NAT. HIST. BULL. SIAM Soc. 51 (1): 37 -6 7， 2003

GLIMPSES INTO FOSSIL ASSEMBLAGES OF THAILAND: CORAL PERSPECTIVES

Henri Fontaine 1， Sirot Salyapongstl ，and Varavudh Suteetho rn2

ABSTRACT

Foss i1 co ra1 s 釘 e widespread in 百 ai1 and. They belong to many geologic a1 interv a1 s，企 om Ordovician Ordovician to Quatemary. Th ey been have studied actively during the last 20 ye 紅 S. 恒leyare

better better knO Wll， even 白ough some 陀 search still needs to be carried out; for instance ，Triassic co ra1 s ，which oc 四 r in abundance at many loc a1 ities of 百凶land ，remain inadequately known. Carboniferous ，Permian and Triassic corals are widespread 泊Th ai1 and. De vonian and Jurassic

cor a1 s are in abundance in restricted areas. Or dovician and Silurian co ra1 s 紅 er 3I官 and remain poorly poorly knO Wll. Cretaceous corals are absen completely t. In τ'h ailand ，the study of cor a1 s is the study study of a long history. One c組 observe Ii ving cor a1 s as well as m 釦 y groups of fossil cor a1 s. Corals Corals provide important information on past environments and help to reconstruct pa1 eogωgraphy; 由ey a1 so provide information on the ages of the rocks in which 出eyare included. included. In this paper ， two De vonian 10c a1 ities 3I官 mentioned the for frrst time; 由ey were discovered discovered in January 2∞ヨ.

Key Key words: Tabulata ，Rugosa ，S 釘omatoporoids ，Scleractinia ，Ordovician ，S i1 urian ，De vonian ， Carboniferous ，Permian ，Tri 蹴 ic ，Jurassic ，Quatemary.

問 TRODUCTION

In In Th ailand ， corals are widespread both in the sea and on the land. In出 e sea 出ey 出丘 ve 泊 warm water and offer diverse and remarkable shapes. Th ey build up fasc 泊a白19 landscapes. landscapes. Coral reefs ， the “rainforests of the sea" for some au 血ors ， are well developed 泊官lai1 and. 百ley form very complex and very diverse ecosystems. 司自biological diversity includes includes commercially important species ， such as fishes ， providing liveliliood to many people 泊百ailand. On land ，corals are dis 凶buted all over 白e coun 町. Th ey are fossils included 泊 rωks 佃 d are commonly easy of access to everybody ，but 出.ey 釘 e not prominent and do not attract immediate attention. 百ley 訂 'e also associated with diverse other fossil species species of plants and animals. Limestone Limestone rich in corals ，especially Permian limestone widespread in Thai1 and ， has been used to produce wonder 白1 marble. Corals 紅 'e visible on m 佃 Y walls of Bangkok ， along along several s佐'eets. However ， because of the tropical climate ， the good polish of the marble marble slowly disappears and the details of the coral structure become oblitera 低d. Nowadays ， many people prefer granite which is more resi 包tant to weathering. In side houses and hotels ，

18 18 a1 1ee de la Chapelle ， 92140 Clanart ，France. E-mail: henri-fontain e@ wanadoo. 仕 2Geologic a1 Survey ， De partment of Min er a1 Reso 町 'ces ， Rana VI Road ，Bangkok 10400 ，百 ail 釦 d Received Received 11 March 2002; accepted 30 October 2002.

37 38 38 HENRI FONTAINE ，SIROT SALYAPONGSE AND VARAVUDH SUTEETHORN

m 訂 ble is well protected. Beautiful corals 釘 'e splendidly preserved and adom m 組 y kinds of of marble pieces (see Figs. lA-B). Before Before travelling about and visiting the cor a1 rock exposures ， one should know some characteristics characteristics of the corals. Th e prim 訂 y pu 叩ose of this paper is to provide general information information on the fossil corals of Th ailand.

ENVIRON 恥lliNT

Corals 訂 e exclusively marine ，sessile organisms which grow as solitary individuals or exhibit exhibit a colonial mode of growth. Most of the colonial corals are restricted to w 紅 m ， shallow shallow and clear sea water. They 訂 e sensitive to temperature ，depth ，s a1 inity ，nutrient supply supply and illumination. Locally ， they build up reefs.τ 'h ey are generally found at depths of of less th 佃 50m ，組d may be exposed to routine wave energy. Although well adapted to environments ， they can be severely damaged by cyclones. A small climatic ch 加 ge may result result in heavy losses. M 叩 y corals are characterized by narrow thermal tolerances ， and are not not cosmopolitan in dis 位ibution. Th ey have limited ability to acclimatize or adapt to the effects effects of climate change. Hence they 訂 ea growing concem of scientists studying present coral coral reefs ，afraid that coral adaptation cannot keep pace with increasing sea temperatures. Mortality Mortality of reefs has been suggested recently in some areas of the world because of bleaching bleaching outbreak ，possibly due to global warming (for instance ， see DENNIS ， 2002). Coral Coral bleaching occurs when coral expels from its body the symbiotic zooxanthellae so important important to its life ， probably because these algae b配 ome toxic to the cor a1 at higher temperatures. temperatures. Mass coral bleaching has been estimated to have destroyed about one-sixth of of the world's coral colonies during the last m 司or occu 町 ence in 1998; 18 percent of reefs were were damaged in Southeast Asia. Despite this severe event ， recovery is occuring. It is difficult difficult to predict what proportion of bleached coral will die. There is much debate over whether whether coral c叩 adapt to increasing temperature ， but so far ，there is little evidence of thermal thermal adaptation. Such adaptation may be too slow to be noticed by observers. In τ 'h ailand in in the Phuket 訂 'ea ， sea temperatures increased at a rate of more than 20 per 100 years between between 1981 and 1999 (BURKE ET AL. ， 2002). The corals of 出e past lived in similar conditions as today. Wh ere they are abundant and and highly diverse in rocks ，they 訂 e often associated with many other thermophilic benthic organisms. organisms. A w 紅 m climate during the Permian is evidenced in West Cambodia near the Th ai border by the accumulation of bauxite. Green a1 gae occur with the cor a1 s and indicate shallow shallow sea depth. Using geochemistry ，it is possible to get some information on the temperatures temperatures of the seas of the past and to confmn 白at corals are commonly good indicators of of warm climates (YASAMANOV ， 1980). Cor a1 s 紅 'e useful in deciphering paleoclimates because because of their environmental sensitivity; they can constrain paleogeographic reconstructions. reconstructions. For instance ，fossil corals of Peninsular Th ailand are different from those of of Central and East Thailand; they show a lower diversity ，especiallY during the Carboniferous Carboniferous and the Permian. These differences fit with the propos a1， by geologists (BUNOPAS ，1981) ，that Th ailand occupies and straddles two different continent a1 blocks Over the past 530 million years (Ma) ， the seas have invaded parts of Th ailand many times. times. Environments were not at all times and places favourable to cor a1 s. For 泊stance ， when sea water was loaded with sand in some areas ，cor a1 s were killed by the sand moving G Ll MPSES INTO FOSS IL ASSEMB LAGES OF TH A IL AND 39

Figur e 1. A ，B . Va se (1 2.5 C I1l hi gh) l1l acl e of Permi an li l1l es tone

co ntainin g a fa sc icll lat巴 co ra l (P se lld ollll oll gia) . On on巴sicl 巴， co rallir es are p巴rp endi clll ar 10

th e sllr face of th 巴 po t. On th e

oth 巴r sicl e，th 巴 co rallit es are

obli qll 巴 to all1l ost parall el to

thi ss urfa ce . Th 巴 S 巴pta are of

two ord ers，long ancl sh or t. Th 巴 co lu l1l 巴Il a is v isibl e in the

C 巴nt 巴r of th 巴 cora llit 巴

b

G Ll MPSES IN TO FOSS IL ASSE MBL AGES OF TH A IL AND 4 1

FigUl 巴s 3A-F. Tablll ata in th e fi elcl ancl in th in s巴cti ons. A: Kilm er ia probl emali ca Man sll y.V 巴ry pec lI li ar coral w ith an ope rclI llI lll of one 10 lhr ee convex plal eS (se 巴 Fo ntain e 巴1 al. 1994 ， p. 49-5 0). Sa lllpl 巴 T 3352 fro m B an Na C hal ianga long lh e road frolll Lalll Na rai 10 Ph elch ab lln. Thi s type 01' co ral

is rar e in Thailand. It ap par entl y occ llr s al lWO olh er loca lili es: n巴ar Wat Thalll W lI a Da eng (160 04'24"N ，1 0 I o26'00"E) ancl al Khao Thalll Yai betwee n LOlll sak ancl Na lll Nao. B: Beclcl ecl lilll es ton 巴 ofL a巴III Ph ap Pha in Sur at Thani Provinc e，ri ch in Sillopora ，a bran chin g Tab lllala

C: C: Pr olom icil e /i llia af f. 1I/ ll11i lO bul α la Yabe & I-Iayasak a; sa mpl e T519 fr olll Khao Thalll Na lll

Ban g，20 klll sO llth of Ph e lchablln. D: P ro lo /'ll ic il elillia simp lex Z h ao & C h巴n; sa mpl e T 3728

lll fro lll Khao C hak C han along th e r oa cl fro lll 1くhok Salllro ng to Takfa . E: P l'O lomicilel illi a CI 乱 .¥'s ilil eca

Z h ao & C hen; sa lllp le T253 1 fr olll Phll Ph a Khao 巴aS l of Ban Phia ，Lo ei Pro vin ce. F: Beclcl ecl lilll 巴slon 巴 atth 巴 fOOI 0 1' Phll Pha Wang along lh e road fro lll Wan g Sap hlln g to Uclo n T hani .T hi s lilll eS lone is ri ch in Pr olO llli cil e /i llia. 42 H ENR I F ON T A INE ，S IR OT SALYAPONGSE AND V ARAVUD H SUTEET H ORN

- h t c 、 b

e

Figure s 4A - E Tab ulat a. A ，Sinopo ra asialica . Tran sv 巴rse s巴ction of a co rallil 巴， sho w in g Ih e mi cro stru clure of th es keleto n. Micro stru ctul 巴 is imp ort3nt to diff erenl iate sO lll e co ral s. B ，C ，Khm er ia pr oiJI ema li ca

from Ban Na C hali ang. D ，C hael eles. Obliqu 巴 sec li on.Salllp le T 385 I frolll Middl 巴 Car bon fi erou s lilll estone in a poncl nea r Ban Na D lI an g. E ，H eli ol iles. Longitllclina l sec li o n. Salllp le T4315 frolll

th e Devonian lilll 巴ston 巴 of Phu Nalll Khoi in No nh eas l Thai lancl. G Ll MPSES lNTO FOSS IL ASSEMB LAGES OF THA IL AN D 43

kl k kl

c

Figlll 巴s 5A - G. Solit ary R lI gosa. Th es 巴 co rals we l巴 ab l巴 to li v巴 in div巴rs巴 envlronm 巴nl s.Th ey are know n at

man y loca liti es 0 1' Thailand .A ，L ophoph yllid iu l1I pendlll u l?1 G raba ll. Tran sve rs巴 sec tion show in g

lh e se pta a nd a so lid co lllm e ll a. Samp le T856 from Ban Phll PI lI along th 巴 road fro m Kan cha nabllri

to Th ong Pha Ph llm .A hori zo n of th e Midd le Permi an sec lion expos 巴d al B an Phll P llI is ralh el

ri ch in soli ta ry co rals. B ，Co ral know n bya s川 gl巴 sec tion，a tr an sve rse se ctio n. Samp le T6 292 from from Phah ong Cave ，a li m 巴stone hill 500 m fa r fr om lh e r oad from Lom s ak toC hum pha 巴. T he lim 巴ston 巴 is ri ch in crin oid s， b ut poor in co rals. C ，D ，Ca ll inoph yllll ll1 sp. Tr ansv 巴rse sec tlon s

ne ar th e bas巴 (4d ) and nea r th e top (4c) of lh 巴 co ral. M idd le Ca rb onif ero uss amp l巴 T6350 from H lI ai Na m S lI ai，n onh of Ban Na Duan g，L o巴iPro vin c巴. T hi s sa mpl 巴 is erod 巴d at it s p巴riph ery.

