Sungaya aeta, Hennemann, 2023

Sungaya aeta View in CoL n. sp.

( Fig. 46-49, 71L, 72P & 73 R-S)

ZooBank: https://zoobank.org/ D37B64B4-3985-45B6-8DEB-9AB2FACDB329

HT, ♀: Philippines, NW-Luzon, Bataan Prov., Ilanin Forest , alt. 100-250m, June 2008 [ RBINS] .

PT, 4 ♀, 5 ♂, 3 eggs: Philippines, NW-Luzon, Bataan Prov., Ilanin Forest , alt. 100-250m, June 2008 [ RBINS] .

PT, 6 ♀, 6 ♂: Philippines, Luzon, Subic, Ilanin forest , ex breeding J. Bresseel 2015 [ RBINS] .

PT, 1 ♀, 1 ♂, 1 egg: Philippines, NW-Luzon, Bataan Prov., Ilanin Forest , alt. 100-250m, June 2008 [ FH, No’s 1455-1, 2 & E] .

PT, 5 ♀: ex Zucht: Rob Krijns 2009, F1 Generation , Herkunft : Philippinen, Luzon , Bataan Prov., Ilanin Forest 100-250 m [coll. FH, No’s 1455-3 to 7] .

PT, 9 ♀, 8 ♂: ex Zucht: Frank Hennemann 2010, F2 Generation , Herkunft : Philippinen, Luzon , Bataan Prov., Ilanin Forest 100-250 m [coll. FH, No’s 1455-8 to 24] .

Differentiation. – Females of this new species most closely resemble those of S.ibalaoi n. sp. butmay bestbe separated bythe shape of the praeopercular organ ( Fig. 47E), which is formed by a distinct, almost semi-circular median excavation at posterior margin of sternum VII (small, shallow and with a pair of tubercles in ibaloi ), the somewhat more distinct median indention of the anal segment ( Fig. 47D) and on average more pronounced medials of the mesonotum, which however are variably in number and size ( Fig. 47 A-B). Moreover, ♀ of aeta have all the limbs comparatively stockier. From ♀ of the type-species S. inexpectata these ♀ may be distinguished by the stockier shape and notably stockier and relatively shorter legs, presence of enlarged mesonotals and semi-circular instead of triangular posteromedian excavation od abdominal sternum VII ( Fig. 47E). Males can merely be differentiated from those of S. ibaloi n. sp. by the somewhat shorter mesothorax, which is only 2.6x longer than the prothorax (3x in ibaloi ) and stronger terminal hook of the vomer, which has

– A. ♀ holotype, dorsal view [ RBINS]. – B . ♀ holotype, dorsolateral view [ RBINS]. – C . ♀ holotype, ventral view [ RBINS]. – D . ♀ paratype, dorsal view [ FH 1455-3 ]. – E . ♀ paratype, dorsolateral view [ FH 1455-4 ]. – F . ♂ paratype, dorsal view [ RBINS]. – G . ♂ paratype, dorsolateral view [ RBINS]. – H . ♂ paratype, ventral view [ RBINS] .

– A. Head, pro- and mesothorax of ♀ paratype with strongly developed mesonotal medials, dorsolateral view (captive reared from Lamao Forest reserve, Bataan Province, Luzon ) [ FH 1455-5 ]. – B. Head, pro- and mesothorax of ♀ holotype with weakly developed mesonotal medials, dorsolateral view [ RBINS]. – C. Mesosternum of ♀ holotype, ventral view [ RBINS]. – D. Terminalia of ♀ holotype, dorsal view [ RBINS]. – E. Terminalia of ♀ holotype, ventral view [ RBINS]. – F. Terminalia of ♀ holotype, lateral view [ RBINS]. – G. Terminalia of ♀ paratype with a large posteromedian lobe on terga VII and VIII, lateral view [ RBINS] .

the sinistral apical corner somewhat protruded and roundly angular ( Fig. 72P). The eggs ( Fig. 73 R-S) represent the most reliable distinctive feature to separate this new species from S. ibaloi n. sp. or S. inexpectata . They clearly differ from those of ibaloi bylackinga posteroventral bump or angle of the capsule, larger micropylar plate, which has the lateral extensions slender and almost parallel-sided (expanded and broadly rounded apically in ibaloi ), circular operculum (oval in aeta ) and notably darker general colour. From those of inexpectata they can be distinguished by the larger dimensions, less angular posteroventral region of the capsule, not indented polar protrusion and much broader and more expanded anterior end of the micropylar plate.

