The Permian of Timor: stratigraphy, palaeontology and palaeogeography

Journal of Asian Earth Sciences 20 (2002) 719±774

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The Permian of Timor: stratigraphy, palaeontology and palaeogeography T.R. Charlton a,*, A.J. Barber b, R.A. Harris c, S.T. Barkham d, P.R. Bird e, N.W. Archbold f, N.J. Morris g, R.S. Nicoll h, H.G. Owen g, R.M. Owens i, J.E. Sorauf j, P.D. Taylor g, G.D. Webster k, J.E. Whittaker g b

a Ridge House, 1 Saint Omer Ridge, Guildford, Surrey GU1 2DD, UK Department of Geology, Geological Research in Southeast Asia, Royal Holloway, University of London, Egham, Surrey TW20 OEX, UK c Department of Geology, Brigham Young University, Provo, UT 84602-4606, USA d Enterprise Oil plc, Grand Buildings, Trafalgar Square, London WC2N 5EJ, UK e Premier Oil plc, 23 Lower Belgrave Street, London SW1W 0NR, UK f School of Ecology and Environment, Deakin University, Rusden Campus, Clayton 3168, Victoria, Australia g Department of Palaeontology, The Natural History Museum, London SW7 5BD, UK h Department of Geology, Australian National University, Canberra, ACT 0200, Australia i Department of Geology, National Museum of Wales, Cardiff CF10 3NP, UK j Department of Geological Sciences, State University of New York, P.O. Box 6000, Binghamton, NY 13902-6000, USA k Department of Geology, Washington State University, Pullman, WA 99164-2812, USA

Received 21 September 2001; accepted 6 December 2001

Abstract The Permian of Timor in the Lesser Sunda Islands has attracted the attention of palaeontologists since the middle of the nineteenth century because of the richness, diversity and excellent state of preservation of its fauna. These abundant fossil data have been compiled and updated for the present account. The Permian rocks of Timor were deposited on the northern margin of Australia. At the present time the northern margin of Australia, in the region of Timor, is involved in a continent±arc collision, where Australia is colliding with the Banda Arcs. As a result of this collision, Permian rocks of the Australian margin have been disrupted by folding and faulting with the generation of mud-matrix meÂlange, and uplifted to form part of the island of Timor. Due to this tectonic disruption, it has proved dif®cult to establish a reliable stratigraphy for the Permian units on Timor, especially as the classic fossil collections were obtained largely from the meÂlange or purchased from the local people, and do not have adequate stratigraphic control. Detailed systematic, structural, stratigraphic and sedimentological studies since the 1960s have provided a ®rmer stratigraphic and palaeogeographic background for reconsideration of the signi®cance of the classic fossil collections. Permian rocks on Timor belong either to a volcanic-carbonate sequence (Maubisse Formation), or to a clastic sequence (Atahoc and Cribas formations) in which volcanics are less prominent. The Permian sequences were deposited on Australian continental basement which was undergoing extension with spasmodic volcanic activity. Carbonates of the Maubisse Formation were deposited on horst blocks and volcanic edi®ces, while clastic sediments of the Atahoc and Cribas formations were deposited in grabens. The clastic sediments are predominantly ®ne-grained, derived from a distant siliciclastic source, and are interbedded with sediments derived from the volcanics and carbonates of adjacent horst blocks. Bottom conditions in the graben were often anoxic. In the present account, events on Timor during the Permian are related to the regional tectonic context, with the northward movement of Australia leading to the amelioration of the climate from sub-glacial to sub-tropical, together with the separation of crustal blocks from the northern Australian margin to form the Meso-Tethys. q 2002 Elsevier Science Ltd. All rights reserved. Keywords: Continent±arc collision; Permian rocks; Timor

1. Introduction The Permian rocks of Timor have yielded the richest Palaeozoic marine fossil fauna in the world, this richness is * Corresponding author. E-mail address: [email protected] (T.R. Charlton).

apparent both in the number of species recorded, and in terms of numbers of individual specimens. Description of the crinoid Timorocidaris sphaeracantha, for instance, was based on a collection of 110,000 specimens (Wanner, 1940a). For this study we have compiled a database of over 1000 species from the Permian of Timor (see Appendix A). Timor also occupied an important palaeogeographical

1367-9120/02/$ - see front matter q 2002 Elsevier Science Ltd. All rights reserved. PII: S 1367-912 0(02)00018-4

720 T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774 Fig. 1. The geographic and tectonic situation of Timor. Horizontal ruled area is the forearc ridge, composed largely of Australian continental margin sediments (including Permian) forming a foreland-fold-andthrust belt; the upper parts of this belt, including Timor, which appear as emergent islands, are in black; toothed lines are thrusts; black triangles are volcanoes of the Banda volcanic arc (from Simundjantuk and Barber, 1996; after Hamilton, 1979). The Ashmore and Sahul Platforms on the Australian NW Shelf are areas of shallow basement

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Fig. 2. Location map for the Permian of Timor showing the principal outcrop areas and selected metamorphic massifs. Smaller outcrops of Permian rocks occur widely within the Bobonaro MeÂlange Complex, notably in the Amarassi, Kapan and Basleo areas.

position during the Permian, being situated between Gondwanan Australia and terranes of present-day mainland Southeast Asia. At the beginning of the Permian, Gondwana was recovering from the Permo-Carboniferous glaciation. Throughout the Permian the Timor±northern Australia region was undergoing active extension, associated with the separation of the Sibumasu Terrane of Southeast Asia from northern Gondwanaland (Metcalfe, 1996). Forming the northern edge of post-breakup Gondwana, Permian Timor provides an important palaeogeographical link between Southeast Asia and Australia (Charlton, 2001). The island of Timor, lying 300 km from the northwest coast of Australia, forms part of the Indonesian archipelago (Fig. 1). The island is separated from the continental shelf of Australia by the 2000 m deep Timor Trough, which marks the deformation front of the Banda island arc subduction and collision system. Timor and adjacent islands therefore form part of the Banda arc±continent collision zone, where an island arc of Asian af®nity is in the process of colliding with the formerly passive continental margin of Australia. Permian rocks in Timor are exposed as thrust slices in a fold and thrust belt in which the more distal parts of the Australian continental margin have been thrust southward over the more proximal shelf (Charlton et al., 1991; Harris, 1991). Permian rocks are also commonly found as isolated blocks within shales which have been mobilised to form a claymatrix meÂlange (the Bobonaro Scaly Clay, e.g. Audley-

