Fossil Biodiversity in the Limestones of Thailand: a Cornucopia of Information About the History of Life
NAT. NAT. HIST. BUL L. SIAM Soc. 53 (1): 33-70 ，2005
FOSSIL BIODIVERSITY IN THE LIMESTONES OF THAILAND: A CORNUCOPIA OF INFORMATION ABOUT THE HISTORY OF LIFE
Henri Henri Fontaine 1， Sirot Salyapongstl and Varavudh Suteethorn 3
ABSTRACT
In In Th ailand ，foss iI s are common and diverse. 百 ey come from bo 血 terrestri aI and m 紅泊E environments.τ 'h ey belong to many t泊施 periods beginning with the Cambrian ，血 us spann 泊g more 血an 500 m iII ion years (白 e Phanerozoic eon). 明lis rich past emerges 台。 m extensive pub Ii shed data and is st iII very in 加 esting to explore. 明lis pub Ii cation concems only foss iI s included in Ii mestone deposited in 白巴 seas of the pas t. Li m 巴stone is widespread 泊百ailand and of various 格的. Marine floras (aI gae) and
faunas 釘宮 in abundance at many Ii mestone exposures 飢 d their skeletons ar 草加 impo 此釦I component component of 血.e limestone. 百ley give a deep-time perspective on the evolution of the Ii fe in 出e seas of 血 .e past. τ'h e Ii mestones of Th aiI and are not res 町icted to 出 e widespread exposures visible on th 巴 land. land. Th ey have been found by hydrocarbon exploration at varjed depths and in different are 鎚. Permian Permian Ii mestone has been reached by we Il s under the Kh orat Plateau and its extent has been determined determined by seismic interpretation; it is widespread (for instance ， see MOURET ， 1994). Th is pub Ii cation is concemed only wi 出 l加 estones exposed at ground surface. Limestone is a gene raI term for diverse types of rocks ，deposited in different environments. Before describing their their biodiversity ，we discuss the orig 泊。f 出e Ii mestones of τ'h a iI and.
Key words: Diversity ，foss iI s，limestone ，m 悩 sextinctions ，paI eog ωgraphy， ，s回 .tigraphy ，百 lailand
L Th庄 ESTONES OF THAILAND
In Th ailand ，limestone (Hi n Pun in 百lai language) is a rock very widespread at the surface surface of the land all over 血e country. It makes up small to very large hills ，which are locally locally numerous ， for instance north of Saraburi between Lopburi and Muak Lek. Beautiful landscapes have been carved by erosion ，producing a marvellous castellated topography. Caves 紅 .e common. In addition to the prominent hills ，smalllimestone outcrops 釘 e scattered at at ground surface.τ 'h ey are not visible in the topography ，but easily seen in the banks of the the rivers as well as in ponds or other holes dug by f，創 mers. Li mestone is known by everybody and m 佃 y hills 紅 e called Kh ao Hin Pun or simply Kh ao pun (= Limestone Hi ll). In other places ， the names show the imagination of the local peοple; they describe the peculi 釘 topography of the limestone (k 紅 st topography; see Figs. 1，4 to 6). Many hills are called Kh ao 百lam or Kh ao Kh uha (= Hill with a Cave). A qualifier qualifier may be added because of 組 underground river (Khao 百四n Nam Lot) ，a peculi 釘
18 18 aIl ee de la Chape Il e，92140 Clamart France 2143 2143 Moo 6， Soi Pinnakom 4，Borommaratchachonnari Road ， Bangkok 10170 ，Th aiI and 30 eo log ic aI Survey ，Dep 釘飢lent of Mi ner aI Resources ，Rama IV Road ， Bangkok 10400 ，τ'h aiI and Received Received 8 November 2004; accepted 16 May 2005.
33 33 34 HENRI FONTAINE ，SIROT SAL Y APONGSE AND V ARA VUDH SUTEETHORN feature feature (Kh ao Suwana Kh uha = Hill of the Gold Cave) ， or the size (Kh ao Th am Yai or Khao Th am Luang = Hill of the Large Cave). Kh ao Laem indicates a pointed hill. Phu Tum means a bud shape; it is a common name in Loei Province. Doi Sam Sao (Hill of the 百rree Triangles) ， between Chiang Dao and Phrao ，displays 3 angular points of limestone standing standing out from its top. Li mestone hills are commonly steep-sided; Khao Pha Daeng is a hill with a red cliff ，Khao Pha Dam is a hill with a black cliff ，Khao Pha Lat is a hill with with an inclined cliff ，Kh ao Lak is a hill surrounded by cliffs and showing a milestone shape. shape. Khao Wong is a hill with a sinkhole at its centre. Wh en the limestone is clearly bedded ，it is compared to a pile of folded clothes (Khao Phap Pha). Kh ao Phap Pha is a common name in Surat 官 lani Province (see Fig. 3). Th e names of other hills are focussed on the flora or the fauna of the hills: Kh ao Ruak (Hill (Hill of a small kind of bamboo) ，Kh ao P耐ik (Hill of the Red Pepper) ，Khao Pha Fai (Hill with with a cliff and cotton) ，Kh ao K wang (Hill of the Deer) ，Kh ao Th am Ik ea (Hill of the Cave of of the Bats) ，Kh ao Lan (Bald Hill ，without vegetation). Many species live in karst topography at at the surface of the hills or in the caves which are common in limestone hills.
Types of Limestone
Li mestone is commonly grey to black ， but locally it may be white ， yellow or red. It is is massive or thinly to thickly bedded (Figs. 1 to 6). is It fine- to coarse-grained. From a chemical chemical point of view ，it is almost pure carbonate of calcium deposited in a clean environment; environment; this is the case at many localities of Th ailand. However ，this is not the rule; clay clay and even sand locally occur in the limestone ，occasionally in such a quantity that the appropriate appropriate names of the rocks become calcareous shale or calcareous sandstone. Limestones are are polygenetic; they were deposited in differing environments which will be described below. below. In these different types of rocks ，fossils are commonly well preserved. Their abundance abundance dis 佐ibutions and their diversities reflect the images of ancient living communities ， with with some fidelity. After After its deposition ，limestone has been transformed by different diagenetic processes which were destructive or constructive. It may have been dolomitized; then ，it displays a fair fair content of magnesium. In the field ，dolomitic limestone is easy to recognize because its its surface is not smooth ， but looks like elephant skin. Dolomitization commonly destroys the the fossils and precise identifications are impossible. Wh en limestone has been heated by intrusion intrusion of igneous rocks ，it is recrystallized and the fossils 紅 e again largely or entirely destroyed. destroyed. In dolomitized and recrystallized limestones ， the original characters 訂 'e obscured or or erased. Th ese rocks are not helpful in reconstructing life of the past and they are locally difficult difficult to date. They are fortunately restricted to small areas of Thailand.
Depositional Depositional Environments
Limestone Limestone has been deposited almost everywhere in τbailand 加 marine environments ， and commonly in distinct subenvironments of shallow seas. An exception has been recognized recognized (IW AI ， 1972 and 1973) ，a few limestone beds at the base of the Kh orat Group in in Northeast Th ailand have been considered lake deposits; they 紅 'e thin ，less 由加 10 cm thick ， with the exception of a bed reaching 2m in thickness. Th is limestone is practically barren barren of fossils. Lacustrine faunas ぽ e commonly of low diversity. Such deposits are FOSS IL BI OD IVERS ITY IN T H E L1 MESTONES OF TH A IL AND 35
Figll re 1. Lim es ton e hill s nea r Na ll1 Pi ya ng Din Wa ter fa ll sO lllh 01' Wa ng Sa phll ng in Lo巴iPro vi nce パlorlh eaS lell1 T hail a nd. Th e li ll1 eS l0ne，Ea rl y Permi an in age ，co nl a ins ll1 any fo rall1 ini fe ra (s ll1 all erf oramini fe raa ncl
fu sulin ace an s). It is al so ri ch in ca Ic isph er es a nd T ll bip /り刊 s frag ll1 enl s; co rals are ral 巴 (FON Tfl INE ti
AL. ，2002 ，a nd in pl 巴parati on)
Fig ur e 2. B edd ed lim 巴stone atK oS iC hang nea rV as hir av lll Brid ge. K oS i C hang is an isla nd 01' lh 巴 north easlel 日 part part of th e Gul f0 1' Th ail and . Th 巴 li ll1 es lone is part ly rec rys talli ze d ancl poo r in foss il s (Sfl LYfl PONGSE ET A L.， 20 02); it is co nsid erecl O rdov i cian in ag 巴 36 HE NR IF ONTA IN E，SIR OT SALY AP O NGSE AN D V A RAVUDH SUTEETI -IORN
Fi gllr es 3. 8 edded lim es ton巴s at two diff ere l1l hill s. a: a: Permian li l1l es ton e hill of S lI rat T ha ni Pr ovin ce in Penin s ular Th ail and . T he lim es ton e is c1 ea rl y bedd e d. II is cO l1l par ed to to a pil 巴 of fold ed c1 01h es (Phap Pha ) and ca ll ed Kha o Phap Pha (Hill of th 巴 Fo lded
C loth 巴s) . lt is loca ll y ri ch in f oss il s ，blll of of a low di ve rsil y . b: Khao Ph a Wak ，a hill hill so uth of lh e road f' rom T ak 1 0 M ae SO I. The li l1l eSlon 巴 is poor in fo ss il s. Thi s hill hill has a cave and is hi sto ri ca ll y int eres tin g; it was a hidin g pla ce 1' 01 so ldi ers dllrin g World War Il FOSSIL BIODIVERSITY IN THE LIMESTONES OF THAILAND 37
Figures 4. Limestone hills at a Buddhist temple before arri ving at the Nam Pi yang Din Waterfall. a: Thickly bedded limestone at a hjll near the temple. b: The same limestone, eroded, at the entrance of the temple. 38 38 H ENR I FO NTA INE ，S IROT S ALYA PO NGSE AN D V A RAVUD I-IS UTE ETHOR N
F ig llr e 5. K hao Thalll Kr ac ha 巴ng ，h ill of san Na ng Sata al巴a in sO ll th 巴rn mo st T hail and ，w ith an u nd 巴rgro und
tu nn 巴1 fo ll owed by a ri ve r. T he li lll es tone is ma ss iv巴 and rec rys ta lli z巴d.
F ig ur e 6. Do i Ph aC hu (18・23'15"N ，100 ・4 8' 55"E ) in Na No i area o f Na n P rov in ce. Ma ss ive lilll esto n巴co nt 白川川g I' oss il s in lll oderate qll antit y: algae ，fa sc icul ate co rals，s po nges ，b ryozoa ns ancl b iva lves (FONTA INE ET A L. ，200 1) FOSS 且 BIODIVERSITY IN THE LIMESTONES OF THAILAND 39 known in lakes of North America ，occurring under different sets of circumstances. In addition addition to the limestone of the Kh orat Gr oup ，caliche ，travertine and ωfa can be mentioned. τbey are te 町'estrial calcareous materials of secondary accumulation ，known at many localities of of Th ailand; 出ey 釘 'e too small in space and too restricted in time to be very interesting. However ，出 is type of rock has recently a町 acted attention at Kh ao Si Siet ， a hill built up by unconsolidated pebbly alluvial deposits ， north of Am phoe Lao Kwan 泊 the northem p釘 t of Kanchanaburi Province. Polished 仕agments of carbonate material formed 泊 these deposits deposits give beautiful pieces of marble ，occasionally containing land snails. In addition to 出at ，fissure fillings 紅 e common at the limestone hills of Th ailand; they locally contain fossils fossils such as skeletons of Quatemary terres 凶 al vertebrates (for instance ， see CH AlMANE E ET AL. ，1996 ，fossils from Kh ao Sam Ngam near Ratb 町 i). 1n 1n the geologic record ，limestone is commonly absent 企om deep sea deposits. However ， it it is occasionally represented by 由加 turbidite beds ，called allodapic limestone. Fossils are not not in their original place ，白 ey were transported from shallow areas and transfer 冗 d to ba 白川 depths. 1n Cen 佐al Th ailand ，this type of rock (Nam Duk Formation) has been described described along the road from Lom Sak to Chumphae ，especially ne 紅 milestone 17.2 km (HELMC 阻&KRAI 阻 ONG ， 1982); it contains a poor allochthonous fusulinacean fauna. Shallow Shallow water marine limestone is extensive in Thailand. It has been commonly deposited deposited in well oxygenated water ，illuminated by day-light ， at suitable salinity and temperature. temperature. Great numbers of fossils with high diversity suggest favourable environments. Th e shallow depth of 白e water is indicated by the occurrence of green algae at many localities. localities. Warm water is suggested by the abundance of 1釘 ge compound corals at many Devonian Devonian to Jurassic localities. Some limestone formations are 白ick; they were continuously deposited during a long t泊施， in the same environment. Th ey consist of accumulations of skeletal material as well as as organically induced accumulations of types other of carbonate material. Some organisms encrust encrust or attach to one another; 白ey locally built up a rigid mass even before fossilization. τbe environment was locally energetic ， with strong waves increasing the supply of food food and oxygen. Many organisms could thrive easily. But after 由自dea 出， their skeletons were were often broken by the turbulent water ， but commonly 泊 not too small 仕agments ，出 us allowing allowing good identifi~ation. In some places ， the fossils and their debris were accumulated in in de 甘ital mounds ，later on consolidated by inorganic cementation. Reefs were commonly wave-resistan t.百 le limestone produced in energetic environments is formed mainly of grains grains and skeletal debris. However ，modem sea-grass is regarded as protecting lime-mud buildups; buildups; the size of the grain must not be always conside 問 d accord 泊g to the water turbulence. turbulence. Environment Environment was more or less quiet in other areas. Li mestone is fine-grained. 1t is locally locally ultrafine-grained ，clay-sized carbonate and it is called micstone. Fossils 釘 e better preserved ， but they are commonly less diverse and less abundan t. They may be almost absent absent where water was confined to an undisturbed environment ， poorly aerated and almost deprived deprived of oxygen. In lagoons ，salinity can become low with influx of 仕esh water ， or become high with evaporation du 出19 合y seasons. High evaporation has produced 血e
deposition deposition of gypsum in several 紅 eas of 百lailand. In In addition to the limestone displaying castellated topographic relief ， other formations consist consist of shale ，siltstone ，sandstone and limestone interbeds or lenses ， with less impressive topographic topographic features.τbe limestone is commonly not visible in the topography ， but it may 40 H 町 RI Fo 悶 'AINE ，SIROT SAL YAPONGSE AND V ARA VUDH S凹宙開O 剛
be 'O f great interest in 白e se 釘 ch f'O rf 'O ssils. In L 'O ei Pr 'O vince ，s 'O me Middle Carb 'O nifer 'O us limest 'O ne lenses 紅 'e algal m 'O unds very rich in algae ass 'O ciated with 'O ther f'O ssils σONTA 町 E ET AL. ， in prep 紅 ati 'O n).