E，So lit ary R lI gosa co ll ec tedf ro日1 a wea th 巴red shale of Lo 巴iProv in ce. F，Ufi l1l ia Sp. Tran sve rs巴 sec ti on show in ga clear bil ateral sy mm 巴lr y:C = ca rdin al Se plll m ，k = CO llnl 巴rse plum ，1 = alal se pta ， kl = cO llnt erl aleral Se pla. M iddl 巴 Penni an sa mpl e T 2286 fro m K o To Nga in orth of Pak

M eng b巴ac h in P巴nin slll arT hai land. G ョ Paradllpl oph yllll l11 Sp.T ran sv 巴rse se cti on. Lowe r Penn ian sa mpl e T3835 from Ba n Na Din D am ar 巴a in Lo巴i Pro vin ce. 44 44 トIE N RI F ON T A INE ，S IR O T S A L 可A PONGSE A N D V A R AVU DH S UT E ET J-JO JW

巴

Figllr es 6A - E_ Fascic lI late R lI go sa_ A ，s ，Woa ge ll oph ylh 川 . T ransve r se sec li on (6a ) and longillldin al sec li on (6b). (6b). Sa l1l ple T5 354 fro l1l Ih e P巴rl1l ian lilll es lon 巴 of Khao Phai ，Saka 巴o Provi n ce. C ， Wao ge ll oph yllll l1l. Tr an sve rs巴 secl 旧 n. Salllp l巴 T5 35 I fro l1l th e P巴rlllian lim eston e of Khao Phai ， Saka 巴o Provinc 巴. D ，E ，C hihiaph ylll ll1l. Lon gitll di nal se cti on (6d ，s ampl e T80) and Iran sve rse sect ion (6 巴， salllp l巴 T59) . Sa mpl es from K ha o Khao belween Sa r ab llri and Lopbllri. G Ll MP SES IN TO FOSS IL ASSEM8 L AGES OF THA IL AN D 45

Figllr es 7 A - D. Ma ss i ve R lI gosa . A ，8 ，I pcip !J ylllllll. T ran sve r se (7a) ancl longilllclina l (7 b) sec tion s. Salllp le

T5675 fr olll Kh ao T halll Yai. C ，I pcip !J yll lllll. Tran sve r se sec tio n. Sa lllpl 巴 T 5672 f rolll Kha o

Thalll Thalll Ya i.D ，Para ip cip !J yllll l'//. Tran sve r se sec l旧 n. S 出 llp l巴 T90 3 I'rolll Khao Thalll S lI a in

P 巴nin slllar Thai la ncl 46 46 I-I EN RI FONTA INE ，S IR OT S AL YAPONGSE AND V ARAVU DH SUTEET I-I ORN

Fi gur es 8A - D. Ma ss ive R lI gosa .A ，Ip ciph y llum la ose ns e. Lon gitud in al sec tion. Sa mpl e T 1 39 from Khao Khi

Nok in Wichianburi ar ea of Cen lT al Th ail an d. s ，I pciph yllu l1/ la ose l/ se. Tran sve r se sec ti on Sampl 巴 T 3 133 from Kh ao Pho 巴m north of C hon Da 巴n. C ，0 ，Mullimurilllls sul eel hom i. Tran s- ver se sec tion s. Samp l es T370 1 C a nd T 3698 0 from Wat Th am S lI a Khu ha at Khao Wong C han

Da 巴ng ・ GLIMPSES INTO FOSS IL A SSEMBLAGES OF THAILAND 47

F igUl 巴s 9A - D . A ーC ，Kepingophylliade . T ran sver se se ction s. A ，Samp l巴 T3994 from Ban Na Duang area ，L o巴l

Provinc 巴 B ，Samp l巴 T4008 from th ee a stern fool of Phu Khao norlh we st of Ba n Na D uang. C ，

Sa mpl e T40LO fr om the ea sl巴rn foo l of Phu Khao nor thw es l of Ban Na Dua ng. 0 ，Multi l11 uri nus

T ran sve rs巴 sect ion. Samp le T4 84 from the north e rn end of Kh ao Som ph ot in L am Na rai area 48 HE NRl Fo 町 A 闘 E ，SIROT SAL Y APONGSE 刷 D V ARA VUDH Su 花町田制 with with the water.τ 'h ey generally do not occ 町泊sandstone. Muddy environments also were usually usually not favourable to corals ， with rare exceptions of some small s泊lple corals. In shale and mudstone ，corals 紅 e absent or rep 児 sented on1 y by scattered solitary corals ，in 血eir living living positions or 回 nsported before burial. In limestone deposited 泊 warm ，c1 ear and shallow shallow water ，corals are 泊 great abundance and high diversity ， and are predom 加佃tly large large compound corals. Th ey secreted skeletons of calcium carbonate ，reaching up to more th 釦 1m 泊 diameter. In some locations they have even bu i1 t up reefs. Wi 白血eir very small soft bodies or polyps ，commonly one to a few mm in diameter ， the the corals secrete skeletons of hig h1 y diverse shapes and sizes; 白ey have produced a very great great quantity of calcium carbonate on 白ee 紅白 since the Ordovician ，自 at is to say during the the last 500 My. All over the wo r1 d， they presen t1 y produce about one billion tons of calcium calcium carbonate per year. Limestone is widely dis 凶buted 泊白.e s回 .tigraphy of 官lailand ， especially 泊 the Ordovician ，Devonian ，Carboniferous ，Pen 凶an ，Triassic and Jurassic sequences. sequences.

CLASSIFICA TION AND BIOSτ 'RA τ1GRAPHY

百le corals belong to Phylum Cnidaria ， Class An thozoa. Our knowledge about them has has grown considerably during the last 50 ye 釘 s and the number of described families ， genera genera and species has grea t1 y increased. In In the field ，corals 街活 very d町icult to study (see photographs of Triassic and Jurassic corals corals in Figs. 2A-B). For precise microscopic observation ，corals must be cut into 白血 longitudinal longitudinal and transverse sections 0.03 mm or less thick (Figs. 3-9). Th en their characteristics characteristics appe 紅 cle 釘 ly if the coral is well preserved (see a complete solitary Rugosa ， about about 310 Ma old ， in Fig. 5E ， coral impossible to identify without thin sections). In In addition to giving information on p出 t environments ，many corals are useful for determining determining the ages of the rocks in which 白ey are imbedded. Some corals display a short stratigraphical stratigraphical range and hence can document the age of a rock. Corals can be used for intemational intemational conelations ，but 曲ey 紅 e limited to warm climate are 邸 covered by shallow and and clear sea water.

1. 1. Paleozoic Corals

Paleozoic Paleozoic corals consist of two c1 asses: Tabulata and Rugosa (or Te 回 .corallia). 百ley became widespread during 也e Or dovician Period and remained important fossils until the end of the Permian.

Tabulata: Tabulata: Th e hard p紅 ts of the Tabulata are slender polygonal or circular tubes (the corallites) corallites) with perforate (with mutal pores) or imperforate w a11 s， divided by horizontal diaphragms diaphragms (血 e tabulae). 百le tabulae 紅 e easy to see only in longitudinal sections; 白ey suggested 白e name of the class. Small radial elements may be present; 血ey 紅 'e commonly reduced reduced to short septa ，spines or squamulae ，visible 泊 transverse sections. Th e coral tubes may grow in approximately parallel position or 血ey may be p釘t1 y or entirely welded laterally laterally to their neighbours. Tabulata are divided into about 60 families and 330 genera. Th e stratigraphic dis 凶bution of these families and genera is variable ， as indicated by H i11 (1 981 ，table 3， p. 49ι505). GLIMPSES GLIMPSES INTO FOSSIL ASSEMBLAGES OF THAILAND 49

The Tabulata 訂 e characterized by their exclusively colonial mode of growth. Th ey are fasciculate fasciculate (shrubby ， with corallites not laterally contiguous) or massive (with contiguous corallites). corallites). Th ey are commonly a few cm to 50 cm in diameter. The mineral present in the skeletons skeletons of Tabulata is calcite. The microstructure of these skeletons is important for the determination determination of phylogenetic relationship between taxa ， but other features of secondary origin origin may cause confusion (OEKENTORP ，2001). In Th ailand ， the Tabulata are in abundance and and very diverse in Devonian rocks. In Loei Pr ovince ， massive corals 釘 'e common 組 d include include genera such as Favosites and Heliolites. Branching corals are also in abundance ， for for instance Thamnopora. Th e Tabulata also occur with moderate diversity in Carboniferous and and Permian sediments.

Rugosa: The hard p訂 ts of the Rugosa are much more complicated than the skeletons of Tabulata. Tabulata. In the skeleton of Rugosa are prominent radial plates called septa (Fig. 5A-D andF ベ}， 6A-D ，7A ーD ，8B-D ， and 9A-D). Th ey are of two orders (major and minor septa) septa) in many corals ， but in others there are three or ，more rarely ， four orders. 百ley commonly show a clear bilateral symmetry. Th e first septum appearing in the youngest p釘 t of the coral establishes the bilateral symmetry. At first ， the cardinal (C) and the counter counter (K) major septa appear in opposite position; then ， the two alar (L) septa appear ， followed followed by the two counter lateral (KL) septa (Fig. 5F). This plan of septal insertion is sporadically sporadically associated with a differentiation in size. For instance ， the cardinal sepωm is shorter shorter than the other m 勾or septa and the major septa of the cardinal quadrants may be thicker. thicker. In longitudinal sections ， small arched plates are visible at the periphery of the corallitβ(for corallitβ(for instance ， see fig. 8A); they are the dissepiments. Tabulae occupy the rest of the the corallite cavity. In the axial part of the corallite ，a columella is present in some genera. The Rugosa consist of about 80 families and 700 genera. The stratigraphic distribution of these these families and genera has been indicated by Hill (1981:7ι94 ，table 1). Th e skeletons of fossil Rugosa 紅 e of calcite ， but was it the primary mineral in the living living corals? Opinions about 出is differ (OEKENTORP ， 2001). Th e Rugosa may consist of a single corallite of conical to cylindrical shape (a trochoid shape shape is shown in Fig. 5E). Some were deprived of dissepiments and simple ，and were able to to live in diverse environments. Th ose with dissepiments were more complex ， and apparently needed needed slightly better environments. Many other Rugosa are compound.τnese indicate very very favourable environments. Some types 釘 e fasciculate with ramose corallites or massive with with contiguous corallites. Studies Studies of stratigraphy reveal that several crises occurred in the history of the Rugosa. For For instance ， seven different crises occurred during the upper part of the Paleozoic ， at the end end of the Lochkovian 400 Ma (million years ago) ， with a strong decline of the Silurian taxa; taxa; at 白e Frasnian-Famennian boundary (365 Ma ，Kellwasser event with the disappearance of of the massive Devonian corals); at the Serpukhovian-Bashkirian boundary (350 Ma ， minor minor extinction with renewal of the coral faunas); at the Moscovian-Kasimovian boundary (305 (305 Ma ，decrease of the coral community); at the end of the Lower Permian (270 Ma ， for instance instance disappearance of 出 e Kepingophyllidae); at the end of the Middle Permian (250 Ma ，disappearance of all the massive Rugosa); and at the end of the Permian (245 Ma ， complete complete extinction of the Rugosa). Th e Heterocorallia differ from the common Rugosa and form their own small order. Th ey 訂 e good markers of the age of the rocks because they appe 訂 ed during the Devonian 50 HENRI Fo 肝 AINE ，SIROT SALY APONGSE AND V ARA VUDH SUTEETHORN

and and disappeared during the Carboniferous. Th ey 釘 e 合agile ，elongate corals consisting of a single corallite of small diameter. Th ey 釘 'e easy to recognize. In Th ailand ， they are abundant abundant in some Lower Carboniferous limestones; two genera (Hexaphyllia and Heterophyllia) Heterophyllia) have been found. In China during the last 20 ye 訂 s ，diverse Heterocorallia have have been described with the creation of new families ， genera and species. 百le validity of many of these taxa remains questionable ， as comparable variability among Hexaphyllia specimens specimens from England has been considered to represent only intraspecific variation. Stromatoporoids: Stromatoporoids: A group of fossils related to Porifera (sponges) must be mentioned because because they are associated with the corals in m 組 y places and sometimes played a great role role in the building of reefs. Th e stromatoporoids originated at the beginning of Middle

