Tisamenus napalaki, Hennemann, 2025

Tisamenus napalaki View in CoL n. sp.

( Fig. 33-35, 46 Q-R)

ZooBank: https://zoobank.org/ D2C6C9D6-F283-4AB1-AAC8-B07F0479670C

Hoploclonia draconinus, Matsumura & Hirayama, 1932 : fig.

(Misidentification)

- Shiraki, 1935: 24. (Misidentification)

- Chen & He, 2008: 360, pl. 4: 4. (Misidentification)

Hoploclonia draconia, Huang, 2002: 84 View in CoL , figs. (Misspelling of draconinus and misidentification)

- Xu, 2005: 335. (Misspelling of draconinus and misidentification)

HT, ♂: Coll. R.I.Sc.N.B., Philippines, N Luzon, Cagayan, Sta Ana , vi.2014, 18°28’N 122°09’E, Leg. I. Lumawig, gift from B. Kneubühler, I.G.: 32.613 [ RBINS] GoogleMaps .

PT, 2 ♀, 1 ♂: Coll. R.I.Sc.N.B., Philippines, N Luzon, Kalinga, vi.2014, Purchased from I. Lumawig, Gift from B. Kneubühler, I.G.: 32.613 [ RBINS] .

PT, 2 ♂: Coll. R.I.Sc.N.B., Palaui Island , ex. breeding, N. Wuyts [ RBINS] .

PT, 5 ♀, 6 ♂, 1 egg: Philippinen, N Luzon Id., Prov. Cagayan, Pampanga, Santa Ana, sealevel, local collector VIII.2012 [ FH, No’s 1519-1 to 9, E1] .

PT, 4 ♀: ex Zucht: Tim Bollens 2019, Herkunft : Philippinen, N-Luzon, Palaui Isl. , Prov. Cagayan, Barangay , San Vicente [ FH, No’s 1519-10 to 13] .

PT, 1 ♀, 7 ♂, 4 ♀ (immature), 22 eggs: ex Zucht: F. Hennemann 2020, Herkunft: Philippinen, N-Luzon, Palaui Isl. , Prov. Cagayan, Barangay, San Vicente [ FH, No’s 1519-14 to 25, E2] .

PT, 1 ♀: San Pablo , Isabela, E-Luzon, leg. I. Lumawig VII.2015, leg. Lumawig 2015 [TB] .

Differentiation. – This large and characteristic species shows the closest affinity to the other species with four mesopleural spines T. draconinus (Westwood, 1859) from Luzon. From draconinus both sexes of this species are distinguishable by the notably larger dimensions, slightly slenderer shape and generally more pronounced armature of the body with all the major spines comparatively longer and slenderer. Moreover, both sexes clearly have two metapleural laterals ( Fig. 33A) instead of only one as in draconinus ( Fig. 17A), which perhaps is the most obvious differential character. Males may also be distinguished by the less evident mesothoracic constriction at the 3 rd mesopleural lateral, characteristic gibbous dorsal swelling of abdominal terga II-V ( Fig. 35I) and much larger pair of posteriors on terga II-III as well as the less distinct ventro-basal swelling of the meso- and metafemora ( Fig. 35 L-M). The eggs ( Fig. 46 Q-R) are larger and somewhat more elongate than those of draconinus , and readily separated by the much more developed sculpturing of the capsule surface and smaller, broader micropylar plate, which is only about three-quarters the length of the capsule (as much as seven-eighths the length of capsule in draconinus ) and has all three extensions notably shorter and wider.

Etymology. – The name of this new species ( napalaki Filipino = inflated) refers to the characteristically inflated abdominal terga II-V in ♂ of this remarkable new species ( Fig. 35I), which is the most colourful in the entire genus, with ♀ showing pretty colour patterns including tones of white, cream, yellow and black and ♂ often with pretty orange to reddish colours.

