum and mesopleurae with a black longitudinal lateral line; abdominal terga II–VI wholly black; Luzon.. .. feruloides

3. Small species (body length <100.0 mm); mesofemora shorter than mesothorax; vomer with base almost parallel-sided and then strongly tapering towards apex ( Fig. 18H ); Mindanao........................................ mindanaense sp. nov.

- Larger species (body length> 110.0 mm); mesofemora as long as mesothorax; vomer triangular and tapering towards apex ( Fig. 6M ); Luzon, Samar & Panay......................................................................... fallax

4. Alae long and reaching to abdominal segment V; Borneo...................................................... 5

- Alae shorter and at best reaching to abdominal segment IV; not Borneo.......................................... 6

5. Large (body length> 130.0 mm); abdominal sterna II-VII with a black posterior marking..................... annulatus

- Small (body length 104.0 mm**); abdominal sterna II-VII with a white pre-posterior marking................ nigricornis

6. Poculum with a distinct but obtuse medio-basal protuberance; protarsus light cream-coloured to whitish................ 7

- No medio-basal protuberance on poculum; protarsus not conspicuously contrastive in colour......................... 8

7. Abdominal terga II–VI 6x longer than wide; apex of abdomen not conspicuously wider than preceding segments with tergum VIII just slightly widening towards posterior; two outer ventral carinae of meso-femora orange ( Figs. 5C–D ); Java & Sumatra ............................................................................................... borrii

- Abdominal terga II–VI 4.5x longer than wide; apex of abdomen swollen and club-like, with tergum VIII strongly inflated and trapezoidal in dorsal aspect (posterior margin almost 2x wider than anterior margin); two outer ventral carinae of mesofemora not of contrastive colour; Peninsular Malaysia, Sumatra, Enggano Id. & Nias Id........................... .. enganensis

8. Head sub-globose with vertex distinctly convex; alae at best reaching halfway along abdominal segment IV; Philippines... 9

- Head elongate-ovoid, narrowed and angular posteriorly and vertex flattened ( Figs. 9B–C ); alae reaching to Posterior margin of abdominal segment IV ( Fig. 9A ); Peninsular Malaysia, Bangka Island & Nicobar Islands....................... incertus

9. Anal segment not widened posteriorly; no black lateral streak on mesonotum..................................... 10

- Anal segment widening towards the posterior with the outer angles deflexed and triangular ( Figs. 15G–H ); mesonotum with an irregular black lateral streak ( Figs. 15D–E ); E-Luzon............................................ matipuno sp. nov.

10. Colourful insects; head of contrastive lighter and different colour than rest of body; tibiae distinctly annulated and dentations of all legs distinct.....................................................................................11

- Mostly green/olive insect ( Figs. 20A–B ); head not of contrastive colouration ( Figs. 20C–D ); tibia only with faintly indicated annulae and dentations of all legs notably reduced and minute; Luzon............................... tubaense sp. nov.

11. Alae reaching no more than halfway along abdominal segment III; abdomen orange and terga II-VII to a variable degree with black lateral markings ( Figs. 13E–F ); costal region of alae dark brown with cream-coloured longitudinal veins; vomer broad with base sub-circular ( Figs. 13I, L ); Mindoro.................................................. maganda sp. nov.

- Alae reaching to abdominal segment IV; abdomen green and terga II-VII with a black transverse posterior band; costal region of alae cream-coloured with brown longitudinal veins; vomer heart-shaped ( Fig. 11G ); Leyte, Panay & Mindanao. .. leytensis

* ♂♂ of S. decoris ( Seow-Choen, 2016) , S. mjoebergi (Günther, 1935) and S. panayense sp. nov. are not known.

** According to Redtenbacher (1908: 451)

Figures

FIGURE 1. Map of Southeast Asia, showing the distributional range of Sadyattes Stål, 1875 (Source: https://aseanup.comfreemaps-asean-southeast-asia)

FIGURE 2. Sadyattes annulatus (Redtenbacher, 1906) ♂ from Borneo, Kalimantan Barat, Mount Bawang [FH 1390-1]: A. Habitus, dorsal view; B. Head and prothorax, dorsolateral view; C. ♀ Detail of tegmina and alar bases, dorsal view; D. Kneeregion of left midleg, anterolateral view; E. Terminalia in lateral view; F. Terminalia in dorsal view; G. Terminalia in ventral view.

FIGURE 3. Sadyattes banwaon sp. nov. ♀ holotype [RBINS]: A. Habitus, dorsolateral view; B. Habitus, dorsal view; C. Habitus, ventral view; D. Knee-region of left midleg, anterolateral view; E. Head and prothorax, lateral view; F. Head and prothorax, dorsal view; E. Terminalia in lateral view; F. Terminalia in dorsal view; G. Terminalia in ventral view.