Charles, 1968; Harris et al., 1998). Because of this structural complexity, very few good stratigraphic sections through the Permian succession have been identi®ed, and the relationship between stratigraphic units has not been adequately resolved. Many of the classic Permian fossil collections from Timor were taken from the meÂlange terrains, and are therefore not well located in terms of the stratigraphic sequence. The Permian succession of Timor is, in most localities, readily divisible into two lithologically distinct successions: a volcanic/shallow marine carbonate succession, generally massive in structure and occupying a high structural position; and a thin-bedded siliciclastic succession, usually strongly folded and occupying a low structural position. Most of the literature, prior to the late 1970s, interpreted the volcanic/carbonate successions as allochthonous, derived from far to the north of the parautochthonous siliciclastic sequences (e.g. Molengraaff, 1913; Wanner, 1913; Brouwer, 1942; Gageonnet and Lemoine, 1958; AudleyCharles, 1968; Barber et al., 1977). It was already recognised by Wanner (1956), however, that the supposed allochthon and the parautochthon contained essentially similar faunas, and subsequently stratigraphic interdigitation between the two successions has been demonstrated (Grady and Berry, 1977; Sawyer et al., 1993). It is now generally agreed that all the Permian successions of Timor developed together in one heterogeneous sedimentary

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environment on the northern continental margin of Australia (e.g. Audley-Charles and Harris, 1990). The Permian is the oldest recognised element in the Timor cover succession; early identi®cations of Carboniferous fossils from Timor (e.g. Beyrich, 1862, 1865; Bather, in Boehm, 1908; Schubert, 1915) have subsequently been reattributed to the Permian. Evidence in support of a more recent identi®cation of possible Late Carboniferous fossils (Lophophyllidium and Archeodiscus, in Sawyer et al., 1993) has yet to be published. Permian rocks in Timor form isolated outcrops through the central parts of both East and West Timor, with the most extensive outcrops being found in the Maubisse and Cribas areas of East Timor and the Bisnain and Kekneno areas of West Timor (Fig. 2). Throughout the island the Permian limestone units frequently form steep-sided isolated karstic mountains (`fatus' in Timor), whilst the clastic successions form gently rolling hill country. 2. History of investigation Beyrich (1862, 1865) was the ®rst to record Palaeozoic rocks from Timor, although these were initially interpreted as Carboniferous in age.They were subsequently identi®ed as Permian by Rothpletz (1891, 1892). These early samples were collected from the vicinity of Kupang, the main city in West (formerly Dutch) Timor, from the Ajer Mati locality. Subsequently Wichmann (1892) reported further collections from this locality, and from the Amarassi region to the southeast of Kupang. The ®rst Permian rocks from Portuguese East Timor were reported by Hirschi (1907) at Sahe Laca (Pualaca). Major contributions to our knowledge of the Permian in West Timor come from several geological expeditions carried out prior to World War II. J.Wanner visited Timor in 1909 and collected a rich Palaeozoic, Triassic and Jurassic fauna (Wanner, 1910a). In 1911 a German expedition under the leadership of Wanner and, concurrently (1910± 1912), a Dutch expedition led by G.A.F. Molengraaff carried out ®eldwork in western and eastern West Timor respectively. A second Dutch expedition was undertaken in 1916 led by H. Jonker, and in 1927 the Permian succession at Basleo was sampled by H. Ehrat and J. Venema (Wanner, 1930a). Finally, the University of Amsterdam Lesser Sunda Islands Expedition in 1937 under the leadership of H.A. Brouwer included a number of regional Ph.D. studies, of which, those by Tappenbeck (1940), de Roever (1940), Simons (1940) were particularly concerned with the Permian in Timor. Further sampling was also undertaken in the Basleo area by J. de Marez Oyens as part of this expedition. In East Timor the Permian was extensively sampled by the oil company geologist F. Weber in 1910/ 11 (Wanner, 1956). An enormous quantity of Permian fossil material was collected during these expeditions. The fossil groups were

systematically analysed by palaeontologists, the results being published in a series of monographs, most notably the series `PalaÈontologie von Timor' (edited by J.Wanner, 1914±1929). Monographs relevant to the Permian fossil taxa are listed in Appendix B and bibliographic details are given in the reference list. In the post-World War II period, the focus of research shifted to East Timor. Grunau (1953, 1956, 1957), Wanner (1956), Gageonnet and Lemoine (1958) published important regional studies, including sections on the stratigraphy and palaeontology of the Permian. In 1961, K. Nakazawa and two students from the University of Kyoto sampled the Permo-Triassic succession of East Timor and described the Permian fossils in several palaeontological papers (Nogami, 1963; Shimizu, 1966; Nakazawa and Bando, 1968). This early work was formalised into the presently recognised stratigraphy of East Timor by Audley-Charles (1968), who also produced a reconnaisance geological map of East Timor at a scale of 1:250,000. In West Timor the only extensive work during this period was undertaken by D. de Waard and students from the University of Indonesia, the main result of relevance to the Permian being reconnaissance geological mapping around the Basleo fossil locality (de Waard, 1955). A further phase of investigation from the 1970s to the 1990s, primarily in West Timor, was undertaken by the Geological Research and Development Centre, Bandung (GRDC, formerly the Geological Survey of Indonesia) in collaboration with the University of London Southeast Asia Research Group. West Timor was mapped at a reconnaissance scale of 1:250,000 by GRDC (the Kupang and Atambua sheets: Rosidi et al., 1981). The Permian Maubisse Formation was adopted as a mapping unit from East Timor, after initially being described as the Bifemnas Formation (Sartono, 1975) and the Kiupukan Formation (Hartono et al., 1975). The Permian clastic succession was mapped as the Bisane Formation (part of the Kekneno Series of older literature). From the London University Group the most signi®cant contributions were an MPhil thesis by Giani (1971) in the Belu area of West Timor, and Ph.D. theses by Cook (1986), concentrating mainly on the Triassic, by Bird (1987) covering the clastic Permian succession of the Kekneno massif, and by Barkham (1993) on the Maubisse Formation The political situation over the past twenty-®ve years has largely prevented systematic geological research in East Timor. Berry and Grady (1981) published an account of the deformation and metamorphism of the Aileu Metamorphic Complex on the north coast, which may include Permian units, but could also include Mesozoic and older Palaeozoic rocks. Grady (1975), Grady and Berry (1977) examined the relationships between the Permian units and the metamorphic basement rocks, and Berry et al. (1984) used conodonts to determine the age for part of the Maubisse Formation. Berry and Jenner (1982) established the tectonic af®nities of Permian volcanic rocks by

