Trichadenotecnum species from Peninsular Malaysia and Singapore (Insecta: Psocodea: 'Psocoptera': Psocidae)

Species of the bark louse genus Trichadenotecnum Enderlein (Insecta: Psocodea) from Peninsular Malaysia and Singapore are revised with illustrations and identification keys. Twenty species are here recognised, with four new species and ten recorded for the first time from this region, together with an unnamed species represented by a single female. The previ-ously described species T. marginatum New & Thornton is not included because its generic assignment is questionable. Females of T. cinnamonum Endang & New, T. imrum New & Thornton and T. sibolangitense Endang, Thornton & New, and the male of T. kerinciense Endang & New are described for the first time. A new species group is defined for T. Endang, Thornton New.


Introduction
The free living Psocodea or "Psocoptera" is now recognized as a paraphyletic assemblage (Yoshizawa & Johnson 2003, 2010, within which Trichadenotecnum Enderlein is one of the most species-rich genera (Lienhard & Smithers 2002;Lienhard 2011). Previous studies on the Oriental fauna have revealed enormous species diversity within this genus (Hong Kong- Thornton 1961, Yoshizawa & Lienhard 2004Indonesia-Endang & Thornton 1992, Endang et al. 2002, Endang & New 2005Japan-Yoshizawa 2001, 2003China-Li 2002;Nepal-Yoshizawa et al. 2007). Peninsular Malaysia is designated as one of the biodiversity hotspots in the world (e.g., Ceballos & Ehrlich 2006). However, the Trichadenotecnum fauna of Peninsular Malaysia has been considered in only a few scattered papers, with only five species (excluding the questionable T. marginatum) recorded from the region to date (New 1975;New & Thornton 1976;New & Lee 1992). Even including the species from Singapore, the total number of Trichadenotecnum known from this region is only six (New & Lienhard 2007), in contrast to the 20 species known from the neighboring island, Sumatra (Endang & New 2005).
Within Trichadenotecnum, several species groups have been proposed based mainly on male and female genitalia. However, considerable morphological gaps among the species groups, or even among some species within a group, render morphology-based phylogenetic estimation and estimation of morphological transformation series difficult. High diversity of Trichadenotecnum in the Oriental Region, and lack of extensive investigation of some areas within the region, suggest that undiscovered species may help to fill such morphological gaps. Faunal surveys of this region may also help in understanding the phylogeny and morphological evolution within this genus.
Remarks. The male specimen examined here is in complete agreement with the original description of T. cinnamonum, except that the hypandrial structures are in mirror images between them (Fig. 2C). Because of the exact similarity, we concluded that the hypandrium illustrated in the original description is drawn from the reverse. This is the first record of this species from Peninsula Malaysia.  Originally, the corniculum group was characterized by the presence of a process arising from the trichobothrial field of the male paraproct (Yoshizawa, 2003). Later, absence of this structure in T. felix Thornton, 1961 was reported, although the hypandrial structure strongly suggests its placement to the corniculum group (Yoshizawa & Lienhard, 2004). The hypandrial structure of T. cinnamonum is more similar to that of T. felix than to species with the trichobothrial process, but T. cinnamonum has a keel on the trichobothrial field which is most likely homologous with the trichobothrial process. This suggests that the trichobothrial process is secondarily absent in T. felix, and its presence is autapomorphic for the corniculum group. The assignment of T. cinnamonum to the corniculum group can be warranted by this structure.