Ordovician Ordovician and were widely dis 汀ibuted until the end of the Devonian.τbey 血en disappe 紅 'ed ， and and reappe 訂 ed from Middle Jurassic to Cretaceous. Th ey are very important in 百lailand ， especially especially because they are major frame builders of the Devonian reefs found in Loei Province. Province.

2. 2. Post-Paleozoic Corals or Scleractinia

百le mass extinction that ended the Permian Period saw the end of the Rugosa ，trilobites ， and and fusulinaceans. Other organisms declined markedly. Corals are almost absent from the Lower Triassic. A large break seems to sep 訂 ate the Paleozoic corals 合om the post-Paleozoic corals corals or Scleractinia. At the beginning of Middle Triassic (An isian) ，Scleractinia appeared and and started to show a great variety of forms belonging to solitary ，fasciculate ， or massive corals. corals. At the end of Mi ddle Triassic (Ladinian) ， they started to build reefs. 百le early phylogeny phylogeny of the Scleractinia is poorly known because of poor fossil records during the Early Early Triassic. EZAKI (2000) ，after describing Permian corals with a putative 訂 agonitic skeleton skeleton in Tunisia in 1997 ，discovered solitary “scleractiniamorphs" in the Middle Permi 叩 of of China (Yabeina zone). Th e mode of septal insertion of these corals was impossible to study ， but six orders of septa are distinctly presen t.“ Septa 訂 'e seem 加gly arranged radially in in circular corallites whereas those in elongate and elliptical corallites may show a bilateral symmetry" symmetry" (EZAKI ， 2000). Ezaki suggested that an initial radiation of Scleractinia-like corals corals occurred during the Permian. 百lis radiation might have been terminated by the mass extinction extinction of the end of 由e Permian ， or it might have survived in a few refuges with relatively relatively good environments. Scleractinia Scleractinia are an extant class known from the Triassic to 出e presen t. τbey are the most important constituents of modem coral reefs. Th ey f: 訂 su 中ass the older corals in diversity ， and comprise a great number of orders ，families ，genera 佃 d species. Th ey are characterized characterized by prominent cyclic insertion of the septa. The first-formed septa 訂 e6 in number. number. A bilateral skeletal s汀ucture is very rare and underlines a phylogenetic relationship between between the Sceractinia and the Rugosa. Th e skeleton is of aragonite. Thr ough diagenesis ， 紅 agonite is easily replaced by co 釘 se calcite and the of microstructure the coral is lost forever. forever. Scleractinia 紅 'e solitary or compound corals (fasciculate and massive). In In Th ailand ，Scleractinia date to different periods from Triassic to the present; they are observable observable in sea and on land. GLIMPSES GLIMPSES INTO FOSSIL ASSEMBLAGES OF THAILAND 51

CORALS OF THAILAND

In In 1920 ，a Carboniferous Tabulata was found at Kh uan Din So in the Phatthalung 訂 'ea of Peninsular of Peninsular Th ailand (REED ， 1920). In 1944 ，a Permian Rugosa species was reported in Northwest Northwest Th ailand (MINATO ， 1944). Later on ， other species were discovered and their number reached 25 in 1969 (PITAKPAIVAN ET AL. ， 1969); they were assigned mainly to Permian ，subordinately to Triassic (5 species) and to Carboniferous (3 species). Since 伽 n， diverse diverse fossil corals have been noticed at m 佃 y localities of Th ailand; they 釘 'e locally in great great abundance and indicate favourable environments in the pas t. Th ey belong to m 組 y time time intervals ，合 om Ordovician to Quatemary. From Ordovician to Jurassic ，more th 佃 250 species species have been mentioned in 官lailand ， belonging to about 150 genera. Additional studies studies should multiply the numbers of genera and species. Even if it were possible to find corals corals in the Cambrian (54 0- 500 Ma) ， the environment was not favourable to corals in 出e Cambrian areas of Th ailand. 百le fossil coral faunas of 百lailand are largely not endemic ， but exhibit similarities with with other faunas of the world. Detailed studies show that ， even within 百lailand ，endemicity and and the predominance of some taxa may v釘 y geographically and according to the period ， under under global and regional controls. Corals Corals are widespread in 官lailand ， but their importance in the sediment varies according to to the environments in which they lived. 官ley predominated in local 釘 'eas and bound together together with other organisms (stromatoporoids in the Devonian in 出e Loei 紅 'ea) ，出 ey built built up rigid frameworks having wave resistance ， forming reefs. In other places ，出 ey appe 釘 only 泊 relatively 出in horizons ， for instance the Carboniferous Lublinophyllum bed (1 m 出ick) of Ban Na Charoen 泊 Northeast 百lailand ， or in some Permian limestone beds north north of Saraburi along the road to Lam Narai ， where they form biostromes. In other areas 白ey 釘 e more scattered ，especially in 白e Permian limestone of Peninsular 百lailand where 出ey con 佐ibute a small fraction of the sedimen t. At some localities ，出 ey show evidence of being being transported by cu 町 ents from their original place of attachment.

Ordovician Ordovician (50 仏-4 35 Ma)

百le oldest coral-bearing strata in Th ailand 釘 'e Ordovician in age ， but the corals are very very rar ・e and remain poorly known. HAHN & SIEBENHUNER (1982: 38) mentioned Middle- to-Late to-Late Ordovician solitary Rugosa at a locality of Northwest τoailand ， west of Chiang Mai and northeast of Ban Mae La No i. Th ese corals were considered belonging to “Me 凶 ophyllidae? or Streptelasmatinae 泊det". 百出 identification did not indicate a precise Paleozoic Paleozoic age ， but the corals were associated with trilobites (Trinucleina) ， bivalves and gastropods. gastropods.

Silurian Silurian (435 -4 10 Ma)

Silurian Silurian corals also remain poorly known and appear to be uncommon in Th ailand. 官lis paucity is p紅 tly explained by poor environments. A few Tabulata (Favosites ，Heliolites) 釦 da Rugosa (R hizophyllum) found in 出e Nam Som area of Northeast Th ailand have been assigned assigned a Silurian age (SAKAGAMI & NAKORNSRI ， 1987). Rhizophyllum ，commonly considered considered to be a typical Silurian coral ，actually r佃 ges from Early Silurian to the end of Early Early Devonian. 52 HEN 悶 FONTA 酎 E， SIROT SALYAPONGSE AND VARAVUDH Su 花町HO 剛

Devonian (410 ー， 360 Ma)

Devonian Devonian rocks are widely exposed in Northeast and West Th ailand ， but with different lithologies. lithologies. 1. 1. Northeast Thailand Devonian sediments are known only in Loei Pr ovince and Nong Bua Lamphu Province. τ'h ey locally display thick limestone bodies. 官 ley are very rich in diverse fossils and corals are are in abundance at many localities (FONTA 町 E ET AL. ， 1981 AND 1990; FONTAINE & TANTIWAN 汀， 1987; FONTAINE ， 1990; FONTAINE & SUTEETHO 悶， 2000) ，which may be divided 加to 由民e areas ， as follows.

Area 1a: Th e area extending in a N-S direction around longitude 101 Q 55'E from the

Mekong River in the vicinity of Pak Chom to the road from Wang Saphung to Udon 百1佃 i exhibits exhibits many Devonian limestone outcrops and appe 釘 s to be the richest in corals. From north north to south ， the main localities are: ne 釘 Ban Kh ok Phai (Pha Tok and Phu Kh ong) ， in Phu Laem area (Phu Laem Huai Na Phung ，Phu Yot Huai Th ian ，at 如 d ne 釘 Phu Laem) ， in in Ban Chiang Klom 釘 ea (Phu Yai east of B 叩 Chi 佃 g Klom ，Phu Wang Pha ，Wat Tham Saeng 百四n Phromawat at Phu Foi Lom) ， east of Ban Chom Noi (6 localities from Ban

Huai Bo Sun to the north to Khao Sam Y ot to the south; at 佃 old quarη ，a good section ， 83 83 m thick ， has been observed) ， and south of B 叩 Chom Noi (Phu Thanon and Huai Nam Suai). Suai). Corals 訂 'e in great abundance and high diversity. Tabulata 釘 'e common and consist of: of: Chaetetidae: Chaetetes yunnanensis ，Chaetet 伊ora; Favositidae: Squameopora ， Emmonsia ， Squameofavosites; Pachyporidae: Cladopora ，Gracilopora ，Striatopora ， Thamnopora nicholsoni ， T. po か'gonalis ， T. sp ， T. tumefacta?; Al veolitidae: Alveolites cf. admirabilis ， A. smithi ， A. sp. ， A. cf. taenioformis ，Crassialveolites crassiformis ， C. crassus ， C. C. sp. ， Caliapora battersbyi ，Scoliopora; Coenitidae: Coenites ，Planocoenites; Heliolitidae: Helio /i tes porosus ，H. sp.; Syringoporidae: Syringopora. Rugosa 釘 'e scattered and include: Cystiphyllidae: Cystiphyllidae: Cystiphyllum vesiculosum; Stauriidae: Dendrostella; Ptenophyllidae: Dohmophyllum involutum and other taxa; Endophyllidae: Endophyllum abditum ， Sinospongophyllum tabulatum ， S. sp.; Phillipsastraeidae: Scruttonia ，Pterorrhiza ， Thamnophyllum stachei ， T. sp.; Stringophyllidae: Solipetra. 官 le Tabulata 佃 d Rugosa are associated associated with stromatoporoids: Actinostroma devonense ， A. sp. ，Amphipora angus ω， A. insignis ， A. pe ヴorata ， A. ramosa ， A. rudis ， A. sp. ，Clathrodic ηon ，Hermatostroma beuthii ， H. H. pustulosum ，Stachyodes ，Stromatopora ，Trupetostroma. All the fossils show affinities with with the Devonian faunas of Viet Nam and South China; 出ey focus mainly on a Givetian- Lower Frasnian age. Th e corals build reefs in association with stromatoporoids. Devonian reef reef building peaked during the Givetian in many parts of the world ，including Th ailand. Th is Devonian area of Th ailand extends to Ban Vang ，a Laotian village on the northem bank of Mekong River. Near this village ，a limestone hill contains Tabulata (Favosites のriacus ，Thamnopora cf. polygonalis ，Syringopora schulzei Heliolites porosus) 佃 dRugosa (Xystriphyllum (Xystriphyllum laosense ，Breviphrentis yunnanense ， Microplasma ronensis). Th ese fossils have have been described by FONT A 刑 E (1 961). Fossils other 出an corals and s甘omatoporoids have have rarely been studied. At Phu Koi (CHAIRANGSEE ET AL. ，1990) ，a smalllimestone hill northwest northwest of Ban Chiang Klom ， conodonts have been isolated and belong to the genera

Belodella Belodella and Neoprionodus; they 紅 'e associated with tentaculitids (Sη liolina). Cherts ne 釘 GLIMPSES GLIMPSES INTO FOSS 江 ASSEMBLAGES OF THAILAND 53 and south of Pak Chom contain radiolarians indicating a Frasnian to Toumaisian age (SASHIDA ET AL. ， 1998). Sandstone near Ban Pak Niem has yielded a palynological material indicating indicating an Uppermost Devo 凶an to Lower Carboniferous age (FONTAINE ET AL. ， 1981).