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Fig. 3. Simpli®ed stratigraphy of the Permian sequences of Timor, as used in this paper, compared with earlier nomenclature. The `V's indicate volcanic horizons. N.B. The Fatu and Sonnebait Series, as used in the publications of the University of Amsterdam's Expedition to the Lesser Sunda Islands (Brouwer, 1942), both include rocks of a wide range of ages.

geochemical analysis. Harris and his co-workers (Harris, 1991; Hunter, 1993; Prasetyadi, 1995; Prasetyadi and Harris, 1996; Harris et al., 1998, 2000; Harris and Long, 2001) re-examined the Permian Maubisse, Atahoc and Cribas formations and the Aileu Complex in their type localities in East Timor. GRDC has published the Dili and Baucau map sheets covering the whole of East Timor at 1:250,000 scale (Bachri and Situmorang, 1994; Partoyo et al., 1995). These maps are based on the mapping of AudleyCharles (1968), with additional ®eld mapping and airphoto interpretation. 3. Stratigraphy The Permian succession in Timor is subdivided into three formations, the Maubisse, Atahoc and Cribas formations (Fig. 3), with a fourth unit, the Aileu Metamorphic Complex on the north coast of East Timor, having a protolith which may, at least in part, be of Permian age (Fig. 4). The Maubisse Formation is composed predominantly of shallow marine carbonates and basic volcanics, whilst the Atahoc Formation and the succeeding Cribas Formation together comprise a predominantly siliciclastic basinal succession, with subordinate volcanics and volcaniclastics, deposited in deeper water. It is possible that the Aileu Complex includes

representatives of both the carbonate and siliciclastic Permian successions. Permian stratigraphic studies in Timor have largely employed the Russian Stage and Substage nomenclature. In this paper we follow the currently accepted Permian chronostratigraphical scale of Jin et al. (1997) (Fig. 3), but in most cases we also indicate the stage/substage nomenclature assigned by the original author as there appear to be considerable discrepancies in the ages of faunal assemblages as determined from the different fossil groups (see in particular Fig. 10 and Sections 5 and 6). 3.1. The base of the Permian in Timor It is now generally accepted that the entire Permian succession of Timor formed part of the Australian continental margin prior to its collision with the Banda Arcs in the Miocene±Pliocene. This implies that the Permian rocks were originally deposited on pre-Permian, Australianaf®nity continental crust, perhaps on Precambrian basement which underlies the Ashmore Platform to the south. Alternatively, considering the Permian palaeotectonic setting of Timor (see Section 8.3), it is possible that the pre-Permian basement of Timor is composed of older Palaeozoic strata, metamorphosed as a result of Permian crustal extension and igneous activity. This possibility awaits further investigation. The metamorphic rocks in East Timor have been called

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Fig. 4. Type areas for the Permian units in East Timor, based on GRDC Geological Map Sheets Dili and Baucau (Bachri and Situmorang, 1994; Partoyo et al., 1995), with modi®cations by R.A. Harris.

the Lolotoi Complex (Audley-Charles, 1968), and in West Timor the Mutis Complex (e.g. Rosidi et al., 1981; Sopaheluwakan, 1990). At several places in eastern Timor, stratigraphic contacts have been described between the Lolotoi Metamorphic Complex and the Permian Maubisse Formation. Giani (1971) reported that the Maubisse Formation overlies the Lolotoi Complex unconformably in the Masin River, at the western end of the Lolotoi `type' massif, on the border between East and West Timor. Stratigraphic contacts have also been recorded between the Maubisse Formation and the Lolotoi Complex at two places on the large Laclubar metamorphic massif in central East Timor (Fig. 4). Wittouck (1937, reproduced in van Bemmelen, 1949, Fig. 240) mapped stratigraphic contacts between limestones, then interpreted as Triassic in age, and ?Permian meta-igneous rocks west of Pualaca. Subsequently, the limestones and associated volcanics were dated as Early Permian, based on bryozoans and fusulinids (Grunau, 1953; Nogami, 1963). Further north on the Laclubar massif, Grady and Berry (1977) reported an irregular, generally shallowly dipping sedimentary contact between the metamorphic complex and overlying Permian limestones and other sediments of Permian age. In this area the Lolotoi Complex is represented by low-grade metavolcanics interbedded with

dark blue-grey phyllites. Overlying the metavolcanics are Permian limestones containing occasional compound rugose corals and other shallow-water faunas. Fragments of the underlying metavolcanics occur in the basal limestones. In West Timor crystalline schist fragments are found within conglomeratic sandstones in the Permian Kekneno Series ( ˆ Atahoc/Cribas formations), but, according to Brouwer (1942), were nowhere found in Permian conglomerates of the Sonnebait Series ( ˆ Maubisse Formation). Conglomerates in the Maubisse Formation are composed predominantly of basic volcanic fragments. This suggests that the basinal areas of West Timor received sediments shed from (presumably) Australian continental basement, whilst the structural highs, upon which the Maubisse Formation accumulated, were primarily composed of basic volcanics. If the observations from East Timor that the Maubisse Formation unconformably overlies metamorphic rocks of the Lolotoi Complex are con®rmed, then parts of the Lolotoi Complex represent upthrust fragments of the Australian continental basement. This has important structural implications as it suggests a basement-involved style of thrust deformation in eastern Timor, rather than the more widely accepted thin-skinned style of thrusting. It also suggests that