Ages
The limest 'O nes 'O f Th ailand bel 'O ng t 'O different ages. Th ey are pr 'O minent from Ord 'O vician t 'O Middle Jurassic ， 'O r 企om 490 t 'O 150 milli 'O n ye 釘 s (Ma) ag 'O. During the Cretace 'O us and the Tertiary (合'O m 135 t 'O 5 Ma) ，there was n 'O actual sea 'O n the present land 'O f Th ailand and there is n 'O marine limest 'O ne 'O f these ages. During Late Tertiary and Quatemary ， m む ine influence sp 'O radically extended t'O l'O w parts 'O f 百 lailand ， with 'O ut imp 'O rtant dep 'O siti 'O n 'O f limest 'O ne. In In Kr abi Pr'O vince 'O f Peninsular Th ailand ，calcare 'O us mudst 'O ne is exp 'O sed at Phra Nang Bay 16 km 合'O m Kra bi t'O wn. It 'O verlies Late Eoc ene c 'O al beds. Very rich 泊 gas 位op 'O ds ， it it is an enchantment f'O rt 'O urists.τ 'h eg 部位op 'O ds bel 'O ng t 'O a few species 'O f 合'esh water m 'O lluses ，acc 'O rding t 'O identificati 'O ns made 泊 1949 (PπAKPAIVAN ET AL. ， 1969). During During the Jurassic (206 t'O 143 Ma) ， sea c'O vered 'O nly the westem p紅 t 'O f 百 lailand ， fr 'O m Mae H 'O ng S'O nt 'O Penins u1 arτ 'h ailand ， while the eastem-n 'O rtheas 旬mpart 'O f百凶land ， especially 出 e Kh'O rat Plateau ，w 錨 a plain with meandering rivers and r'O aming din 'O saurs. Jurassic Jurassic limest 'O ne is present 'O nly in 出， e westem p制'O f the c'O un 町. It l'O'O ks fresher 白血 the the 'O lder limest 'O nes. It builds up 1紅 ge hills 泊 Mae S'O t and Umphang areas. It is l 'O cally present present in Kanch 如 ab 町 iand Mae H 'O ng S'O n 紅 'eas. At many l'O calities ，it is rich in diverse ssils; f'O ssils; algae ，f 'O raminifera ，c 'O rals ，brachi 'O p'O ds ，ga 紺 op 'O ds ，bivalves and amm 'O nites have been n 'O ticed and 'O ccasi 'O nally described. Triassic Triassic (248 t 'O 205 Ma) limest 'O ne is presently kn 'O wn 泊 many 釘国s 'O f 百国land ， from the pen 加sula t 'O n'O rthem and eastem 百 lailand. Because Triassic limest 'O ne is n'O t different 合'O m the Permian limest 'O ne at a first gl 佃 ce ，it was 儲 signed t 'O Permian 泊血e past past in s'O me 釘 eas 'O f central ， eastem and s'O uthem 百 lailand. Because it is rich in diverse f'O ssils ，it has been p 'O ssible t 'O ascertain the exact age 'O f the Triassic limest 'O ne 'O f several areas areas during recent pale 'O nt 'O l 'O gical research σONTA 町 E & VACHARD ， 1981; FONTA 町 EET AL. ， 1993; AMPORNMAHA ， 1995; FONTAINEET AL. ，2001 and 2003). In the Lamp 如 g Basin 'O fn 'O rthem 官lailand ，出e Triassic sequences 紅 'e m 'O re 血佃 3000 m thick and have been kn 'O wn f'O ra l 'O ng time (PITAKPAIVAN ，1955). 百 ey c 'O ntain significant carb 'O nate facies 'O f different different lith 'O l'O gic types. Th e limest 'O ne approximately c'O rresp 'O nds t 'O af 'O urth 'O f the t'O凶 secti 'O n. Its fauna (bivalves ，brachi 'O p'O ds ， and amm 'O nites) has been described. It bel 'O ngs t 'O L 'O wer (Upper Gri esbachian 'O r the l'O wer part 'O f the L 'O wer Triassic) ，Middle (L 'O wer and Upper Anisian) and Upper (Middle Carnian) Triassic (CHONGLAKMANI ， 1981; CHONGLAKMANI & OUDOMUGSORN ， unpublished rep 'O rt: 14 pふ Permian limest 'O ne (290 t 'O 247 Ma) is very widespread all 'O ver the c'O un 町， even after the the c'O rrecti 'O n 'O f the ages 'O fs 'O me limest 'O ne 'O utcr 'O ps e汀 one 'O usly assigned t'O出 e “Permian Ratburi Ratburi Limest 'O ne" 泊出，e pas t.百 le Permian limest 'O ne mainly bel 'O ngs t'O L 'O wer and Middle Permian. Upper Permian limest 'O ne is rare and d'O es n'O t build thick f'O rmati 'O ns; it is is kn 'O wn at a few l'O calities 泊 Lampang and Nan 泊 N 'O rth 官lailand ，泊血e Klaeng 釘 'ea 泊 East Th ailand ，and fr 'O m Phangnga 佃 da few 'O ther l'O calities ('O nly l'O wer pぽ t 'O fUpper Permian) Permian) in Peninsular 百 ailand. 百 le Upper Permian limest 'O ne is n'O t diverse in f'O ssils; it 釘 m 'O unces 白e greatm 部 s extincti 'O n 'O f the end 'O f 血 e Permian. It is ass 'O ciated with shale FOSSIL BIODIVERSITY IN THE LIMESTONES OF THAILAND 41
Table Table 1. Divisions of the Mesozoic. Times are the beginning of the periods in Ma.
Tertiary Tertiary 65Ma Cretaceous Cretaceous Upper 99Ma Lower 144Ma Tithonian Tithonian 151Ma Upper = Malm Kimmeridgian 154Ma Oxfordian Oxfordian 159Ma Callovian Callovian 164Ma Bathonian Bathonian 169Ma Jurassic Jurassic Middle = Dogger B 吋ocian 176Ma Aalenian Aalenian 180Ma Toarcian Toarcian 190Ma Pliensbachian Pliensbachian 195Ma Lower = Li assic Sinemurian 202Ma Hettangian Hettangian 206Ma Upper Norian 221Ma
C 倒 nian 227Ma Triassic Triassic 恥I{ iddle Ladinian 234Ma Anisian Anisian 242 民I{ a Lower Olenekian 245Ma lnduan lnduan 248 勘1a
containing containing peculiar brachiopods ， the Lyttoniidae. Lower and Middle Permian limestones are are commonly very rich in diverse fossils with prominent assemblages of fusulinaceans and and corals. Th e Lower Permian ，known for a long time at Khao Th am N 創 n Maholan in Loei Loei Province ， extends south to the Nam Piyang Din Waterfall 釘 ea according to new findings findings (FONTA 町 E ET A L.， 2002; new data). Carboniferous Carboniferous (354 to 289 Ma) and Devonian (417 to 355 Ma) limestones are r訂 e in Peninsular Peninsular and West Thailand. ln Loei region in Northeast Thailand ， they are widely exposed exposed at many localities (FONTAINE ET AL. in preparation). Carboniferous Carboniferous limestones are less widespread than the Permian limestones. Th ey are in in evidence in Eastem Th ailand (Klaeng ，Sa Kaeo and Kabinburi areas) ， in Central Th ailand (Ban (Ban Bo Nam area east of Lam Narai ，Noen Maprang and Chon Daen areas west and southwest southwest of Petchabun) ，泊 Northeastem 官lailand (at many localities of Loei Province) ， in in Northwestem Thailand (between Mae Hong Son and Chiang Dao as well as north of Chiang Chiang Dao) ， in Westem Th ailand (northwest of Kanchanaburi) and in the southem part of of Peninsular Th ailand. Fusulinella has been very recently discovered in a limestone lens included included in shale near Nam Piyang Din Waterfall (new data). Accordingly ，this new locality (17 003'21 "N ， 101 044'45"E) belongs to an age corresponding to upper Middle Carboniferous; the the Carboniferous was previously unknown in that area. In Thailand ，Lower Carboniferous is is known at a greater number of localities than Middle and Upper Carboniferous. Lower Carboniferous Carboniferous fossils have been collected from limestone in Peninsular Th ailand as early 42 42 匝到RI FO N1・ A 別 E ，SIRIσr SALYAPONGSE AND VARAVUDH Suτ 'EETH ORN
Table Table 2. Di visions of the Paleozoic
Upper Upper = Lop 泊gi 佃 Changhsingian 252Ma Wuchiapingian Wuchiapingian 256Ma Midian Midian 259Ma Mi ddle = Guadalupian Murgabian 262Ma Permian Permian Kubergandian 264Ma Kungurian Kungurian 266Ma Lower Lower = Cisuralian Art inskian 269Ma S紘 marian 282Ma Asselian Asselian 290Ma Upper Upper =Upper Pennsylvanian Gshelian 296Ma Kasimovian Kasimovian 303Ma Mi ddle=Lower Pennsylvanian Moscovian 311Ma Carboniferous Carboniferous Bashkirian 323Ma Se 甲ukhovian 327Ma Lower Lower = Mississippian Visean 342Ma Tournaisian Tournaisian 354Ma Upper Upper Famennian 364Ma Frasnian Frasnian 370Ma Devorian Devorian Middle Givetian 380Ma Eifelian Eifelian 391Ma Emsian Emsian 400Ma Lo wer Pr agian 412Ma Lochkovian Lochkovian 417Ma Pri doli 419Ma
Siluri 組 Ludlow 423Ma Wenlock Wenlock 428Ma Ll andovery 443Ma Ashgill Ashgill 449Ma Caradoc Caradoc 458Ma andeilo Ll andeilo 464Ma Ordovician Ordovician Ll anvirn 470Ma Ar enig 485Ma Tremadoc Tremadoc 490Ma Upper Upper Cambrian Cambrian Middle Lower Lower 543Ma FOSSIL FOSSIL BIODIVERS 汀 Y IN THE LIMESTONES OF THAILAND 43 as as 1899. Since then ， other localities have provided Lower Carboniferous fossils ， but Middle and Upper Carboniferous limestones remain unknown in 由is p紅 t of 出e country. In In 1969 ，it was believed 血at no definite Devonian fossil had been found in 官lailand (P 汀雌PAIVAN ET AL. ， 1969). In fact ，as 均 le fossil had been mentioned in 1961 (FONT AINE ， 1961 ， p. 216); it was at a locality in Nong Bua Lamphu Pr ovince. Since 1980 ， diverse Devonian fossils have been collected from limestone widespread in the Loei region (Lo ei and Nong Bua Lamphu Provinces).τbey consist primarily of corals and stromatoporoids σONTA 町 E ET AL. ， 1981; FONT AINE & T.釧 TIWA 町 T， 1987; FONTAINEETAL. ，1990). Devonian limestone limestone is less important and widespread in West Th ailand from Chiang Mai to Satun 組 d it does not contain the diverse faunas of the Lo ei region. Devonian conodonts have been been reported in Mae Sariang area where they are associated wi 血 fishes. West of 官loen ， Devonian limestone has been described in Mae Ping National Park (BURREπ ET AL. ， 1986). 1986). Devonian trilobites and conodonts have been collected from limestone of Satun Pr ovince (LONG & BURRETT ，1975) ，trilobites and tentaculites (HAHN & SIEBENHUNER 1982) 1982) as well as conodonts (SAVAGE & SARDSUD ，2003) 仕om calcareous rocks of 百long Pha Phum 紅白 in Kanchanaburi Province ，tentaculites from a limestone lens along Huai Song Song in 血e National Park north of 官lO ng Pha Phum (FONTAINE ET AL. ， 1988). Silurian Silurian (443 to 418 Ma) is of relωively short duration. Li mestone is not prominent in 血e Silurian sequences of Thailand. It has been mentioned in Northeast ，West and Peninsular Peninsular Th ailand. In 1969 ，Silurian fossils were known only in shale (町 TAKPAIV AN ET AL. ， 1969). Ordovician Ordovician (490 to 444 Ma) limestone is widespread in western 百lailand ，from Mae Hong Son to the Malaysian border. It has provided diverse fossils. In 1969 ，cephalopods ， gastropods ， brachiopods and trilobites were already reported in the Ordovician of southern 百lailand (PπAKPAIVAN ET AL. ， 1969). Later on ，Lower to Middle Ordovician limestone with with argillaceous layers has provided a diverse fauna in Satun Province ，Th ailand; 血e fauna fauna included trilobites ，nautiloids ，gas 住opods ，brachiopods and os 回 .cods. 百le limestone had been deposi 制 in peritidal to deeper waters (WONGWANICH ET AL. ， 1990). Other localities localities have been described. 百le ages indicated in Tables 1 and 2 are those of the bases of the stages. 百 ley are average average ages ， based on slightly different results obtained from different isotopic systems. Paleontological Paleontological investigations are focussed on well-preserved sediments ， and absolute ages
訂 'e obtained from volcanic ashes or layers. Nevertheless ，results from the s創 ne geologic horizons horizons can provide a better understanding of the rapidity of the evolution of the living world. world. Additional research on radiome 出cda 由港 of volcanic layers is continuously 也lproving the the precision of the ages.