Area lb: Devonian rocks are exposed in a second 紅 'ea extending in a N-S direction around longitude longitude 102 0 10 ・E from the Mekong River (ne 紅 Ban Muang: 18 0 10'14"N ，102 0 09'30"E) to to Ban Chok Chai (1 70 35'22"N ，102 0 l1 '27"E) southwest of Nam Som. Li mestone is less prominent; prominent; shale ， marl and sandstone are more widespread.τbese sedimentary rocks 紅 e older older than the limestone of the first area; they belong to Lower Devonian ，maybe down to to Upper S i1 urian. At Ban Muang and Ban Noi ，Tabulata 訂 e 泊 abundance. Th ey include avosites F avosites goldfi 凶 si 佃 d Heliolites sp. At B 佃 Muang ，they 釘 'e associated with Lancicula ， an an alga common in Lower Devonian. Near Ban Na Ngiu ， a village S-SW of Ban Muang ， a limestone has yielded a fragment of Favosites associated with Renalcis ，an algo-microbial S釘uc 旬re. Further south at Ban Na Kh ae (1 7054'25"N ，102 0 04'58"E) ， Tabulata (Favosites cf. cf. sηriacus ， F. cf. ottiliae) and solitary Rugosa are associated with brachiopods (Spiriferacea ando 出ers) 凶釘gillaceous limestone and shale. At B 佃 Yuak (1 70 40'48"N ，I02 0 09'58"E) ， shale shale with black limestone lenses contains Tabulata (Emmonsia in abundance ， Squameofavosites) ，solitary Rugosa (Cystiphylloides fongi) ，brachiopods (A t. η 'pa and others). At the bottom of a pond near Ban Chok Chai ，a limestone is rich in foss i1 s: Renalcis ， Tabulata Tabulata (Favosites nitidus or alpina? ，Squame o. 向vosites ，Heliolites) ，solitary Rugosa ， 柑 omatoporoids. At the wat of Ban Chok Chai ， limestone is again rich in corals. Two new discoveries of January 2002 have extended this Devonian area towards the south. south. Phu Pha Duang (1 7~6'54"N ， 102 0 16'36"E) ， south of Ban Chok Chai 佃 d1 km north of of Ban Wichit Pat 血ana ， is a limestone hill with a1 紅 ge cave. The limestone is gray and thickly thickly bedded. It is partly recrystallized and foss i1 s appear to be uncommon. However ， some scattered foss i1 s have been noticed at 白e westem side of the hill; they consist of Tabulata ，solitary Rugosa ，組 dS 甘omatoporoidea. Th ey have not been studied ye t. An other limestone limestone hill (1 7~6'45"N ， 102 0 16'49"E) ， about 800 m east of Phu Pha Duang ，exhibits sim i1 ar foss i1 s.

Area lc: South of the road from Wang Saphung to Udon 官lani ，Devonian limestone is exposed exposed in an area elongated in a NW-SE direction (FONTAINE & Su 百 E四 ORN ，2000). From nor 白west to southeast ，it bu i1 ds up a few hills: Phu Lao (1 70 15'24"N ，102 0 03'20"E) ， Kh ao Pha Cham Nam ，Phu Khr ao and Phu Kh ao (1 70 13'50"N ，102 0 03'55"E). It crops out at at the bottom of a pond near Wat Pakittisophanaram (1 70 14'48"N ，102 0 02'40"E). Foss i1 s consist consist of Tabulata (Heliolites ，Alveolites ，Thamnopora and other genera) ，Rugosa and Stromatoporoidea. Stromatoporoidea.

2. 2. West Thailand

百us area includes the Peninsula ，and 紅 eas from Kanchanaburi to Mae Hong Son and Fang. Fang. Devonian seas were widespread ， but not favourable to corals. Environments were muddy and sandy ， with rare deposition of limestone containing conodonts 阻 d tentaculites; corals corals have not been found so 伽 (JAEGER ET AL.， 1969; HAHN & SIEBENHUNER ， 1982; LONG & INGAVAT ， 1989; WONGWANICH ET AL. ， 1990). 54 HENRI FONTAINE ，SIROT SAL Y APONGSE AND V ARA VUDH SUTEETHORN

Carboniferous Carboniferous (360 ー295 Ma)

Th ailand was mostly covered by seas during the whole Carboniferous ， with local and temporary temporary emergences. For instance ，am 釘 ine regression occurred in Northeast Th ailand at at the end of the Lower Carboniferous; it allowed the deposition of coal (組曲 racite) 泊 Ban

Na Duang 紅白 and gypsum in Loei-Wang Saphung area. A shale exposed west of Ban Loeng ，38 km northeast of Loei as 出e crow flies ， is rich in continental plant imprints. Th ese fossils have not been studied yet. 百leyapp 訂 ently belong to the Carboniferous (new data). data). After 血is regression ，出 e sea came back and corals reappe 釘 ed. Th e corals of Thailand belong to several 紅 'eas and to different time intervals.τ 'h ey 紅 e most prolific in central and northeastem Th ailand.τ 'h ey are tl1 e richest coral faunas so far known in tl1 e Carboniferous of Southeast Asia. 官 ley were particularly abundant during tl1 e younger younger part of tl1 e Lower (Visean Carboniferous to Early Serpukhovian). After decimation at at the end of the Lower Carboniferous ，new coral faunas replaced the old ones during Middle and Upper Carboniferous; 白ey were less widespread. In In Peninsular Thailand ， environments were not favourable to corals during the Carboniferous; Carboniferous; only tiny Rugosa wi tl1 0ut dissepiments are expected there. A Tabulata (Cladochonus (Cladochonus cf. michelini EDWARDS & HAIME) has been reported in Phatthalung area (REED ， 1920). The Carboniferous is divided into two p訂 ts as follows.

1. 1. Lower Carbon 俳rous

Corals 訂 e abundant at several Lower Carboniferous localities of East ，Central ，Northeast and Northwest Th ailand.

Area la: In eastem Th ailand ，Lower Carboniferous limestone is known in two areas: no 託h of of Klaeng ，and in tl1 e Kabinburi area. North of Klaeng or east of Si Racha ，a diverse assemblage assemblage of corals belonging to the end of Lower Carboniferous has been found re 四 ntly east east of Khao Yai Mo Noi (13 022'30"N ，10I 041'15"E; FONTAI 問&SALYAPONGSE ， 1997). It It consists of a few tubular Tabulata ， large solitary Rugosa (apparently Kueichouphyllum) ， common fasciculate Rugosa (Siphonodendron cf. irregulare 釦 d Solenodendron cf.furcatum) and ， in abundance ，Heterocorallia (Hexaphyllia). Th e corals are associated wi tl1 algae and diverse diverse faunas. Near Kabinburi ， only a solitary Rugosa has been collected 合om a limestone qu 釘 ry; it has been described as belonging to Koninckophyllum (SUGIYAMA & To 町 YAMA ， 198 1). Area lb: In Central Th ailand between Pichit and Phetchabun ，severallimestone exposures containing containing Lower Carboniferous corals are distributed from Noen Maprang to Chon Daen over over a distance of more than 50 km (FONT A町 E ET AL. ， 1991). At Khao Hin Pu n in Chon Daen area (1 60 07' 30"N ，100 0 42'40''E) ，limestone is bedded and contains scattered Tabulata (Syringopora) (Syringopora) and solitary Rugosa (Koninckophyllum) ， associated with foraminifera ， brachiopods brachiopods and crinoids. It belongs to Lower-Middle Visean and is a little older than the limestone limestone of many other localities. At Khao Pha Th ap Pon south of Noen Maprang (1 6~9'59"N ， 100 0 40'20''E) ，a section of limestone overlying shale and sandstone is easy to observe. observe. Th e corals 訂 e well preserved and belong to Tabulata (Syringopora ，Chaetetidae) ， diverse diverse solitary Rugosa (Cyathaxoniidae: Cyathaxonia; Laccophyllidae: Syringaxon ， GLIMPSES GLIMPSES INl ひ FOSSIL ASSE 勘也 LAGES OF THAILAND 55

Amplexocarinia; 加 tiphyllidae: Rotiphyllum; Hapsiphyllidae: Hapsiphyllum ，Za phrentites ， Allotropiophyllum; Allotropiophyllum; Cyathopsidae: Caninia ， Melanophyllum; Aulophyllidae: Arachnolasma cylindricum ， A. equiseptatum ， A. sp. ， Kueichouphyllum sinense) ，fasciculate Rugosa thostrotionidae: (Li thostrotionidae: S伊honodendron irregulare ， S. chonglomense ， S. cf. petalaxoidea ， Solenodendron Solenodendron furcatum; Axophyllidae: Lonsdaleia congjiangensis ， L. cf. duplicata) ， massive massive Rugosa (Li thostrotionidae: Lithostrotion decipiens; Axophyllidae: Actinocyathus floriformis ， A. sp.) and Heterocorallia (Hexaphyllia). 百 are ley associated with algae (common Koninckopora) ，foraminifera ， bryozoans and diverse brachiopods. They show more simi 凶 ities with 由e Chinese and European forms than with the Aus 佐alian counte 中町ts; for 泊stance ，Lithostrotion is not known with certa 泊ty in Aus 甘alia (Webb 1994). 百le fossils of of Kh ao Pha Th ap Pon 釘 'e also present at other localities of Noen Maprang 倒閣， locally with with other corals: Kizilia ，Siphonodendronjunceum ，S. pauciradiale ，Ax ophyllum Vaughani and Dorlodotia cf. mengi.