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parts of the Lolotoi Complex in East Timor may not be equated with the Mutis Metamorphic Complex in West Timor, which has been demonstrated to represent basement derived from the pre-collisional Banda forearc on the basis of its association with Late Cretaceous-Tertiary island arc volcanics and mantle ophiolite (e.g. Earle, 1983; Sopaheluwakan, 1990). 3.2. Maubisse Formation 3.2.1. The Maubisse Formation in East Timor Audley-Charles (1968) de®ned the Maubisse Formation in East Timor (after the Maubisse Series of Gageonnet and Lemoine, 1958), with a type locality in cliffs which form part of the Ramelau Range to the southwest of Maubisse village (Fig. 4). A detailed study of the structure and stratigraphy of the Maubisse Formation in the Maubisse area has been conducted more recently by Hunter (1993). At its type locality the Maubisse Formation consists of three mappable facies: limestone and calcareous shale, basalt, and volcaniclastics (Hunter, 1993). Limestone, forming precipitous cliffs due to its resistance to weathering, is the most obvious unit. From a distance limestone cliffs of Permian or Triassic age are dif®cult to distinguish, because they are both commonly covered by iron-stained aragonite deposits; many of the areas mapped originally by AudleyCharles (1968) as Maubisse Formation, including outcrops at the village of Maubisse itself, have proved to be composed of Triassic Aitutu Limestone (Hunter, 1993). The bulk of the Maubisse limestone at its type locality consists of poorly sorted and altered biocalcarenites containing skeletal grains, lime mud pellets, lithoclasts, and volcanic clasts (including volcanic glass) all enclosed in a lime mud matrix (Hunter, 1993). Calcite clasts frequently contain remnant structures of faunal origin. The bioclasts show little or no abrasion. The fauna includes brachiopods, crinoids, molluscs, ammonites, and rugose corals. The matrix is commonly recrystallized to neomorphic spar, and contains iron oxides, giving the Maubisse limestone its distinctive red color. The iron oxides are frequently concentrated along bedding-parallel pressure solution seams, which diverge around bioclasts. Individual limestone layers up to 4 m in thickness are mostly very hard. Collectively, these layers form massive successions of up to 60 m thick, as in the cliffs SW of Maubisse village, de®ned as the type locality by AudleyCharles (1968). However, the lateral continuity of each limestone succession is limited to less than a few hundred meters. The pod-like shape of the massive limestone facies in the Maubisse area and throughout much of Timor is interpreted as a combination of primary reef-type deposition (de Waard, 1957; Hunter, 1993) and secondary structural disruption (Barber et al., 1986; Harris et al., 1998). Interbedded with the thick layers of mostly coarse limestone are pink biocalcarenite and red to pink calcareous shale beds 3±15 cm thick. These beds occasionally have

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cross and parallel current laminations and are rich in fossils (Hunter, 1993). The greater part of the Maubisse Formation in East Timor consists of basalt, with minor ma®c intrusive rocks, syenite and tuffaceous material. Basalt commonly occurs as amygdale-rich, spilitic, pillow and sheet ¯ows. One of the clearest contact relations between basalt and limestone of the Maubisse Formation is found near the NE crest of Lacouse Mountain, where pillow lavas are overlain by fossiliferous limestones, with a basal conglomerate of basalt pebbles, dipping around 358 to the north. Another outcrop near Memo (SW of Maliana) has boulder-sized blocks of basalt encased in pink Maubisse limestone with brachiopod, coral, and crinoid debris. Intrusive rocks and alkalic volcanic material, rich in potassium feldspar, are abundant in the Hatu Builico valley and form the greater part of Monte Tata Mai Lau. Like the limestones, the basalt facies of the Maubisse Formation also lacks lateral continuity. The strong lenses of limestone and basalt have remained mostly intact during deformation, as the weaker shale and volcaniclastic material ¯owed around them. The mechanical heterogeneity and variable deformational response of the Maubisse Formation commonly cause it to dismember into fault-bounded blocks of basalt and limestone encased in a remobilized clay-rich matrix (Harris et al., 1998). Even where the Maubisse Formation is mostly intact, as at the type locality, most of the basalt and limestone units show evidence of shear along their contacts, while the associated shale and volcaniclastic units are intensely disrupted (Hunter, 1993). Near its type locality the Maubisse Formation is dominated by a volcanic-rich basal unit composed of basalt and volcanically derived clastics. The basal unit is overlain by and grades laterally into carbonate, calcareous shale and volcaniclastic material (Hunter, 1993; Harris and Long, 2001). The volcaniclastic rocks are green to grey ash- and clastic-rich sedimentary rocks, up to 40 m in thickness, interbedded with both the basalt unit and the calcarenites (Hunter, 1993). This unit is found directly to the NW of Maubisse village where it overlies a thick sequence of carbonate. Locally the volcaniclastics are tuffaceous, but in most places they show clear evidence of current deposition, in the form of grading and cross bedding. Fining-upward sequences, up to several meters in thickness, are characterised by a poorly sorted conglomerate of subangular, vesicular, basalt fragments up to 10 cm in diameter, at the base, grading upward into progressively ®ner-grained, cross laminated units and shale. Thin sections show basaltic and felsic volcanic rock fragments and calcite cement (Hunter, 1993). The occurrence of pillow lavas and current structures indicate a mostly subaqueous volcanic environment. Hunter (1993) interpreted the well-preserved and diverse faunal assemblages as indicating mostly low-energy, open marine conditions during limestone and shale deposition.