The female specimen examined here differs significantly from the description of the paratype female given by Endang & New (2005); this clearly shows that they belong to different species. The paratype female of T. cinnamonum also looks significantly different from the holotype male in forewing markings (heavily spotted in female paratype whereas much more sparsely spotted in holotype male). In contrast, the female identified here as T. cinnamonum is more similar to the holotype male of this species in the forewing markings (Fig. 1BC). In addition, the female genital structures of the specimen examined here (Fig. 3) are very similar to those of T. felix and T. imrum, both of which are members of the corniculum group. In contrast, the genital structures of the paratype female, especially the absence of the posterior lobe of the external valve of gonapophyses, are more similar to the species of the spiniserrulum group (e.g., Trichadenotecnum sp.My19 or T. malayense, treated below: Figs 14 and 17, respectively). Therefore, we conclude that the female paratype of T. cinnamonum was misidentified as that species, and we here provide the first description of the female genital structures of T. cinnamonum. New & Thornton (Figs 1D,4,5) Trichadenotecnum imrum New & Thornton, 1976: 414. Specimens examined: [Singapore] 1 male, Bukit Timah forest, 24 hr survey, section trap 7-9 am, 26.x.1969, DHM (MHNG); 1 male 1 female, Bukit Timah forest, beating foliage, 1.xi.1981, DHM (MHNG); 1 male, Bukit Timah Nature Reserve, between Summit hut and Car Park, along Tangga Rengas trail, Rock Path and Lower Path, 110-140 m, beating, 17.xii.1987, CL (SEHU).

Trichadenotecnum imrum
Description of female genitalia. Egg guide long (Fig. 5A), much longer than basal width, slightly narrowing toward apex, distal margin only slightly arched, medially and laterally with longitudinal membranous regions; body of subgenital plate rather small, posterior margin with short and broad membranous region next to egg guide, anteromedially with broad membranous region. Gonapophyses (Fig. 5B). Ventral valve long; dorsal valve with long distal process; posterior lobe of external valve small. Spermapore plate ( Fig. 5C) lightly pigmented anteriorly and around spermapore, symmetrical.
Remarks. The male specimens examined here match very well with the original description of this species, including genital structures (Fig. 4). This species was originally described from a single male from Selangor, Malaysia, and is here recorded from Singapore for the first time. The female of this species is described here for the first time.
This species is apparently closely related to T. cinnamonum but can be distinguished from the latter by the flattened epiproct (Fig. 4A), presence of the small projection placed left to the circular-plate on the hypandrium (Fig. 4C), and absence of the keel on the trichobothrial field ( Fig. 4A) in male and the shape of the subgenital plate ( Fig. 5A) in female. Furthermore, this species is very similar to T. felix in male and female genital structures, as also mentioned for T. cinnamonum. These three species apparently compose a monophyletic group within the corniculum group based on having the bifurcated posterior process on the phallosome (Figs 4D). They also share the following character state: the trichobothrial field is divided into two regions ( Fig. 4A) (Thornton, 1961). The empty space of the trichobothrial field, observed in T. felix and T. imrum, corresponds to the keeled area of T. cinnamonum and other species of the corniculum group ( Fig. 2A). As mentioned above, this situation strongly suggests that the presence of the trichobothrial process/keel is autapomorphic for the corniculum group and plesiomorphic within this group; its absence corresponds to an incomplete reversal leaving the empty area on the trichobothrial field (Fig. 4A).  Trichadenotecnum pardus Badonnel ( Fig 1E) Trichadenotecnum pardus Badonnel, 1955: 231. Trichadenotecnum pardidum: New, 1975 [Not Trichadenotecnum pardidum Thornton, 1961: 16.] Specimen examined. 1 female, Singapore, Botanic Gardens, Rain Forest, beating, 25m, 16.xii.1987, CL (MHNG). Remarks. See Yoshizawa (2001) and Yoshizawa & Lienhard (2004) for diagnosis of this species.
Thorax. Prothorax blackish brown. Mesonotum mostly brown, scutum with yellowish white bands along posterolateral margins of anterior lobe, at middle from anterior to posterior ends of scutum, and along anterior margin of lateral lobe. Metanotum brown, median part and anterior region of lateral lobe of scutum yellowish white. Meso-and metapleuron brown except membranous regions.
Legs. Mostly blackish brown, ventral surface of distal half of fore femur and tip of hind femur white, tibiae paler.

) (SEHU & UKM).
Description. Male. Head. Yellowish white in ground color; vertical markings blackish brown, each marking touching with neighbors; with pair of triangular blackish brown markings anterior to vertical markings; orbital markings blackish brown; coronal suture black; epicranial suture bordered with blackish brown band dorsally; frons with central pair of brown bands reaching to oceller region dorsally, and with lateral pair of blackish brown broader band not reaching to epicranial suture dorsally, central and lateral bands fused with each other ventrally; eye dark gray, IO/D= 0.6; ocelli white, ocellar field black; gena mostly blackish brown, with pair of oval whitish areas arranged vertically; postclypeus mostly blackish brown ventrally except ventrolateral corner white, dorsal region white with four rows of blackish brown spots; anteclypeus blackish brown. Antenna brown, scape and pedicel blackish brown. Mouthparts brown.