Area lc: In Northe 錨 t 官lailand in the Lo ei region ， marine sediments belong 泊g to Lo wer Carboniferous 釘 e exposed in 伽 'ee areas: in Pak Chom 訂 'ea at Ban Sa Ngao 佃 d Ban Pak Niem ， at Pha Chom Nang along the road from Chiang Kh an to Pak Chom ，and 加 BanNa Kl ang 釘 ea at a few localities. Near B 佃 Sa Ngao (1 80 04'N ，10I 0 57' 30"E) ， limestone is exposed exposed along Huai Sa Ngao and extends to 白e west at ground surface south of Ban Pak Niem (FONTA 凶 E EI AL. ， 1982). It contains solit 町 Rugosa (Kueichouphyllum sinense ， Arachnolasma equiseptatum ，Yuanophyllum) ，Rugosa (Solenodendron furcatum ， Siphonodendron Siphonodendron petalaxoidea ，Hexaphyllia and Heterophyllia). Th ese corals are associated with with calcispheres and foraminifera. 百le Ban Sa Nga o- Ban Pak Niem limestone is well dated dated and belongs to the Upper Visean. At Pha Chom Nang ， limestone contains scarce solitary solitary corals (Gangamophyllum) associated with a poor assemblage of foraminifera belonging belonging to a Visean age. Near Ban Na Klang ， 15 localities have yielded Lower Carboniferous Carboniferous corals (FONT A町 E & Su 四 ETHO 剛， 2000). Ne 紅 Phu Pha Wiang no 巾 of Ban Na Klang (1 70 25'05"N ，102 0 05'00"E) ， solitary Rugosa (Arachnolasma and Kueichouphyllum) Kueichouphyllum) have been found and indicate a Late Visean age. At Ban Non Th awon east east of Ban Na Kl ang ，corals 紅 e more diverse and consist of Tabulata (Syringopora) and Rugosa (Lithostrotion ，Solenodendron ，Hexaphyll ω ，Heterophyllia). At Kha o Pha Se 組 6 km south of Ban Na Klang ，corals 釘 'e few; they belong to Tabulata (Chaetetipora) and Rugosa (Ro ゆhyllum ， Lonsdaleia cf. congjiangensis). Other localities of Ban Na Kl ang 釘 ea contain rare corals.

Area ld: In Northwest Th ailand ，Carboniferous limestones 紅 'e exposed at some localities between between Phrao (Chiang Dao area) 組 d Mae Hong Son. Corals have been found 泊 Visean limestones limestones exposed between Ban Mae Lana and 由e Burmese border (CARIDRO 汀 EIAL. ， 1987; 1987; FONTAINE ET AL. ，1993). 百ley consist of Clisiophyllum near the Burmese border (1 90 41'N ，98 0 13'40"E) ， of Arachnolasma and Hexaphyllia associated with Koninckopora (an (an alga) northeast of Ban Mae Lana (1 90 35 'N， 98 0 13 ・35"E). Th ese localities have been assigned assigned to 佃 Upper Visean -S e叩ukhovian age.

2. 2. Middle-Upper Carbon 俳rous Corals Corals have been found in Middle-Upper Carboniferous sediments in Central ，Northeast 佃 d Northwest Th ailand. 56 H 四 RIFoNTA 別 E， SIROT SALYAPONGSE AND VARAVUDH Su 百 ETHO 附

Area 2a: In Central Thailand between Lam Narai and 官 1ep Sa 出it ，Middle-Upper Carboniferous Carboniferous corals have b田 n found 泊 a small area ne 紅 Ban Bo Nam (1 5017' 40"N ， 101 017 ・15"E; FONTAINE E1' AL. ，1999). 百1ey 紅 'e few ，scattered 泊 the rock. Th ey consist of Tabulata Tabulata (Multithecopora) ，solitary (Caninia ，Bothrophyllum ，Amygdalophylloides) ， fasciculate fasciculate (not identified) and massive (Petalaxis siamensis) Rugosa. 百 are 1ey Moscovian- K 踊加ovi 叩加age. At Wat Kh ao Takaeng south ofBan Bo Nam (1 50Q4 '20"N ，10I O I9'40"E; FONT AlNEE1' AL. ，1996) ，a Gshelian limestone has yielded a solitary Rugosa (Sestrophyllum).

Area 2b: In no 巾 e槌 t官1ailand ，Mi ddl e- Upper Carboniferous corals occ 町 atm 佃 y localities belonging belonging to five 釘 eas.

2bl: 2bl: An important area is elongated 泊 aN ー S direction along longi 制 de 101 0 45'E 企om 出e Mekhong Ri ver to Kh ao 百1am Nam Maholan south ofWang Saphung. 官 1e localities w i1l be mentioned from north to south. East-southeast of Phu Pha Maen (1 7054'06"N ， 101 048'12"E) ，a limestone contains a few solitary Rugosa (Arachnolasma) 佃 da few foraminifera foraminifera in c1 uding Eblanaia. An Upper Toumaisian-Lower Visean age has been suggested suggested because of 出.e occurrence of Eblanaia. During a recent fieldwork (J anu 紅 Y 2002) ，a few solitary Rugosa have been found at the top of Phu Pha Maen (1 7054'58"N ， 101 045'49"E) ，solitary and fasciculate Rugosa at other localities to the south (1 7049'56"N ， 10 10 104 7' 48"E; 17 049'03"N ，101 048'36"E; 17 048'56"N ，101 048'43"E); these fossils have not been studied yet. Tabulata (Multithecopora ， Chaetetes) and fasciculate corals (Para かtvophyllum? ， Opiphyllum cf. intermedium) occ 町 in Upper Bashkirian limestone exposed exposed in a small valley (Huai N 倒的 3 km east of Ban Tat So. At km 13 of the road 針。 m Ban 官 1at to Pak Ch om ， Tabulata (Chaetetes nagaiwaensis ，Donetzites ，Multithecopora cf. choiana) ，solitary Rugosa (Sestrophyllum) and massive Rugosa (Petalaxis kita 初 miensis) have have been discovered in another Upper Bashkirian limestone. Near Nam Th ao reservoir (1 7035'N ，10I 052'30"E; FONTA 町 E E1' AL. ，1994) ， Moscovian limestone contains Tabulata (Chaetetes (Chaetetes nagaiwaensis ，Multithecopora) ，solit 釘 y Rugosa (Lophophyllidium ， Caninia l伊oensis) and massive Rugosa (Petalaxis siamensis). At Pha 百1a ，corals are represented by common Chaetetes and Kionophy l/ um in a Bashkirian limestone. At Phu Bo Bit near and and east of Loei (1 7029'00"N ，I0104 7' 00"E) ， Multithecopora and Koninckocarinia have been been mentioned also in a Bashkirian 1加 .estone. At Ban Pha Noi ，a section is visible along Huai Nam Puan and ranges from Upper Moscovian to Permian. A few Tabulata (M ultithecopora) 釦 d solitary corals (Caninia lipoensis ，Amygdalophylloides ， Rotiphyllum and and others) have been collected 合om this section. East of Wang Saphung along the road to to Udon 官1ani ，a few Tabulata (Multithecopora) 佃 d solitary corals (Caninia cf. tien i) have have been noticed 泊 Moscovian beds. In a probably Upper Moscovian limestone at 血e northem side of Phu Ki Kai ， Tabulata (Multithecopora) ，solitary (Lophophyllidium ， Amygdalophylloides ，Sestrophyllum) and fasciculate (Lublinophyllum?) Rugosa have been found found (FONTA 町 EET AL. ，1991). West of Kh ao 百四n Nam Maholan at the wat ，rare solit 釘 y Rugosa occur 泊 a Gshelian limestone. South of Kh ao 百四n Nam Maholan between Ban Pha Kh ao and B 佃 Somsak Phatthana (1 7001'57"N ，I01 058'45"E; FONTA 町 E E1' AL. ， 1995) ， shale and limestone beds exposed at the bottom of a pond contain solitary (Pseudotimania ， Caninia cf. tieni ，Caninophy l/ um indosinense ， C. som ωiense) 如 d fasciculate (Lublinophyllum) (Lublinophyllum) Rugosa; 白ey belong to Upper Moscovian. 2b2: 2b2: In Ban Huai Som Tai area east of Phu Kr adung ，Mi ddl e- Upper Carboniferous sediments GL 品1PSES INTO FOSSIL ASSEMBLAGES OF THAILAND 57

occur occur widely ，but 紅 e not easy to observe because they 釘 e commonly covered by alluvium (FONTAINE ET AL. ， 1995). At the bottom of a pond 1 km east of Ban Huai Som Tai (1 60 53'00"N ，10F59'55"E) ，fossils are in abundance in interbedded shale and limestone; they they belong to foraminifera ，corals ，bryozoans ，brachiopods ，echinids (spines) and crinoids. Corals Corals are diverse and belong to Tabulata (Multithecopora cf. yohi) ，solitary Rugosa (Caninophyllum (Caninophyllum indosinense ， C. somtaiense) and massive Rugosa (lvanovia sigilla ω). 百le age age of 由is locality is Early Kasimovian. At another Middle Carboniferous locality ne 訂 B 叩 Wang Lan (1 60 52'30"N ，102 0 01'20"E) ，limestone blocks in a pond contain rare small solitary solitary Rugosa (Amygdalophylloides). Northwest of Ban Wang Lan (1 60 53'00"N ， 102 000'30"E) ， limestone lenses in shale belong to Lower Moscovian and contain solitary Rugosa (Caninophyllum).

2b3: 2b3: In Ban Na Duang 紅 ea (1 70 30'N ，101 0 58'E) ，Moscovian to Gshelian sediments have been been observed in a river (Huai Nam Suai) and in more than 10 ponds (FONTAlNE ET AL. ， 1991; 1991; FONTA 別 E& Su 百 ETHORN ， 2000); they consist of shale and limestone ，locally with a minor interbed of volcanic rock. Corals have been found in Moscovian limestone beds; 出ey belong to Tabulata (Chaetetes ，Multithecopora) ，solit 釘 y Rugosa (Cyathaxoniidae: Cyathaxonia; ・Cyathopsidae: Caninia simpliseptata; ・ Bothrophyllidae: Caninophyllum domheri ， Pseudotimania aff. mosquensis; Aulophyllidae: Sestrophyllum carinatum) ， fasciculate fasciculate Rugosa (Lublinophyllum thailandicum) and massive Rugosa (lvanovia rareseptata rareseptata ).

2b4:Phu 百四nPha 百lang is an isolated Upper Bashkirian limestone hill 10 km southeast of of Pak Chom (FONTA 町 E& SUTEETHORN ， 1988). It is surrounded by spilitic basalt and is supposed supposed to overlay 血is basal t.百 le limestone is rich in fossils including Tabulata (Chaetetes lungtanensis). lungtanensis).