Description

♀ ( Fig. 33)

Form and colouration. – Size large (body length64.0-69.0 mm); general form moderately slender and elongate; the elements of armature strongly developed and mostly spinose, the anterolateral spines of the triangular mesonotal area compound ( Fig. 35A); legs elongate and slender with rather weakly developed but spinose rather than dentate armature. Body surface unevenly granular. Colouration very variable and complex for the genus, often with unusual ochre, orange and red tones and patterns. General colour mostly ranging from fairly plain drab or ochre over various tones of brown and fuscous to almost black; median portion of abdominal terga II-VII often cream, ochre to clay coloured with the lateral surfaces dark brown to blackish. Meso- and metasternum lighter in colour than dorsal body surface and buff to almost clay coloured with the three small mesosternals each marked by a blackish spot ( Fig. 33G, 34B). Pro-, meso- and metanotum as well as abdominal terga I-II with an ochre, orange or reddish medio-longitudinal streak that is laterally bordered by a narrow blackish line; this broad on pronotum;narrow but covering most of disc of triangular area onmesonotum, narrow on metanotum and median segment and widened into an almost circular marking on abdominal tergum II. Femora mid to dark brown or almost black and flecked with ochraceous tones ventrally. Largest spines of head and body with apical half dark brown and tipped with orange. Antennae uniformly black but becoming rather greyish towards the base.

Head. – Rectangular, longer than wide with the genae weakly widening towards the posterior. The supra-orbital large and spinose with a few small tubercles around base; occipitals and median coronals small and tubercular; the lateral coronals sub-spinose, conical and notably larger than median coronals, the latter slightly larger than occipitals ( Fig. 35A). Eyes small, hemispherical and their diameter corresponding to about 0.4x length of gena. Genae supplied with 2-3 low, nodose to tuberculate gulars. Antennae slender with all joints except IV longer than wide and consisting of 26 joints; scapus weakly triangular in dorsal aspect, pedicellus about half the length of scapus and almost cylindrical; III slightly longer than pedicellus, IV much shorter, following increasing in length up to XV, then slightly decreasing in length with the terminal antennomere much elongated and slightly longer than preceding two joints combined.

Thorax. – Pronotum sub-quadrate; triangular area moderately indicated with margins behindthe transverse mediansulcusbounded by small tubercles; anterolaterally with a strong bifid spine, whose second spike usually is somewhat longer than the anterior one; behind the bifid anterior spine with one or two spiniform tubercles ( Fig. 35A). Mesothorax fairly slender, slightly widening towards the posterior, about 2.1x longer than prothorax and with posterior portion 1.3x wider than anterior margin. Mesonotum weakly sub-trapeziform with lateral margins obliquely convergent towards the posterior and with a very shallow median narrowing, about 2.1x longer than width at anterior margin; the triangular area roughly attaining middle of notum, longer than wide, disk shallowly concave and the almost straight margins tubercular with the anterolateral angles protrudedinto a composite, 3-4 spinose crest that has the median spike notably longer than the others, all acutely spinose; the anterior margin of triangular area with a dentiform median protrusion ( Fig. 35A). Posterior portion of mesonotum with a broad and shallowly granulate medio-longitudinal bulge, that is faintly indicated in the triangular area; pre-posteriorly with a distinct, bi-spinose swelling. Mesopleurae somewhat expanding towards the posterior and narrowed at anterior margin with four large and spinose laterals; the antero-lateral small than the following, the second and third represented by long and acutely pointed spines and the slightly smaller mesopleural with a variable number of small tubercles around the base. Metanotum sub-rectangular anda little longer than wide witha slight median narrowing, surface with the same granulose medio-longitudinal bulge and pre-posterior bi-spinose swelling seen on mesonotum. Metapleurae with two prominent and spinose laterals, the anterior one of which is smaller; metapleural small, tubercular and the supra-coxal angle with a strong more or less compound supra-coxal spine that is accompanied by a variable number of spiniform tubercles around its base. Mesosternum tri-carinate, the lateral carinae somewhat irregular, converging towards the posterior and supplied with three shallow node-like mesosternals ( Fig. 35B); metasternum with a n obtuse medio-longitudinal carina and a few faint indications of metasternals laterally ( Fig. 34G).

