The Pleidae (Hemiptera, Heteroptera) of Thailand, with the descriptions of two new species and a discussion of species from Southeast Asia Author Cook, Jerry L. Department of Biological Sciences, Sam Houston State University, Huntsville, TX 77341, USA jcook@shsu.edu Author Sites, Robert W. Enns Entomology Museum, University of Missouri, Columbia, MO 65211, USA Author Vitheepradit, Akekawat Department of Entomology, Kasetsart University, Bangkok 10900 Thailand text ZooKeys 2020 973 35 68 http://dx.doi.org/10.3897/zookeys.973.54026 journal article http://dx.doi.org/10.3897/zookeys.973.54026 1313-2970-973-35 0D8FEF96BC1B437E81ABD70BFCDB314A 302E15180DB756F5B144807DC8875EF9 Paraplea liturata (Fieber, 1844) Figures 12 , 13, 14 , 15 , 16-17 , 24C = Ploa liturata Fieber, 1844: Entomol. Monogr. p. 19. = Plea liturata : (Kirkaldy, 1904): Wien. Entomol. Zeit. 23: 129. = Plea metiadusa Distant, 1910: Fauna Brit. India 5: 337. = Plea rufonotata Distant, 1914: Rech. Sci.N.- Caledonie 2: 387. = Plea quinquenota Paiva, 1918: Rec. India. Mus. 14: 29. = Plea fasciata Horvath , 1918: Ann. Mus. Nat. Hung. 16: 144. Plea (Paraplea) liturata : Esaki and China 1928 : Rev. Esp. Entomol. 4: 166 (subgenus description). Paraplea liturata : Drake and Maldonado-Capriles 1956 (elevation to genus) 51: 53. Remarks. Paraplea liturata shares some of the same taxonomic history as P. frontalis since both were described in the same paper. Paraplea liturata was described as Ploa liturata by Fieber (1844) for specimens collected in the East Indies. This original description was not only brief but did not capture an important set of markings that is commonly present on specimens of P. liturata , even though the description relied almost completely on coloration. Fieber noted the typical two dashes between the eyes and the pair of lateral and single midline spots on the posterior margin of the pronotum. However, two spots on the anterior part of the pronotum were not listed in the description nor on his plate. The figure does show the hemelytral banding that is common in many specimens. Although types were not designated by Fieber, subsequent researchers were able to associate the description with many specimens of this common species, although none of these were those used by Fieber. Placement of this species into the genus Paraplea followed the same sequence as reported for P. frontalis . Kirkaldy (1904) moved this species into the genus Plea but made no comments on the species. Distant (1906) translated Fieber's description but did not further report on this species. Distant (1910) described Plea metiadusa from Calcutta, India and reported that it had no maculations, but this species was still determined to be a variant of P. liturata by Lundblad (1933) . In doing so, Lundblad (1933) commented on the variability of the pronotal spots that are most commonly five in number but can range from none to seven. Distant (1914) also described Plea rufonotata but did not associate it with P. liturata . The description of P. rufonotata from New Caledonia was the first time that a pronotum with five spots was described, which now appears to be the most common state for P. liturata . Again, Lundblad (1933) made the association of these species as being synonyms. Horvath (1918) described Plea fasciata from specimens from Batavia, Java (Indonesia) but this was considered a strongly-colored example of P. liturata ( Lundblad 1933 ). Plea quinquenota ( Paiva 1918 ) was described from a single specimen from Inle Lake, Yawnghe State (now in Myanmar) and was not examined by Lundblad (1933) , but was still synonymized with