2bS: 2bS: In Ban Na Charoen 訂 ea south of the road from Wang Saphung to Udon Th ani ， several several localities have been found (FONT A町 E ET AL. ，1982 ，1991 ， 1994). At the foot of a small small hilll km south Ban Na Charoen (102 0 01 '30"N ，17 0 11 '30"E) ，a limestone bed contains ， in in abundance ， very large Moscovian fasciculate corals (Lublinophyllum thailandicum) ，up to to 1m in diameter; it was discovered during a road construction in 1980. In a pond 2 km east east of Ban Na Charoen ， Tabulata (Multithecopora) and solitary Rugosa (Caninophyllum indosinense) indosinense) occur in an argillaceous limestone. At a small hill south ofthe pond ， an Upper Carboniferous Carboniferous limestone contains massive Rugosa (lvanovia sigillata). Near Ban Sam Luang 6 km southeast of Ban Na Charoen ， an Upper Kasimoviar トLower Gshelian limestone is is rich in Lublinophyllum thailandicum. A Moscovian limestone near Th am Pha Noi ，3 km northeast northeast of Ban Na Charoen ， contains Caninophyllum and Lublinophyllum thailandicum. At Ban Samboon ，5 km south-southwest of Ban Na Charoen ，solitary corals are common in in an Upper Moscovian limestone. Area 2c: In Northwest Thailand north of Chiang Dao ， only a few Tabulata and solitary Rugosa have been noticed in Late Kasimovian-Early Gshelian limestone 2 km north of Ban Na Wai (CARIDROπ ET AL. ， 1987). 58 E也NRI Fo 悶 AINE ，SIROT SAL Y APONGSE AND V ARA VUDH SUTEETHORN

Permian (295-245 Ma)

Pennian Limest 'O ne is widespread in Th ailand and its castellated t'O p'O graphy ad 'O ms m 佃 y areas. N 'O tt 'O'O far 企''O m Bangk 'O k ，it is well kn 'O wn near Ratburi and in 出eSar 油田t and L 'O pburi 紅 eas. Between Chiang Mai 佃 d Mae H 'O ng S'O n ， it displays typical k釘 8t t'O p'O graphy with sinkh 'O les ，caves ，undergr 'O und rivers ，rivers disappearing and reappe 釘加g. It It is an enchantment f'O r the visit 'O r. All 'O ver 官凶land ，many names 'O f limest 'O ne hills indicate indicate karst t'O p'O graphy: Kh a 'O Lak (hill with a milest 'O ne shape) ，Kh a'O百四 n 'O r Kh a 'O Kh uha (hill with cave) ，Kh a'O百四 n Nam L 'O t (hill wi 血 undergr 'O und river) ，Kh a 'O W 'O ng (hill (hill with sinkh 'O le) ，Kh a 'O Laem (p 'O inted hill) and S 'O 'O n. 百 1e limest 'O ne exp 'O sures 'O f Thailand r佃 ge 合om the base t 'O theωP 'O f 出e Pennian if if we c'O nsider the c'O un 町 in general. As a matter 'O f fact ，their dis 位ibuti 'O n is heter 'O gene 'O us stratigraphically stratigraphically as well 出 ge 'O graphically (F ONTAINE EI' AL. ， 1994). At the end 'O f the Pennian ， the Rug 'O sa became extinct al 'O ng with the tri1'O bites and fusulinids 泊 a very 加 p'O rtant mass extincti 'O n even t. Th e disappearance 'O f 血 e Rug 'O sa is easy t 'O 'O bserve in 官1ailand. 百 1ese c 'O rals died 'O ut after several milli 'O n years 'O f decline. 百lI s crisis was m 釘 ked by a decrease 泊 diversity in tw 'O main events: frrst ， the disappearance 'O f all the massive massive Rug 'O sa at the end 'O f the Midian (the upper part 'O f Middle Permian); and sec 'O nd ， the the disappearance 'O f all the 'O ther Rug 'O sa at the end 'O f the Upper Pennian.

Lower Permian

In In Th ailand ，L 'O wer Pennian limest 'O ne is n'O t very c'O mm 'O n. Li mest 'O ne bel 'O nging t 'O 恥 l'O werp 制'O fL 'O wer Pennian (Asselian and Sakm 釘 ian) is p制 icularly rare ， but it is in evidence evidence in a few are 部. C 'O rals have been f'O und in 伽 ee areas 'O fN 'O rtheast 百 1ailand. Th ey

紅 'e als 'O present 泊 Cen 住al 百 1ail 組 d 泊 at least 'O ne l'O cality. In In N 'O rtheast Th ailand ， s'O litary c 'O rals and massive c 'O rals (Keping 'O phyllidae) have been c'O llected 台。 m a few l'O calities (FONTAINE EI' AL. ， 1991). At the e部 tem f'O'O t 'O f Phu Kh a 'O 6 km n'O rthwest 'O f Ban Na Duang ， an impo 託釘1t assemblage 'O fc 'O rals 'O cc 町 S 泊佃Asselian -S akmarian limest 'O ne. Th ec 'O rals c 'O nsist 'O fs 'O litary Rug 'O sa (Cyath 'O psidae: Caninia Caninia lingwuensis ，Pseudozaphrentoides mapingense; B 'O thr 'O phyllidae: Bothrophyllum? ， Caninophyllum; Geyer 'O phyllidae: Amygdalophylloides) and massive Rug 'O sa (Keping 'O phyllidae: Antheria cf. polygonalis ，Nephelophyllum). In the Ban Na D 泊 Dam 釘 ea ， an 'O ther Asselian -S akmarian limest 'O ne c 'O ntains s 'O litary Rug 'O sa (Cyathaxonia ， Paraduplophyllum ，Amplexocarinia ，Calophyllum ，Lophophyllidium pendulum ，

Pseudozaphrentoides) ，fasciculate Rug 'O sa (Dens た， olumnophyllum tha i1 andicum) and massive Rug 'O sa (Anj 同ctophyllum). West 'O f Chumphae ， Keping 'O phyllidae (Anj 均 ctophyllum ， Antheria) Antheria) have been f'O und 2 km s 'O utheast 'O f Wat Sama はi 百 1am al 'O ng the r'O ad t'O血 e Center Center f'O r Wildlife C 'O nservati 'O n. Keping 'O phyllidae are ac 'O ral family previ 'O usly described 泊 China fr 'O m the Maping Limest 'O ne (Upper Carb 'O nifer 'O us-L 'O wer Permian). In In centralτ 'h ailand ，血e red limest 'O ne 'O f Kh a'O百四 n Rusi Laat n'O rthwest 'O f Ban Bung Samphan c 'O ntains p'O'O rly preserved s'O litary and massive Rug 'O sa. At Kh a 'O Ch 'O n 百 1'0 n'O地 -n 'O rthwest 'O fBan Na Ch aliang ， s 'O litary c 'O rals (Pavastehphyllum) 'O cc 町泊an Asselian- Sakmarian Sakmarian limest 'O ne. In In the upper p制'O f the L 'O wer Pennian 'O fL 'O ei regi 'O n， Tabulata (Protomichelinia) ぽ e relatively relatively widespread and in abundance at severall 'O calities. Phu Pha Wang (17 0 19 ・01"N ， GLIMPSES 削 TO FOSS 止 ASSEMBLAGES OF THAILAND 59

102 "0 1'40"E) along the road from Wang Saphung to Udon Th ani is a good example; 由is hill hill contains Protomichelinia laosen 抑制d P. simplex. South of Wang Saphung at Kh ao Pha Fai (1 7002'30"N ，I01 044'38"E) ， very rare massive Rugosa are present in an Art inskian limestone; limestone; they appe 釘 to belong to Wentzellophyllum.

Middle Permian

Corals Corals are known in Middle Permian limestones ，which 紅 'e dis 剖 buted all over 百四land. 百ley 釘 e 泊 abundance and great diversity in East ，Central ， Northeast and North Th ailand. 百ley are associated with large assemblages of fusulinaceans; these faunas are comple 飽ly different different from those in Australia. Fusulinaceans are unknown in the Permian of Australia. In In Peninsular 官lailand. corals are different 仕om those of the rest of Th ailand; they are less diverse. diverse.

Peninsular Peninsular Thailand: Limestone ranges from the end of Lower Perr 国組to Middle Permian and 血e beginning of Upper Perr 凶an. 官le ex 佐'eme top of the Permian h出 not been evidenced paleontologically paleontologically yet ， seemingly because of dolomitisation. 百le limestone contains particular and moderately diverse faunas. Corals occur at many localities of Kanchanaburi ，Ratburi ， Phetburi ，Pr achuab Kh irikhan ，Chumphon ， Surat Thani ，Phang Nga ，Phatthalung 組 dTr 加 g are 邸 (FONTAINE & S札 YAPONGSE ，2001). 百ley are rarely diverse and not usually abundan t. Their Their age is Murgabian to Midian. Phissadarn Cave (1 0045'27"N ，99 013 ・57"E) is a limestone limestone hill north of Chumphon; it contains a quite rich coral assemblage consisting of Tabulata Tabulata (Sinopora) up to 30 cm in diameter ， rare solitary and common massive Rugosa (Paraipciphyllum (Paraipciphyllum thailandicum ， P. kulvanichi). In fact ，corals ar 芭 scattered 泊血e limesωne. Th ey did not appe 紅 as important builders. In Peninsular Th ailand ，Tabulata 紅 'erep 陀 sented by widespread Sinopora. Solitary Rugosa without dissepiments (Lophophyllidium ， Amplexocarinia ， Ufimia) have been found in 白e Trang 訂 ea; Lophophyllidium pendulum ， Amplexocarinia ，and Paracaninia occur at Ban Phu Plu (14 005'45"N ，99 016'05"E) ， Lophophyllidium Lophophyllidium at Kh ao Laem Dam. Solitary Rugosa with dissepiments (Pavastehphyllum and Iranophyllum) are present at several localities: Khao Laem Dam (14 048 ・07"N ， 98 035'19"E) ，Kh ao Lan (13 033'44"N ，99 036'12"E) ，3 localities in Prachuab Khirikhan ，and a few localities in 血e Chumphon ， Surat Th ani ，Phang Nga and Pha 凶lalung 釘 e拙. Colonial corals corals exhibit a geogr ョphic dis 仕ibution similar to 血at of the solitary corals with dissepiments. Th ey belong to massive corals (mainly Para 伊c伊hyllum ，locally Wentzelella or Chombungia) and rarely fasciculate corals (Khao Yoi). Wentzelella has been found at a single locality:

Ko Ma (6 045'34"N ，99 045'22 官)， an island of Satun area; it has been identified 槌 Wentzelella megastoma (FONTAINE Er AL. ，1979). 官 le corals of Peninsular 官 lailand do not include genera genera such as Ipciphyllum and Pseudohuangia which 釘 e so common in Central Th ailand. Th e environment was not favourable to the development of many organisms. At Kh ao Lan (13 033 ・44"N ，99 036'12"E) near Chom Bung ， massive corals are rare and display growth bands bands at the foot of 出 e hill; higher up ，出 ey 釘 e common without showing grow 白 bands (FONTA 別 E& JUNGYUSUK ，1997). 官lI s fact has been interpreted as indicating a climate change ，from a seasonal climate at the foot of 出 e hill with different growth rates during 白e seasons to a continuous warm climate at the top of 出 e hill with a more regular growth. Th e base of the Lower Permian of Peninsular Th ailand contains pebbly mudstone which has has been interpreted by several geologists as a glacial deposi t. Corals 紅 e completely unknown in 血is part of the Permian. Other faunas also underline 山 peculiarities of the 60 HE NRl Fo 附 AINE ，SIROT SALYAPONGSE AND VARAVUDH SUTE 町田附

Permian Permian of Peninsular 官 lailand (F ONTA 町 E EI AL. ， 1994).