Abdomen. – Median segment distinctly trapezoidal with anterior margin weakly rounded; the medio-longitudinal carina shallow and with two small nodes pre-posteriorly. Segments II-III roughly equal in length and width and rectangular, IV-VII slightly decreasing in length andV-VII notably narrowing; II and III about 2x wider than long, VII only1.6x wider than long andnarrowest of all segments. All terga with two weakly indicated and closely spaced, parallel medio-longitudinalcarinae; II-IV withprominentsecond-paired posterior spines ( Fig. 34 D-E) and a variably but small and conical posterior mesal; the second paired posteriors only represented by small spinose tubercles on V.Sterna II-VII only with weakly indicated posterior medials, otherwise simple and minutely granular ( Fig. 34G); praeopercular organ formed by a two shallow, converging bulges that posteriorly terminate in a small swelling ( Fig. 35F). Tergum VIII A. PT, dorsal view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon – arrows pointing to the two metapleurals that readily distinguish this species from T. draconinus ) [ FH 1519-5 ]. B. PT, dorsal view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [ FH 1519-14 ]. C. PT, dorsal view (from Santa Ana, Cagayan Province, N-Luzon) [ FH 1519-13 ]. D. PT, dorsolateral view (captive reared from San Vicente, Barangay, Palaui Island , Cagayan Province, NLuzon) [ FH 1519-14 ]. E. PT, dorsolateral view (from Santa Ana, Cagayan Province, N-Luzon) [ FH 1519-5 ]. F. PT, lateral view (captive reared from San Vicente, Barangay, Palaui Island , Cagayan Province, N-Luzon) [ FH 1519-13 ]. G. PT, ventral view (captive reared from San Vicente, Barangay, Palaui Island , Cagayan Province, N-Luzon) [ FH 1519-13 ]. H. PT, live captive reared specimen (PT) from San Vicente , Barangay , Palaui Island , Cagayan Province, N-Luzon. I. PT, live captive reared specimen (PT) from San Vicente , Barangay , Palaui Island , Cagayan Province, N-Luzon .

trapezoidal in dorsal view, IX obtusely tectate with an obtuse, triangular posteromedian protrusion that is formed by the two medio-longitudinal carinae ( Fig. 35C). Anal segment strongly declining with the lateral margins obliquely convergentin posterior half and with a fairly well developed medio-longitudinal carina; the posterior margin broadly bi-lobed with a shallow median indention ( Fig. 35 D-E). Epiproct weakly tectate longitudinally with an indication of a medio-longitudinal carina and slightly cucullate ( Fig. 35C), the lateral margins almost parallel-sided and the posterior margin roundly angular ( Fig. 35 D-E). Subgenital plate fairly elongate, notably projecting beyond epiproct, navicular, distinctly carinate in posterior half with the apex narrowed and pointed ( Fig. 35F).

Legs. – All slender and elongate with the teeth rather moderately developed and fairly spinose; the two apical ventral teeth of meso- and metafemora rather small and those on the dorsal carinae of the meso- and metafemora distinctly spinose and the largest elements of leg armature. Profemora almost as long as mesothorax, mesofemora notably shorter; metafemora somewhat projecting over posterior margin of abdominal segment IV and metatibiae reaching to posterior margin of tergum VIII. The one or two basal spines on the two dorsal carinae of meso- and metafemora notably arched towards baseof femur.All tibiae unarmed dorsally, mesotibiae only witha few weakly indicated denticles ventrally and metatibiae with about eight acute teeth on two ventral carinae. Basitarsi rather elongate, almost as long as following three tarsomeres taken together.

