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ABSTRACT—Three productidine brachiopods of Haydenella, Paraplicatifera, and Compressoproductus are recovered from the Tak Fa Formation (Wordian, ...
J. Paleont., 83(5), 2009, pp. 804–810 Copyright ’ 2009, The Paleontological Society 0022-3360/09/0083-0804$03.00

MIDDLE PERMIAN PRODUCTIDINE BRACHIOPODS FROM CENTRAL THAILAND (THE INDOCHINA TERRANE) WITH PALEOBIOGEOGRAPHIC IMPLICATIONS MASATOSHI SONE,1 CHONGPAN CHONGLAKMANI,2

AND

ANISONG CHITNARIN3

1

School of Engineering and Science, Curtin University of Technology, Sarawak Campus, 98009 Miri, Malaysia, ,[email protected].; 2 School of Geotechnology, Suranaree University of Technology, 30000 Nakhon Ratchasima, Thailand; and 3School of Biology, Suranaree University of Technology, 30000 Nakhon Ratchasima, Thailand

ABSTRACT—Three productidine brachiopods of Haydenella, Paraplicatifera, and Compressoproductus are recovered from the Tak Fa Formation (Wordian, Middle Permian) of the upper Saraburi Limestone Group exposed at Khao Wong of central Thailand (the western margin of the Indochina Terrane). The latter two genera are new to the Permian of Thailand, and the new species Paraplicatifera thaica is proposed herewith. Some taxonomic and nomenclatural problems in relation to the three genera are discussed. The assemblage suggests endemism for a Middle Permian marine faunule of the Indochina Terrane.

INTRODUCTION

brachiopods (suborder Productidina), Haydenella granti Yanagida and Nakornsri, 1999, Paraplicatifera thaica n. sp., and Compressoproductus sp., are described from the Middle Permian Tak Fa Formation of the upper Saraburi Limestone Group, exposed at Khao Wong of central Thailand. The latter two genera are new to the Permian of Thailand, and it is the first time Paraplicatifera Zhao and Tan, 1984 is confirmed outside China. The generic diagnosis for Paraplicatifera is emended based on the new material. The sampling area is situated within the Early–Middle Permian Khao Khwang Platform, an outer carbonate platform developed along the western margin of the Indochina Terrane. Forty species of brachiopods, seven of which are new, were known previously from the Khao Khwang Platform (Pe´rez-Huerta et al., 2007; Yanagida, 1964; Yanagida and Nakornsri, 1999; Yanagida et al., 1988), suggesting the potential existence of a diverse Middle Permian fauna. All examined specimens of the three taxa, including the type series of Paraplicatifera thaica n. sp., are deposited in the Geology Program, the National University of Malaysia (Universiti Kebangsaan Malaysia), with registration numbers prefixed UKM-F.

T

HREE PRODUCTIDINE

FOSSIL SAMPLING LOCALITY AND AGE

Khao Wong, where the current material was collected, is a large limestone hill, located about 18 km north of Khok Samrong town and is just southwest of Ban Nong Muang village along Route 1 (Fig. 1). It consists of medium- to thickbedded, dark gray limestone intercalated with gray shale lamina. Brachiopods, corals, bryozoans, and foraminifers (both fusulinoids and smaller foraminifers) are abundant at many exposures. Brachiopods were sampled at three localities: KW1, KW4 (N15u12914.50, E100u38959.90), and WTS2 (N15u12909.60, E100u40901.10). The first two localities are near each other and are accessible from the north of the hill. KW1 is near the entrance to the temple Wat Khao Wong, and KW4 is accessible via a footpath from the temple. WTS2 is near temple Wat Tham Santisuk, east of the hill. Strata at these localities have strikes and dips of 25u/10uW (KW1), 0u/ 20uW (KW4), and 330u/10uW (WTS2). The brachiopod beds are fusulinoid-rich, biomicritic wackestones. No conodonts have been found at Khao Wong (Metcalfe and Sone, 2008).