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In East Timor major outcrops of the Maubisse Formation are found in the Ramelau Range, extending both southwestwards and northeastwards from Maubisse, Lacouse Mountain on the border between East and West Timor and the Legumau Mountains in the eastern part of East Timor. In the areas examined by Gageonnet and Lemoine (1958) and Audley-Charles (1968), similar lithologies are found to those described by Hunter (1993) from the type locality, although Gageonnet and Lemoine (1958) also distinguished a white crinoidal limestone facies, without associated volcanics, containing Spirifer and Productus, to the west of Manatuto. Wanner (1956) described similar facies from the Pualaca and Hatu Dame areas, and listed other Permian carbonate/volcanic localities then known. Descriptions of localities for the Maubisse Formation from Pualaca and elsewhere in East Timor are given by Grunau (1953). In most of these localities volcanic units, particularly basalt, are the most abundant component. 3.2.2. The Maubisse Formation in West Timor In West Timor equivalents of the Maubisse Formation were placed within the Fatu and Sonnebait Series by geologists of the University of Amsterdam Lesser Sunda Islands expedition (e.g. Brouwer, 1942) (Fig. 3). The Fatu Series, named after the characteristic Timorese landform of steepsided isolated hills, typically composed of massive limestone, was interpreted as a high structural element, including massive limestones of Permian (Maubisse Formation), Triassic and Tertiary age. The Sonnebait Series, composed largely of more argillaceous material, corresponds approximately to the Bobonaro Scaly Clay MeÂlange Complex of more recent literature (e.g. Harris et al., 1998). De Waard (1957), however, recognised that some of the carbonate buildups in the supposed Fatu Series formed bioherms within the Sonnebait Series. A comprehensive study of the Maubisse Formation has been carried out by Barkham (1993) in two areas of West Timor: the Bisnain area, immediately to the east of the classic Bitauni fossil locality; and the Laktutus area near the border with East Timor. The Bisnain area illustrates some fundamental characteristics of the Maubisse Formation and its relationships to the Atahoc and Cribas formations (Fig. 5). The structure of the Bisnain area is complicated by numerous minor wrench and normal faults, but essentially the area is divided into two structural domains, separated by a southward-directed thrust. The northern domain, and the larger part of the southern domain, are composed of basalts and carbonates of the Early Permian Maubisse Formation, but the central part of the study area near Bisnain village consists of Late Permian± Triassic limestones (mainly the Permian Oemofai Member described in Section 4, and the succeeding Triassic Aitutu Formation). Basinal Permian successions of the Atahoc and Cribas formations outcrop in the extreme south of the study area (Fig. 5). Barkham (1993) identi®ed nine facies associations

(members/beds) within the Maubisse Formation at Bisnain, seven of which are essentially contemporaneous, dated or interpreted as of Artinskian±Kungurian age. The remaining two are the Late Permian±Triassic Oemofai Member, and the Hoeniti Member, which is dated as late Sakmarian from brachiopods, but may continue up into the Artinskian. These seven or eight Artinskian±Kungurian members of the Maubisse Formation, together with the Kungurian part of the Cribas Formation to the south, show a regular progression of depositional environments (from north to south): Nekmelalat Member: Planar unpillowed basalts, succeeded by bryozoan and crinoid packstones and volcaniclastic sediments. Manufui Lavas: Pillow basalts, feeder dykes and interbedded tuffaceous sediments (Fig. 6). Melelat Beds: A diagenetic facies association, from the hydrothermal alteration of carbonates and calcareous claystones. Hoeniti/Banfafor members: A relatively deep water (below fair weather wave base, above storm wave base), shallowingupward sequence of bryozoan-dominated wackestones and grainstones (Hoeniti Member), passing along strike and stratigraphically upwards into crudely laminated calcareous muds with interbedded turbidites or storm-generated deposits (Banfafor Member) (Structural break at thrust, and interruption by Late Permian±Triassic succession) Bitauni Beds: A terra rossa palaeosol, developed unconformably on Artinskian±Kungurian marine limestones (Fig. 6). Bisnain Beds: Massive bryozoan-dominated, mud-matrix limestone, passing laterally into well bedded crinoidal packstones and grainstones. Interpreted as a mud mound passing laterally into a winnowed ¯anking facies. Khali Beds: Cyclic crinoidal rudstone±packstone±wackestone, ®ning upward and then coarsening upward, overlying a dolomitised, eroded and bored base. Interpreted as cycles of transgression±regression with emergence at regressive maxima (Fig. 6). Cribas Formation: Varicoloured siltstones and shales, wackestones, pillow lavas and sandstones. The lower part of the section, of Kungurian age, is composed predominantly of limestones and marls contains abundant bivalves (Atomodesma exarata) and small brachiopods, and was deposited on the outer part of the inner shelf (20±50 m water depth based on the Atomodesma bivalves, if not reworked into deeper water). This passes upwards into ?Late Permian±Early Triassic shales with interbedded volcaniclastic horizons and pillow lavas. This sequence shows an overall progression from a volcanic high in the north to a basinal succession in the south. The cyclical Khali Beds occur at the boundary between the volcanic-carbonate platform and the predominantly clastic basin. Barkham (unpublished report, 1986) logged a rather poorly exposed stratigraphic section over 1000 m thick, but due to the cyclical nature of the succession he was not able

T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774

Fig. 5. Simpli®ed geological map of the Bisnain area, West Timor (after Barkham, 1993).

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Fig. 6. Selected measured sections (schematic for the Khali Beds) in the Permian Maubisse Formation of the Bisnain area, West Timor (data from Barkham, 1993). Locations are shown on Fig. 5 Sections are at various vertical scales.

to determine to what degree (if at all) the sequence had been tectonically thickened. Barkham (1993) recognised at least three sedimentary cycles in the Khali Beds, typically 10±12 m thick, with the following upwards succession of sedimentary environments (Fig. 6): High energy beach or bar calcareous sandsÐBrachiopod and crinoid gravel banksÐInner shelf, slowly accumulating organic mudsÐShallow marine, high energy crinoidal sands/gravelsÐTransgressive basal lagÐPalaeokarst (below unconformity). This small-scale cyclicity, combined with the substantial thickness of the Khali Beds, suggests low- (third-order?) transgressive±regressive cycles within an overall prograding (second-order?) regressive sedimentary wedge. Brachiopods within the upper part of the third-order cycle were dated as Sterlitamakian (late Sakmarian) by Archbold and Barkham (1989), but the shells were described as crushed and incomplete, and may have been reworked into slightly younger sediments. Barkham (1993) interpreted the Khali Beds as probably Artinskian±Kungurian in age. A conodont sample from this unit yielded an Artinskian age (Section 6.5). A further indicator of regression within the Bisnain succession is the development of a palaeosol on Artinskian±Kungurian marine limestones in the Bitauni Beds (Fig. 6). This is most likely the same regressive cycle as interpreted in the Khali Beds. The