Thorax. Prothorax blackish brown. Mesonotum mostly brown, scutum with yellowish white bands along posterolateral margins of anterior lobe, at middle from posterior end to middle of anterior lobe of scutum, and along anterior margin of lateral lobe. Metanotum brown, median part and anterior region of lateral lobe of scutum yellowish white. Meso-and metapleuron brown except membranous regions.
Legs. Mostly blackish brown, ventral surface of distal half of fore femur and tip of hind femur white, tibiae paler.
Forewing ) gradually extended from body of subgenital plate, basally constricted, anterior margin rounded, ventral surface with slight swelling basally; body of subgenital plate widely sclerotized, posteriorly with strongly projecting triangular processes, postero medially with triangular membranous region continuing to anterior trapezoidal membranous region. Gonapophyses (Fig. 9B). Ventral valve long. Dorsal valve with long distal process. Posterior lobe of external valve broad but weakly projecting; internal lobe short. Spermapore plate (Fig. 9C) broad and short, triangular in shape, anterior margin with small notch, broadly pigmented but darker around spermapore and median region of anterior margin.
Remarks. This species is close to T. yatai, described above, but differs by the shape of male epiproct (Figs 6A, 8A) and female subgenital plate (Figs 7A, 9A). T. kojimai differs also from T. yatai in the possession of the posterolateral projections on the subgenital plate in the female and the shapes of the lateral clunial arm and epiproct lobe in male. In having a pair of posterolateral projections on the female subgenital plate (Fig. 9A), T. kojimai is similar to T. adika and T. paradika Endang & New, 2005 from Indonesia. However, T. kojimai differs clearly from T. adika, by the shape of the lateral clunial arms, epiproct lobe, and distal part of the hypandrium. T. paradika is known from female only, and it differs from T. kojimai by the shapes of the posterolateral projection on the subgenital plate and the external valve of the gonapophyses.  Trichadenotecnum adika Endang, Thornton & New (Figs 1H,10) Trichadenotecnum adika Endang, Thornton & New, 2002: 166. Specimen examined. 1 male, Singapore, See Soon, Rifle Range Swamp Forest, beating, 18.xii.1987, CL (MHNG).
Remarks. The specimen examined here agrees well with T. adika in male genital structures. In contrast, it is significantly smaller (forewing length 1.8 mm) than the male holotype of T. adika from Java (forewing length 2.6 mm). The small form could provisionally be interpreted as a lowland form (cf. Bergmann's rule): all Indonesian specimens were collected between 700 and 1460 m (Endang et al., 2002;Endang & New, 2005). Therefore, we identified the above specimen as T. adika. This is the first record of this species from the region. Trichadenotecnum endangae Yoshizawa & Lienhard,sp. n. (Figs 1I,11,12) Holotype. Male, Malaysia, Gunung Brinchang, Cameron Highlands, Pahang, 11.iii.2003, K. Yoshizawa (UKM).
Thorax. Prothorax blackish brown. Mesonotum blackish brown, medially with trifurcated white marking. Metanotum blackish brown, medially with white spot. Meso-and metapleuron brown except for white membranous regions. Legs. Mostly blackish brown, distal half of ventral surface of fore femur and distal tip of hind femur white, tibiae paler.
Forewing (Fig. 1I). Extensively covered with tiny spots, dense basally from basal band, sparse distally from basal band. Distal spot in cell a1 distinct, basal spots obscure. Opposing spots in cell r obscure. Basal band narrow but distinct, broadly interrupted above M-Cu fork, narrowly interrupted above CuP vein. Distal band narrow but distinct. Spot on roof of cell m3 faint, only distinct at top of areola postica. Submarginal spots distinct in cells r3 to m2, obscure in other cells. Marginal clouds almost absent. Hindwing hyaline; veins brown.