Based on the bedding orientation, the sequence in Khao Wong is interpreted as more or less stratigraphically equivalent to limestones exposed around the temple Wat Khiri Nakharattanaram about 2 km to the southeast, where seven fusulinoids, Nankinella? sp., Neofusulinella saraburiensis Toriyama, Kanmera, and Ingavat, 1969, Neofusulinella lantenoisi Deprat, 1913, Verbeekina verbeeki (Geinitz, 1876), Pseudodoliolina pseudolepida (Deprat, 1912), Sumatrina annae Volz, 1904, and Parafusulina gigantea (Deprat, 1913), were reported by Toriyama and Pitakpaivan (1973). Toriyama and Pitakpaivan (1973) suggested an early Middle Permian age for the assemblage. However, in modern view the association of N. lantenoisi, V. verbeeki, P. pseudolepida, S. annae, and Laosella gigantea (Deprat, 1913) should be regarded as a fauna in a higher level of the Permian, that is, an upper Murghabian to Midian (e.g., Leven, 1997). The fusulinoids P. pseudolepida, Chusenella cf. C. cambodgiensis (Gubler, 1935), and Minojapanella sp. were identified from Locality KW4 of Khao Wong. This assemblage is considered to be related to, and the same age as, the above Wat Khiri Nakharattanaram fauna. Nevertheless, index fusulinoid genera for the basal Midian Stage, Yabeina Deprat, 1914 and Lepidolina Lee, 1934 (see Leven, 2003, p. 277), have not been found in the Tak Fa Formation; therefore, the current brachiopod-fusulinoid sequences in Khao Wong are regarded tentatively as late Murghabian (Wordian) in age, although a possibility of Midian age cannot be excluded. SYSTEMATIC PALEONTOLOGY

Suborder PRODUCTIDINA Waagen, 1883 Superfamily PRODUCTOIDEA Gray, 1840 Family PRODUCTELLIDAE Schuchert, 1929 Subfamily PRODUCTININAE Muir-Wood and Cooper, 1960 Tribe CHONETELLINI Licharew in Sarytcheva et al., 1960 Genus HAYDENELLA Reed, 1944 Discussion.—There is disagreement over the familial-group classification of Haydenella. The need to restudy this genus was highlighted by Muir-Wood and Cooper (1960), who placed Haydenella in the Marginiferidae Stehli, 1954. Grant (1976) then suggested a linoproductid affinity for the genus, based on the nature of silicified cardinal processes seen in some Pakistani species. Jin and Hu (1978) disagreed and placed the genus in their new subfamily Haydenellinae under

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and Cooper, 1960, p. 225). However, conspicuous, regular concentric lines are displayed over the entire shells of Haydenella granti Yanagida and Nakornsri, 1999 from Khao Hin Kling, central Thailand; this adds new information to the genus. HAYDENELLA GRANTI Yanagida and Nakornsri, 1999 Figure 2.1–2.4

FIGURE 1—Locality map of Khao Wong limestone hill in central Thailand, indicating the three fossil sampling localities KW1, KW4, and WTS2. Inserted tectonic subdivision map of mainland Southeast Asia after Sone and Metcalfe (2008, fig. 1).

the family Chonetellidae Licharew in Sarytcheva et al., 1960. Brunton et al. (2000) also placed Haydenella under the tribe Chonetellini, and this is followed in this study. Here, it is argued that Haydenella is not a member of the Linoproductoidea, because the genus lacks spines at the hinge. Haydenella was thought to have no rugae on the body (e.g., Brunton et al., 2000) or have only fine growth lines (Muir-Wood

Haydenella granti YANAGIDA AND NAKORNSRI, 1999, p. 127, pl. 29, figs. 1a–d, pl. 30, figs. 2–8. Material examined.—Two ventral valves UKM-F570 and UKM-F571 from localities KW4 and KW1 of Khao Wong, central Thailand. Discussion.—The available shells are decorticated, and thus the body surface appears rather smooth. The ornament of radial costae and feeble concentric lines is weakly preserved, and the ears are well ornamented with even rugae, typical for Haydenella. The new shells from Khao Wong are referable to Haydenella granti, known previously from the correlative limestone sequence at Khao Hin Kling (Middle Permian) in the south of Petchabun, central Thailand. This identification is indicated by their similar transverse outline, moderately convex profile, flattened venter, and relatively thick costellae. The tendency of anteriorly fusing costellae for H. granti characterized by Yanagida and Nakornsri (1999, p. 127) is also exhibited by the present specimens. The regular, tight and fine concentric lines seen in the Khao Hin Kling H. granti is however not observable in the present decorticated shells. Productus kiangsiensis Kayser, 1883 (p. 185, pl. 26, figs. 6– 11), the type species of Haydenella, from the Wuchiapingian of South China, also has a weak to moderately convex ventral profile. It, however, tends to have a semiovate to elongate outline with a relatively short hinge line, and thus it can be differentiated from H. granti. Another Thai form reported as Haydenella kiangsiensis (Kayser) from near Ban Phot, north of Khao Hin Kling, by Yanagida (1964, p. 8, pl. 2, figs. 1a–e, pl. 3, figs. 4a–e) has a shorter hinge line and higher ventral convexity than H. granti. Subfamily OVERTONIINAE Muir-Wood and Cooper, 1960 Tribe COSTISPINIFERINI Muir-Wood and Cooper, 1960 Genus PARAPLICATIFERA Zhao and Tan, 1984, emend. herein Type species.—Paraplicatifera regularis Zhao and Tan, 1984; upper Chihsia Formation (late Kungurian), Hunan, South China. Included species.—1) Productus (Marginifera) typicus elongatus Huang, 1932, Maokouan, Guizhou, South China; 2) Neoplicatifera costata Ni in Yang et al., 1977, Maokouan, Hubei, South China; 3) Neoplicatifera lata Tan and Ding in Liu et al., 1982, Maokouan, Hunan, South China; 4)