Kungurian carbonates of the basal Cribas Formation (a distal facies equivalent of the Khali Beds?) pass upward into ?Middle/Late Permian±Triassic ®ne-grained siliciclastics, which represent a subsequent phase of transgression. A further facies association described by Barkham (1993) is the Tuke Beds of the Oinlasi area, some 9 km south of Basleo. No in situ outcrops of this facies have been found, but it occurs as numerous boulders, particularly in the headwaters of the Noil Tuke river. The dominant lithology is a fusulinid rudstone/grainstone in which the fusulinids have a parallel orientation, are well sorted, and show sedimentary structures such as imbrication and cross-bedding. Although these fusulinid limestones are rich in individuals, only a few species are represented (Thompson, 1949) which are dated as Early Permian, probably Artinskian. The tops of some of the limestone boulders show vertical burrows or borings in®lled by red micritic mud. The only other Maubisse Formation lithologies from the Oinlasi area are crinoid-ossicle grainstones, bryozoan packstones with Tubiphytes, and bryozoan-rich ¯oatstone, all of which contain fusulinids. All lithologies except the ¯oatstones are partially dolomitised. Barkham (1993) interpreted this facies association as representing sheets or banks of mobile gravel deposited above fair weather wave base. The gravel was probably deposited in a tidally dominated environment with breaking

T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774

729

Fig. 7. Simpli®ed geological map of the Kekneno area, West Timor, showing the distribution and relationships of the Permo-Triassic rocks, including the Kasliu fossil locality, slightly modi®ed from Bird and Cook (1991).

waves, accounting for abraded fusulinid tests, but also in more offshore environments, where the water energy was still strong enough to winnow argillaceous material, but not strong enough to rework the larger bioclasts. In the Kekneno area, Bird (1987), Archbold and Bird (1989) described the Kasliu Member of the Maubisse Formation from outcrops on a structural/topographic high immediately east of the Kekneno basinal area (Fig. 7). This is particularly signi®cant as it is one of the few areas of uncondensed in situ outcrop accurately dated to the Late Permian. This member consists of crinoidal dolomites, minor red marls and shales, bryozoan packstones and wackestones, volcaniclastic sands, and at the top of the sequence, amygdaloidal pillow lavas. The crinoidal limestones, bryozoan packstones and particularly the volcaniclastics contain a rich brachiopod fauna, dated by Archbold and Bird (1989) as Chhidruan (Wuchiapingian). The Kasliu fauna shows a close relationship to the brachiopod faunas at the Basleo and Amarassi fossil localities (Section 5).

3.3. Atahoc Formation The Atahoc Formation was de®ned by Audley-Charles (1968) from the Cribas anticline in north-central East Timor (Figs. 2 and 4). It corresponds to the lower Cribas Series of Grunau (1956) and Gageonnet and Lemoine (1958). The Atahoc Formation conformably underlies the Cribas Formation of Audley-Charles (1968), the latter being the equivalent of the upper Cribas Series of earlier workers. The base of the Atahoc Formation is not exposed in the Cribas area, nor has it been recognised elsewhere. Based on structural arguments, however, Charlton (2002) has interpreted the Cribas Anticline as a basement-cored structure, and this interpretation suggests that basement, possibly the Lolotoi Metamorphic Complex, may directly underlie the Atahoc Formation in its type area. In the type area the Atahoc Formation has an estimated stratigraphic thickness of 600 m. It is largely a monotonous series of ®nely laminated, well lithi®ed black shales. The lowest exposed member is a hard, unfossiliferous pink and

730 T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774 Fig. 8. Selected stratigraphic sections of the Permian Atahoc and Cribas formations in West Timor. Measured sections from the River Bisnain are after Barkham (1993) and summary sections from the Kekneno area after Bird and Cook (1991). Sections are at various vertical scales.

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731

Fig. 9. Palaeogeographic model for the Permian of West Timor, based on models by Barkham (1993) for the Bisnain area, and by Bird and Cook (1991) for the Kekneno area.

grey, massive, quartz sandstone with some thin black shale intercalations. The overlying shales are mainly silty, with numerous calcareous and clay-ironstone nodules and some thin limestone beds. The limestones are beige, grey and red, usually fossiliferous, with debris of crinoids, rare trilobites, cephalopods, corals and bryozoans. A few thin cherts are interbedded with the limestones, and thin beds of hard, micaceous quartz sandstone and siltstone are also developed, particularly towards the top of the formation. The top of the formation is designated arbitrarily at the top of an amygdaloidal basalt 3±7 m thick, which occurs below the ®rst appearance of Atomodesma bivalves that typify the succeeding Cribas Formation. The age of the Atahoc Formation was interpreted as Sakmarian in its type area based on an ammonoid-trilobite fauna, comparable to the Somohole Beds of West Timor (Grunau, 1953, 1956; Gageonnet and Lemoine, 1958). Audley-Charles (1968) considered the formation a marine ¯ysch sequence, but found no ®rm indicators for water depth. The occurrence of calcareous concretions and black pyritic shales was taken as a possible indicator of stagnant anoxic conditions. Elsewhere in East Timor the Atahoc Formation has been recognised only in the Loiquero area at the extreme eastern end of the island (Fig. 2). There it consists of hard grey slates, with pink and grey, hard, massive quartz sandstones (Audley-Charles, 1968). A Permian age is indicated by the occurrence of the bivalve Merismoptera sp. cf. M. macroptera (Grunau, 1956). In West Timor the Atahoc Formation has been recognised by Bird (1987) in the Kekneno area (Fig. 7) and Barkham (1993) in the Bisnain area (Fig. 5). In the Kekneno area of north-central West Timor, Bird (1987) described the Atahoc Formation as comprising two members, the Naibalan Member and the succeeding Noil Bisone Member (Fig. 8). The Naibalan Member, with an estimated stratigraphic thickness of 240 m in its type locality, consists predomi-