Measurements of paratype s [mm]. – Body 64.0-69.0, pronotum 5.0-5.3, mesonotum 11.0-11.5, metanotum 6.0-6.2, median segment 2.7-2.8, profemora 11.1-12.0, mesofemora 10.3-10.6, metafemora 13.8-14.6, protibiae 11.7-12.7, mesotibiae 10.8-11.2, metatibiae 14.9-16.0, antennae 25.0-27.0.

♂ ( Fig. 35)

Form and colouration. – Size rather large (body length 44.0-50.0 mm), general form fairly slender and elongate, the elements of armature prominent andessentially like in ♀ but comparatively more developed;abdominal terga II-V with a gibbous swelling ( Fig. 35I) and the legs broader and stouter than in ♀ with the metafemora moderately incrassate and more heavily armed tan in ♀ ( Fig. 35L). Colour essentially like in ♀ but the medio-longitudinal streak along dorsal body surface often continued up to abdominal tergum VIII and more commonly orange to red than in ♀. Terminal antennomere buff.

Head. – Like in ♀ but the anterior supra-orbital notably longer ( Fig. 35N) and the eyes proportionately larger with their diameter corresponding to about 0.5x the length of gena. Antennae proportionally longer and almost reaching to posterior margin of metanotum.

Thorax. – Pronotum essentiallyas in ♀ but less trapeziform inoutline and the bi- or trifid anterior much longer; like in ♀ the median spike usually the longest. Mesothorax 2.2x longer than prothorax and slightly narrowed anteriorly and medially. Mesonotum sub-trapezoidal with the lateral margins slightly converging towards the middle and then with a slight pre-posterior widening, about 2.2x longer than width at anterior margin; the triangular area relatively more elongate than in ♀ being about 1.6x longer than width across anterolateral angles and roughly reaching to middle of notum; the lateral margins strongly raised, obtusely granular and the anterolateral angles protruded into a large bifid spine whose first spike is small and sub-spiniform and the second a large and upright spine ( Fig. 35N). Posterior portion of mesonotum with a distinct medio-longitudinal bulge that is weakly indicated in the concave interior region of the triangular area; pre-posteriorly with a prominent conical swellingthatbears two lateral directed spines. Mesopleurae weakly narrowed medially with elements of armature like in ♀ but more spinose, much longer and the mesopleuralwith less small tubercles arounds its base; the second and third mesopleural laterals laterally projecting by at least half the width of mesothorax. Metanotum longer than wide, narrowed anteriorly and with the lateral margins convex in posterior half; surface with the same medio-longitudinal bulge and bis-spinose pre-posterior median swelling seen on mesonotum. Metapleurae with armature like in ♀ but the spines all much longer and slenderer. Mesosternum tri-carinate, the median carina prominent and the lateral carinae very close to lateral margins of sternum ( Fig. 35M). Metasternum with a medio-longitudinal keel and metasternals indicated by a few small and uneven granules ( Fig.35G).

Abdomen. – Median segment distinctly trapezoidal in outline with the anterior margin angular and the anterolateral areas somewhat impressed. Segments II-VII distinctly decreasing in length a very slightly narrowing, II-IV sub-quadrate and scarcely longer than wide, V-VII wider thanlong and VII narrowest of all segments; terga II-V with the posterior half strongly swollen gibbous; the second paired posteriors merely present on terga II and III and represented by strong spines, on IV merely indicated as small tubercles ( Fig. 35 H-I). A medio-longitudinalcarina only seen inanterior portion of II-V and more definite over entire length of VI-IX. Sterna II-III with a distinct medio-longitudinal keel that is less obvious on IV, just weakly indicated on V andmissing on VI-VII.TergaVIII and IX transverse andtrapezoidalin outline with an obtuse dentiform posteromedian protrusion;the posterolateral angles of VIII and IX somewhat digitiform. Anal segment distinctly trapezoidal, somewhat declining towards the posterior, the lateral margins angular with an obtuse tooth at the angle and the posterior margin shallowly concave with the outer anglesobtuselyprotruded ( Fig. 35J). Epiproctbroadlytransverse, shield-shaped and rounded ( Fig. 35J). Vomer shorter than width at base, the ventral surface with some irregular transverse ridges, the outer margin somewhat convex and the terminal hook prominent, triangular and somewhat sinistral directed. Poculum rather shallowly cup-shaped with the posterior flange very broad, weakly bi-labiate and emarginate medially ( Fig. 35K).