E 踊 t Thailand: Many limestones have yielded fossils ranging 企'om Kubergandianω Mi dian ， it it is to say belonging to 血e whole Middle Permian (FONTA 町 E&S 札 YAPONGSE ， 1997). Corals Corals have been found at severallocalities between the road 合om Sa Kaeo to Chanthaburi and the Cambodian border. At Kh ao Chakan (1 3"3 9'50"N ，102 0 05'E) ，there 紅 'e only a few solitary solitary corals: Khmerophyllum cf. cambodgense (SUGIYAMA & TORIYAMA ，1981) 出 well 拙 at Kh ao Bang Phrik (1 30 15'30"N ，102 0 18'E; Tachylasma ，Lo phocarinophyllum; FONTA 別 E & SAL Y APONGSE ， 1997). At other localities belonging mainly to Midian ， corals are somewhat more diverse. Th ey consist of a few Tabulata (Sinopora) ，fasciculate Rugosa (Waagenophyllum) (Waagenophyllum) and massive Rugosa (Parawentzelella ，Multimurinus). Parawentzelella ， agenusp 路 V ぬIUsly des 叫 bed 泊血e Mi dian 耐nesωne ofthe Sisophon area in West Cambodia ， has has been found again at some localities of East Thailand: Khao Phai (1 30 40'20"N ， I01 0 57' 55"E) ，Kh ao Sung- 悶 lao Ta Ngok at the Cambodian border ，Kh ao Chongkhaeb north north of Kh ao Ta Ngok. Central Central Thailand: Middle Permian limestone is widespread and is rich in many kinds of fossils. fossils. It had been deposited 泊 shallow and w 釘 m water; changes in sea level even led to 由.e temporary deposition of clastic beds containing continen ta1 plants around Saraburi and Phetchabun. Phetchabun. Corals 釘 'e prolific and diverse at a very great number of localities ne 釘 Muak Le k，Saraburi ，Lopburi ，Lam Narai (in p紅白ul 紅Kh ao Somphot 佃 d Kh ao Wong) ，Kh ok Sar 町 ong ，Tak Fa ， Wichianburi (Kh ao Khi Nok) ，Nong Phai and Phetchabun. Th ey are especiallyabundant 血血.eMurgabi 佃.百 ley consist ofTabulata (Protomichelinia ，Sinopora) ， solitary solitary Rugosa (Lophophyllidium pendulum ， L. zaphrentoidea ， Pseudozaphrentoides mapingense ，Verbeekiella? ，Pavastehphyllum ，Laophyllum ， Iranophyllum splendens) ， fasciculate fasciculate Rugosa (Yatsengia asiatica ，Waagenophyllum kueichowense ，Chaophyllum ， Pseudohuangia chiuyaoshanensis ， P. counilloni ， P. thailandica ， P. cf. aberrans ， Chihsiaphyllum Chihsiaphyllum kanmera i) and massive Rugosa (Ipciphyllum laosense ，1. elegans ，1. subelegans ，I. irregulare ，1. phadaengense ，1. saraburiense ，Paraipciphyllum magnificum ， Crass 伊'arietiphyllum sattayaraki ， C. tenue ， Multimurinus regularis ，M. khmerianus ，M. fontainei ，M.frechi ，M. suteethorni ，Wentzelophyllum thailandicum). A very peculiar coral (Khmeria (Khmeria problematica M 佃 suy) ，previously described in West Cambodia ， has been found south south of Phetchabun in Ban Na Chaliang area (FONTA 町 E ET AL. ， 1994); it is associated with with Codonofusiella ，a fusulinace 阻泊dica 曲19 a Murgabian to Late Permian age. Between Central Central and Northeastτ11ail 佃 d，Kh ao Th am Yai is a very large limestone hill with an important important cave (16 0 56'40"N ，101 "3 0'35"E) ， well visited by to 町 ists. It displays as 回 tion 700 m 血ick rang 泊g 仕om Murgabian to Midian ， very e錨 Y to observe (FONTA 町 E& SALYAPONGSE ，2001). 百le lowerp 副 of the section is rich in massive Rugosa (Ipciphyllum ， Multimurinus). Multimurinus).

Northeast Northeast Thailand: Permian corals 釘 'e known in a1 釘 'ge 釘 ea west of Chumphae 佃 d 泊 Lo ei region (FONT A sI沼町 AL. ， 1994; FONTA 町 E EI AL. ， 1995). West of Chumphae ，corals 紅 'e common 泊 Murgabian limestone along the road from Lomsak to Chumphae: at the northem end of Khao Ruak 33.7 km from Chumphae (Sinopora ，Lophophyllidium ， Ipciphyllum Ipciphyllum subelegans ， Multimurinus fontainei) ， at Phu Sam Phak Nam 28 km 合om Chumphae (Sinopora ，Lo phophyllidium ，Ipciphyllum ，Multimurinusfrechi) and at Sak Ch ay quarry quarry 25 km f旨om Chumphae (Protomichelinia laosensis ， P. globosa ， P. simplex). At the GLIMPSES INTO FOSSIL ASSEMBLAGES OF THAILAND 61 southeastern southeastern foot of Khao Hin King 35.5 km from Chumphae ， solitary Rugosa (Lophophyllidium) (Lophophyllidium) have been found in a Kubergandian limestone. Along the road from Khon San to Chulabhom Dam ， at Khao Dat Fa 22 km from Khon S加 ， Sinopora occurs in in small numbers. At Wat Samakki Tham 23 km from Kh on San ， corals are more diverse and consist of rare Tabulata (Protomichelinia ，Sinopora) ，solitary Rugosa (Lophophyllidium) and fasciculate Rugosa reaching 1m in diameter (Pseudohuangia cf. chiuyaoshanensis). At Khao Pha Phung 7 km north of Wat Samakki Th am ，a new genus of colonial Rugosa (Phaphungia) (Phaphungia) has been found; it is associated with Tabulata (Sinopora) ，solitary corals and other other massive Rugosa (Paraipciphyllum magnificum). In the Loei-Wang Saphung 陀 gion ， Middle Permian limestone is exposed east of Loei and northwest of Wang Saphung. East of of Loei ， Tabulata (Sinopora) and massive Rugosa (/pc 伊hyllum laosense and 2 other species of of Ipciphyllum) have been found at Ban Sam Yaek; Tabulata (Sinopora) ，solitary Rugosa (P αvastehphyllum) ，fasciculate Rugosa (Pseudohuangia chiuyashanensis) and massive Rugosa (Crassiparietiphyllum sattayaraki) at Ban Nam Suai Th a Saat; Tabulata (Sinopora) ， solitary solitary Rugosa (Lophophyllidium) ， fasciculate Rugosa (Chihsiaphyllum vacuum ， Pseudohuangia) Pseudohuangia) and massive Rugosa (Crassipariet 伊hyllum sattayaraki) at Ban Nam Suai Th a Sawan. F紅白er to the east around Nam Som ， Tabulata (Sinopora) ，solitary Rugosa (Pavastehphyllum) 組 d rare massive Rugosa probably belonging to Ipc 伊hyllum have been observed observed at Phu Pha Kup and at a neighbouring small hill ，Th am Suwana Kh uha. Northwest of of Wang Saphung ， corals are rare at Khao Pha Phung and Khao Pha Bao Pha Sao. Th ey are are associated with Codonofusiella ，a fusulinacean unknown so far east of Loei; they belong belong to Upper Murgabian or Midian. North Thailand: Permian corals have not been found in great abundance; they are widely scattered scattered in the rock (YANAGIDA ET AL. ， 1988; FONTAINE ET AL.， 1994; new data). Th ey have been collected mainly in Nan area from a Late Kubergandian ・Early Murgabian limestone limestone at Khao Pha Toob and Khao Pha Sing. In Lampang 訂 ea ， they are poorly known. At Kh ao Pha Sing 12 km north of Nan ，corals are rare; they are solitary Rugosa (La ophyllum) and fasciculate Rugosa (Chihsiaphyllum cf. kanmerai). At Kh ao Pha Toob 9 km north of Nan ， corals are more common ，especially fasciculate corals (Pseudohuangia cf. aberrans ， Chihsiaphyllum Chihsiaphyllum cf. kanmerai). Th ey include also some solitary (Laophyllum) and massive (Wentzelella) (Wentzelella) Rugosa. Northwest Thailand: Corals are not widespread. Th ey are known: in the Fang area ， between Chiang Dao and Mae Hong Son (at Sop Pong ，Ban Mae Lana and Ban Mae Suya) and in the Mae Sariang 紅白 (MINATO ， 1944; FONTAINE & SUTEETHORN ， 1993; FLUGEL ， 1997; 1997; HAHN & SIEBENffi 別 ER ， 1982). Th e coral identified by MINATO (1 944) as a specimen of of Wentzelella subtimorica belongs actually to Ipciphyllum laosense (M 町 ATO & KATO ， 1965). 1965). It had been collected from Fang area between Ban Doi Hua and Ban Ai about 80 km north-northeast of Chiang Ma i. Fasciculate Rugosa (Waagenophyllum) and Tabulata (Sinopora) (Sinopora) have been found north of Sop Pong. South of Ban Mae Lana ， fragments of Waagenophyllum have been collected from a dolomitic limestone. At Ban Mae Suya ， massive massive colonies of Multimurinus are common; they are associated with fusulinaceans indicating indicating a Late Murgabian age. HAHN & SIEBENHUNER (1 982) have cursorily mentioned Verbeekiella ， Yatsengia and Waagenophyllum at two localities between Mae Sariang and Tha Song Yang. 62 H 問問Fo 悶 AU 唱E ，SIR'σr SALY APONGSE AND V ARA VUDH Su 百四百ORN

Upper Permian

Upper Permian limestone has been evidenced in East and North 官lailand. It has yielded yielded very few corals.

East Thailand: at Wat Kh ao Kaew (1 30 06 'N， 102OZ4'20"E) ，solitary corals (Paracaninia)

occur 泊 a limestone containing Colaniella 佃 d prob 油 ly belon 仰 gωWuchiapingian. No 巾 east of Kl aeng ，Palaeofusulina ，a Changshingian fusulinacean ， has been found at two small small limestone hills near Wat Kh ao H 恒(1 20 52'30"N ，I01 0 48'45"E; FONTA 凹 E EI' AL. ， 1999); 1999); no coral was observed. North Thailand: solitary Rugosa (Lophophyllidium) and fasciculate Rugosa (Liangshanophyllum) (Liangshanophyllum) have been collected 企om limestone at Ban Cham Ka northeast of L 倒npang and at Phra 官lat Muang Kh am southeast of Lampang; they 釘 'e associated with a Wuchiapingian microfauna (FONTAINE & VACHARD ， 1988). At Huai 刊誌west of Ban Pang Kh o along the road 企'om Lamp 叩 g to Ngao ， an Upper Permian sequence of shale ， calcareous calcareous shale and argillaceous limestone contains rare small solitary Rugosa (Asserculinia) associated associated with abundant Leptodus (brachiopods). North of this locality at Doi Pha Phlung ， Waagenophyllum aff. virgalense has been 民 :ported 泊 Changshingian limestone (Y ANAGIDA EI' EI' AL. ， 1988; Is 田 BAS 田町AL. ， 1998).

Triassic Triassic (245-205 Ma)

Triassic Triassic limestones have been evidenced recently at many localities of 百lailand. In由 e past ， they were considered Permian in age because 血ey are similar to Permian limestones if if fossils are not taken 泊to accoun t. Th ey range from Lo wer Triassic to the lower p釘 t of Upper Triassic (Carnian); the lower part of the Norian may be locally presen t. Triassic Corals Corals have been found at several limestone exposures 泊 Peninsular ，East ，North ，and West Th ailand. Th ey have never been collected from Lo wer Triassic rocks. A little more than than 20 species have been mentioned. Solitary forms (Montlivaltia ，Distichophyllia ， Conophyllia ，Margarophyllia) ，fasciculate and massive colonies (Thecosmilia ，Volzeia ， Cyathocoenia ，E か'sastraea ，Margarosmilia ， Thamnasteria) have been identified mainly from reports on regional geology (Pπ'AKPAIVAN EI' AL. ， 1969; YANAGIDA EI' AL. ， 1988; Ao ACHI EI' A L.， 1993; FONTA 町 E EI' AL. ， 1993). Th ey have not been studied in detai l.百 ley need to be revised. Some identifications 釘 'e actually debatable. Peninsular Peninsular Thailand: Solitary ，fasciculate and massive corals have been observed at Kh ao Phanom Wang ， a hill 9 km northwest of Ph atthalung ，where 血ey 釘 e 泊 abundance( Ao ACHI EI' EI' A L.， 1993; FONTA 町 E EI' AL. ， 1993). Solitary corals have been found at Khao Kh uha south south of Phat 白alung where 出ey 釘'e associated with Aulotortus σo 悶 A 町 E ET AL. ， 1993). A Carnian age has been proposed for the two localities. 百le Triassic limestones of Phatthalung Phatthalung r佃 ge from late Early Triassic to early Late Triassic (AMPo RNMAHA ， 1995).