nantly of dark shales, interbedded with ®ne quartz sandstones and occasional detrital limestone horizons. The shales, which comprise about 60% of the member, are primarily dark grey, but in part red, green and black, laminated, carbonaceous, and pelitic, locally silty with normally graded beds 2±3 cm thick. Slump folds and large ironstone concretions occur occasionally. The sandstones, which form about 30% of the member, are ®ne-grained, well sorted quartz arenites, carbonaceous in part and commonly pyritic. Beds up to 2 m thick are sharply de®ned, generally massive to slightly normally graded, sometimes showing basal ¯ute and groove casts. Limestones form the remaining 10% or less of the member, consisting of thin (5±30 cm) bioclastic packstones and wackestones. Fragments of gastropods, bivalves, crinoids, brachiopods, corals and unfragmented small pelagic foraminifera are present. There are also abundant well-preserved ammonites, which date the sequence as Sakmarian and possibly latest Asselian. The Noil Bisone Member (Fig. 8) consists almost entirely of dark grey and black shales similar to those in the Naibalan Member. These are uniformly pelitic and carbonaceous, with large iron-rich calcareous nodules. Poorly preserved ammonoids and fragments of trilobites occur rarely, but are not age-diagnostic. Slump folds are common, locally passing into debris ¯ows. Thin intercalations of limestones and ®ne sandstones/siltstones are present. The limestones are generally mudstones or wackestones, containing brachiopod and crinoid debris. The member has an estimated thickness of about 300 m. Palaeocurrent data from the Naibalan Member indicate a bidirectional NNW±SSE trend, suggesting a narrow, approximately E±W elongated basin with northerly and southerly sedimentary sources. Palaeocurrents in the Noil Bisone Member are more unidirectional toward the NW. The muddy, carbonaceous nature of the sediments and the lack of a benthic fauna implies anoxic substrate conditions, and a depositional environment largely isolated from clastic

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input (Fig. 9). Debris ¯ows and slump folds suggest tectonically triggered instability. The lower part of the Atahoc Formation in the Bisnain area, described by Barkham (1993) (Fig. 8), is very similar in lithology and age to the Naibalan Member described by Bird (1987) in the Kekneno area. It consists of black shales with a rich ammonoid fauna, and minor interbedded sands and grainstones. Above the shale unit at Bisnain is a volcanic unit comparable to, but much thicker than, the basaltic volcanics at the top of the Atahoc Formation at its type locality in East Timor, with pillow basalts, massive basalts and interbedded limestones. Both the lower sedimentary section and the succeeding volcanic unit are about 80 m thick. 3.4. Cribas Formation The Cribas Formation (Audley-Charles, 1968) is the upper part of the Cribas Series of Grunau (1956) and Gageonnet and Lemoine (1958). In its type locality in the Cribas Anticline in East Timor, the Cribas Formation overlies the Atahoc Formation conformably (Fig. 4), the boundary being placed arbitrarily at the top of a basaltic horizon, but marked palaeontologically by the ®rst appearance of Atomodesma bivalves which are characteristic of this formation (Audley-Charles, 1968), and by the last appearance of trilobites, rare but characteristic of the Atahoc Formation (Gageonnet and Lemoine, 1958). Upward the Cribas Formation is apparently transitional into Ladinian (Middle Triassic) siliciclastics and carbonates, although no fossils of Early±early Middle Triassic age have been reported from the Cribas area. Audley-Charles (1968) considered the Cribas Formation to be Late Permian in age, and this is supported by the occurrence of the bivalve Atomodesma variabilis which also occurs at Basleo. However, the additional occurrence of the Kungurian Atomodesma exarata suggests an overall late Early±Late Permian (and younger?) age range for the Cribas Formation in its type area. As in other places in Timor (see Section 4), the Permo-Triassic boundary section is apparently represented by a sparsely fossiliferous condensed stratigraphic section. In its type section the bulk of the Cribas Formation is composed of shales and silty shales with common calcareous and clay-ironstone nodules. At the base are black and blue-grey shales, micaceous siltstones and greenish, ®ne quartz sandstones. The overlying strata are mainly shales and micaceous shales, locally red and green. Bedded limestones are rare near the base of the formation, but in the upper part bedded biomicarenites and calcilutites become more common. Upward, calcareous nodules become less common and non-carbonate arenites become rare. The type section is about 500 m thick (Audley-Charles, 1968). The environment of deposition for the Cribas Formation was interpreted by Audley-Charles (1968) as broadly similar to the Atahoc Formation, possibly marine pro-delta shelf deposits and delta slope beds. The occurrence of calcareous

nodules in black, occasionally pyritic shales was taken as indicative of locally stagnant anoxic conditions. The composition of the clasts in the arenite beds suggests derivation from an acid igneous source. Audley-Charles (1968) recognised the Cribas Formation in East Timor outside its type area, in the Loiquero, Veru, Bazol, Viqueque and Aliambata anticlines (Fig. 2), although the characteristic Atomodesma bivalve was not identi®ed at Bazol or Viqueque; Brunnschweiler (1977) re-interpreted these last two occurrences as Triassic ¯ysch, equivalent to the Babulu Formation of West Timor (Giani, 1971; Bird and Cook, 1991). Hunter (1993) discovered Cribas Formation shale, arenite and minor skeletal lime packstone structurally underlying the limestones of the Maubisse Formation in low-lying areas around Maubisse village (Fig. 4). In outcrops immediately to the north and south of Maubisse Village Atomodesma was found throughout these units. In the Maubisse region shales of the Cribas Formation commonly form purple, red or green layers up to 10 cm thick (Hunter, 1993). These are interbedded with brown and grey, ®ne to very ®ne, sandstone layers. Detrital mica is abundant and phyllite-grade metamorphic mica is found along the thrust plane at the top of the unit. Overall, the shale-rich part of the Cribas Formation is 300±400 m thick and grades upward into coarser clastic units (Hunter, 1993). Arenite beds up to several meters in thickness are found throughout the upper-part of the shale unit (Hunter, 1993). They are characterized by ®ning-upward sequences, with a basal massive sandstone unit encompassing shale clasts up to 3 cm in diameter with locally boulder-sized clasts. These are overlain by parallel-laminated and cross-laminated, ®ne to very ®ne sandstone. The sandstone is poorly sorted and the grains are subrounded, with framework grains composed of basaltic rock fragments, carbonate and mono- and polycrystalline quartz grains and feldspar (Harris et al., 2000). These sandstone units are similar to other Permian siliciclastic material analysed throughout Timor (Harris et al., 2000), which plot within the continental block provenance ®eld of Dickinson (1985). The upper part of the Cribas Formation includes thin beds of light brown skeletal packstone with cryptosome bryozoans and Permian ammonites (Hunter, 1993). Hunter (1993) interpreted the interbedded shale and sandstone of the Cribas Formation as part of a distal submarine fan complex, prograding into an open ocean. Only the upper parts of Bouma sequences (C, D and E) are found in most units. The immaturity of sandstone grains and textures indicates a proximal source, and limited residence time on the shelf before redeposition into a basin by turbidity currents. Phosphatic grains found within the sandstone also support the interpretation of initial deposition in shallow water conditions, followed by redeposition. The dominance of sandstone in the upper part of the Cribas may re¯ect a coarsening upward sequence associated with basin in®lling (Hunter, 1993).