Legs. – All stockier with all teeth comparatively more developed than in ♀. Pro- and mesofemorashorter than mesothorax, metafemora reaching about half the way along abdominal segment V and metatibiae at best slightly projecting beyond tip of abdomen. Ventral teeth more spiniform than those on dorsal carinae; the two basal spines on the dorsal carinae of the meso- and metafemora somewhatarched towards the base of femur. Ventro-basal portion of metafemoraweaklygibbose and the twoouter ventralcarinae armedwitha variable number of variably sized but distinct spiniform teeth; the ventral carinae of metatibiae armed with about 12-18 unevenly sized, acute teeth ( Fig. 35L).Tarsi like in ♀.

Measurements of holotype [mm]. – Body 45.8, pronotum 4.1, mesonotum 8.8, metanotum 5.0, median segment 2.6, profemora 9.7, mesofemora 8.3, metafemora 10.4, protibiae 9.8, mesotibiae 8.6, metatibiae 11.0, antennae 20.0.

Measurementsof paratypes [mm]. – Body 44.0-50.0, pronotum 3.8-4.6, mesonotum 8.7-9.3, metanotum 4.1-5.1, median segment 2.5-2.8, profemora 8.6-9.0, mesofemora 7.0-8.0, metafemora 9.3-10.5, protibiae 8.7-9.0, mesotibiae 6.8-8.7, metatibiae 9.7-11.0, antennae 18.0-21.0.

Variability. – This remarkable species shows slight variability in the armature of the head, body and limbs in both sexes, where the most decisive variability is seen in the size and shape of the pronotal and mesonotal anteriors. The most striking variability is seen in the colourationof both sexes, which is illustrated in Fig. 33-35. Both sexes range in general colour from drab or ochre over various tones of brown and fuscous to almost black and the dark yellow, orange to almost red distinctive medio-longitudinal streak of the meso- and metanotum is variable in its intensity. Males may also occur in almost wholly orange to red morphs (e. g. Fig. 34 H-I).

Egg ( Fig. 46 Q-R)

Large for the genus; capsule ovoid, slightly and somewhat narrowed posteriorly, oval in cross-section with the lateral surfaces almost parallel-sided in median portion and with a fairly distinct posterior indention of polar-area; higher than wide with dorsal surface bulgier than ventral surface; capsule about 1.5x longer than wide. A slight constriction just below the somewhat inflated and minutely tubercular anterior margin. Surface all over covered with an irregular meshwork of ridges that are densely supplied with fringy to setose excrescences; the fringy carinae only wanting in the very antero-dorsal and lateral portion of capsule. The capsule otherwise densely and minutely granular. Micropylar plate rather small and just scarcely more than 0.7x the length of capsule; Y-shaped with the median portion broad, gently tapering and clearly not attaining anterior of capsule; the two posterolateral extensions somewhat slenderer, rather short and on lateral surfaces roughly reaching axis of egg capsule. Posterior portion 70° Vshaped and with a small but distinct bowl-shaped micropylar cup in centre. Outer margin of plate marked by a rim of fringy excrescences, the inner portion somewhat inflated, bordered by short fringy protuberances along the outer margin and only with a few scattered protuberances interiorly; the fairly narrow space between outer margin of capsule and the raised interior portion forming an un-sculptured indented rim. Median liner distinct and formed by a fringy bulge that reaches as far as to the polar-area. Operculum inserted into capsule at an angle of about -5° and very slightly oval in