P. liturata based on the illustration of the new species perfectly matching the typical form of P. liturata . Along with the taxonomic clarifications, Lundblad (1933) also redescribed the species. Below is a supplemental description of the species, incorporating specimens from Thailand. Holotype . None designated. Type locality. Fieber (1844) reported that the description was made from specimens collected by Dr. Helfer in the East Indies but the exact location in South or Southeast Asia is unknown. Supplementary description. All measurements are in millimeters and were taken from 221 adult specimens from throughout Thailand as reported in the distribution portion of this paper. Specimens used for this description are deposited at UMC and SHSU. Body size . Total length, 1.27-1.68 (average 1.48) in Thailand specimens. Fieber (1844) reported total length of "approximately 2 mm." Distant (1910) gave the size of his later synonymized P. metiadusa from India and his later synonymized P. rufonotata from Caledonia as 2. The specimen used to describe the later synonymized P. quinquenotata was listed as 1.5 ( Paiva 1918 ). Anderson and Weir (2004) reported a length of 1.8-2.0 for this species in their key and reported it from Northern Territory and Western Australia. Color . Color may be quite variable among individuals within a population. Base color of body ranges from tan (Fig. 12A, B ) to white (Fig. 12D ). Darker bands on the sides of the hemelytra are common but not observed in all individuals. When present, bands are darker than the base color, ranging from light orange-brown (Fig. 12D ) to tan (Fig. 12A ) to brown (Fig. 12B ). Punctures are sometimes a darker shade (Fig. 12A ). Honeycombing matches the base color. Most specimens have five characteristic dark spots on the pronotum (Fig. 12B ); two on the anteromedial portion (Fig. 12 B ), two on the posterolateral region (one on each side) (Fig. 12B, D ), and one on the posteromedial region (Fig. 12B, D ). Conversely, these spots are sometimes faint or absent in some individuals and at some locations. Figure 12. Paraplea liturata specimens from Thailand. A female in lateral view with typical coloration, B dorsal view in that shows dark banding and honeycombing. C frontal view of specimen in B. D mele showing an alternatively colored form in lateral view. Head . Head (Fig. 12C ) colored with base body coloration, ranging from white to brown. Many specimens with a vertical light-colored bar between eyes. If present, bar can be thin (Fig. 12C ) to wider, sometimes occupying nearly a third of width between eyes. Two dark spots common on face, one on each side between central bar and eyes (Fig. 12C ). Eyes in dried specimens range from red to gold. Mouthparts usually darker that the rest of head. Antenna three-segmented, usually hidden from view below eye. Head size similar among Thailand specimens, head width at widest point including eyes 0.62-0.79 (average 0.72), head width at narrowest point between eyes, 0.31-0.43 (average 0.40), OI 47-58 (average 53). Pronotum (Fig. 12B ). Base color ranging from white to light brown and honeycombing apparent in most specimens; most specimens with five dark spots on pronotum (91% of Thailand specimens with five spots, ~ 9% with three spots and lacking anterior pair, less than 1% with no spots); a shallow puncture in center of each cell of honeycomb and under high magnification a minute hair can usually be seen coming from each pore; with slight bulge posteriorly, wider than long, width 0.66-0.92 (average 0.79); pronotum length 0.33-0.57 (average 0.46); PI 39-68 (average 58). Wings . Complete to posterior; punctures equally spaced but not generally in rows (0.03 in diameter) (Fig. 12A, D ); underlying honeycomb structure usually present; claval suture distinct, complete; scutellum with punctures smaller (0.01 in diameter), more widely spaced than other punctures, scutellum base color often golden but sometimes dark brown and often darker than hemelytra; honeycombing absent from scutellum, making it appear somewhat transparent; lateral view shows distinct horizontal dorsal profile and near vertical aspects posterior profile with a transition of nearly 90° (Fig. 12A, C ); scutellum slightly wider than long (Fig. 12B ), length 0.28-0.43 (average 0.35); width 0.31-0.50 (average 0.40); SI 103-139 (average 116). Hind wings membranous, fully developed, completely concealed by hemelytra. Legs . Legs with numerous hairs and small spines (Fig. 13 ), hairs prevalent on apical half of prothoracic and mesothoracic tibiae, without numerous long hairs as found on tibia and tarsus of many Paraplea . Typical leg measurements: prothoracic leg coxa 0.05, trochanter 0.09, femur 0.42, tibia 0.35, first tarsomere 0.04, second tarsomere 0.09,pretarsal claw 0.08; mesothoracic leg coxa 0.06, trochanter 0.10, femur 0.41, tibia 0.28, first tarsomere 0.03, second tarsomere 0.13, pretarsal claw 0.08; metathoracic leg coxa 0.05, trochanter 0.10, femur 0.39, tibia 0.49, first tarsomere 0.02, second tarsomere 0.13, third tarsomere 0.18, pretarsal claw 0.11. Figures 13, 14. Paraplea liturata. 13 prothoracic leg above, mesothoracic leg in the middle and metathoracic leg below. 14 profile of the ventral keel with the anterior end (thoracic) to the top and ventral to the left. Median ventral keel . Thoracic portions distinctly separate, prothoracic keel generally rounded, two posterior thoracic segments serrated; abdominal keel with first two segments fused, teeth more pronounced posteriorly (Fig. 15 ). Several minor differences between Thailand specimens reported here and those reported by Lundblad (1933 : fig. 42C-E). Figure 15. Ovipositor of Paraplea liturata . Scale bar: 0.05 mm. Characters of female . Ovipositor most commonly as in Fig. 15 . Ovipositor roughly rectangular in shape but with apical gonapophysis 1 slightly wider; five distinct teeth along posterior border (apical row) plus two teeth on ventral border (ventral 1 and 2); two rows of small teeth away from apex, three primary, three secondary, and occasionally one tertiary (not shown in Fig. 15 ); bottom secondary tooth larger and extends slightly beyond end of ovipositor, making it sometimes appear as being along posterior margin; three to five long hairs on ventral side of region where gonapophyses 1 and 2 meet; Subgenital plate slightly longer than wide (Fig. 16 ), length ~ 0.30, width ~ 0.26, lightly rugose in basal half followed apically by a series of pits, dark region in center near apex, pair of tufted hairs on each side near apex. Characters of male . Aedeagus bulbous and somewhat asymmetrical in typical fashion of family; operculum (subgenital plate) as in Fig. 17 , most of surface slightly rugose, longer than wide, length ~ 0.27, width ~ 0.20. Figure 16, 17. Genital plates of Paraplea liturata . 