East Thailand: Solitary and fasciculate corals have been noticed 創出ree Triassic limestone exposures: exposures: the frrst at Kh ao Cha ・ang On north of Kl aeng (13 0 12'30"N ，10I 0 39'30"E); the second second at Kh ao 百lep N 加 it Banphot south of Aran yapathet (13 0 37' 30"N ，102OZ 7' 30"E); and the third at Khao Noi Si Chomphu near the Cambodian border (1 30 34'40"N ， 102 0 32'E; FONTA 町 E& SALYAPONGSE ， 1997). GLIMPSES GLIMPSES INTO FOSSIL ASSEMBLAGES OF 百IAILAND 63

North Thailand: 町 rAKP AIV AN Er AL. (1969) mentioned solitary and fascicula 低 Scleractinia 加 Phrae Pr ov 泊ce at 血e 佃 timony mine at Pha Kh an ，Am phoe Lo ng. Thecosmilia was reported reported at Huai Rong Sak no 巾 of Ngao (YANAGIDA Er AL. ， 1988). In Nan Province ， Triassic Triassic limestone is exposed 泊 several ar 回 s (FONTAI 阻 Er AL. ， 2001). Solitary ，fasciculate and massive corals have been found in Ban Huai Kaet are a; YANAGIDA Er AL. ，(1 988) mentioned mentioned Elysastraea ，Montlivaltia and Thecosmilia at this locality. Solitary and a few fasciculate fasciculate corals have been noticed in Na Noi 釘 'ea and between Pong and Santisuk (FONTAINE Er AL. ， 2001). West Thailand: Triassic limestone is widespread and builds up two north-trending series of of hills in Uthai 官lani area (F ONTAINE Er AL. ， 2000). Hi lls of the westem series display well-preserved well-preserved fossils and corals have been observed at five hills. Th ey contain Upper Triassic Triassic microfossils. At Kh ao Pathawi (1 5"2 8'20 ・'N ，99 045'40"E) ，solitary ，fasciculate and massive massive corals have been found. Large solitary corals reaching 6 cm in diameter and 20 cm 泊 leng 血 are common 泊 the middle part of 出 .e hill; fasciculate corals up to 1m in diameter diameter are in abundance at the northem end of 出 e hill. North of Kh ao Pathawi ，solitary and fasciculate corals have been observed at Kh ao H 加Th oen and Kh ao Phra North; at Kh ao Kachi ，corals are very rare. South of Kh ao Pathawi at Kh ao Pu n，as 泊gle fasciculate coral coral has been noticed. At 血e eastem series of hills along longitude 99 0 58'E ， limestone is recrystallized. recrystallized. Nevertheless ，corals are still visible at the surface of the rock at a few hills: Kh ao Ngo ，Kh ao Rum ，Kh ao Lai ，Kh ao Pun (same name as another hill of the westem series) ， small hill northeast of Kh ao Hin Phloeng ， and Kh ao K 蹴 o. 百ley 紅 e poorly preserved; to to the naked eye ，出 eyappe 釘 similar to those in the hills of the westem series.

Jurassic Jurassic (205-135 Ma)

Marine sediments of Jurassic age are known only in West Th ailand. Th ey are exposed in in several areas from Peninsular Th ailand to Mae Hong Son. Th ey have been 血e subject of of a recent PhD 曲目is (MEEs OOK ， 1994). Shallow marine conditions existed over most of West Th ailand. Mudstones ，siltstones and sandstones are widespread. Limestone is prominent only 泊 Mae Sot ，Umphang and Kanchanaburi areas. Fossils 紅 'e abundant and diverse; they consist consist of algae ，foraminifera ，corals ，sponges ，bryozoans ，brachiopods ，bivalves ，g槌 tropods and and ammonites (F ONT A削 E&SU 田町iO剛， 1988; B 臥 UVAIS & FONTAINE ， 1993; MEEs OOK ， 1994). 官ley indicate mainly Toarcian ， Aalenian and Lo wer Bajocian ages. However at Pa La 百la south of Umphang ，corals are associated with two Late Jurassic fossils: Cladocoropsis mirabilis ，a calcisponge ，and Salpingopore l/ a pygmaea ， an alga. Th e marine Jurassic 抑制 of of West 官lailand can be corr 芭lated wi 出 the non-marine Jurassic of Northeast 百lailand ， belonging belonging to 血e “Kh orat Group" and extending from Chiang Rai to Nakhon Phanom and achinburi. Pr achinburi. Corals Corals are sporadically present from Kanchanaburi to Mae So t.百 ley may be only a few scattered fragments of corals as in Kanchanaburi area (at Huai Si Sa Ho) or 血eymay be rich assemblages as in some areas of Umphang and Mae Sot where they consist of soliωry ，fasciculate and massive specimens. About 50 species have been 飽 cog 凶zed in 官 lailand [F ONT A 副 E& S U"TEE四 ORN (1 988) with a chapter written by Beauvais; BEAUVAIS & FONTA 町 E (1 993)]. 64 HENRI FONTAINE ，SIROT SALYAPQNGSE AND VARAVUDH Su 百四百ORN

h 出 e area 'O f Kl'O官 1'0 west 'O f Umphang ，c 'O rals are in abundance and 血 ey c'O nstitute the the richest Jurassic c 'O ral fauna kn 'O wn 泊百聞land. 百 1ey d 'O n'O tf 'O rm 佐ue reefs. Alth 'O ugh 出 ey 紅 e 泊 a 住'emend 'O us number ，出ey d 'O n'O t j'O in 'O ne an 'O ther and hence d 'O n'O t build up a rigid 合amew 'O rk. Th ey c 'O nsist 'O f: Epistreptophyllum giganteum ，Montlivaltia numismalis ， M. tenuiradiata ，M.fragilis ，M. cf. ca ηophyllata ，Calamophylliopsisflabellum ，C.flabellum var. var. compacta ， C. klothoensis ， C. etalloni ， C. sp. ，S.か 'losmilia tenuis ，S. michelini ，S. corallina ， S砂lina girodi ， S. deluci ，Cladophyllia thailandica ， C. dichotoma ，Lochmaeosmilia radiata ， Latomeandra Latomeandra ramosa ， L. sp. ， Dermosmilia laxata ， D. crassa ， D. sp. ，Fungiastraea pseudoarachnoides ，Microsolena exigua ，M. sp. ，Litharaeopsis fontainei ，Thamnasteria sp. ，Actinaraea granulata ，Kobyastraea subagaricites ， K. bourgeati ， K. haradai ，Thamnoseris frotei ， Haplaraea sp. ，lntersmilia exigua ，Allocoeniopsis bernensis ， La tiphyllia langi ， Clausastraea Clausastraea edwardsi ，Andemanastraea densisepta ，Araiophyllum sp. Near Ban U The Khi s'O uth 'O f Umphang and Ban Pa La Tha ， s 'O litary c 'O rals (Cyathophyllopsis ，Epismilia) have been c'O llec 飽d 合om a single l'O cality. Calamophylliopsis has has been f'O und n 'O rthwest 'O f Ban Pa La Th a. In In the Mae S'O t area ，c 'O rals are rare at D 'O iD 泊 Chi n 'O rthwest 'O f Mae S 'O t; they 釘 e litary. s'O litary. Five t 'O 10 km east 'O f Mae S'O t，there 釘 'e als 'O small numbers at a few l'O calities ne 紅 Ban Pha De ，al 'O ng Huai Pha Y 'O and al 'O ng Huai H 泊 F 'O n.

Near Ban Huai Ya-u 25 km e拙 t 'O f Mae S'O t (as 白 e crow flies) ，many Jurassic c 'O rals have have been f'O und recently but have n'O t been studied ye t.官 1ey are mainly fasciculate and massive massive c'O l'O nies. 百 1e fasciculate c'O rals 紅 e in abundance; 出 ey are 30 t 'O 50 cm 泊 diameter. Th e massive c 'O rals are smaller and less c'O mm 'O n; they reach a maximum diameter 'O f 'O nly 15 15 cm. 百 1e Jurassic c 'O rals 'O f Th ailand display s'O me similarities with Jurassic c 'O rals through 'O ut Asia (Ind 'O nesia ， Japan ， Philippines ， Sarawak) ， westem Eur 'O pe (England ， France ， Switzerland) ，M 'O r'O cc 'O， Madagascar ， and the Mi ddle East. H 'O wever ，血ey sh 'O w peculiarities 佃 d 百凶land can be c 'O nsidered a dis 由1ct subpr 'O vince within Asia.

Cretaceous Cretaceous (135 -6 5 Ma)

Th ere is n 'O marine Cretace 'O us 泊官1ailand 'O r 泊 Camb 'O dia ，La'O s and Vietnam. In S 'O utheast Asia ， marine Cretace 'O us is kn 'O wn 'O nly in 白 e Philippines ， East Malaysia ，and In d 'O nesia. Acc 'O rdingly ， there is n 'O Cretace 'O us c 'O ral at all in 司副land.

Tertiary Tertiary and Quaternary (65 Ma to present)

On land far from sea ，Tertiary sediments fill small t 'O m 'O derately large b錨泊s; they 釘 e n 'O n-marine. Even at 白 .e b'O tt 'O m 'O f 血 e Gulf 'O f 官同land ，Tertiary -Q uatern 訂 y sediments are are fluvi 'O・ lacus 凶ne at 血 eir base and bec 'O me marine 'O nly at 曲et 'O p (ALDERSON EI AL. ， 1994). 1994). During the Quaternary ， sea level changes were 加 p'O rtant and sedimentati 'O n was n'O n-m 紅加et 'O marine in the c 'O astal z 'O nes. Th e latest Pleist 'O cene-t 'O -Middle-H 'O l'O cene marine marine sediments have been described (f 'O r 泊 stance) in the S'O ngkhla 紅国 (C HAlMANE E ， 1987) 1987) 'O r加白.e Bangk 'O k plain wi 白血'O ccurrence 'O fa l-m thick 'O ys 飽rbank (5 ，500 years ld) 'O ld) at Wat H 'O i in Am ph 'O e La t Lum Kae 'O， 40 km n 'O rth 'O f Bangk 'O k (CHONGLAKMANI EI AL. ，1983 ，DH 回貼DILOK ，1987). Acc 'O rdingly ，c 'O rals sporadically ∞C 町 in varying numbers. GLIMPSES INTO FOSSIL ASSEMBLAGES OF THAILAND 65

Coral fragments form up to 30 percent of the beach sand of Samui and Phangan Islands (KOHPINA & CHIEMCHINDARATANA ， 1997). In the Gulf of 官lailand and the Andaman Sea ，cor a1 reefs occupy about 1，800 km 2.

ACKNOWLEDG 勘ffiNTS

百le study of Th ailand cor a1 s has been carried out with the permission of the Directors of the Geological Survey of Th ailand; the authors are indebted to them for their continuing encouragement. Dr Sompoad Srikosamatara ，Mahidol University ，gave judicious advise for the production of 血is paper; the authors are very grateful to him. Dr Warren Y. Brockelman ， Editor of the Natur a1 History Bulletin of the Siam Society ，must be thanked for his interest and kind help in editing.

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