BIODIVERSITY
Disturbance Disturbance of present environments can reduce the biodiversity of 血e present-day ecosystems ecosystems and become a challenging problem of more th 佃 academic interes t. At a few localities localities of Th ailand ，geologists and other scientists have observed the negative results of mangrove destruction ， leading to seashore erosion and loss of land for farmers ， loss of habitat habitat for some species and loss of reproduction area for other species. Mangroves 紅 'e a
旬pe of vegetation growing under the s位'ess of complex tidal movements. Th ey are well 暢 44 HEN 阻 FONTAINE ， SIROT SALYAPONGSEAND VARAVUDH SU'I宮 E叩 O悶
known in present-day tropical zones ， but are geographically res 凶C旬 d.τbey are difficult to to recognize 泊 sediments older than the Cenozoic because 出.ey are easily destroyed. In Singapore ， loss of habitat by an 血ropogenic changes has produced a wide range of local extinctions extinctions (BROOK EI' AL. ， 2003). Fish stocks are declining ， at least in some seas ，p 釘 tly but but not entirely ，due to overfishing. In some 紅 'eas ，fish stocks have failed to recover when fisheries fisheries were closed. In 2002 off the Oregon coast ， the upwelling of water poor in oxygen killed killed great quantities of crabs and fishes (0 貼 NTHAM EI' AL. ， 2004). Some sp 民 ies of deer and birds (moa of New Ze aland ，dodo of Mauritius) have disappeared. 百le worldwide decline decline of coral reefs calls for an improved understanding of the ecological processes that underlie underlie reef resilience (B 乱 LWOOD EI' AL. ， 2004). About 70% of North American large manlmal species were lost at the end of the Pleistocene ，a response to climate ch 組 ge (GUTHRIE ， 2003). Vegetation locally changes in composition. Increas 泊.g carbon dioxide 加 出.e atmosphere gives a competitive advantage to fast growing plant species at 由e expense of of the native vegetation. Scientists have discovered 出at knapweed grown 泊 a sterile liquid secretes secretes a chemical (catechin) 台om its roots 白紙 kills other plants ，allowing 血is species to to invade the Am erican West (Ac 国間ACH ， 2004). In 1982 ， an imported alga (Caule 中 a taxifolia) taxifolia) st 制 ed to invade the Mediterranean sea bottom south of France and reached It alian and Spanish coasts by 1992. 団副 uy toxic ，it does not interest herbivores. It disperses rapidly ，eliminating many other plant and animal species (ME 町 ESZ ， 1999). A scenario for 2050 foresees import 佃 t shifts in 血e dis 住ibution and abundance of species because of climate climate fluctuation (THOMAS EI' AL. ， 2004). Highly endangered species will be most vulnerable vulnerable to loss of their environments. Natural mechanisms can change productivity and population population dynamics and 田 osystems 釘 e vulnerable to fluctuations. Wildlife conservation becomes a complex problem ，especially when it spreads from one coun 町 into another (SRIKOSAMAT 組 A& SUTEE 叩 O 悶， 1994). 百le present-day ecologists can test how the organisms 釘 e affected by each other and by 白eir environments. They can meas 町 e and evaluate evaluate the physical factors con 釘olling the environments.τbey can map pattems of biodiversity. biodiversity. Paleontologists lack 白紙 luxury; their analysis is a more imprecise 釘t.官時 ir conclusions conclusions are partly based on comparisons with present-day 加 alogous examples; as elsewhere elsewhere in geology ， the present is the key to the p出t. For instance ，it is difficult to know with with a great precision 白e comple 飽 food web of an ecosystem of the p錨t. However ， our understanding understanding of the evolution of the living world has increased and our vision of paleoecology paleoecology has progressed. Studies Studies of the remote past show successions of floras and faunas on a time scale of millions millions of years. 官 ley give a deep-t 加 e perspective on biological events. All the limestones of 官 lail 姐 dwe 陀 not deposited in exactly the s創ne environments. Biodiv ぽ sity has multiple facets. facets. In order to give a general pict 町 'e of 出is biodiversity ，information will be given on the the diversity of the fossils ，出 eir s回 tigraphic r佃 ges ， the changes in their populations with geographic geographic dis 甘ibution ，and the dramatic events of their history.
The Fossils
τbe frrst 泊lportant discoveries of fossils 泊百lailand date back to 1899. A few molluscs and a foraminifer were found in Chon Daen 釘 'ea 加 Central Th ailand by an officer of the Forest Forest Department; 出ey were studied by the British Museum and were assigned to Carboniferous. Carboniferous. Th ey actually belong to Lower Permian. 百le report of 曲is frrst discovery FOSSIL FOSSIL BIODIVERSITY IN THE LI 恥IESTONES OF THAILAND 45 was published in the Bangkok Times Weekly Mail of 7 August 1900 and was forgotten. ln 1924 ， N. B. Garrett ，interested in old newspapers ， found the report and suggested that it be be reprinted in the Journal 01 the Siam Socie 砂 (vo l. 18: 63 -64). Also in 1899 ，Lower Carboniferous Carboniferous fossils (cephalopods ，bivalves ，brachiopods ，trilobites and a fragment of Tabulata) Tabulata) were collected from Phatthaluog area by a Cambridge expedition; later 00 ， they were were described (REED ， 1920). Th e great majority of paleontological studies have been carried out during the last 50 years. 百ley have established the biostratigraphy ofτ 'h ailand and allowed intemational correlations correlations over very long distances. Among Thai geologists involved in the first paleontological paleontological studies ， K. Pitakpaivan is well known in particular because of his publications on fusulinaceans. In 1966 ， he described fusulinids collected prior to 1957 and mentioned “ the future possibility of using fusulines in distinguishing faunal zones in the differentiation of of the Rat Buri limestone" (PπAKPAIVAN ，1966 ， p. 4). Other geologists have been partly forgotten ， although" they 紺 ongly stimulated paleontological research in the 1950s and 1960s ，for 加stance V. Sethaput ， S. Buravas ，M. Veeraburas ， D. Bunnak ， S. Kaewbaidhoon and and A. Hongnusonth i. 百le aim of this paper is not to give long boring lists of the fossils found so far in Th ailand ， but to illustrate the diversity of the fossils with occasional mention of the names of of particular fossils.
Algae Algae
Algae 釘 e locally in great abundance. They are more common 白an indicated in 白e published published literature ， because they have been studied only occasionally. Green algae (Dasycladaceae) (Dasycladaceae) indicate a shallow water environment because 白ey need strong light to survive. survive. Some algae are good stratigraphical markers. Tubiphytes is a problematical type difficult difficult to 出 sign to a systematic group; it is sometimes regarded as an alga. lt is widespread and in abundance in Carboniferous and Permian limestones of Th ailand. It occasionally acts acts as an incrusting and binding organism. lt is prominent at many localities. Renalcis ， 釦 0 白er problematical form ， has been found in Devonian limestone at two localities of Ban Chok Chai area in Northeast Th ailand (FONTA 悶 E ET AL. ， 1990). It is common in the Devonian of Westem Australia and is known in Europe and Canada. A Jurassic alga ， Holosporella siamensis Pia ，was described as early as 1930; it had been been collected at the Burmese border ne 釘 Mae Sot from a bed supposed to be of Upper Triassic Triassic age (PIA ，1930) ， an age not confirmed by recent studies. Later on ，this alga was found found at Jurassic localities in 百lailand and other countries. lt mainly belongs to Middle Jurassic; Jurassic; it has been rarely mentioned in Upper Jurassic (BASSOULLET ， 1988). Algae 訂 "e widespread in the Triassic limestones of different areas of Th ailand ， from Peninsular Thailand to East and Central Thailand. They belong to several genera (VACHARD ， 1988; ADACHI ET AL. ， 1993; FONTA 削 E ET AL. ， 1996). Algae Algae have recently been discovered at Kh ao Ma (14 0 09'OO"N ，99 ・07' 40"E; samples T8540 to T8547) ，a limestone hill 186 m in elevation 5 km northwest of Amphoe Sai Yok and the new Mahidol University campus. They 釘 e locally the dominant framework constituent constituent of the limestone of this hill. Th ey app 釘 "ently belong to a single species.τ 'h ey are are not associated with foraminifera in the material which has been collected so far; see Figure Figure 10d (preliminary information before the detailed study). 46 HENRI Fo 悶 'AINE ，SIROT SALYAPONGSE AND VARAVUDH SUTEE 四 0 悶
Permian Permian a1 gae are diverse and abundant at many loc a1 ities. 百1ey have been studied in detail detail in Saraburi where 64 species have been described 但NDO ，1969) ，but 血ey occur a1l over over Th ailand and have been mentioned in many publications ，occasion a1 1y with precise identifications identifications and photographs. Mizzia is a green a1 ga found at many Permian loc a1 ities of of Th ailand. In In Middle Carboniferous rocks 泊出e Loei area ，a1 gae (Beresella 佃 d others) are very abundant abundant and 釘 e the dominant framework constituent of the limestone. During their life ， 出.ey grew erect 釦 d probably looked like a meadow of sea grass. Some 釘 'e presently still in in their living positions. Most have been broken and 血e debris have accumulated to form a1 g a1 mounds (Fontaine et a1.泊 prep 紅 ation). 百1e alga Koninckopora has been found in Northeast ，East ，C 印刷1 and Northwest 官1ailand. It has a quite narrow stratigraphic a1 range ，belong 時 to 由e upper p制 of Lower Carboniferous ， mainly Mi ddle-Upper Visean.
In In the Lower Devonian of Ban Muang 加 Nong Bua Lamphu Province ，gr 田l n a1 gae (La ncicula) are loc a1l y in abundance. 百us type of a1 gae is not very widespread 泊 the Devonian of the world ， but is known in seve ra1 coun 凶 es from Austria to Austr a1 ia.
Radiolarians Radiolarians
Devonian ，Carboniferous ， Permian and Triassic radiolarians have been extracted 仕om chert chert and siliceous sh a1 e at sever a1 loc a1 ities of northem ，northeastem and southem 百凶land. Studies Studies of radiolarians 合om limestone 紅 e rare exceptions.
Radiolarians Radiolarians have been recovered from a thin-bedded limestone at Kh ao Chiak ne 紅 Pha 曲 alung ，southem 百1ailand. 百1ey were well-preserved ，associated with conodonts (SASHIDA&I ∞， 1992); 白.ey 泊.dica 低血age ranging from Lower Tri 蹴 ic (l atest Ol eneki 佃) to to the lowermost Mi ddle Triassic (e 釘Ii est An isian). Lower Triassic radiolarians have a1 so been been found at another limestone locality of Phatth a1 ung 紅 'ea (SAS 凹 DA EI' AL. ， 1998).