Etymology. – Named after the Aeta people, who live in various parts of the island of Luzon and are regarded as being among the earliest known migrants or inhabitants of the Philippines.

Description

In addition to the dried type-specimens, the colouration is described from live captive reared specimens and photos of live wild and captive reared specimens.

♀ ( Fig. 46 A-E)

Form and colouration. – Of average size (body length 71.0-84.0 mm) and shape for the genus with moderately developed body sculpturing and armature; body surface unevenly tuberculate. Colouration strongly variable and often complex in captive reared specimens (see notes on variability below) but the wild type specimens basically plain testaceous to dark drab. The holotype ( Fig. 46 A-C) with a roundish and velvety black lateral marking on abdominal terga II-IV, and a large irregularly shaped velvety black lateral marking on tergum VIII. The largest paratype with the posterior portion of the metanotum and metapleurae as well as the median segment greyish drab and abdominal terga IV-V light cream-coloured and another paratype with an ochre pre-apical marking on meso- and metafemora. Cephalic and thoracic tubercles mostly tipped with ochre.

Head. – Scarcely longer than wide, broadest at the eyes and with the vertex strongly tumescent and posteriorly somewhat projecting over anterior margin of pronotum ( Fig. 47 A-B); eyes projecting hemispherically and diameter of eye corresponding to 2x the length of gena. Frons somewhat inflated and with rather prominent conical supra-antennals, that are accompanied by some smaller tubercles anteriorly; the posterior portion of frons rather impressed; supra-orbital about equal in size to supra-antennal. Genae only set with a variable number of small gular tubercles. Anterior portion of vertex with a variable number of small occipital medials, the post posterior pair of which is strongly enlarged, prominent and rather spiniform; anterior pair of occipitals notably enlarged but somewhat smaller than the posterior occipital medials and about 2-3 smaller occipitals present in central portion of vertex. Median and lateral coronals large and largest elements of cephalic armature; the lateral coronals comparatively smaller than the median ones although. Antennae consisting of 26 joints and reaching to posterior margin of abdominal segment III; joints first gradually increasing and then decreasing in length towards tip of antennae; the median ones strongly elongated. Scapus compressed dorsoventrally and oval on outline with an obtuse medio-longitudinal carina dorsally.

Thorax. – Pronotum ( Fig. 47 A-B) relatively large, longer and wider than head, wider than long and with a distinct almost semi-circular pre-median excavation at lateral margins, the lateral margins notably deflexed in posterior half; transverse median sulcus moderately impressed, short and curved. Surface unevenly and sparsely tuberculate.Anterior margin only with a small median pair of tubercles and the antero-lateral pronotals very small. In anterior half with a prominent pair of obtuse, conical anterior pronotals and a small pair of medial pronotals present just in front of transverse sulcus; posterior portion with two somewhat pronounced pairs of conical medials, the posterior pair of which is comparatively larger and more spinose; posterior pronotals and inter-posteriors prominent and similar in size and shape. Mesothorax distinctlytrapezoidalin dorsal aspect; mesonotum weakly narrowed anteriorly and parallel-sided in posterior two-thirds, about 2.4x longer than pronotum andabout 2.15x longer thanwidth at posterior margin. Surface of mesonotum unevenly tuberculate and with a row of somewhat enlarged but unequally sized tubercles along lateral margins and the pre-medians, medians and postmedians more or less enlarged and unequal in size; the posterior mesonotals rather low but composite and with an arched lateral ridge that is covered with rather node-like tubercles ( Fig. 47 A-B). Mesopleurae with a marginal row of unequally sized tubercles, the antero-, medio-lateral and two supra-coxals slightly enlarged; mesopleural obtusely conical and moderately sized. Metanotum somewhat narrowed towards the anterior with the posterior margin widely concave; one or two median pairs of tubercles slightly more pronounced and the posterior metanotals notably larger than the posterior mesonotals with the central tubercle distinctly spiniform and the lateral ridge more pronounced. Metapleurae with about seven blunt laterals and some smaller tubercles in between; the median supra-coxal indistinct and the metapleural composite but small. Both, meso- and metapleurae with a further row of small tubercles along lower margin. Meso- and metasternum ( Fig. 47C, 71L) with a fine but distinct medio-longitudinal carina that is densely set with shiny node-like tubercles (this keel and tubercles gradually disappearing in anterior half of mesosternum); otherwise set with a few rather low tubercles, which are unequal in size and somewhat more pronounced on mesosternum.