FIGURE 4. Sadyattes borrii Stål, 1875 ♀ (holotype of Nesiophasma zanus Hennemann, 1999 [RBINS]): A. Habitus, dorsal view; B. Habitus, dorsolateral view; C. Head and prothorax in lateral view, showing the typical cream-coloured and crested probasitarsus; D. Knee-region of left hindleg, posterolateral view; E. Terminalia in lateral view; F. Terminalia in ventral view; G. Terminalia in dorsal view.

FIGURE 5. Sadyattes borrii Stål, 1875 ♂: A. Habitus of specimen from Sumatra, dorsolateral view [FH 1523-1]; B. Habitus of specimen from Sumatra, dorsal view [FH 1523-1]; C. Knee-region of left midleg, anterolateral view [FH 1523-1]; D. Dorsolateral view of tegmina and alae of specimen from Java [ZMUH]; E. Head and prothorax of specimen from Java, lateral view [ZMUH]; F. Head and prothorax of specimen from Sumatra, dorsal view [FH 1523-1]; G. Six terminal abdominal segments of holotype in lateral view (arrow indicating the posteromedian swelling of tergum V) [RBINS, © Jéróme Constant]; H. Terminalia of specimen from Sumatra, lateral view [FH 1523-1]; I. Terminalia of specimen from Sumatra, dorsal view [FH 1523-1].; J. Terminalia of specimen from Sumatra, ventral view [FH 1523-1]. Al = ala, Mes = mesonotum, Met = metanotum, Pha = phallus, Teg = tegmen.

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 7. Sadyattes feruloides (Westwood, 1859) comb. nov. ♀ holotype [NHMUK, © Francis Seow-Choen]: A. Habitus in dorsolateral view; B. Terminalia in dorsal view; C. Terminalia in ventral view; D. Left midleg in posterolateral view.

FIGURE 8. Sadyattes incertus (Redtenbacher, 1906) comb. nov. ♀ from Peninsular Malaysia, Perak [ZMUH]: A. Habitus, dorsal view; B. Habitus, dorsolateral view; C. Head and prothorax, lateral view; D. Head and prothorax, dorsal view; E. Mesofemur in ventral view; F. Terminalia in ventral view; G. Terminalia in dorsal view; H. Terminalia in lateral view.

FIGURE 9. Sadyattes incertus (Redtenbacher, 1906) comb. nov. ♂ from Peninsular Malaysia, Kuala Kangsar [ZMUH]: A. Habitus, dorsolateral view; B. Head and prothorax, dorsal view; C. Head and prothorax, lateral view; D. Terminalia in lateral view; E. Terminalia in ventral view; F. Terminalia in dorsal view; G. Two terminal abdominal segments in ventro-caudal view. (Cerci missing). Pha = phallus, TP =thorn pads

FIGURE 10. Sadyattes leytensis (Zompro, 1997) comb. nov. ♀ Philippines, Mindanao [FH]: A. Habitus, dorsolateral view [FH 0601-9]; B. Habitus, dorsal view [FH 0601-9]; C. Knee-region of right midleg, posterolateral view; D. Terminalia in lateral view [FH 0601-9]; E. Terminalia in dorsal view [FH 0601-2]; F. Terminalia in ventral view [FH 0601-2]; G. Head and pro- and mesothorax, lateral view [FH 0601-9]; H. Head, pro- an mesothorax, dorsal view [FH 0601-2].

FIGURE 11. Sadyattes leytensis (Zompro, 1997) comb. nov. ♀ Philippines, Mindanao Island, Mount Busa [FH 0601-1]: A. Habitus, dorsal view; B. Habitus, dorsolateral view; C. Head and prothorax, lateral view; D. Knee-region of left midleg, anterolateral view; E. Terminalia in lateral view; F. Terminalia in dorsal view; G. Terminalia in ventral view. Pha = phallus.

FIGURE 12. Sadyattes maganda sp. nov. ♀ paratypes [OC]: A. Habitus, dorsal view (olive green specimen); B. Habitus, dorsal view (buff specimen); C. Habitus, dorsal view (specimen with white medio-longitudinal dorsal stripe and paired black posterolateral mesonotal and metanotal markings); D. Head and prothorax, lateral view; E. Mesofemur, dorsal view; F. Kneeregion of left midleg, anterolateral view; G. Terminalia in lateral view; H. Terminalia in dorsal view; I. Terminalia in ventral view.