Measurements. B 2.4, Fw 3.5, Hw 2.6. Female. General morphology almost as in male. IO/D = 2.5. Genitalia. Egg guide (Fig. 12A) broad basally, strongly constricted to middle and slightly broadened to slightly arched distal margin, laterally with membranous regions; body of subgenital plate broad, anteriorly with broad membranous regions and anteromedially with deep V-shaped membranous region. Gonapophyses (Fig.  12B). Ventral valve long. Dorsal valve with long distal process. External valve lacking posterior lobe, narrow. Spermapore plate (Fig. 12C) with strongly pigmented area posteriorly. Remarks. This species is very close to T. bromoense Endang, Thornton & New, 2002 from Java, Indonesia, but differs from it by the shape of the paraproctal lateral process (Fig. 11A). In the original description, the long hypandrial process of T. bromoense arises from the left side. However, in ventral (external) view, the process arises from the right side in T. endangae (Fig. 11C), as well as in close relatives such as T. godavarense New, 1971 andT. sclerotum New, 1978. Therefore, it is very likely that the illustration of the hypandrium of T. bromoense is drawn in dorsal (internal) view or reproduced in reversed position.
Trichadenotecnum hammani Endang, Thornton & New (Figs 1J,13) Trichadenotecnum hammani Endang, Thornton & New, 2002: 161 Specimen examined. Remarks. This species was originally described from Java, Indonesia, and is here recorded from Peninsula Malaysia for the first time. The specimen examined here matched very well with the original description of this species, especially the shape of external valve of gonapophyses (Fig. 13B) that is characteristic for this species.   Remarks. This species is known only from a single female. Genital structures (overall shape of subgenital plate, long ventral valve and small external valve of gonapophyses, and overall shape of spermapore plate: Fig. 14) are very similar to those of T. endangae, suggesting their close relationship. However, body size (body length = 1.7 mm, forewing length = 2.4 mm) and forewing markings (Fig. 1IK) are significantly different between them. Because T. sp.My19 is known only from a female, we prefer not to name it here. Remarks. This species was described from a single female obtained from Gombak, Malaysia (New & Lee, 1992). The female specimen examined here is in good agreement with the original description (Figs 1L, 15). Although the present material does not include the male of this species, it was described by Endang & New (2005) from Sumatra, which clearly shows that this species is closely related to T. falx Yoshizawa, 2001 from Japan and T. isseii Yoshizawa & Lienhard, 2007 from Nepal due to the possession of the asymmetrical and sickle-shaped hypandrial median tongue. The hypandrial median tongue illustrated by Endang & New (2005) is left-skewed whereas that of the other species is right-skewed. The illustration given by Endang & New (2005) is very likely drawn in dorsal (internal) view or reproduced in reversed position (Yoshizawa et al., 2007). Trichadenotecnum malayense New (Figs 1M,16,17) Trichadenotecnum malayense New, 1975: 257;New & Lee, 1992: 156;Endang, Thornton & New, 2002: 156;Endang & New, 2005: 24. Specimens examined. Remarks. This species was originally described only on female specimens from Malaysia, and the male of this species was later identified from Sumatra, Indonesia by Endang & New (2005). The female genital structures examined here (Fig. 17) are in complete agreement with those described by New (1975), and male terminal structures of the specimens examined here (Fig. 16) agree very well with the description given by Endang & New Trichadenotecnum sibolangitense Endang & New (Figs 1N,18,19) Trichadenotecnum sibolangitense Endang & New, 2005: 27. Specimens examined.
Description of female genitalia. Egg guide (Fig. 19A) short, shorter than basal width, ventrally membranous and dorsally widely sclerotized, distal margin concave; body of subgenital plate broadly sclerotized, anteromedially with wide membranous region, posterior margin with pair of long, narrow and sharply pointed lateral processes. Gonapophyses (Fig. 19B). Ventral valve not exceeding external valve; dorsal valve broad, with deep pouch posteromedially, distal process long; external valve with broad membranous region anteriorly and with flap-like expansion posteriorly, posterior lobe broad, internal lobe narrow. Spermapore plate (Fig. 19C) symmetrical, pigmented around spermapore.