Abdomen. – Median segment about 2x wider than long with anterior margin widely rounded; near posterior margin with five short posterior spines. Segments II-V almost uniform in length and width and on average 1.2x wider than long (III-V usually somewhat inflated in specimens that are in full eggproduction), only VI-VII somewhat decreasing in length and width.Terga II-VII with all of the typical elements of armature only represented by small tubercles, the fiveposteriorssomewhat more pronouncedalthough;these terga also with a fine butacute medio-longitudinal carina, that becomesincreasingly more pronounced towards VII where it may occasionally be protruded into a small, rounded posteromedian lobe ( Fig. 47G). Sterna II-IV with a fairly strong anterior pair of spinesand a notablysmallerpair of posterior tubercles; the posterior pair indistinct on V-VII. Praeopercular organ formed by a broad and almost semi-circular median excavation at posterior margin of sternum VII, whose margin is notably inflated and down-folded ( Fig. 47E). Terga VIII and IX with the medio-longitudinal carina obtuse and often protruded into an obtuse and low dentiform posterior projection;VIII notably longer than wide and narrowing towards the posterior, IX subquadrate in dorsal aspect. Anal segment narrowing posteriorly, descendant and obtusely tectate medio-longitudinally; the posterior margin with a distinct concave median excavation and the outer angles rather obtuse ( Fig. 47D); lateral margins concave medially. Cerci small, narrowing and compressed laterally ( Fig. 47 F-G). Epiproct about as long as anal segment, broad, weakly tectate longitudinally and just weakly narrowing towards a broadly angular apex ( Fig. 47E). Subgenitalplate moderately long, lanceolate and distinctly keeledin the apical half; the apex gradually narrowed, pointed and slightly surpassing tip of epiproct ( Fig. 47 D-G).

Legs. – Of moderatelength and ratherstocky for the genus withallcarinae armed except for the dorsal carinae of the tibia. Ventral carinae of tibiae only with small dentations, although those of the metatibiae are more numerous, notably more pronounced and acutely pointed. Basal flexure and constriction of profemora well developed, the two anterior carinae each with three large, triangular teeth (the first one notably smaller than the two apical ones); the two posterior carinaewith five teeththatstronglydecrease in size towardsthe base of femur and the dorsal ones of which are comparatively larger. Exterior ventral carinae of meso- and metafemora basically with about six acute teeth that increase in size towards the apex of femur; the two dorsal carinae with about five much broader teeth; several smaller intercalated teeth present on ventral carinae, especially on metafemora. Medioventral carina of meso- and metafemora moderate and irregularly granular ( Fig. 47C). Tarsi about half as long as corresponding tibia; basitarsi elongate almost as long as proceeding three tarsomeres taken together.

♂ ( Fig. 46 F-G)

Form and colouration. – Size and shape typical for the genus (body length 51.0- 61.5 mm) with sparse body armature but distinct and spinose posterior meso- and metanotals. Body surface minutely granular and unevenly tuberculate. Colouration variable in captive reared specimens but much less than ♀ although. The wild examples rather uniformly testaceous brown ( Fig. 46 F-H) and all with a ± distinct ochre medio-longitudinal dorsal streak on thorax ( Fig. 49D). Median portions of meso- and metasternum reddish ochre ( Fig. 48G).

Head. – Basically, as in ♀ but thevertex more strongly tumescent andall of the main elements of armature comparatively larger, but otherwise with notably less smaller tubercles; the supra-antennals simple and much smaller and the gulars indistinct or wanting( Fig. 48F, H). Antennae as in ♀ but relatively longer and reaching to posterior margin of abdominal segment IV.