Foraminifera Foraminifera
τ'h is group of protozoans is so diverse 白紙 it is difficult to give a complete information about about them. Th ey rank high 泊 terms of usefulness for age determination. Th ey have been studied studied by sever a1 p a1 eontologists and have been described in a1 紅 ge number of papers; the the main au 血ors are: Pitakpaivan ，Toriy 創 na ，Igo ，In gavat ，Vach 紅 d，Bassoullet ，Ueno ， Charoentitira t. Pen 叫叩 foraminifera ，including sm a1l er foraminifera and 釦sulinaceans ，
have have been extensively studied 泊 Centr a1百 1ailand and a1 so 泊 other p釘 ts of 百1ailand. Carboniferous ，Triassic and Jurassic foraminifera are known 加 a much sm a1l er number of publications. publications. Foraminifera of other ages ar 芭 almost unknown. In the Lo wer Pa1 eozoic a1 1 over over the world ，出.ey are ac 旬a1 1y not diverse and 釘 e mostly simple tubular or globular forms. forms. In Middle-Upper Devonian ，血 ey show a frrst radiation. No important assemblage has has been noticed so far 泊 the Devonian of Th ailand. C a1 cispheres 釘 'e a group of nU crofossils 白at have been described as “single-chambered foraminifera"; they are widespread in the Carboniferous Carboniferous and Permian limestones of 百1ailand. Jurassic Jurassic foraminifera were described for the frrst time 泊百1ail 組 d 泊 1976. 百1ey were included included in limestone samples collected from the northem p制 of Kanchanaburi Pr ovince (HAOEN & KEMPER ， 1976; KEMPER ，1976). Later on ，Jurassic foraminifera were discovered in in Mae Sot and Umphang are 出 (BASSOULLET ， 1988 and 1994). 百1ese forarninifera belong FOSSIL FOSSIL BIODIVERSITY IN 百 IE Ll MESTONES OF THAILAND 47 to 出e following genera: Gutnicella ，Haurania ，Mesoendothyra ，Timidonella ，Bosniella ， Spiraloconulus Spiraloconulus and other less important genera. 百ley suggest Lower-Middle Jurassic ages. Timidonella Timidonella has also been mentioned in Jurassic limestone of Mae Hong Son area. Triassic Triassic foraminifera are small and c創 mot be easily noticed in the field. Because of that ， they have not been considered valuable for many ye 釘 S.τbeir recent discovery at several several localities of 百lailand has changed our ideas about these fossils. Th ey have been found found ne 釘 Nan (FONTAINE Ef AL. ，2001) ， in Kanchanaburi Province (KEMPER Ef AL. ， 1976) ， in Uthai 百四姐紅白 of Central Th ailand (FONTAINE Ef AL. ，2000) ， in K1 aeng 紅 'ea (FONTA 町 E & V ACHARD ， 1981) and near 恥 Cambodian border 泊 East Th ailand (FONT AINE Ef AL. ，1996) ， at sever a1 10calities (southwest of S<;>ngkhla ，泊 Trang ，Phatth a1 ung ，Phangnga and Chumphon areas) of southem 百 ailand (FONTAINE Ef AL. ， 1993). Permian Permian foraminifera are widely distributed 加百lailand. 百 ley are very diverse and they 訂 'e good index fossils. Th ey have been described in many papers published during the last last 40 ye 紅 s. As early as 1969 ， 41 species offusulinaceans were already known in 百lailand (Pπ'AKPAIVAN Ef AL. ， 1969). In con 回 st with other foraminifera which are small ，some fusulinaceans fusulinaceans are easy to see in the field with the naked eye. Wh en limestone is polished ， they they may adom some pieces of beautiful marble. In Peninsular and West Th ailand ， Permian fusulinaceans fusulinaceans are not diverse and are distributed 泊 o叫yap ぽ t of the Permian. Elsewhere ， they 釘 e present in a1 1 the Permian ，especi a1 1y from Asselian to Midian (Lower Midian is more widespread than Upper Midian) ， and display a high diversity. Th ey provide a precise biostratigraphic biostratigraphic zonation of the Permian. Th ey 紅 'e common in the Lower Permian of Centr a1， Northeast ， North and Northwest Th ailand. Th ey 釘 e abundant in the Mi ddle Permian a1 10verτbailand. A veηgood section ，rich 師向sulinaceans ranging 合om 白e upper part of of the Lοwer Permian (Misellina horizons) to a1 most the end of the Mi ddle Permi 組 (Colania (Colania douvillei horizon) ， has been described from the Saraburi 釘 ea ， mainly at Kh ao Prong Prong Pr ab and partly at Kh ao Khao 佃 d Kh ao Inl ot (see table 4 in the following paragraph on 出 e stratigraphic r佃 ges of the fossils); eleven biozones have been reported (TOR 町 AMA ， 1976; 1976; TORIYAMA & KANMERA ， 1979; To 則 YAMA ， 1984). Th e top of Middle Permian (Upper (Upper Midian) contains Lepidolina ，a genus found at sever a1 10c a1 ities of Sakaeo Pr ovince (PITAKPAIVAN & lN OAVAT ， 1980) and at Kh ao 百四n Yai east of Lomsak in Northeast 百 ailand (FONT A町 E Ef AL. ， 2002). During Upper Permian ，fusulinaceans are known only 泊 a few sm a1 1 釘 eas of North and East 官lailand; 白ey 釘 e small. Outside the fusulinaceans group ，a sm a1 1 foraminifera ，recently described as Sphairionia Nguyen 1989 and widespread 加百lailand ， deserves special mention because it indicates a high horizon of 出e Permian near near the boundary between Middle and Upper Permian. In March 2003 between Kanchanaburi Kanchanaburi and Th ong Pha Phum ，白 is genus was found in 血e bedded limestone of Kh ao Yai at Wat Th am Phromm a1 ok (sample T8514; 14 ・13'05"N ，99 ・08'15"E; new data) in in the area of Mahidol University campus. is It associated with Tubiphytes. At 血e top of a limestone hill (1 4・05'42"N ，99 ・16'09"E; samples T8519 and T8520; new data) near 出e Buddhist Buddhist Temple of Ban Phu Plu along the road from Kanchanaburi to Th ong Pha Phum ， the the limestone beds are packed with Rectost 伊ulina ，a small elongated foraminifer which indicates indicates an Upper Permian (Wuchiapingian) age ， but is not perfectly preserved. At the foot foot of the same hill about 30 m below the Rectostipulina horizon ，limestone beds are rich in in fusulinaceans including Eopolydiexodina (FONTA 町 E ET AL. ，1988 ， p. 35-36). Genus Rectost 抑 lina Jenny-Deshusses 1985 is a very peculiar foraminifer (F ONTAINE & NOUYEN ， 1989). 1989). It has been found at five localities of the Sai Yok ー官long Pha Phum 紅白; it has 48 HENRI FONTA 別 E ，SIROT SALY APONGSE AND V ARA VUDH S U1宮町'H ORN
als 'O been c 'O llected fr 'O m Kh a 'O Kh釦 B 叩 Dai n 'O rtheast 'O f Pr achuabkhirikhan ，where it is ass 'O ciated with 'O ther imp 'O rtant f'O raminifers: Hemigordiopsis ，Codon o_ かsiella ，Dagmarita and Paradagmarita. Pu blicati 'O ns 'O n the Carb 'O nifer 'O us f 'O raminifera 'O f百lailand 釘 efewer 白血publicati 'O ns 'O n Permian f 'O raminifera. They st 釘 t in 1972 with a descripti 'O n 'O f Middle-Upper
Carb 'O nifer 'O us f 'O raminifera f'O und 泊 limest 'O nes 'O f the Wang Saphung 訂 'ea 泊 L 'O ei Pr 'O vince (IGO ， 1972; see Table 3 in the f'O ll 'O wing lines 'O f this text). Later 'O n， Middle-Upper Carb 'O niferous f 'O raminifera were f'O und at m 叩 y 'O ther l'O calities 'O fL 'O ei Pr 'O vince as well as as L 'O wer Carb 'O niferous f'O raminifera; 40 species have been rep 'O rted in the l'O wer p釘 t 'O f Middle Middle Carb 'O niferous: Upper Bashkirian (VACHARD ， 1990; FONT AlNE ET AL. ， in prep 卸値'O n). L 'O wer ， Middle and Upper Carb 'O nifer 'O us f 'O raminifera have been disc 'O vered at many calities l'O calities between Mae H 'O ng S'O n and Chiang Da 'O邸 well as n'O rth 'O f Chiang Da 'O. F'O raminifera ，ass 'O ciated with c 'O rals and bel 'O nging t'O出 e upper part 'O fL 'O wer Carb 'O nifer 'O us (Visean) ，'O ccur n'O rth 'O f Klaeng 泊 East 百lailand. Qnly L 'O wer Carb 'O nifer 'O us f 'O raminifera have have been f'O und in Peninsular and West 官 lailand. F 'O raminifera (chernyshinellid) bel 'O nging t 'O the l'O wer part 'O fL 'O wer Carb 'O nifer 'O us (T 'O urnaisian) have been c 'O llected fr 'O m al 'O cality in in a nati 'O nal park n'O rth 'O f Kanchanaburi (FONTAINE & VACHARD ， 1987).
Corals Corals
C 'O rals 釘 e 泊 abundance at many limest 'O ne exp 'O sures in 官lailand. 百ley 釘 el 'O cally c'O n佐ibut 'O rs t 'O the building 'O ft 'O p 'O graphic pr 'O minences 町 reefs ，f'O r instance in the Dev 'O nian
'O fN 'O rtheast Th ailand. In 'O ther 釘 'eas ，也ey are restricted t 'O limest 'O ne beds ，f 'O r 泊stance the Mi ddle Carb 'O niferous bed 'O f Ban Na Charoen in N 'O rtheast 百lailand rich in Lublinophyllum and s'O me Permian limest 'O ne beds al 'O ng the r'O ad fr 'O m Saraburi t 'O Lam Narai (near Km 10). 10). At 'O ther l'O calities ，they 釘 'e scattered 泊 the limest 'O ne. 百leir dis 釘ibuti 'O n in time and space space has been presented in a recent paper (FONT A町 E ET AL. ， 2003). h 出 e past ，c 'O rals 'O f the Atlantic were r，紅 'ely c 'O nsidered endemic t'O也 is 'O cean; 血 .ey were were said t 'O bel 'O ng t'O血 e same lineages as the c 'O rals 'O f the Pacific because 'O fm 'O rph 'O l 'O gical similarities. similarities. An alyses 'O f rnit 'O ch 'O ndrial and nuclear genes have sh 'O wn 白紙 this assumpti 'O n was inc 'O rrect; many Atlantic c 'O rals bel 'O ng t 'O a previ 'O usly u町 'ec 'O gnized lineage (Fu KAMI ET AL. ，2004). 百 le microstructures 'O fc 'O ral skelet 'O ns actually substantiates this c'O nclusi 'O n. F 'O ssil c 'O rals must be studied acc 'O rding t'O their m 'O rph 'O l'O gy ， but als 'O acc 'O rding t'O血 eir micr 'O structure with 'O ut f'O rgetting p 'O ssible diagenetic changes.
Jurassic Jurassic c'O rals 紅 e in great abundance at s'O me l'O calities 'O f the Mae S'O t and Umphang areas; areas; they l 'O cally reach 1m in diameter (BEAUVAIS ， 1988; BEAUVAIS & FONTA 町 E， 1993). 1993). Th ey have been studied by L. Beauvais ，especially 泊 amaj 町 publicati 'O n 'O f 1988; 49 species have been described. They were c 'O llected fr 'O m a fine-grained limestone ， characteristic characteristic 'O fa m 'O derately'quiet envir 'O nmen t. Triassic Triassic c 'O rals have been f'O und 合om Peninsular t'O n'O rthern 百lail 阻 d. In N 'O rth 百 ailand ， three three species have been rec 'O rded fr 'O m 血 e Kang Pla Limest 'O ne at 白e Ban Pha Kh an antim 'O ny mine in Phrae Pr'O vince (PITAKPAIVAN ET AL. ， 1969). Triassic c'O rals 釘 e abundant and and diverse at s'O me l'O calities ne 釘 U 出 ai 百四泊 in central Th ailand (FONTA 町 E ET AL. ， 2000) ，'O fN 如釘'ea 泊 n'O rthern 百lailand (FONTA 町 E ET AL. ，200 1) and 'O f Sakae 'O Pr'O vince ne 釘 the Camb 'O dian b'O rder (FONT A町 E & SAL Y APONGSE ， 1997). From Kh ao Phan 'O m W 佃 g，ab 'O ut 9 km n 'O rthwest 'O f Pha 凶lalung in Peninsular Th ailand ，s 'O litary ，fasciculate FOSSIL FOSSIL BIODIVERSITY IN THE LIMESTONES OF THAILANO 49
and massive corals have been considered important builders of an Upper Triassic (C 訂 nian) limestone limestone (AOAC 田町AL. ， 1993). Permian corals have been found all over Th ailand and have been observed at more than than 100 localities. 百ley are abundant and diverse at many localities of Central ，Northwest ， No 凶1 ，Northeast and Southeast Th ailand. Th ey locally consist of large fasciculate or massive colonies ，reach 泊g1m in diameter (FONTAINE ET AL. ， 1994; FONTA 町 E，1999). 百 ey 紅 e less less diverse in Peninsular Th ailand (FONTAINE ， 1989). Outside Peninsular 百 ailand where corals corals are unknown in Lower Permian ，出 ey range from Lower to Upper Permian with a decline decline after the end of Middle Permian. More 出an 90 species of Tabulata and Rugosa have have been identified.