FIGURE 13. Sadyattes maganda sp. nov. ♂: A. Habitus of holotype, dorsolateral view [RBINS]; B. Habitus, of holotype, dorsal view [RBINS]; C. Habitus of paratype, dorsolateral view [FH 0278-2]; D. Habitus, of paratype, dorsal view [FH 0278- 2]; E. Abdomen of paratype with mostly orange terga IV-VII in lateral view [OC]; F. Abdomen of paratype with almost wholly black terga IV-VII in lateral view [OC]; G. Terminalia of holotype in lateral view [RBINS]; H. Terminalia of holotype in dorsal view [RBINS]; I. Terminalia of holotype in ventral view [RBINS]; J. Terminalia of paratype in lateral view [FH 0278-2]; K. Terminalia of paratype in dorsal view [FH 0278-2]; L. Terminalia of paratype in ventral view [FH 0278-2]; M. Head and prothorax of holotype, lateral view [RBINS]; N. Head and prothorax of paratype, lateral view [FH 0278-2]; O. Lateral view of tegmina and alae of paratype [OC]; P. Knee-region of left midleg of holotype, posterolateral view [RBINS].

FIGURE 14. Sadyattes matipuno sp. nov. ♀ holotype [RBINS]: A. Habitus, dorsal view; B. Habitus, dorsolateral view; C. Habitus, ventral view; D. Head and prothorax, lateral view; E. Head and prothorax, dorsal view; F. Mesofemur in ventral view; G. Terminalia in ventral view; H. Terminalia in dorsal view; I. Terminalia in lateral view.

FIGURE 15. Sadyattes matipuno sp. nov. ♂ paratype [FH 1266-2]: A. Habitus, dorsolateral view; B. Habitus, dorsal view; C. Habitus, ventral view; D. Head, prothorax and anterior half of mesothorax, lateral view; E. Head, prothorax and anterior half of mesothorax, dorsal view; F. Terminalia in lateral view; G. Terminalia in dorsal view; H. Terminalia in lateral view. Pha = phallus, TP =thorn pads.

FIGURE 16. Sadyattes mindanaense sp. nov. ♀♀ (paratypes) showing intraspecific variability [FH]: A. Habitus of green specimen, lateral view [FH 0630-1]; B. Habitus of green specimen, dorsal view [FH 0630-1]; C. Habitus of brown mottled specimen with large white portions on meso- and metanotum, lateral view [FH 0630-46]; D. Habitus of brown specimen with a whitish medio-longitudinal dorsal streak, dorsolateral view [FH 0630-16]; E. Habitus of brown specimen with a whitish mediolongitudinal dorsal streak, dorsal view [FH 0630-16]; F. Habitus of green specimen with white meso- and metanotal markings, dorsal view [FH 0630-52].

FIGURE 17. Sadyattes mindanaense sp. nov. details of ♀♀ (paratypes) [FH]: A. Terminalia in lateral view [FH 0630-1]; B. Terminalia in lateral view [FH 0630-44]; C. Terminalia in dorsal view [FH 0630-1]; D: Terminalia in dorsal view [FH 0630-16]; E. Terminalia in ventral view [FH 0630-44]; F. Head and prothorax of green specimen with faint dark green postocular streak, lateral view [FH 0630-1]; G. Head and prothorax of green specimen with black dotted postocular streak, lateral view [FH 0630- 52]; H. Head and prothorax of mottled specimen with black dotted postocular streak, lateral view [FH 0630-20]; I. Right midleg lateral view; J. Knee-region of right midleg, posterolateral view (specimen with a slender and rather spinose ventro-apical tooth); K. Knee-region of right midleg, posterolateral view (mottled specimen with a broad and triangular ventro-apical tooth); L. Knee-region of right midleg, posterolateral view (green specimen with a broad and triangular ventro-apical tooth).

FIGURE 18. Sadyattes mindanaense sp. nov. ♂♂ (paratypes) [FH]: A. Habitus, lateral view [FH 0630-7]; B. Habitus, dorsolateral view [FH 0630-21]; C. Habitus, dorsal view [FH 0630-21]; D. Knee-region of right midleg, posterolateral view; E. Terminalia in lateral view; F. Terminalia in dorsal view; G. Terminalia in ventral view; H. Ventral view of two terminal abdominal segments (enlarged); I. Head and prothorax of specimen with faint dark green postocular streak, lateral view [FH 0630-7]; J. Head and prothorax of specimen with faint dark green postocular streak, dorsal view [FH 0630-7]; K. Head and prothorax of specimen with distinct black postocular streak, lateral view [FH 0630-21]. Pha = phallus.

FIGURE 19. Sadyattes panayense sp. nov. ♀ holotype [NHMUK]: A. Habitus, dorsolateral view; B. Habitus, dorsal view; C. Habitus, ventral view; D. Head and prothorax, lateral view; E. Head and prothorax, dorsal view; F. Knee-region of left midleg, posterolateral view. G. Terminalia in lateral view; H. Terminalia in dorsal view; I. Terminalia in ventral view.

FIGURE 20. Sadyattes tubaense sp. nov. ♂ holotype [RBINS]: A. Habitus, dorsolateral view; B. Habitus, dorsal view; C. Head and prothorax, lateral view; D. Head and prothorax, dorsal view; E. Terminalia in lateral view; F. Terminalia in dorsal view; G. Terminalia in ventral view. 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].