FIGURE 2—Haydenella granti Yanagida and Nakornsri, 1999 from the Tak Fa Formation at Khao Wong, central Thailand. 1, immature ventral valve UKM-F570 from Locality KW4, ventral view; 2–4, possible mature ventral valve UKM-F571 from Locality KW1; ventral, anterior, lateral views, showing a row of spine ridges along the bottom of the lateral flank to disk margin.

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FIGURE 3—Paraplicatifera thaica n. sp. from the Tak Fa Formation at Khao Wong, central Thailand. 1–3, ventral valve UKM-F572 from Locality KW1; posterior (indicating the positions of lateral spines in row along the margin of the disk and flank and of thinner spines on the ears), anterior, and lateral views; 4–9, ventral valve UKM-F573 from Locality KW4, posterior (4, 5), middle, anterior, and lateral (8, 9) views; 10–12, dorsal external mould UKM-F574 from Locality KW1, direct, anterior, and lateral views; 13–19, holotype, ventral valve UKM-F575 from Locality KW4, posterior (13, 14), lateral (15, 16), middle, and anterior (18, 19) views. Ventral spine base arrangement indicated with small circles in the brighter images 5, 9, 14, 16, and 19.

Paraplicatifera sangzhiensis Zhao and Tan, 1984, late Chihsian, Hunan, South China; 5) Costispinifera costatiformis Ding in Zhang et al., 1983, Maokouan, Gansu, Northwest China; 6) Paraplicatifera thaica n. sp., Wordian, central Thailand. Diagnosis (emended).—Medium sized Costispiniferini. Subquadrate to transverse corpus with elongate trail. Moderately to strongly geniculate. Typically subcylindrical outline in some species. Ventral trail costate, spinous. Ventral visceral disk weakly inflated, with small umbo. Rugae stronger than costae

on disks of both valves, regular or weakly undulating. Anterior ribbing moderate to coarse. Dorsal valve with similar ornament to ventral valve; dimples scattered over disk; median sulcus weak to almost absent. Spines in ventral valve only, quincuncially arranged. No subperipheral ridge. Discussion.—Paraplicatifera Zhao and Tan, 1984 is a relatively recently recognized genus, separated from Neoplicatifera Jin, Liao, and Fang, 1974, and thus it has not yet been widely documented. In this study, six species, other than the type species, are confidently included in Paraplicatifera as listed above.