Carboniferous Carboniferous corals 釘 'e locally in abundance. Th ey consist of dendroid Tabulata and solit 釘 y，fasciculate ，and massive Rugosa. 官1ey are known in limestones r佃 ging from Lower to Upper Carboniferous in the Loei region (FONTAINE ET AL. ，1991 ， 1995). Th ey occur occur in Lower Carboniferous of Central Th ailand (west and southwest of Phetchabun) ， of East East 百1ailand (Klaeng 釦 d Kabinburi areas) 佃 d of Northwest Th ailand (Amphoe Pang
Mapha and Ban Na Wai 釘 'eas). One group of corals is easy to recognize; it belongs mainly to to Hexaphyllia ，a genus indicating the upper p釘 t of Lower Carboniferous. Middle-Upper Carboniferous Carboniferous limestone lenses have also provided corals east of Lam Narai in Central 官1ailand. In In the Lo ei region (Loei and Nong Bua Lamphu Provinces) of Northeast Th ailand ， Devonian limestone is widespread and builds up many hills. It contains diverse coral faunas faunas belonging mainly to Lower and Middle Devonian. Th ese corals consist of branching (Thamnopora ，Cladopora ，etc.) and massive (Favosites ，Alveolites ，Heliolites ，etc.) Tabulata ， solitary solitary (Sol 伊etra ，Si 初nos 申1J on 噌1昭g叩o. 'P 卸hμYμl !y 伽ωI1 um肌爪7η 1丸， et に仏.よc b勘r叩 chi 加ng (ω Dend 砂，ro ωste ωeU 伽la ，Thamnophyllum ， etc.) etc.) and massive (Phillipsastraeidae ，Endophyllum and others) Rugosa (FONTA 別 EET AL. ， 1981; 1981; FONTAINE ET AL. ，1990). 官1ey 釘 e associated with stromatoporoids which are in abundance abundance at many localities. In In Northeast Th ailand ，Silurian corals (Tabulata and solitary Rugosa) have been reported from two localities north of Nam Som: 5 kn1 south of Ban Nong and ne 紅 Ban Na Tum 33 33 kn1 south of Ban Nong (SAKAOAMI & NAKORNSRI ， 1987). A few coral specimens have been reported in Middle-Late Ordovician of Chiang Mai 釘 'ea (HAHN & SIEBENHUNER ， 1982). Corals associated with trilobites and brachiopods have been mentioned in 組 Upper Ordovician limestone of 官lO ng Pha Phum 紅 'ea (HAOEN ET AL. ， 1976).
Sponges
Sponges 釘 e important and widespread organisms of the presen トday seas ， with an estimated estimated 10 ，000 species alive today. Th ey were also abundant in the p錨 t. According to their their present systematics ， they include Stromatoporoidea. Calcareous Calcareous sponges 紅 'e common in many limestone exposures of 百lailand. Some authors authors consider 白紙Ch aetetidae (加出 e past assigned to the Tabulata: corals) and Solenopora (red (red alga in many old publications) 創泡 sponges (for 加S飽nce ，s田 R 田町G ， 2004). In百 1ai1 and ， Ch aetetidae have been described at m 組 y localities belonging to Devonian and Carboniferous. Solenopora ， as ciriginally defined ，is known only in the Ordovician and possibly the Lo wer Silurian. Silurian. Th e “ Solenopora" specimens found 泊Th ailand are apparently not authentic 50 HENRI Fo 町 AJNE ，SIROT SALY APONGSE AND V 線 AVUDH S U1百四百 O附
Solenopora. Solenopora. Th ey bel 'O ng t 'O ages y'O unger 血an Silurian; they have been f'O und at several Triassic Triassic l'O calities.τ 'h ey may actually be red algae; 出ey need t 'O be re-identified. Many kinds 'O f calcare 'O us sp 'O nges have been n'O ticed ，especially 泊 the study 'O f thin secti 'O ns fr 'O m limest 'O ne samples 'O f Th ailand. Th ey have been curs 'O rily menti 'O ned 血 diverse diverse publicati 'O ns ，with 'O ut detailed study. Th ey are abundant 血 s'O me limest 'O nes and because because 'O f 白紙， s'O me auth 'O rs have described “sp 'O nge limest 'O nes" ，f 'O r instance 泊 the Triassic Triassic 'O f Phatthalung 紅 ea (ADAC 凹 Ef AL. ， 1993). A sp 'O nge species (Cladocoropsis mirabilis) mirabilis) has been rep 'O rted at a few Jurassic limest 'O ne exp 'O s町 'es s'O uth 'O f Umphang (BEAUVAIS ， 1988). Several species 'O f Upper Permian sp 'O nges have been discriminated at a limest 'O ne l'O cality 'O f Phrae Pr'O vince in N 'O巾官 lailand and the limest 'O ne has been c'O nsidered “reefal" (SENOWBARI-DARYAN & INGAVAT- 抱 LMC 阻， 1993). 百 e sponges were ass 'O ciated with f 'O raminifera including Colaniella ，bry 'O z'O加 s，ga 柑 op 'O ds and echin 'O derms. Str 'O mat 'O p 'O roids are very abundant and diverse in the Dev 'O nian limest 'O ne 'O f the L 'O ei regi 'O n; 出ey are imp 'O rtant framew 'O rk builders 'O f the Dev 'O nian reefal s佐uc 伽res 'O f 血is regi 'O n. 百ley are dendr 'O id 'O r massive. Wh ere they 'O ccur ，出 ey pr 'O vide precise inf 'O rmati 'O n 'O n 血e age 'O f the limest 'O ne exp 'O sures (FONT A町 E Ef AL. ， 1990). Th el 'O cal abundance 'O f出e slender slender stick-like Amphip 'O ra suggests 白e presence 'O f dense “ 'Amphipora mead 'O ws". In In additi 'O n，s 位omat 'O p'O r'O ids have been menti 'O ned at several Ord 'O vician l'O calities 'O f Peninsular Peninsular Th ailand (WONGWANICH & BURRE'π ， 1983; BURRETT & STAIT ， 1985). S佐omat 'O p'O r'O ids have been c 'O nsidered t 'O be the main reef-building 'O rg 佃 isms in the Dev 'O nian 'O f Central Eur 'O pe.
Bryozoans
Bry 'O z'O ans are c'O mm 'O nf 'O ssils at many l'O calities. Th e fan-shaped Fenestellidae 創 e easy easy t 'O rec 'O gnize 泊 the field; 出eyappeared 泊 Middle Ord 'O vician and disappeared at 白e end 'O f 白e Permian. Carb 'O nifer 'O us and esp 田 ially Permian bry 'O z'O ans 'O f 百 ailand were actively studied between between 1964 and 1980. 百ley were c'Ol1 ected fr 'O m shale ，si1i ce 'O us rock ，calcare 'O us rocks and 'O ccasi 'O nally limest 'O ne ，especially 'O f Peninsular 官 lailand (f 'O r instance ， see SAKAGAMI ， 1968; 1968; Pπ'AKPAIVAN Ef AL. ， 1969). Bry 'O z'O ans 'O ccu 凶 ng 泊 calcare 'O us facies 紅 'ec 'O mm 'On1 y associated associated with brachi 'O p'O ds and crin 'O ids; they are less str 'O ngly ass 'O ciated with fusulinaceans (SAKAGAMI ， 1976).
Brachiopods Brachiopods
Brachi 'O p'O ds are mainly inltabitants 'O f 白e shall 'O w sea b'O tt 'O m.τ 'h ey 釘 'e imp 'O託釦 t ssils f'O ssils 'O f Th ailand ，but 出.ey have been c 'O llected m 'O re 'O ften 合'O m clastic r'O cks 出an fr 'O m limest 'O ne (see Fig. 8). Because 'O f 白紙， they are menti 'O ned 'On1 y briefly 加出is paper. Brachi 'O p'O d spines have been n 'O ticed in many limest 'O ne 出泊 secti 'O ns fr 'O m different Pale 'O z'O ic l'O calities. Jurassic Jurassic brachi 'O p'O ds are especially abundant in mar 匂 limest 'O nes and limest 'O nes 'O f Umph 佃 g 紅 ea (MEESOOK ， 1994). Lytt 'O niidae 釘 e an interes 出19 gr 'O up 'O f brachi 'O p'O ds that have been f'O und in the Upper Permi 佃'O fN 'O rth and East Th ailand ， but in shales cl 'O se t'O Upper Permian (Changhsingian) c 'O ntaining Palae φlsulina ，a fusuline indicative 'O f 由et 'O p 'O f the Permian. Brachi 'O p'O ds FOSS IL B IODIVERSITY IN THE Ll MESTO NES OF THA ILAND 5 1
Figul 巴 7 . Ko Si Chang. Fo ss ils vi sib l巴 al th es urfac e of a lillles lon 巴巴xposure ， but difficu lt to ex tr acl
Figu re 8. Nea r K hao Than in Saw i Di str ict of C humph on Pr ov in ce ，a long th 巴 sea shore ( 10 ・IO'25"N ， 99' IO' 53"E). Brachiopod s are easy to s巴巴 and coll ect in the shale and calca reou ss hal e 52 H EN RI FO NTA IN E，S IROT SA LY APO NGSE AN D V ARAVU DH SUTE ETHOR N
9a 9a
9b 9b
9c 9c
Figure s 9. Cora ls see n in th 巴fielcl atth 巴sllrfa c巴of lim es tone expo slll 巴 s. a: horn shap ecl solit ary cora l. b: branching co ral. c: ma ss iv巴 cora l;co rallit es at 巴 pr ismatic ， se pta ancl co lum 巴lI a are clearl y visibl 巴.
very very peculiar in sh ape wit h an 巴longate bilobate out lin e and an anterior incision have been fo und in shal巴 nort h of Nan in Nort h Thailand (MEESOOK & WANAPEERA ，19 83). They
b巴long to Permianella ，加 1 Upper P 巴rmian gen us known from Japa n to Ir an . This group of brachiopod s has been assig n巴d to Lyttonioidea si nce its first discovery in Afg hanistan. Middle Pelmian brachjopods hav 巴 b巴巴 n d iscovered in a calcareo us mudston 巴 south -so uthwest of of Phetchabun (YANAG IDA ，1 964). In Pe ninsul ar Thailand ， Early Permian brachiopods h ave be 巴n fo und in pebb ly mudstones at s巴vera l lo ca liti 巴s; th 巴yh av 巴 also b巴en co ll ected FOSSIL FOSSIL BIO Dl VERS IT Y IN THE Ll M ESTONES OF THAILAND 53 from sa ndstone 21 km north of C humphon. From a ge ner al point of view ，bra c hiopods h ave provid 巴d information on the ages of rock 巴xpo sure s of P巴ninsular Thailand. Th eir diver sity is low in thi s pa 1' t of th 巴 co untry and they sho w strong affiniti 巴s with Gondwana faunas ，indicating that th ey thriv ed in a coo lt 巴mp erat 巴巴nviTonm 巴nt (SHI & ARCHBOLD ，
1995). 1995). Middle Ca 1' bonif ，巴 rous bra chiopod s of Loei ar ea in Northeast Thail and have been collected collected from shal es and tuffac 巴ou ss hal es. Middl 巴 Ordovician brachiopod s hav 巴 be en desc ribed in calca 1'巴 ou s shale from Satun P 1' ovince. Brachiopod s from so me P巴rmian lime ston 巴 outcrops of Penin sular Thailand hav 巴 rec 巴IV 巴d some att 巴ntion with four paper s d巴vo ted to systematic studies ，fo 1' in sta nce the lime s tone of Khao Phrik in R atburi area (Y ANAGIDA ，1970 ; se 巴 th 巴 ov 巴1' Vl 巴w of WAT ERH O USE ， 1981). A 1' ich br ac hiopod fauna ha s b巴巴n described from Khao Tham N am Maholan in Loei Provinc e (YANAGIDA ，1967 ). Middle Permian and Low erD e vonian brachiopod s have bee n found in lim es ton 巴s of Thong Pha Phum ar 巴a (HAGEN & K EMPE R， 1976 )
10 c 10d
Figure s 10 . Abundant alga 巴 occasio nall y assoc iat ed with a few fu sulin es. a:B er ese llid a lgae (I ongitu c1 inal an c1 口an sve rse sec tion s) belong in g 10 the Mi c1 dl e Carboniferous lim es ton e of Lo ei Province. Sample
T7644 T7644 from Phu Bo Bil (17' 30' 14 "N ，1 0 10 46' 1 "E). b: A lgae (Dasyc lac1 acea ns;tran sv 引 se sec ti o ns) in in a lim esto ne belon ging 10 Ih e top of Middl 巴 Permian. Sample T626 0 from Kh ao Th am Yai 0 (I 6 56'41"N ，101 ・30'50"E ) b巴tween Lom sak a nd Na m Nao. c: Middl 巴 Ca rboniferou s a lgae of a lim esto n巴 cro ppin go ut 1 km so uth of Ban Saa t (1ア39'09"N ，lO nO '27"E; sa mpl 巴 T7 816) in Loe i
Pro v in ce. d:A lgae in a bedded lim es tone of th 巴 top of Khao Ma in th ea rea of Mahidol Univer sit y 0 ca mpu s (W09'00"N ，99 07' 40"E ; sa mpl e T8542). 54 H ENR I FONTAINE ， SIROT SALYAPONGSE AND V ARAVUD HS UTEET HORN
Fi gllr es 11 . Lim es ton es pac k ed w ith fusu1in es . Thi s typ巴 of 1im 巴stone ve ry ri ch in fllsu 1in es is co mm on in Thai 1and . a: Th 巴 fll su1in es in c1 ud e Ru zhence νites a nd indi ca tea Lo we rP e nnian (Ass 巴1i an) age . Thin sec tion from sa mp1 e D54 2 co ll 巴cted from Phll Sam Ph a (l T04'37"N ，101 ・59'57"E). b: Li l1l eston e ri ch in fu su1in es in c1udin g a grea t numb er of D lIlI baru/a sp巴cim ensa nd be10ng in g to th e top of
Mi c1c1 1e P巴rmi an (Upp 巴r Mi c1 ian ). T hin sec tion fro l1l sa l1l p1e T6261 co ll ec ted from Khao Tha l1l Yai (16・56'41"N ，101 ・30'50"E) betw 巴en Lom s ak an d Na m Nao . Se e FONTA INE ET AL . (2002) FOSSIL FOSSIL BIODIVERSITY IN THE LIMESTONES OF THAILAND 55
Mollusca Mollusca
Mollusca Mollusca presently live in a great variety of environments: sea ，streams ，lakes ， ponds and and even land. They are common fossils ，but 白ey 訂 'e more easily studied in shale than in limestone. limestone. Th ey have been mentioned at many limestone outcrops of Th ailand ， but often without without identification. Gastropods: Gastropods: Th ey occur at m 叩 y limestone localities ， but they are not easy to ex 仕act from this type of rock. Th ey occur ， for inst 佃 ce ， in the Jurassic of Klo Tho (BASSOULL 町， 1994) ， in the Triassic near the Cambodian border and in Uthai Th ani area (FONTA 町 EET L.， AL.， 1996) and the Triassic of Phatthalung area (Ao AC 田 ET AL. ，1993) ， in the Permian of Chon Daen area (CHONGLAKMANI & FONTAINE ，1992) ， in the Ordovician of Th ong Pha Phum (HAHN & SIEBENHUNER ，1982) ， in Ordovician black calcareous shale of Satun Province Province (PITAKPAIVAN ET AL. ，1969). An Early Ordovician Gondwana snail (Peelerophon) is is very common in the Shan- Th ai Block. Gastropods are commonly more difficult to use for for stratigraphy 白血 other fossils such as the foraminifera. Bivalves: Bivalves: Unbroken bivalves shells ， conjoined or disjointed ， are well-preserved at some localities. Th ey have been recognized as good index fossils ，especially for the Triassic and and the Jurassic ， but they have been collected commonly from rocks other th 佃 limestone.