Remarks. This species was originally described from Sumatra, Indonesia based on a single male and is here recorded from Peninsula Malaysia for the first time. The male specimens examined here agree exactly with the original description except for the hypandrial structures being in mirror image (Fig. 18C). Given the exact similarity of each structure, we concluded that the illustration of the hypandrium of this species presented in Endang & New (2005) is drawn in dorsal (internal) view or reproduced in reversed position. This species is apparently closely related to T. alinguum Endang, Thornton & New, 2005 and, in T. sibolangitense, the right side of the hypandrium is strongly expanded, as illustrated in Fig. 18C. Given the lack of the hypandrial median tongue and the triangular shape of the hypandrium, this species is considered to be closely related to T. arciforme (Thornton, 1961;Yoshizawa & Lienhard, 2004). In contrast, these species are clearly different in the condition of the epiproct: strongly expanded over the clunium in T. sibolangitense (Fig. 18A) whereas almost flat and covered by the clunial flap in T. arciforme. The state as observed in T. arciforme is unique even among the tribe Ptyctini, in which Trichadenotecnum is classified. Because of this highly unusual feature, Yoshizawa & Lienhard (2004) did not assign T. arciforme to any species group, although they pointed out that T. arciforme is possibly closely related to T. apertum. The strongly expanded epiproct as observed in T. sibolangitense and similarities of the hypandrial structure between this species and T. arciforme corroborate the idea that T. arciforme (hypandrial median tongue absent and ventral valve of gonapophyses short) is closely related to T. apertum and its relatives, and that the highly specialized condition of the epiproct-clunium interface has to be considered as an autapomorphy of T. arciforme.
The female of T. sibolangitense is described here for the first time. As shown in Fig. 19A, it is characterized by possession of a pair of sharp processes arising from the posterior margin of the subgenital plate. Similar structures have been reported from some species such as T. suwai Yoshizawa & Lienhard, 2007 from Nepal or T. laticornutum Endang et al., 2002 andT. waykananense Endang &New, 2005 from Indonesia. The former species is phylogenetically distant (belonging to the spiniserrulum group) from T. sibolangitense so that presence of these processes can be recognized as convergence. In contrast, although the species-group assignments of T. laticornutum and T. waykananense have not been proposed in the original descriptions (Endang et al., 2002;Endang & New, 2005), their close affinity with T. sibolangitense is very likely.  Trichadenotecnum santosai Endang & Thornton (Figs 1O,20,21) Trichadenotecnum santosai Endang & Thornton, 1992: 364. Specimens examined.
Remarks. This species was originally described from Bali (type locality) and Lombok, Indonesia and is here recorded from Peninsula Malaysia for the first time. The specimens examined in this study match very well with the original description, including male terminalia ( Fig. 20; Note: original figure of hypandrium in mirror image, probably drawn in dorsal (internal) view or reproduced in reversed position) and female genitalia (Fig. 21). This species is very closely related to T. auritum Yoshizawa & Lienhard, 2004 and, based on male and female genital structures, the close affinity of these species with T. apertum Thornton, 1961 is undoubted (Yoshizawa & Lienhard, 2004).
The female of this species resembles T. medium and its relatives in having a short anteromedian sclerotized band on the subgenital plate (Fig. 23A). However, the hypandrial structure (Fig. 22C) and the short ventral valve of the gonapophyses (Fig. 23B) strongly suggest that this species is closely related to T. apertum and its relatives. The hypandrial median tongue is relatively well developed (Fig. 22C), as observed in T. majus and its relatives, representing a plesiomorphic condition within the majus group. As mentioned by Endang et al. (2002), T. quadrispinosum is very closely related to T. soenarti. Trichadenotecnum cornutum Endang & New (Figs 1Q,24) Trichadenotecnum cornutum Endang & New, 2005: 28. Specimens examined.