SONE ET AL.—PRODUCTIDINE BRACHIOPODS FROM THAILAND Neoplicatifera is admittedly the closest genus, whose type species is Plicatifera huangi Ustritsky in Ustritsky et al., 1960 [originally Plicatifera? minor (Schellwien) sensu Huang (1932)] designated by Jin et al. (1974). Neoplicatifera huangi (Ustritsky) has feeble costae on the trail, whereas species of Paraplicatifera possess coarser and stronger anterior ribs. This is the main and possibly only significant difference between Neoplicatifera and Paraplicatifera. Nevertheless, there may be a transitional form between the two genera in regard to the coarseness of anterior ribs. Zhao and Tan (1984) suggested that Marginifera sintanensis Chao, 1927 is such a form. Note that M. sintanensis was included by Zhao and Tan (1984) in Paraplicatifera, but it is here reinterpreted to belong to Neoplicatifera, as will be discussed later. One might argue that a difference in the coarseness of anterior costae is gradational and thus it is a specific variation rather than a generic character. However, the anterior ribbings between the type species of the two genera, namely, Neoplicatifera huangi and Paraplicatifera regularis, appear highly distinct. On balance, the full generic status for Paraplicatifera is accepted in this study. The family-group classification of both Paraplicatifera and Neoplicatifera has generated some disagreement. Zhao and Tan (1984) originally placed Paraplicatifera in the Marginiferidae. Brunton et al. (2000, p. 436), however, relocated the two genera in the tribe Costispiniferini of the subfamily Overtoniinae. Waterhouse (2002, p. 15–16), on the other hand, placed Neoplicatifera in his new subfamily Spinomarginiferinae, while Paraplicatifera was retained in the Costispiniferinae. In this study, Paraplicatifera and Neoplicatifera are considered closely related to each other and they are placed in the same tribe Costispiniferini after Brunton et al. (2000). Paraplicatifera ranges in age from the upper Chihsian (late Kungurian) to the Maokouan. Neoplicatifera is most common in Maokouan sediments of South China, but these occurrences may still include specimens of Paraplicatifera. As defined, Paraplicatifera appeared in the Kungurian, earlier than Neoplicatifera (see Zhao and Tan, 1984). The co-occurrence of the two genera has never been confirmed. The dorsal valve of Paraplicatifera regularis or other congeneric species was not known. Brunton et al. (2000, p. 436) assumed it spinose. However, the dorsal external mould of the present new species from Thailand confirms the absence of spines (Figs. 3.10–3.12), and thus it is interpreted that Paraplicatifera lacked spines in the dorsal valve. Costispinifera Muir-Wood and Cooper, 1960 is also a small genus with an elongate trail and strong, undulating rugae on the visceral disk and is admittedly related to Paraplicatifera. A prime difference from Paraplicatifera is the possession of numerous fine dorsal spines. Costispinifera is common in the Roadian–Wordian of West Texas. It has also been reported sporadically from the Permian of China but is often represented by inadequately described specimens (e.g., Feng and Jiang, 1978, p. 252; Hu, 1989; Jin et al., 1979, p. 90; Li et al., 1986; Liao, 1980). Therefore, the Tethyan occurrence of true Costispinifera requires confirmation, with reference to Paraplicatifera or Neoplicatifera. Costispinifera is perhaps more remotely allied to Paraplicatifera and Neoplicatifera. The marginiferan Spinomarginifera Huang, 1932 appears superficially similar to Paraplicatifera and/or Neoplicatifera. A key difference is the absence of a subperipheral rim or ridge in the latter two costispiniferin genera.

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PARAPLICATIFERA THAICA new species Figure 3.1–3.19 Diagnosis.—Medium to large Paraplicatifera. Widest at hinge line. Ears slightly extended. Rugae weakly undulated. Anterior ribbing moderately coarse but shallow. Ventral trail short to moderate for genus. Ventral umbo weakly inflated, with obtuse beak. No ventral sulcus and dorsal fold. No dorsal spines. Description.—Medium to large size for genus; outline subquadrate to transversely subrectangular at posterior and elongate at anterior, widest at hinge line at maturity; profile fairly evenly curved, gently geniculate at anterior of ventral disk. Ears slightly extended, sturdy. No ventral sulcus and dorsal fold present. No subperipheral structure like rim or ridge in either valve. Interior of both valves not known. Ventral valve only moderately geniculate at maturity (see Fig. 3.15 for Holotype UKM-F575); umbo low and rounded; beak short, obtuse, not projecting beyond hinge line; trail longer than disk. Ornament with rugae and costae; rugae initially stronger than costae on disks, weakly undulated; costae shallow, coarseness varying from shell to shell, irregular in strength and width, occasionally bifurcating at anterior of disk but reuniting towards anterior of trail; trail costate; spines over whole valve, seemingly slightly slanted on disk more erect on trail, quincuncially and radially arranged, principally on crests of ribs; thick spines in row along border between ears and lateral slopes, thinner spines near hinge; spine bases weakly inflated, often disturbing rugae on disk (see Fig. 3.1, 3.5, 3.9, 3.14, 3.16, and 3.19 for arrangement of ventral spine bases). Dorsal valve gently curved in profile; disk slightly concave (see Fig. 3.10–3.12); trail short; similar ornament to ventral valve in reverse over whole valve; dimples scattered over disk, particularly strong dimples in row along lateral margins demarcating ears and disk, reflecting ventral spines; no spines present. Etymology.—After Thai, an abbreviation of Thailand. thaicus, -a, -um [adj.]. Types.—Holotype UKM-F575, ventral valve; three paratypes UKM-F572–574, two ventral valves and one dorsal external mould; Wordian age, Tak Fa Formation; type localities KW1 and KW4 of Khao Wong. Other material examined.—13 specimens from Khao Wong, three UKM-F576–578 (from KW1), eight UKM-F579–586 (KW4), and two UKM-F587, 588 (WTS2). Occurrence.—No other occurrence than Khao Wong. Discussion.—The new species from Thailand shows shallow but coarse costae over the trail, and thus is assignable to a species of Paraplicatifera Zhao and Tan. Its trail is relatively shorter than those of some other congeneric species. The type species Paraplicatifera regularis Zhao and Tan, 1984 (p. 26, pl. 1, figs. 15–17) from the upper Chihsia Formation of South China is also medium-sized (its holotype is 15 mm wide), and has a transverse shell with similar anterior ribbing to Paraplicatifera thaica n. sp. It, however, has a more extended ventral umbo and more regular rugae than the new species. Another Chihsian form, Paraplicatifera sangzhiensis Zhao and Tan, 1984 (p. 27, pl. 1, figs. 18–20), is a tiny (holotype 8 mm wide), subcylindrical form with strong and regular rugae, and can be readily distinguished from the new species. The Maokouan species, Paraplicatifera elongata (Huang, 1932, p. 23, pl. 1, fig. 14a–c) [originally in Productus (Marginifera)], from South China was once assigned to Neoplicatifera by some authors, but was relocated to Paraplicatifera by Zhao