Thorax. – Pronotum as in ♀ butall spinesnotably morepronouncedand spiniform,theposterior inparticular representedbylargeuprightspines;onlythe inter-posterior pair smaller thanin ♀ ( Fig. 48F, H). Mesothorax slender, more or less uniform in diameter and notably swollen posteriorly ( Fig. 48F); about 2.6x longer than prothorax. Mesonotum weakly widened anteriorly and posteriorly and 4.3x longer than width in median portion; surface sparsely and minutely tuberculate and only with a few somewhat enlarged tubercles (in posterior portion in particular) as well as a small and spiniform pair of anterior mesonotals; the posterior mesonotals represented by a pair of strong, upright spines that have some small tubercles around the base ( Fig. 48F). Metanotum distinctly narrowed in anterior half; tubercles somewhat more pronounced than on mesonotum but the posterior metanotals roughlyidentical insize andshape to posterior mesonotals. Meso- and metapleurae strongly deflexed posteriorly and onlywith about five slightlyenlarged, obtuse tubercles; two supra-coxalsof metapleurae strongly enlarged and spiniform; the meso- and metapleural rather small and conical. Meso- and metasternum with a shallow medio-longitudinal carina that is marked by shiny, node-like tubercles that are clustered along the mid line; these forming a rounded wart-like cluster pre-posteriorly; otherwise sparsely set with some low tubercles ( Fig.48G).

Abdomen. – Notably narrower than thorax. Median segment with anterior margin convex and surface with some obtuse paired tubercles medially. Segments II-V roughly uniform in length, rectangular and about 2x longer than wide, II slightly shorter and narrowing towards the posterior; VI noticeably shorter than preceding and VII scarcely more than half the length of III-V and slightly widening towards posterior. All with surface merely supplied with some paired granules or nodes as well as a small but spiniform pair of second paired posteriors, which are distinct on II and III but become increasingly indistinct towards VII. Sterna II-VII with a fine medio-longitudinal carina; II with a pair of anterior and posterior pair of

Faunitaxys, 11 (71), 2023: 1 – 135. 85

– A. Terminalia of ♂ paratype, lateral view [ RBINS]. – B. Terminalia of ♂ paratype, dorsal view [ RBINS]. – C. Terminalia of ♂ paratype, ventral view [ RBINS]. – D. Terminalia of ♂ paratype, dorsal view (captive reared from Lamao Forest reserve, Bataan Province, Luzon) [ FH 1455-17 ]. – E. Terminalia of ♂ paratype, ventral view (captive reared from Lamao Forest reserve , Bataan Province, Luzon) [ FH 1455-19 ]. – F. Head, pro- and mesothorax of ♂ paratype in dorsolateral view [ RBINS]. – G. Meso- and metasternum of ♂ paratype [ RBINS]. – H. Head, and prothorax of ♂ paratype in dorsolateral view (captive reared from Lamao Forest reserve , Bataan Province, Luzon ) [ FH 1455- 24 ] .

– A. ♀ in dorsal view. – B. Closeup of head and prothorax of ♀ in A. – C. Lateral view of ♀ shown in A. – D. ♂ in lateral view. short, rather obtuse spines, which are indistinct on III, almost wanting on IV and missing on V-VII. Terga VIII and IX broader and shorter than preceding, VIII trapezoidal and IX notably wider than long; both with the medio-longitudinal carina terminating in a low and obtuse, protrusion posteriorly; posterolateral angles of IX slightly deflexed. Anal segment notably longer than IX, declining and narrowed posteriorly, the posterior margin deeply indented medially and broadly bi-lobate; the rounded outer angles supplied with a few minute teeth ventrally ( Fig. 48B, D). Epiproct fairly large, shield-shaped and projecting notably beyond posterior margin of anal segment ( Fig. 48B, D). Cerci fairly large, compressed dorsoventrally and oval in outline. Vomer basically triangular in shape with the apex strong, acutely triangular and notably dextral directed ( Fig. 72P). Poculum large, bulgy and cup-shaped ( Fig. 48A) with the vertical posterior portion tectate and the posterior margin weakly bi-lobed ( Fig. 48E, 72P), indented medially and notably projecting over base of anal segment.