Remarks. This species was originally described from Sumatra, Indonesia based on a single male and is here recorded from Peninsula Malaysia for the first time. The specimens examined here are in good agreement with the original description of T. cornutum (Fig. 24). However, Endang & New (2005) described the phallosome of T. cornutum as open posteriorly, whereas, in the presently examined specimens, this structure is closed (Fig. 24D). However, the figure of the strongly deformed phallosome in the original description clearly shows that it was broken before being illustrated. Apart from the posteromedian part, the phallosomal structure of the present specimens is also in good agreement with the original illustration of T. cornutum. Therefore, we concluded that the difference between the present specimens and the original description corresponds to an artifact, and that the specimens examined here are conspecific with T. cornutum. Examination of the complete phallosome showed a very particular, long posteromedian spine, which is uniquely observed in this species within the genus Trichadenotecnum.
Because the female of this species remains unknown, identification of its close relatives among the majus group is difficult. Presence of the median tongue resembles T. majus and its relatives, but it represents a plesiomorphic condition within the group. T. cornutum resembles T. soenarti, for which both sexes are known, in male hypandrial structures (median tongue present) and in the structure of the phallosome (posterior margin strongly extended ventrally). Their close affinity is undoubted so that T. cornutum is also likely to be closely related to T. apertum and its relatives, as well as T. soenarti. Trichadenotecnum anomalum Yoshizawa & Lienhard,sp.n. (Figs 1R,25) Holotype. Male, Singapore, Bukit Timah forest, 24 hr survey, section trap 5-9 am, 26.x.1969, D.H. Murphy (MHNG).
Description (note that the condition of the specimen is bad). Male. Head. Yellowish in ground color; vertical markings and orbital markings obscure; coronal suture black; epicranial suture faintly bordered with brown band dorsally; frons with faint .markings; eye black, IO/D= 1.1; ocelli white, ocellar field brown; gena mostly yellowish; postclypeus with seven rows of spots dorsomedially; anteclypeus brown. Antenna pale brown. Mouthparts brown.
Thorax. Prothorax brown. Mesonotum yellowish, lateral lobe of scutum with faint marking, anterolateral corners of scutellum brown. Metanotum yellowish, lateral lobe of scutum with faint marking, lateral corner of scutellum brown. Meso-and metapleuron pale brown except for white membranous regions.
Legs. Mostly pale brown, foretibia and tarsi of all legs darker. Forewing (Fig. 1R). Hyaline, lacking almost all marking elements characteristics for Trichadenotecnum except following: opposing spots in cell r distinct, fused to single marking; basal band only distinct around Rs-M fusion and in cell cua.
Terminalia. 8th sternum with pair of sclerotized portions (Fig. 25AC). Clunium (Fig. 25A) with broad lateral arms, about parallel sided and with strong dorsal expansion apically. Epiproct (Fig. 25AB) strongly expanded dorsally and anteriorly over clunium, posteriorly with small conical projection medially; dorsal margin of epiproct lobe broad and posterior surface convex. Paraproct (Fig. 25A) with small distal lobe and with short and almost straight distal process directed posteriorly. Hypandrium (Fig. 25C) asymmetrical, without median tongue but with tongue-shaped projection medially near anterior margin, anterolaterally with pair of areas densely covered with tiny denticles, distally with two processes, one arises medially (= right process), acute conical in shape, the other arises left to this process (= left process), broad with diagonally truncated distal end; right posterolateral corner with denticulated longitudinal keel and with broadly expanded plate with serrated distal and internal margins. Phallosome (Fig. 25D) with broad distal margin slightly concave medially, about parallel sided and narrowing to pointed anterior end, lacking anterior process.

Female. Unknown.
Remarks. This species is very unusual in its forewing markings (Fig. 1R). Although density is variable, the forewings of all known Trichadenotecnum species are broadly spotted ( Fig. 1A-Q, S-U). Especially, in less heavily spotted species, six characteristic submarginal spots are always clearly visible, and this is considered to be an autapomorphy of the genus. In contrast, the forewings of T. anomalum are mostly transparent except for some markings in the basal half (Fig. 1R). The condition of the holotype specimen is poor, with many body markings faded. However, the basal band of the forewing can be seen clearly in the holotype so that absence of the marking characteristics of Trichadenotecnum, such as the submarginal row of spots, is probably not due to the poor condition of the specimen. The 9th tergite of T. anomalum possesses posterolateral arms (Fig. 25A), a character which is uniquely observed in Trichadenotecnum among Psocidae. Other diagnostic characters of the genus, such as the triangular areola postica, can also be seen (Fig. 1R). Therefore, the species is here considered as a member of Trichadenotecnum. The species seems to be closely related to T. apertum and its relatives because of the absence of the hypandrial median tongue and other resemblances of the terminal structures. The unique forewing markings are probably autapomorphic to this species.  Trichadenotecnum rachimi Endang & Thornton (Figs 1S,26) Trichadenotecnum rachimi Endang & Thonton, 1992: 363. Trichadenotecnum arciforme: New & Lee, 1992 [Not Trichadenotecnum arciforme Thornton, 1961: 11;Yoshizawa & Lienhard, 2004: 141. (Hong Kong)] [Not Trichadenotecnum arciforme: Thornton, 1984: 157. (Indonesia)] Specimens examined.