A. PT, closeup of head, pro- and mesonotum (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, NLuzon) [ FH 1519-13 ]. B. PT, closeup of head, pro- and mesosternum (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [ FH 1519-13 ]. C. Terminalia in lateral view. D. Terminalia in dorsal view. E. Terminalia in dorsal view [ FH 1519-5 ]. F. Terminalia in ventral view. [ FH 1519-5 ]. G. Live captive reared couple (PT) from San Vicente, Barangay, Palaui Island , Cagayan Province, N-Luzon (dark brown ♂ with orange medio-longitudinal dorsal streak). H. Live captive reared almost wholly orange ♂ (PT) from San Vicente, Barangay, Palaui Island , Cagayan Province, N-Luzon. I. Live captive reared almost wholly orange ♂ (PT) from San Vicente, Barangay, Palaui Island , Cagayan Province, N-Luzon .

A. Dark brown PT with contrastive orange medio-longitudinal dorsal streak, dorsal view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon – arrows pointing to the two metapleurals that readily distinguish this species from T. draconinus ) [FH 1519-21]. B. Buff to ochre PT with weakly indicated yellow medio-longitudional dorsal streak, dorsal view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [FH 1519-18]. C. Dark russet PT with a very narrow red medio-longitudinal dorsal stripe, dorsal view (from Santa Ana, Cagayan Province, N-Luzon) [FH 1519-9]. D. PT, dorsolateral view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [FH 1519-21]. E. PT, dorsolateral view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [FH 1519-18]. F. PT, lateral view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [FH 1519-21]. G. PT, ventral view (captive reared from San Vicente, Barangay, Palaui Island, Cagayan Province, N-Luzon) [FH 1519-21]. H. PT, dorsal view of abdomen [FH 1519-19]. I. PT, dorsal view of abdomen, showing the typical inflated, gibbous posterior portion of the basal terga [FH 1519-19]. J. PT,terminalia indorsalview. K. PT,terminalia inventralview [FH1519-9]. L. PT,anteroventralview of left hind leg. M. PT, closeup of head, pro- and mesosternum. N. PT, closeup of head, pro- and mesonotum.

outline. The outer marginwith a narrow collar of fringy protuberances and in the middle portion with a distinct circular rim of fairlylong fringes; the centre indented. Colour dark blackish brown to almost with all the fringy protuberances of the capsule, micropylar plate and operculum ochraceous to dark olive. Measurements [mm]: Length incl. operculum 4.4, length 4.2, width 2.8, height 3.0, length of micropylar plate 3.1.

Remarks. – A specimen recorded from the island of Kotosho southeast of Taiwan and misidentified as “ Hoploclonia draconinus ” and illustrated by Matsumura & Hirayama (1932) actually is an immature ♀ of this new species, which is distributed in the very northeastof Luzon.Although there are no records of T.napalaki n. sp. from the small islands situated in the Luzon Strait, namely the Babuyan and Batanes Islands, a natural occurrence on Kotosho Island is very unlikely and can most certainly be associated to a meteorological event like a typhoon (see remarks on T.draconinus for more information).

In June 2016 Albert Kang ( Singapore) collected specimens on the island of Palaui and passed these on to Thierry Heitzmann ( Philippines), who was able to breed the stock in captivity and sent eggs laid by the F1-generation to Europe. These have given rise to a culture that is since maintainedand popular amongEuropean breeders because of the remarkably pretty colouration. The species has been distributed as Tisamenus sp. ‘Palaui’ but has not yet been given a Phasmid Study Group culture number.

Distribution. – N-Luzon: Province Cagayan (Santa Ana [RBINS – type locality; photographic record by Albert Kang: https://inaturalist.ca/ observations/130746863]; Papanga, Santa Ana [FH]; San Vicente, Palaui Island [FH]); Province Kalinga, Balbalan [RBINS]).