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and Tan (1984). Its ventral valve is sulcate and strongly geniculate, differing from the new species. Two other Maokouan species, Neoplicatifera costata Ni in Yang et al., 1977 (p. 349, pl. 139) and Neoplicatifera lata Tan and Ding in Liu et al., 1982 (p. 183, pl. 131) from Hubei and Hunan, respectively, are assignable to Paraplicatifera, judging from their strong and coarse anterior costae and subcylindrical form. N. lata has similar macro-ornament to that of P. thaica, but has very thick dimples on the dorsal disk, and its ears seem shorter than those of the Thai species. Costispinifera costatiformis Ding in Zhang et al., 1983 (p. 278, pl. 95, figs. 4–6, 16) from the Maokouan of Northwest China unequivocally belongs to Paraplicatifera, and, so far as defined, it is the closest form to P. thaica. The Chinese species is a little smaller than the Thai species, and its rugae appear more regular and stronger. Two Himalayan shells described by Diener (1915, p. 82, pl. 9, figs. 1a–2c, not pl. 8, fig. 13) as Marginifera spinocostata Abich, 1878 and Marginifera cf. M. spinostata from the Zewan Formation (Changhsingian) of Kashmir possibly belong to Paraplicatifera, interpreting from their subcylindrical ventral shells with relatively coarse costae on trails. If so, they are the sole Late Permian representative of Paraplicatifera. The Kashmir forms have longer trails than P. thaica. Marginifera sintanensis Chao, 1927 (p. 156, pl. 16, figs. 13, 14) from the Maokouan of South China has been assigned to a species of Neoplicatifera by most authors (e.g., Jin and Hu, 1978; Liang, 1990, p. 181; Liu et al., 1982). Zhao and Tan (1984, p. 31), however, argued that the state of its anterior costae is intermediate between Neoplicatifera and Paraplicatifera, and included it in their latter genus. It is judged here that the costae of M. sintanensis are still fine enough to be in Neoplicatifera, so the species is excluded from Paraplicatifera. Superfamily LINOPRODUCTOIDEA Stehli, 1954 Family LINOPRODUCTIDAE Stehli, 1954 Subfamily STRIATIFERINAE Muir-Wood and Cooper, 1960 Discussion.—Jin and Hu (1978, p. 115) originally placed their subfamily Compressoproductinae under the Striatiferidae Muir-Wood and Cooper, 1960. Brunton et al. (2000, p. 546) reclassified the two subfamilies Striatiferinae and Compressoproductinae into the Monticuliferidae Muir-Wood and Cooper, 1960. However, the Monticuliferidae is now considered to be a rather specialized linoproductoid group (Sone and Leman, 2005, p. 601; Waterhouse, 2001, p. 28), and it should not include the above two subfamilies. The Striatiferinae is placed the family Linoproductidae. Furthermore, in opposition to the new Treatise of Brunton et al. (2000), a rather traditional classification of a familial-group, the Striatiferinae including Striatifera and Compressoproductus, is here favored after Cooper and Grant (1975), Jin and Hu (1978), and Waterhouse (2001). Genus COMPRESSOPRODUCTUS Sarytcheva in Sarytcheva et al., 1960 Type species.—Productus compressus Waagen, 1884; the Wargal Formation of the Salt Range, northern Pakistan (5Compressoproductus morahpressus Waterhouse and Piyasin, 1970 as a new replacement name). P. compressus Waagen is a common linoproductid name but is an invalid name, because it is a junior primary homonym of P. compressus Say in James, 1823. However, the valid substitute name C. morahpressus has been seldom used. Details of this nomenclatural circumstance were outlined by