Remarks. Judging from the wing markings and female genitalia, the specimens listed above were identified as T. rachimi. This species was described originally from Lombok, Indonesia, and is here recorded from Peninsular Malaysia for the first time. The females examined here are also undoubtedly conspecific with the females recorded under the name of T. arciforme from Peninsular Malaysia by New & Lee (1992). However, the Malaysian females are clearly different from the female of T. arciforme described from Hong Kong (Yoshizawa & Lienhard, 2004). Female T. arciforme recorded from Bali and Lombok, Indonesia (Thornton, 1984) differs from both Malaysian and Hong Kong species recorded under the name.
The short ventral valve of gonapophyses ( Fig. 26B) strongly suggests that these females are closely related to T. apertum and its relatives. Among the species examined here, the females of T. anomalum and T. cornutum are unknown to date. T. cornutum is significantly larger than T. rachimi and has different forewing markings ( Fig. 1Q and S) so that its conspecificity is less likely. T. anomalum and T. rachimi are significantly different in their forewing markings ( Fig. 1R and S), although T. anomalum is known only from a single male and its intraspecific variation cannot be evaluated.  serrated internally; right process keel-like, densely covered by denticles ventrally. Phallosome (Fig. 27D) broad on posterior margin with shallow concavity in middle, broadened laterally, narrowing to somewhat truncated anterior end, without anterior process.
Remarks. The species was originally described from Sumatra, Indonesia based only on female specimens and is here recorded from Peninsula Malaysia for the first time. The male of T. kerinciense is also described here for the first time. Female genitalia (Fig. 28) match very well with the original description of this species, especially in the characteristic body part of the subgenital plate. Judging from the male hypandrium structure, especially the unmovable median tongue (Fig. 27C), its placement within the majus group is unquestionable. The hypandrial median tongue is well developed and the ventral valve of gonapophyses is long, both characters are widely observed in other species group and thus are plesiomorphic conditions within the majus group.
[Malaysia] 1 male, Gnung Berembun, Path 3, Cameron Highlands, Pahang, 14.vii.2003, HK.etal (canopy fogging, St. 6). Remarks. The species was originally described from Java, Indonesia, later also recorded from Sumatra, Indonesia, and is here recorded from Peninsula Malaysia for the first time. The variation of the hypandrial structure, three (Javanese) or four (Sumatran) hypandrial spikes, has been pointed out by Endang & New (2005). The presently examined specimen has four hypandrial spikes. Judging from the original hypandrial illustration published by Endang et al. (2002: fig. 333), missing of some hypandrial parts on the left side of the specimen seems evident; on that side only one spike is figured whereas two on the other side. It is likely that the variation pointed out by Endang & New (2005) is actually an artifact caused by dissection or preparation.
The hypandrium of this species (Fig. 29C) is unique, very different from that of any other known species. Absence of the hypandrial median tongue may indicate its close affinity with T. apertum and its close relatives. In the original description of this species, the phallosome is illustrated in highly deformed condition, but the present examination clearly shows that this species has the pseudo-parameres on the phallosome (Fig. 29D). The presence of this structure is a synapomorphy of the sexpunctatum, medium and New World species groups (see Yoshizawa et al., 2008) so that assignment of T. krucilense into the majus group is excluded. However, this species cannot be assigned to the sexpunctatum, medium or New World species groups because of lack of apomorphies characterizing any of these species group so that a new species group for its own is proposed here. Females of this species are unknown to date.