Claraia Claraia and Halobia 紅 'e well known fossils ofthe Lower (Claraia) and the Upper (Halobia) Triassic. Triassic. In the Jurassic of Th ailand ， they are abundant and diverse; 39 bivalve species have have been identified ，from Mae Hong Son to Nakhon Si Th ammarat (MEESOOK ， 1994). In Mae Hong Son area ，20 species have been found at the same locality of Ban Pa Lan whereas whereas 17 species have been reported 3 km east of Ban Klo Tho. In Peninsular Th ailand ， a few Jurassic bivalves have been found at Chumphon River mouth in a calcareous sandstone. South South of Nakhon Si Th ammarat ， rare Jurassic bivalves were collected from interbedded sandstone sandstone and mudstone. Bivalves occur in some Jurassic limestones ， for instance in Mae Sot Sot area. Cephalopods: Th ese fossils are an important element of marine faunas since the Ordovician. Ordovician. Th ey 訂 e good index fossils ， but they are commonly difficult to collect 仕om limestone limestone even though they occur in this type of rocks. Nautiloids 釘 'e useful fossils throughout throughout the Ordovician carbonates of Peninsular ，westem and northwestem Th ailand. As early as 1958 ， they have been reported in southem Peninsular Thailand (KOBAYASHI ， 1958; 1958; PITAKPAIVAN ET AL. ， 1969; WONGWANICH & BURRE 甘， 1983). The Ordovician was a period of rapid and great differentiation for nautiloids. Ammonoids have been described at at many localities of Th ailand; they have been collected mainly from shale ，siltstone and calcareous calcareous shale or calcareous siltstone. Th ey w i1l be briefly mentioned in this paper because because some specimens have been found in calcareous rocks ，訂gi1l aceous limestone ， and exceptionally exceptionally pure limestone. Jurassic ammonites have been found in the Mae Sot area ， mainly mainly in calcareous shale (SATO ， 1961; BRAUN & JORDAN ，1976). An ammonite belonging to to Dumortieria has been collected from a Jurassic calcareous bed 3 km southeast of Kl o Tho (BASSOULLET ， 1994). Later on ，Jurassic ammonites have been more systematically collected collected from many localities of Mae Hong Son ，Mae Sot ，Umphang and Chumphon (Kh ao Lak) areas; they have been identified to generic level (MEESOOK ， 1994). Upper Permian Permian ammonoids have been described at Doi Pha Phlung in North Thailand while other Permian Permian ammonoids have been found in Loei and Muak Lek areas. Eight upper Lower to lower lower Middle Permian species of ammonoids have been reported in limestone at Kh ao 56 HE NRl FONTAINE ，SIROT SALYAPONGSE AND VARAVUDH Sm 官邸百O 剛
Nong Hoi ，a hill near Ban Hua Kr ok and 3 km nor 仕l-northeast of Amphoe Muak Le k (GLENIS 百 R ET AL. ， 1990; LIENGJARERN & ZHOU ， 2002). Carboniferous ammonoids have been reported 泊 Northwest (Ban Mae Lana) 佃 d Northeast (Loei) 官lailand (l SHIBASHI ET AL. ， 1997). Lower Carboniferous ammonoids were collected from limestone ofPeninsular 百lailand as early as 1899; later on ，出 ey were described (REED ， 1920). Belemnites ， cephalopods belonging to Mesozoic ， remain practically unknown in Thailand. “τ'h e reason why belemnites have not yet been found in the Jurassic of 官 lailand is 白紙 the 百団 Jurassic ranges 合om Toarcian to Early Bajocian 佃 d during this period belemnites belemnites were not abundant in Southeast Asia and Australasia" (MEESOOK ，1994 ， p. 163). 163).
Arthropods
Arthr opods ar 官加vertebrates of highly varied forms. 百 leir stratigraphical range is not homogeneous. It may be short to some extent; it may be long extending from Cambrian to to presen t. Th ey include groups very interesting for paleontology and stratigraphy.
Ostracods occ 町 at many limestone localities.τ 'h ese fossils 釘 'e minute bivalved crustaceans ，ranging 企om Lower Ordovician to present. 百ley ぽ e benthic faunas ，still common 加 the present-day seas. 官 ley 釘 e quite good indicators of the differing marine environments. 百ley have been rarely identified in 百lailand. A few ostracods have been mentioned mentioned in the Perr 凶an ofNan Pr ov 泊ce ，at Carboniferous -P ermian localities of Northwest Thailand ， at a probably Carboniferous locality north of Ban Th a Rua ， at a Lower Carboniferous Carboniferous locality of 百lO ng Pha Phum 紅 ea ， at a probably Late Devonian-Early Carboniferous Carboniferous locality east of Ban 官la Song Yang (H Al町& SIEBENHU 阻 R ， 1982). Th e assemblages 紅 e poor ，restricted to 1 or 2 genera. In many papers concerning other localities ， 出ey have been mentioned without identification in limestones of different ages. Near Chon Daenat 血e top of Kh ao Sak (westem end of Kh ao Sap Laeng) ，ostracodes 訂 e in abundance in in Lo wer Permian beds (FONT A 町 E ET AL. ， 1999). Well-preserved os 回， cods have been recently recently extracted 合om Permi 佃 limestones of the Loei-Phetchabun region. 百ley have been identified; they belong to seven genera (CHITNAR 町 ET AL. ， 2003). Trilobites Trilobites include good index fossils. Th ey r釦 ge from Cambrian to the end of the Permian and have been found at many localities of Th ailand. Th ey have been noticed 加 Permian 1加 estone. 百ley have been collected from Middle-Upper Carboniferous shales of
Loei Loei area and a new genus (Thaiaspis) was described 邸 early 酪 1961 by T. Kobayash i. Lo wer Carboniferous 凶 lobites have been repo 此ed 泊 Pha 凶lalung 紅 'ea (REED ， 1920). Early Devonian (probably Emsian) trilobites were collected from a 白le grained limestone 泊 Satun Satun Prov 泊ce (FOR' 百 Y ， 1989). A Si 加 ian trilobite has been collected 針。 m a calcareous mudstone northwest of Nam Som in Loei region (KOBAYASHI & SAKAGAMI ， 1989). Trilobites Trilobites have been described in Middle Or dovician bedded limestone of Satun Pr ovince ， in in Upper Cambrian sandstone of Tarutao Island (KOBA YAS 田&HAMADA ， 1984). Ins 目 ts 瑚 rted to colonize the land during the Devonian and perhaps earlier. For Carboniferous Carboniferous and Lo wer Permian continental deposits ，出 ey have been considered good 抑制igraphic markers and interesting for correlations between continents because 血ey could fly fly very far. In百 lailand ，出ey 紅 'e known as fossils in a black shale at Ban Kh uan Kun between Kr abi and Tr 卸 g. 官 ley 釘 'e common ，but 泊 a continen ta1 Mesozoic (Jurassic) rock. rock. In sects are teπ'es 凶 al organisms and 釘 'e unknown so far in the limestones of 百lailand. FOSSIL FOSSIL BIODIVERSITY IN THE LIMESTONES OF THAILAND 57
Echinoderms
Echinoderms Echinoderms are exclusively marine animals. Many of them 釘 'e attached to the sea bottom; bottom; others are f民e-moving an 出lalS. Urchins 訂 e armoured by an extemal skeleton box-like ， often of exceptional beauty ， but but easily broken by waves. Their fossils are generally reduced to small fragments and are practically practically impossible to study. However ， strong movable sp 凶es may be present on the outside outside of the skeleton; such spines 釘 'e common in the Middle Carboniferous of Loei Pr ovince. Th ey are well preserved ， but have not been studied ye t. Urchin spines have also been been mentioned in Triassic limestone near the Cambodian border (FONT A町 E ET AL. ， 1996). Crinoids Crinoids (or sea lilies) 紅 e very common in the limestones of Th ailand. Fragments of their their stems 釘 'e even the main constituent of some limestones ， then called “crinoidal limestones". limestones". Th ey have con 凶 buted 1釘 ge and important quantities of lime carbonate. From stratigraphical a stratigraphical point of view ，they 釘 'e not very interesting or e槌 Y to use.
Graptolites Graptolites
Graptolites Graptolites are commonly known in shales ， but occasionally occur in limestone. Graptolite Graptolite morphology was described in 1890 after etching undistorted specimens out of limestone.τ 'h ey ranged from Cambrian to Lower Carboniferous and reached their climax during during Ordovician and Silurian. 百ley have been mentioned at severallocalities of Th ailand.
Conodonts
Conodont remains are very small ，tooth-like and plate-like. Th ey c創 mot be seen in the field field with the naked eye. Th ey consist mos t1 y of calcium phosphate and they may be better preserved 出an other fossils. Th ey are very valuable for dating the rocks in which 血ey occur. occur. Th ey went through phases of rapid evolution and 訂 e good index fossils. Th ey 釘 e widespread widespread in layers ranging from Upper Cambrian to the end of the Triassic ，that is to say from 500 to 203 million ye 訂 s ago. In百 lailand ， conodonts have been identified or described at at many localities belonging to different geological intervals. Conodonts Conodonts have been extracted from limestones of several 紅 eas of Th ailand ，from the the peninsula to the north. Ordovician ，Silurian ，Devonian ，Carboniferous ， Permian and Triassic Triassic conodonts have been mentioned at many localities of Nor 白百lailand. 百ley belong to to many species; 22 Lower Devonian species have been identified at the same locality of ChiangM 泊 area (HAI 制&SIEBE Nl町 NER ， 1982; CAREY ET AL. ，1995). In Northeast 百lailand ， conodonts conodonts have been collected 企om the Upper Carboniferous-Lower Permian limestone of Kh ao 官lam Nam Maholan in Loei Province (100 ， 1974). In Northwest Th ailand ，Upper Silurian Silurian conodonts have been found in limestone outcropping between Tat Sador Waterfall and Ko Luong Waterfall ， west of 百loen (BURRE 甘 ET AL. ， 1986). Upper Devonian (Late Famennian) Famennian) conodonts have been obtained from thin limestone beds along the road from Mae Sariang to Ban Mae Sam Laep (LONO ， 1993). In In Kanchanaburi Province at two localities (6 km west of Si Sawat and between Bo Ngam and Sake) ， limestone has yielded a Middle Triassic (Anisian) conodont assemblage (也 MPER ET AL. ， 1976). In Th ong Pha Phum area ，a sequence containing conodonts is continuous continuous from the Upper Devonian into the Carboniferous (HAOEN & KEMPER ， 1976). 58 H 悶悶 Fo 悶 AD 司E，SIROT SALYA 問 NGSE AND V ARA VUDH Su 花町田聞
It It has recently yielded Upper Frasnian conodonts (SA VAGE & SARDSUD ， 2003). Upper Silurian Silurian and Lo wer Ordovician conodonts were also found 泊百lO ng Pha Phum 創明(HAGEN & KEMPER ， 1976). Northeast of Amphoe Sang Kl aburi ， 16 species of Toumaisian (Lοwer Carboniferous) Carboniferous) conodonts have been identified at the same loc a1i ty 仔弘 HN & SIEBENHUNER ， 1982). 1982). Conodonts Conodonts have been obtained from a limestone in Satun Pr ovince; they indicate an E 紅 ly Devonian age (LONG & BURREπ ， 1989). In Phatthalung 紅白， Triassic conodonts were were recovered 企'om several localities; 出ey indicated Early to Late Triassic ages (IGO EI AL. ， 1989; AMPORNMAHA ， 1995).