Legs. – Long and slender, mesofemora longer than mesothorax and metatibiae projecting notably beyond apex of abdomen. Generally, as in ♀ but all teeth, but the ventral dorsal ones of the meso and metafemora much smaller and the ventral ones more spinose in shape. Ventral carinae of tibiae only with weakly indicated denticulations in apicalportion.Tarsimuchslenderer than in ♀ with basal joint notably longer than proceeding three tarsomeres taken together.

Variability. – Females of this species show some morphological variation but remarkable chromatic variability in particular. Morphologically, the variability mostly relates to the size and shape of the cephalic and thoracic armature and size as well as the shape of the posteromedian excrescence of abdominal tergumVII ( Fig.47 F-G).The latter can be almost wholly wanting, or at the other extreme is represented by a fairly distinct, rounded lobe. Slightly variability is also seenonthe posteromedianprotrusionof tergumVIII,butthisis much less than inVII.Thethoracic elementsof armature thatshow the mostnotable variability or the pre-median and median mesonotals ( Fig. 47 A-B). Considerable variability is seen in captive reared ♀, which range from plain buff to dark brown and almost black but may also be of very complex colour patterns that comprise various tones of black, brown, beige, white and green. The green morphs however seem to be caused by breeding conditions such as very high humidity and little or white light and have not yet been found in the wild. According to Dräger (2013:5) the green colour in nymphs disappears after a few skin sheds if the humidity and lighting are changed. The black lateral markings on abdominal terga III, IV and VIII are variable in shape but are missing in about two-thirds of the specimens examined. Occasionally, specimens have a white to cream coloured medio-longitudinal streak along the dorsal body surface ( Fig. 46D) and quite frequently there is an irregularly shaped whitish pre-apical transverse band on all femora ( Fig. 49A, C). Specimens often have the posterior portion of the metanotum and metapleurae as well as the median segment pale cream in colour or abdominal terga V-VII partly to wholly whitish or cream in colour. Some specimenshave the basesof allfemoraaswellas thetibiae wholly cream-coloured. In the wild the dark coloured varieties are found closetothecoast,whereas the whitefleckedvarietiesareonlyfoundinthe inland forests ( Dräger, 2013: 5; basedonpersonal communication Thierry Heitzmann).Notably less variability is seen in ♂.

Egg ( Fig. 73 R-S)

Large and of typical shape for the genus; capsule bulgy, strongly constricted anteriorly and the polar area moderately constricted and obtusely conical with an obtuse central protrusion. Surface smooth and densely, minutely pitted. Micropylar plate weaklyraised abovecapsule surface, sculpturedlike capsule; shape broadly and invertedly T-shaped with the median portion strongly broadened and rounded and the two lateral extensions rather narrow, parallel-sided, very weakly up-curved and reaching ca. 60% along lateral surfaces of capsule; posteromedian portion widely excavated and with a fairly distinct bowl-shaped micropylar cup. Outer margin of plate broad and inflated. Median line an obtuse bulge that reaches about half the way to pole of capsule. Operculum circular, flat and pitted like capsule with a very slight centralprotrusion; opercular angle ca. -5°. Colour dark grey; the outer margin of the micropylar plate, anterior margin and polar protrusion of capsule and the operculum blackish. Measurements [mm]: Length 4.8-4.9, width 3.4-3.5, height 3.7-3.8, length of micropylar plate 3.0.

Remarks. – The type specimens collected at Ilanin Forest, Morong Municipality, Bataan Province in June 2008 have given rise to a first culture, that is still successfully maintained throughout Europe. Culture stock from throughout the Bataan Province and referred to as “Lowland”, “IlaninForest” or“LimayLowland” wasintroduced toEurope onseveral occasions,but these appear to have been crossbred ( Dräger,2013).

In captivity this species is very easy to rear in humid conditions and average temperatures of 22-27°C. Various plants are accepted as alternative food, including bramble and raspberry ( Rubus spp. , Rosaceae ), roses ( Rosa spp. , Rosaceae ), hazel ( Corylus avellana , Betulaceae ), hornbeam ( Carpinus betulus , Betulaceae ), beech ( Fagus sylvestris , Fagaceae ), oak ( Quercus robur , Fagaceae ) and ivy ( Hedera helix , Araliaceae ). As for all members of Obrimini a substrate is necessary to allow ♀ to lay their eggs.

Distribution. – Western Central Luzon, Bataan Province (lowland habitats).