FIGURE 4—Compressoproductus sp. from Locality KW4 of Khao Wong, the Tak Fa Formation, central Thailand. Ventral valve UKMF589; 1–3, ventral, lateral, and posterior views.

Sone (2006), who placed an ICZN Application to conserve the name P. compressus Waagen. A decision has recently been made by the International Commission on Zoological Nomenclature (2008) for not conserving P. compressus, and thus C. morahpressus stands as a valid name for this nominal species. Nevertheless, under Article 67.1.2 of the ICZN Code, the name of the type species remains unchanged even when it is a junior homonym; that is, P. compressus Waagen, albeit an invalid name, remains the type species name of Compressoproductus. COMPRESSOPRODUCTUS sp. Figure 4.1–4.3 Discussion.—A tiny, probably immature, form of Compressoproductus came from Locality KW4. It is a ventral valve UKM-F589, 11.5 cm long and 13.1 cm wide with a narrow, 4.1 cm-wide, hinge. This form is much smaller than the two Late Permian representatives of Compressoproductus from the Salt Range of northern Pakistan, namely, Compressoproductus morahpressus (nomen novum for Productus compressus Waagen) from the Wargal Formation and Compressoproductus mytiloides (Waagen, 1884) from the Chhidru Formation. Within the Indochina Terrane, two other linoproductoids are known in the correlative Middle Permian Sisophon Limestone of western Cambodia. First, Termier and Termier (1970) reported a small shell as Compressoproductus djulfensis (Stoyanow). Note that Productus djulfensis Stoyanow, 1910 (imprint 1909) from Armenia is now the type species of Sarytchevinella Waterhouse, 1983. The Cambodian form shows the regular rugose ornament, so it is more likely a species of Compressoproductus rather than Sarytchevinella. Termier and Termier (1970, fig. 10) illustrated it with body spines, which are not seen in the present Thai shell. Second, Mansuy (1914, p. 20, pl. 6, fig. 6) reported another linoproductoid as Productus? sp., and considered it related to C. morahpressus and C. mytiloides. It, however, has a broad venter with regular rugae, and is better assignable to another genus, Permundaria Nakamura, Kato, and Choi, 1970, rather than Compressoproductus.

SONE ET AL.—PRODUCTIDINE BRACHIOPODS FROM THAILAND PALEOBIOGEOGRAPHIC IMPLICATIONS

The three genera are common in Middle Permian strata. Haydenella and Paraplicatifera are known only from the warm-water Tethys Sea. Previously, a Middle Permian brachiopod fauna of the Saraburi Limestone was interpreted to be strongly similar to a South Chinese Cathaysian fauna (Yanagida, 1964; Yanagida and Nakornsri, 1999). There is no doubt that the present assemblage holds Cathaysian affinity in generic composition. Nevertheless, the two species Haydenella granti and Paraplicatifera thaica n. sp. are only endemic to the Middle Permian of the western Indochina Terrane, and specific linkage to a coeval Maokouan fauna of the South China Terrane cannot be made. Endemism of Middle Permian biota in the Saraburi Limestone is also apparent with an ostracod fauna (see Chitnarin et al., 2008). The present brachiopod assemblage may reveal part of an insufficiently understood endemic fauna, which flourished on the shallowwater margin of the western Indochina Terrane during Middle Permian time. ACKNOWLEDGMENTS

Fieldwork for this study was supported financially by I. Metcalfe (University of New England) from his ARC grant. Kitsana Malila (then of the Suranaree University of Technology, currently PTTEP) is thanked for his assistance in the fieldwork. S. Butts, Shen Shuzhong, and K. Dewing are much appreciated for carefully reviewing the manuscript. REFERENCES

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