16 female 17 mele. Scale bar: 0.1 mm. Distribution. In Thailand, Paraplea liturata appears to be mostly a peninsular species on the southwest side of the country, although there are two records of it in the eastern region of Thailand; one in Sakon Nakhon Province, which was reported by Chen et al. (2006) , and one by RWS and AV in a pond in Ubon Ratchathani Province (Fig. 24C ). It is also known from Australia (Anderson and Weir 2004), India ( Paiva 1918 . Lundblad 1933 ), Indonesia ( Lundblad 1933 , Nieser and Chen 1999 ), Malaysia ( Fernando and Cheng 1974 ), Myanmar ( Paiva 1918 ), New Caledonia ( Lundblad 1933 ), Philippines ( Lundblad 1933 , Nieser and Chen 1999 ), and Taiwan ( Mitamura et al. 2018 ). Material examined. Chai Nat Province : Amphur Neon, Kham, Ban Wang Kor Hai, 14°57.934'N , 99°50.668'E , 24 V 2004, Vitheepradit & Prommi, L-710 (1 specimen UMC). Chumphon Province : Amphur Sa Wi, Tumbon Kron, 10°17.499'N , 99°05.530'E , 21 VI 2006, Vitheepradit & Prommi, L-967 (4 specimens UMC). Kalasin Province : Phu Pan National Park, 7 VI 1998, Sites, Simpson & Vitheepradit, L-165 (3 specimens UMC, 1 specimen SHSU). Krabi Province : Amphur Mueang, Klong Muang Beach, pond, 8°02.979'N , 98°45.540'E , 13 m, 8 VIII 2005, Sites, Vitheepradit, Simpson & Prommi L-862 (8 specimens UMC); Amphur Mueang, Nopphorat Thara Beach, pond, 8°02.625'N , 98°48.517'E , 8 V 2005, Sites, Vitheepradit & Prommi, L-805 (1 specimen UMC); Amphur Nuea Khlong, Tumbon, Nuea Khlong, Ban Paga Sai, pond, 8°02.619'N , 99°01.144'E , 27 m, 8 VIII 2005, Sites, Vitheepradit, Simpson & Prommi, L-864 (2 specimens UMC); Amphur Nuea Khong, Pan Paga Sai, pond, 8°02.619'N , 99°01.144'E , 27 m, 13 VI 2006, Vitheepradit & Prommi, L-951 (1 specimen UMC); Tumbon Klong Kanarn, Ban Klong Kanarn, pond, 8°01.045'N , 99°00.450'E , 37 m, 9 VIII 2005, Sites, Vitheepradit, Simpson & Prommi, L-867 (2 specimens UMC). Phang Nga Province : Amphur Mueang, Tumbon Na Pring, pond, 8°31.750'N , 98°32.001'E , 5 I 2006, Sites, Vitheepradit & Prommi, L-887 (2 specimens UMC); Amphur Mueang, Tumbon Na Prig, pond, 8°31.750'N , 98°32.001'E , 12 VI 2006, Sites, Vitheepradit & Prommi, L-946 (4 specimens UMC); Amphur Takua Thung, Tumbon Krasom, Ban Bang Mak, pond, 8°24.553'N , 98°27.434'E , 12 VI 2006, Sites, Vitheepradit & Prommi, L-945 (1 specimen UMC); Amphur Takua Pa, Tumbon Bang Nai Si, Ban Bang Yai, pond, 08°25.950'N , 98°23.192'E , 20 m, 8 VI 2006, Sites, Vitheepradit & Prommi, L-927 (1 specimen); Amphur Thai Mueang, Tumbon Na Teoy, Ban Bang Klee, 8°18.655'N , 98°17.552'E , 19 m, 2 VI 2005, Sites, Vitheepradit & Prommi, L-825 (8 specimens UMC); Amphur Thai Mueang, Tumbon Na Teoy, Ban Bang Klee, 8°18.655'N , 98°17.552'E , 19 m, 5 I 2006, Sites, Vitheepradit & Prommi, L-886 (11 specimens UMC); Amphur Thai Mueang, Tumbon Na Teoy, Ban Bang Klee, 8°18.655'N , 98°17.552'E , 9 VI 2006, Sites, Vitheepradit & Prommi, L-935 (11 specimens UMC). Phatthalung Province : Amphur Tamot, Tumbon Loh Jak Kra, 7°20.244'N , 100°01.285'E , 44 m, 3 VI 2004, Vitheepradit & Prommi, L-736 (1 specimen UMC). Phuket Province : Amphur Thalang, Jud peat swamp, UV pan trap, 8°07.930'N , 98°18.156'E , 24 m, 7 I 2006, Sites, Vitheepradit & Prommi, L-899 (92 specimens UMC, 5 specimens SHSU); Amphur Thalang, Jig peat swamp, 8°08.772'N , 98°17.892'E , 23 m, 7 I 2006, Sites, Vitheepradit & Prommi, L-906 (44 specimens UMC, 3 specimens SHSU); Amphur Thalang, Laem Yot peat swamp, 8°08.325'N , 98°17.927'E , 28 m, 29 V 2005, Sites, Vitheepradit & Prommi, L-807 (1 specimen UMC); Amphur Thalang, Mai Khoa peat swamp, 8°07.461'N , 98°18.193'E , 19 m, 7 I 2006, Sites, Vitheepradit & Prommi, L-900 (42 specimens UMC, 1 specimen SHSU); Amphur Thalang, Tumbon Mai Khao, Moo Ha, 8°10.718'N , 98°17.611'E , 23 m, 31 V 2005, Sites, Vitheepradit & Prommi, L-816 (1 specimen SHSU); Amphur Thalang, Tumbon Mai Khao, Moo Ha, pond, 8°10.718'N , 98°17.611'E , 23 m, 5 I 2006, Sites, Vitheepradit & Prommi, L-889 (17 specimens UMC). Songkhla Province : Amphur Rataphum, Tumbon Kampangphet, 07°08.030'N , 