Fishes Fishes
Fragments Fragments of fishes skeletons have been noticed at severallocalities of Th ailand. Th ey have have been studied only occasionally; for instance ，some fragments have been recently observed observed in Permian limestone of the Phetchabun area ， but not studied. In In Lo ei Pr ovince ，a single tooth (bradyodont) has been found in Lower Carboniferous limestone limestone of Pha Chom Nang on the side of the road 合om Chiang Kh an to Pak Chom (INGAVAT & J，制 VIER ，198 1). Two teeth (bradyodont and sh 釘 k) have been extracted 仕om the the Middle-Upper Carboniferous (considered Permian 泊 the past) limestone of B 佃 Na CharoenσANVIER ，198 1). At road cuttings (near km 27) along the road south of Mae Sariang to Ban Mae S創 n Laep on the Burmese border ， abundant vertebrate and conodont assemblages have been found found in bluish-grey Devonian (Emsian and Famennian) bedded limestones. Th ey are dominated dominated by small chondrichthyan (sh 釘 k) remains ， but include also other fish remains; 白ey contain cosmopolitan and endemic elements. 百ley suggest a proximity to the wes 旬m m 釘 g血 of east Gondwana (BURRETT ， 1989; LONG & INGAVAT ， 1989; LONG ， 1990 and 1993). 1993).
Other Vertebrates
In In 1988 ，an ichthyosaur was discovered by C. Chonglakmani in a weathered dolomitic limestone limestone at Kh ao Th ong 14 km north of Pha 曲 alung. It w 錨 associated wi 出 poorly preserved preserved ammonites. 百le limestone belongs to Lower Triassic. Prim itive ichthyosaurs have have been rarely found in the world. Th e specimen of Th ailand has been described as a new genus 佃 da new species: Thaisaurus chonglakmanii (MAz町 EI AL. ， 1991). Li mestone has not yielded so far other vertebrates than fishes and ichthyosaurs ， but diverse diverse vertebrates 釘 'e known at several localities of the continen ta1 Mesozoic Kh orat Gr oup of Northeast Th ailand and Dicynodon has been found in Upper Permian sandstone near near Luang Prabang (Laos) ，a simil 釘 Permian terres 凶al sequence occ 町出19 northwest of Loei Loei in Nor 仕leastτ 'h ailand.
STRA TIGRAPHIC RANGES OF 百IE FOSSILS
Many groups of plants and animals lived only for a short time on 出e geologic t加 e scale.τ 'h eir evolutionary development has had an impo 此ant significance in 白 e geological FOSSIL BIODIVERSπY IN THE LIMESTONES OF THA 乱AND 59
Table Table 3. Biozonation of the Middle-Upper Carboniferous of W 叩 g Saphung 紅白
Huai Huai Kap Tin Huai Luang B釦 Pha Noi 百lam Nam M 油olan
Triticites Triticites oz 仰 lai/ Paraschwagerina Paraschwagerina yanagidai yanagidai Protriticites Protriticites tethydis Fusulina Fusulina pulchella Hemifusulina Hemifusulina thaiensis Beedeina Beedeina paradisten ω Profusulinella Profusulinella prisca tlmanica tlmanica Profusulinella Profusulinella parva
Table Table 4. Biozonation of the Permian north of Saraburi
Kh ao Pr ong Pr ab Kh ao Kh ao Kh ao Irn ot Colania Colania douv il/ ei Neoschwagerina Neoschwagerina haydeni Af ghanella schencki Presumatrina Presumatrina schellwieni Presumatrina schellwieni Neoschwagerina Neoschwagerina simplex Neoschwagerina simplex Maklaya Maklaya sethaputi Maklaya Maklaya pamirica Maklaya Maklaya saraburiensis
Misellina Misellina COl 許agaspira Misellina Misellina otai/M. cf. termieri Monodiexodina Monodiexodina ship ωni history history of the world. Th ey provide keys to decipher the history of the deposition of the sediments. sediments. Th ere 訂 'e very good stratigraphic markers at 血， e genus and even the species levels levels because 出ey indicate immediately the age of the rock in which 由ey are included. Foraminifera ，brachiopods ，住 ilobites ， conodonts and others are well known for tha t. In addition ，some organisms could live 泊 different environments and their fossils therefore occ 町泊m 加 y rock outcrops over the world.τ 'h ey allow precise biozonations of sed 泊lentary sequences sequences (see Tables 3 and 4 for examples) as well as interbasinal to inte 町 egional correlations. correlations. Some important events ， of different orig 恒久 have punc 加 ated the org 佃 isms evolution. evolution. Th e initial flowering of animal life on 血ee 紅白 occu 町'ed during the Cambrian ，a period period moderately known in Th ailand because rocks of 白is age are exposed only in very small small areas. Fossils have been found mainly in clastic rocks. 60 60 HENRl FONTAINE ，SIROT SAL Y APONGSE AND V ARA VUDH SUTEETHORN
Ordovician Ordovician and Silurian limestones have provided fossils at severallocalities of Th ailand; see see the paragraph on “Th e Fossils". In Peninsular Th ailand ，biozonation is more detailed. Limestones Limestones have been 児島町ed to Tremadoc and Ar enig in Tarutao Island. On the mainland in in Satun Province ，a succession of limestones and clastic rocks ranges from Middle Ordovician Ordovician (Caradoc) to Silurian (WONOWANICH ET AL. ， 1990); it is conformably overlain by a Devonian sequence of carbonates and clastic rocks. Fossils are diverse and belong to trilobites ，graptolites ，nautiloids ，brachiopods ，gastropods and ostracodes. A thick Siluro- Devonian Devonian limestone sequence has been described west of Thoen in Northwest Th ailand (BURRETT ET AL.， 1986). In In the Loei region ， Devonian limestone is rich in diverse fossils ，indicating Early and Middle Middle Devonian ages and probably extending to the lower p訂 t (Frasnian) of the Upper Devonian. Devonian. A sharp change in the environment after the Frasnian led to the disappearance of of limestone deposition and the extinction of diverse faunas ， probably because of the Kellwasser Kellwasser Crisis ，a prominent perturbation observed in many countries. In West Th ailand ， shaly shaly limestone has yielded Frasnian conodonts ， but the search for Famennian conodonts has has been so far without resu 1t s (SA VAOE & SARDSUD ， 2003). The history of the Carboniferous of Loei region is complex ， with sharp changes in the environments. environments. The Toumaisian consists of chert ，shale ，sandstone and conglomerate. Chert has has yielded Famennian to Toumaisian radiolarians. During the Visean ， limestone was deposited deposited with its diverse and rich marine fossil assemblages. After the Visean ， the water disappeared disappeared because of a lowering of sea leve l. Continental plants developed and their imprints imprints presently occur in shale and siltstone belonging to the boundary between Lower
and and Middle Carboniferous. Coal is locally present ， but not in very 1紅 ge quantity. Gypsum occurs occurs in other areas of Loei region. During the late part of the Bashkirian ， sea invaded again again the Loei region and was inhabited by diverse floras and faunas ，until the end of the Carboniferous Carboniferous (FONTAINE ET AL. ， 1995 and in preparation). In Moscow Basin ，a sharp sea-
level level drop has also been obse 円 ed during the Bashkirian and has been considered a response to to Gondwanan glaciation (ALEKSEEV ET AL. ， 2004). Biozonation ranging from Moscovian to to Gshelian (100 ， 1972 and other recent publications). The Carboniferous strata of Loei region region do not build prominent and sharp topography as the Permian limestone of the same reg lO n. In In Northwest Th ailand ， the Carboniferous is complete. It is largely similar to that of the the Loei region. Th e Permian of the world is divided into about 30 fusulinacean ， arnmonoid or conodont biozones biozones (JIN ET A L.， 1994); 出e division of the Carboniferous is similar. In Th ailand ， fusulinaceans fusulinaceans have been studied more extensively than ammonoids and conodonts. Other fossils fossils were restricted to particular environments; they cannot be used for global correlations. Corals ， for instance ， needed w 訂 m water to thrive; they are useful only for the p紅白 of the world world which were not too far from the equator at the studied period. Permian limestone is is widespread in Thailand and belongs to different parts of the Permian. It is locally packed with with fusulines; see figs. 1la-b as well as the following table 4.Biozonation ranging from the the upper part of Lower Permian to Lower Midian (TORIYAMA ， 1984). Kh ao Khao and Khao Imot are two hills ne 訂Kh ao Prong Prab. Th e3 hills are about 20 km from Saraburi ， and and belong to a series of hills with strong karstic topography ， called the “Kh ao Kh ad Formation" Formation" and extending from Lopburi to Pak Chong. FOSSIL FOSSIL BIO Dl VERSITY IN THE LIMESTONES OF THAILAND 61
Some fossils are very simple and do not receive the deta i1 ed study they deserve. Their study study is more difficult; it must be based on oriented sections and on well preserved specimens. specimens. If the differences between the specimens ar 泡 not noticed ， the foss i1 s are considered to to belong to very long periods. An exception to that is a very simple foss i1 known at m 佃 y Permian Permian loc a1 ities of τbailand; it is easy to recognize in different types of sections. Called Sphairionia ，it has been described for the first time only in 1989 (NGUYEN ， 1989). It is a globular globular form ， small in size (0.5 to 1. 5 mm in diameter) ，consisting of a single chamber ， with with bottleneck-shaped openings. At fI rst known only in Southeast Asia ，it has been discovered discovered in other p釘 ts of the world and has a wide geographic di 紺 ibution. It is presently known at sever a1 localities of Cambodia ，Th ailand ，Malaysia ，Parnir ，Transcaucasia and Turkey; Turkey; it has been considered a marker of the Lower Midian ， belonging to the upper part of of Middle Permian (PRON 町 A， 1996). 百le oldest foss i1 s of a group 紅 'e commonly more simple th 組曲e younger ones; evolution evolution leads to more complex s佐uct 町田. However ，empiric a1 data do not a1 ways follow 出 is principle. In 1912 (C R Acad Sc. 154: 1548-1550) ， Deprat studied the fusulinaceans of of Vietnam ， Laos and Yunnan ， and found some “primitive" fusulines. He established the new Genus PaLae φ， Isu Li na and assigned it to Early Carboniferous because it looked primitive. Now ，we know that this genus is the youngest ， belonging to 由e top of the Permian and corresponding corresponding to a period of decline of the fusulines before the great mass extinction of the the end of the Permian. 百le frrst Triassic fossils found in 百lailand were discovered 8 km southeast of Chiang Rai; Rai; they were identi fI ed as ear 匂 as 1923. In 1952 ，Triassic foss i1 s were found in sh a1 e in Lampang Prov 加ce (PITAKPAIVAN ，1955). 官lI s new discovery stimulated new research in that 釘 ea; highly fossiliferous limestone was found. After 1960 ，Triassic sediments ，largely marine ，became evident in m 釦 y parts of Th ailand. Th ey have yielded diverse faunas (for 釘 n泊ぜera ，corals ，brachiopods ，biv a1 ves and ammonites) associated with a1 gae; 曲ey range range from Lower Triassic to Carnian ，loc a1 1y to Lower Norian.τbey have been divided into into sever a1 formations. Carbonate facies 釘巴 common.
Th e Jurassic is divided into formations ranging from Upper ，Hettangian to C a1 10vian (top (top of Mi ddle Jurassic) ， with the Toarcian and the A a1 enian more widespread 白an the other other parts of the Jurassic (MEESOOK ， 1994). Th e occuηence of Upper Jurassic remains very very doubtful and ， if true ， has a strongly limited geographic dis 佐ibution.