100°18.437' E, 30 m, 2 VI 2004, Vitheepradit & Prommi, L-732 (9 specimens UMC). Surathani Province : Amphur Ban Na Ban Huay Hamg, 8°08.925'N , 99°18.215'E , 24 m, 20 VI 2004, Vitheepradit & Prommi, L-781 (3 specimens UMC); Amphur Phunphin, Tumbon Boh Rai, 8°53.866'N , 98°08.961'E , 10 m, 7 VI 2004, Vitheepradit & Prommi, L-751 (1 specimen SHSU); Amphur Ban Na, Ban Tha Rau Tai, 8°56.567'N , 99°15.130'E , 4 m, 20 VI 2004, Vitheepradit & Prommi, L-782 (3 specimens UMC). Trang Province : Amphur Sikao, Tumbon Mai Fard Ban, Klong Maeng, pond, 8°30.170'N , 99°20.541'E , 6 m, 10 VIII 2005, Sites, Vitheepradit, Simpson & Prommi, L-868 (2 specimens UMC, 1 specimen SHSU). Uthai Thani Province : Amphur Mueang, Tumbon Nam Serm, 15°20.886'N , 100°02.120'E , 24 V 2004, Vitheepradit & Prommi, L-713 (3 specimens UMC). Ubon Ratchathani Province : Amphur Nam Khoon, Ban Non Yang, pond, 14°28.176'N , 104°53.782'E , 190 m, 10 IV 2004, Sites & Vitheepradit, L-613 (10 specimens UMC). Discussion. There is considerable variation in P. liturata if this is a single species. Lundblad (1933) mentioned this variation when synonymizing P. fasciata , P. metiadusa , P. quinquenotata , and P. rufonotata with P. liturata . The main basis for Lundblad synonymizing these species was the similarity of the abdominal keel. The drawings in his paper ( Lundblad 1933 ) show similarities but there are also subtle variations. The Thailand specimens are also somewhat different from those in the Lundblad drawings. These data preserve the question as to whether this is a variable species or multiple species. Several of the species Lundblad synonymized had been described as having a length of 2 mm, although he stated that this was a small species varying between 1.3 to 1.7 mm. This size range of Lundblad's specimens coincides with that of specimens from Thailand; however, it still does not account for those described from India, New Caledonia, and Australia. The original description of P. liturata by Fieber (1844) listed the length imprecisely as "approximately 2 mm" and it is uncertain where in the East Indies these specimens were collected or how accurately that measurement was made. Likewise, some distinct differences in coloration and characters occur in specimens from the various regions. A character that links all of these specimens into one species is the general state of having five spots on the pronotum. In many populations there can be specimens without these markings although the majority of specimens tend to always have five black pronotal spots. Thus, with a single specimen it may not be possible to rely on this trait but with a series of specimens it is easy to determine the species as P. liturata , as it is now defined. There appears to also be some consistency in the characters of the ovipositor. In many of the Thailand specimens, the ovipositor appears remarkably like that figured by Lundblad (1933 : fig. 42H) from Java (Indonesia) which is consistent with the ovipositor being a reliable character for species recognition (see Sublett and Cook 2015 ); however, a small number of specimens lack some or all of the secondary and tertiary teeth. Since the ovipositors of specimens from India, New Caledonia and Australia are unknown, there is still a question as to if specimens from these regions are actually P. liturata. Like P. frontalis , P. liturata could be a species complex. Both these species need additional study to determine their status. Ecology : The habitat of P. liturata in Thailand is typical for the majority of pleids. This species was often found in ponds, in shallow water with vegetation.