PALEOGEOGRAPHY
Since Since the 1970s ， the geologic a1 history of Southeast Asia has been reviewed in 曲e ∞ ntext of the plate-tectonic 出eory (METCAL 回 2002). Th ailand is presently considered to be be made up of an amalgamation of at least 2 main blocks derived from Gondwan a1 and at different different periods: the Shan- Th ai Block derived in the late Early Perrnian and the In dochina Block Block derived in the Devonian. The position of the boundary between these two blocks is still still the subject of controversy. 百 e Shan- Th ai Block has been supposed to be part of 白e Gondwan a1 and until its d 負担 g at the end of the Early Perrnian. It consists of Peninsular 百lai1 and and extends to 血e north 泊to My 佃 m 紅 (and even to 出e Baoshan Block of west Yunnan 泊 southwest
Ch 泊a; see for instance W ANG Ef AL. ，2001) 佃 d to the south into Peninsular M a1 aysia. Its 62 HENRI FONT AINE ，SIROT SALY APONGSE AND V ARA VUDH SUTEETHORN faunas faunas are very different from those of the Indochina Block especially those of the Carboniferous Carboniferous and 恥 Permian. Cold water environments have been found in the Lower Permian Permian and are well documented (see below). 百le “Shan 一司 lai" name was proposed by BUNOPAS & VELLA (1978) as the Shan_: 百lai Craton which extended 血rough a large part of of Th ailand. Th ey described the north-south trending geological structures of Thailand and tried tried to incorporate them into a new plate tectonics vision of Southeast Asia. Later on ，it has has been suggested to replace “ Shan-Thai" by the n創 ne “Sibumasu" ，indicating more clearly clearly the geographic extension of the bloc k. Th e Indochina Bl ock (= Indosinian Craton of BUNOPAS & VELLA ， 1978) includes the eastem eastem part of Thailand and extends to Cambodia ，Laos ，Vietnam and the eastem part of Peninsular Peninsular Malaysia. It s faunas indicate warm water environments throughout the Late
Paleozoic. Paleozoic. Limestone is widespread in the Carboniferous and the Permian exposur 邸l ofthe In dochina Block. It is unknown in the Middle-Upper Carboniferous as well as the lowest part part of the Permian of the Shan ー官lai Block. In In the past ， the Permian limestone of Th ailand was called the “Ratburi Limestone" all over over the countη(BROWN ET AL. ，1951). Th e term “Ratburi Li mestone" is presently restricted to to limestone exposures of Peninsular and West Th ailand. Limestone of the other p紅白 of Thailand Thailand is called “Saraburi Limestone" (BUNOPAS ， 1981). The differences between these two limestones 訂 e easy to observe. Fusulinaceans may be in great number in the two limestones ， but they consist of one or two species in the Ratburi limestone while they are much more diverse in the Saraburi Limestone. Corals occur in the two limestones ， but their diversity diversity is much lower in the peninsula. Among bryozoans ，Fistul 伊ora is quite common in in Peninsular Th ailand where it is represented by 12 species ，some of which 紅 e known in Timor (SAKAGAMI ， 1976). Permian corals of Peninsular Th ailand are quite different from those those of central and eastem Thailand (FONTA 町 E & SALYAPONGSE ，200 1). Limestone is absent absent from the lower part of the Lower Permian in the peninsula; accordingly ，it is impossible impossible to find fusulinaceans and corals (in particular Kepingophyllidae) in the lowest part part of the Permian. In addition to that ， pebbly mudstones are extensive in this part of the Permian Permian in the pen 凶sula. They contain dropstones supposed to have been rafted by marine ice. ice. The largest block is a granitic boulder ，approximately 1m in size ， found in Phuke t. A trondhjemite boulder of northwest Peninsular Malaysia has yielded a Precambrian age (STAU 押田&MANTAJ 汀， 1981; TANTIWANIT ET AL. ， 1983). Th e pebbly mudstones have been been considered glacial deposits. These beds contain cool-water fauna (WATERHOUSE ， 1982). 1982). From a stratigraphical point of view ， they are below the Ratburi Li mestone. In Gondwanaland ，a severe glaciation began at the end of Lower Carboniferous and reached its its maximum near the Carboniferous-Permian boundary. In Thailand ，tiaces of glaciation have have been observed mainly at the base of the Permian. The upper part of the Carboniferous has has not been clearly evidenced in Peninsular and West Th ailand. Th e Ratburi Limestone co 町田ponds to an improving climate. At the foot of a limestone hill in Chom Bung 紅 ea ， corals corals are few and display growth bands ， whereas corals at the top of the hill are very common and devoid of bands (FONTAI 阻&JUNGYUSUK ， 1997). Th e presence of growth bands bands appears to indicate a seasonal climate with cold and warm seasons in an older period of of the Permian. The absence of grow 白 bands suggests regul 紅 growth in a continuously warm climate at a younger time. Th e Shan- Th ai climate became w 紅 mer after the rifting of of this block from Gondwanaland and its rapid drifting northward. Provincialism is strongly indicated indicated by a small foraminifera ，Shanita ， which has been reported at several upper FOSSIL FOSSIL BIODIVERSITY 別 THE LIMESTONES OF THAILAND 63
Middle Middle to lower Upper Permian localities of Peninsular Th ailand 組 d is known to the north 泊 eastem My 佃 mar up to westem Yunnan. Shanita is unknown so f，訂泊 the In dochina Block. Block. Up to now ，Eopolydiexodina ，a fusulinid ， has been reported only from Kanchanaburi Pr ovince. Neoschwagerina ，Colania ， Verbeekina and Lepidolina are fusulinid genera common in the In dochina Block ， but apparently absent from the Shan- Th ai Bloc k. Corals such such as Ipciphyllum and Pseudohuangia ， widespread on the In dochina Block ，紅'e unknown in in the Shan- Th ai Block. The Permian of Australia is extensive; it locally offers complete m 紅白e sequences. It does not contain fusulinids or compound corals (Rugosa) ， but only smaller smaller foraminifera (CRESP 町， 1958) and solitary Rugosa. During Middle and Upper Permian ， the Shan"': 百lai Block is already far from Gondwanaland. In In Northwest Th ailand ， the discovery of widespread fossiliferous Carboniferous and Permian Permian limestones has been unexpec 制 (FO 附 AINE Ef AL. ， 1993); in the past ，a few fossil identifications identifications had been mentioned ， but without paleogeographic conclusion. Th is pぽ t of 百四land was believed to belong to the Shan ー官lai Block and its geology had to be compared to 白紙 of Peninsular Th ailand. Th e newly discovered faunas were diverse. 百ley showed similarities similarities with the faunas of the In dochina Block. Th ey indicated several horizons of the Carboniferous Carboniferous and of the Permian. Later on ，fossils were collected and described from localities localities north of Chiang Dao ，組d they 釘 'e again quite different from those reported from the the Shan- Th ai Block (UENO & 100 ， 1997). A conclusion of all the paleontological studies C但ried out in Northwest 百lailand appe 訂'S very clear: limestone has been deposited almost continuously continuously from Lower Carboniferous to the end of 出e Permian. 百lI s contrasts with Peninsular Peninsular and West Th ailand ，where Middle and Upper Carboniferous limestone remains unknown as well as Early Permian limestone. Ordovician Ordovician fossils (nautiloids ，gastropods ，brachiopods ，stromatoporoids) found in τbailand have suggested that the Shan- Th ai Block was 叫 acent to Aus 凶 ia because of close close faunal similarities ， also noticed for fossils (trilobites) found in the Upper Cambrian of of 官lailand (BURRETT & STAIT ， 1985). Lower-Middle Ordovician limestone is known in Peninsular Peninsular Th ailand. Upper Ordovician limestone has !U so been mentioned at few localities ， but but without strong evidence of 白is age; it is possibly absent as it is in Australia (METCAL 回， 1992). 1992). Early Devonian conodonts of Satun Pr ovince of Peninsular 百lailand include species on1 y known in Australia and South China (LONO & BURRETT ， 1975). Comparison of the Paleozoic Paleozoic paleontology and stratigraphy of the Shan- Th ai Block with those of Australia shows remarkable similarities; some fossil genera have been fou i1d only in Shan- Th ai Block Block and Australia. Shan- Th ai and Australia were still in close proximity at the end of the the Devonian (BUR 阻甘Ef AL. ， 1990).
EXT 町 CTIONS
Th e history of life on the e紅白 has been puncωated ， at a global scale ，by many 叩 isodes of rapid extinction and periods of more or less r叩id ori 伊lation of new forms. Extinction Extinction is a fundamental phenomenon of nature ， involving the loss of entire groups. Some periods saw enormous radiation of animal diversity ， but at other times ，deterioration of of conditions led to catastrophic extinctions. Biological recovery has sometimes been slow. Corals Corals are poorly known in the Lower Triassic ， during a period of five million years. After the the important crisis of the end of the Pe ロnian (see below) ， the diversity of the faunas was 64 HE NRl Fo 悶 AI 阻. SIROT SAL Y APONGSE AND V ARA VUDH SUTEETHO 剛
rest 'O red by 'O nly ab 'O ut 5m i1l i'O n years later ， app 紅 'ently fr 'O m refuges which sustained panchr 'O nic 'O rganisms as well as L 但釘us taxa. Mi crobial mats ， m 'O stly 'O f bacteria and algae ，c 'O mm 'O nly bec 'O me abundant 'O n the sea bed after an extincti 'O n. Th ey c'O uld 白rive better better with 'O ut gr ，但泊 g pressure fr 'O m multicellular 'O rganisms (f 'O r instance see SHEEHAN & IIAR 即 S， 2004). S住'O mat 'O lites are abundant at s'O me 百凶l'O calities c'O rresp 'O nd 加gt 'O the end 'O f the Permian and the beginning 'O f the Triassic. Data ab 'O ut f 'O ssils and inf 'O rmati 'O n 'O n m 'O dem ec 'O systems can lead t 'O a better understanding 'O f extincti 'O ns and their c 'O nsequences. A well-kn 'O wn mass extincti 'O n 'O ccurred at the end 'O f the Cretace 'O us (65 milli 'O n ye ぽ S ag 'O) in b'O血 terres 出 al and marine envir 'O nments ， with the expiry 'O f the din 'O saurs 'O n1 佃 d and the disappearance 'O f the amm 'O nites 泊血e sea. In Thai1 and ，there is n 'O marine Cretaceous and n 'O marine Tertiary dep 'O sits; the marine bi 'O tic crisis cann 'O t be 'O bserved in 血 .e field. At 出 e Triassic-Jurassic b'O undary ，203 m i1l i'O n ye 紅 s ag 'O， a widespread m 邸 s extincti 'O n
'O ccurred als 'O泊 m 創担e 佃 d terrestrial envir 'O nments. Thi s catas 住ophic event is n'O t easy t 'O
'O bserve in detail in Th ailand because marine Jurassic sediments 紅 'e restricted t 'O 'O nly a few 釘 e部'O fwestem 百lai1 and. 百le transiti 'O n 合'O m Triassic t 'O Jurassic is n'O tc 'O mm 'O nly exp 'O sed; but but the lists 'O f the f'O ssils 合om different l'O calities indicate str 'O ng changes 合om 'O ne peri 'O d t 'O the 'O ther peri 'O d. A mass extincti 'O n ended 'O f 血 e Permian ab 'O ut 250 milli 'O n ye 訂 s ag 'O， which w 部 pr 'O bably 出 em 'O st severe 'O f all times and c 'O nsisted 'O f several phases. It elinlinated alm 'O st 90 percent 'O f all species. It caused m 'O re ravages in marine envir 'O nments and especially af おcted the faunas 'O f the shelf seas. 百le marine envir 'O nments suffered bad changes 泊 出 .eir ec 'O systems ， with a large dr 'O p 泊 sea level ， higher salinity ，imp 'O rtant accumulati 'O ns 'O f 'O rganic matter during the Late Permian and an 'O xic c'O nditi 'O ns. Because the Triassic and Permian are well represented in 百lailand ，出is dramatic event is e剖 Yt 'O 'O bserve. It sp 釘med a few m i1l i'O n years. It was n'O ta simple catas 住ophe ， but at least ad 'O uble catas 位ophe which can can be studied 泊 τ'h ailand ne 紅 the Camb 'O dian b'O rder 'O r 泊血en 'O rthem part 'O f the c'O un 町. At the end 'O f the Middle Permian (end 'O f Midian 'O r Capi 伽 ian) ，血e large fusulinids fusulinids and many massive Rug 'O se c 'O rals died 'O ut because 'O f their vulnerab 出 ty t 'O h 'O stile envir 'O nmen ta1 changes. 百le skele ta1 wall 'O f the 1訂 'ge fusulinids is peculi 紅; it is called called a keri 'O theca. Lepidolina ， the last 1釘 ge fusulinid ，displayed abn 'O rmal gr 'O w 出 s bef 'O re its its extincti 'O n (FONTA 町 E EI AL. ，2002 ， p l. 8 ，fig. D). 官lI s is 血 e frrst crisis very easy t 'O 'O bserve in 曲e field. 百le rest 'O f the fusulinids and the c 'O rals disappeared 'O nly at the end 'O f the Permian; these last fusulinids were small in size ，with 'O ut keri 'O theca.τ 'h e 佐il 'O bites are are alm 'O st unkn 'O wn 面白e Upper Permian 'O f 百lailand; they disapperu 芭d entirely fr 'O m e紅白紙the end 'O f the Permian. 百lis is the sec 'O nd crisis which has been very imp 'O rtant ， wi 血 the disappearance 'O f many 'O ther gr 'O ups. S'O me 'O ther animal gr 'O ups survived ， such as
白.e c'O n'O d'O nts which disappe 紅 'ed 'O nly at the end 'O f the Triassic. Other Other extincti 'O ns were less spectacul 眠 alth 'O ugh 血 .ey were important and n 'O ticeable m 白e faunal successions of the Pale 'O zoic. Fr 'O m the base of the Dev 'O nian (410 Ma) t'O血 e end 'O f the Permian (250 Ma) ，5 important extinctions 'O ccurred. Th e base 'O f the Dev 'O nian is is marked by the disappe 紅 ance 'O f the main Silurian faunas. During Upper Devonian at 出e
end 'O f the Frasnian (370 Ma; Kellwasser Crisis) ，many 'O rg 佃 isms living in shall 'O w seas such such as corals or str 'O mat 'O p'O r'O ids disappeared; this fact is easily visible 泊 L 'O ei Pr ovince. Th e end 'O fL 'O wer Carb'Onifer~)U s (320 Ma) is marked by rapid changes 'O f faun 邸. At the end 'O f Lo wer Permian ， changes in 出 ec 'O mp 'O sition of the faunas are visible. FOSSIL BIODIVERSITY 町 THE LIMESTONES OF THAILAND 65
Th e number of extinctions and appe 訂佃ces of species ，as seen 泊 many sedimentary rock sequences ， is commonly not spectacul 低 Extinctions are usually slow ，but continuous. 官lI s fact has allowed us to establish stratigraphical scales ，which become more and more precise after detailed paleontological studies.
ACKNOWLEDGMENTS
Th e authors are deeply indebted to Dr Sompoad Srikosamatara ，Mahidol University ， for his very interesting suggestions. Th ey have also a great pleas 町 'e to acknowledge the S甘ong support and 白e encouragements of Mr Somsak Potisat ，Director General of 血e Dep 釘 tment of Mineral Resources as well as the help of the Di rectors of the Geological Survey of Th ailand and 出e geologists of 由民 organization. Dr Prinya Pu tthapiban ，Mahidol University ，gave a chance to visit fossiliferous localities of Kanchanaburi Pr ovince. Th e authors express their sincere thanks to Dr Warren Y. Brockelman ，editor of the Natural History Bulletin of the Siam Society ，for his continuous interest and his kind and strong help in editing.
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