T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774

In West Timor the Cribas Formation has been described from the Kekneno area by Bird (1987) (Figs. 7 and 8), and from the Bisnain and Laktutus areas by Barkham (1993) (Figs. 5 and 8). As in East Timor, the Cribas Formation overlies the Atahoc Formation conformably. In the Kekneno area reddish limestones are common near the base of the Cribas Formation, and these mark the ®rst appearance of the bivalve Atomodesma exarata. Bird (1987) took the base of these limestones to mark the boundary between the two formations. The Cribas Formation is at least 400 m thick in the Kekneno area. Bird (1987) divided the Cribas Formation in the Kekneno area into two members, the Matpunu and Tutlap Members, and the informal Nunmaliup Beds (Fig. 8). The Tutlap Member overlies the Matpunu Member, but the relationship between these and the undated Nunmaliup Beds is less certain. Locally, the Nunmaliup Beds overlie the Tutlap Member, but elsewhere they interdigitate. Lithological differences between these members/beds are slight however, considering the lithological variability within the members on a ®ner scale. As described by Bird and Cook (1991), the Cribas Formation in the Kekneno area consists of thin-bedded sandstones, siltstones, shales, limestones and marls (Fig. 8). The sandstones consist of cm-bedded sets up to 1 m thick. They are very ®ne grained and rarely graded. Bed bases are planar and occasionally erosive; bed tops are usually ripplelaminated. The siltstones are more thinly bedded and are grouped into thick uniform sets. The beds again have sharp bases and are laterally very continuous and uniform. The siltstones are prominently laminated, showing a range of ripple forms from plane-laminated through wavylaminated to cross-laminated, with aggradational climbing ripple cosets. Horizontal burrowing is common, and the bivalve Atomodesma exarata is abundant and well preserved, although not found in life position. The shales are thick, dark grey and sometimes carbonaceous, similar to those of the Atahoc Formation. Red limestones and marls occur as units up to 20 m thick, particularly near the base of the formation. These are predominantly bioclastic wackestones, the bioclasts largely consisting of Atomodesma, but also solitary corals, crinoids, gastropods and encrusting foraminifera. The sandstones and siltstones were deposited from weak, low density, perhaps storm-generated turbidity currents, with the intervening shales the product of background pelagic sedimentation and biogenic reworking in a restricted marine environment. Kaufmann and Runnegar (1975) suggested that Atomodesma palaeocommunities lived at water depths of 20±50 m, although as the bivalves are not preserved in life position, this is a minimum water depth. The limestones probably accumulated as shallow banks of bioclastic detritus, bypassed by the siliciclastic turbidites (Fig. 9). At Bisnain, as at Cribas and Kekneno, the Cribas Formation overlies the Atahoc Formation conformably (Barkham,

733

1993). Again the boundary is de®ned by the appearance of Atomodesma which, as in the Cribas Anticline, coincides with the top of a volcanic unit. The base of the Cribas Formation at Bisnain is also marked by limestone beds, as it is at Kekneno. In the Bisnain area the Cribas Formation has a measured stratigraphic thickness of about 150 m (Fig. 8). At Laktutus the Cribas Formation is only poorly exposed. It is interpreted as having been deposited in a broadly similar environment to the same formation at Bisnain, although at Laktutus the predominance of ®ne-grained deposits and ripple-laminated sands indicates that this area was more distal to sedimentary sources (Barkham, 1993). 3.5. Aileu Metamorphic Complex The Aileu Metamorphic Complex, on the north coast of East Timor around Dili (Fig. 4), is composed of slates, quartzites, psammitic and pelitic schists, calc-schists, marbles, amphibolites and ultrama®c rocks (Berry and Grady, 1981; Prasetyadi, 1995; Prasetyadi and Harris, 1996). The metasedimentary rocks include fossils of Permian and Mesozoic age (Brunnschweiler, 1977). No rocks older than the Permian have so far been identi®ed. A Permian age for the metamorphic rocks was ®rst suggested by Gageonnet and Lemoine (1958) based on poorly preserved bivalves, similar to those found in the Cribas Series (Atahoc/Cribas formations). A Permian age was more clearly demonstrated by Brunnschweiler (1977) who recorded an ammonoid fauna in purplish calc-schists from the southern part of the complex. Brunnschweiler (1977), Barber et al. (1977) recorded recrystallised limestones containing Palaeozoic crinoids along the north coast and Barber et al. (1977), Prasetyadi and Harris (1996) document strained crinoid stems and Palaeozoictype brachiopods in deformed limestones to the south of Aileu. As described by Barber and Audley-Charles (1976), Barber et al. (1977), and illustrated on the map by Prasetyadi and Harris (1996), the Aileu siliciclastic sequence shows a pattern of northward coarsening, indicative of a classic offshore to landward transition in sedimentary facies. About 5 km north of Maubisse village, near the Daissol River (Fig. 4), slates dominate as limestone and volcaniclastic material of the Maubisse Formation pinch out. The E± W-trending slate belt of the Aileu Complex is very homogeneous and extends northward for 10 km across the Ermera±Aileu anticline to near Remexio, where quartzite is interbedded with phyllite. Another transition is found just east of Dili where quartzite beds dominate (Fig. 4). Most of the compositional transitions become apparent across northdipping thrust faults that have foreshortened the original facies relations (Prasetyadi and Harris, 1996). The regional outcrop pattern and sedimentary facies distribution in the Aileu area suggests that these units occupied an approximately E±W trending basin, perhaps a

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graben or half-graben (Prasetyadi and Harris, 1996). The southern boundary of the basin is formed by a major graben-bounding fault, downthrown to the north, with carbonate clastics shed from the Maubisse area, which was located in the southern footwall block. Siliciclastic sediments were sourced primarily from the northern margin of the basin. The clastics are interbedded with crinoidal limestones; but because of the metamorphism it is not apparent whether these are in situ shallow marine carbonates or calcareous turbidites. 3.6. Stratigraphic relationships The relationship between the Maubisse Formation and the contemporaneous Atahoc and Cribas formations has long been a subject of discussion. The predominantly siliciclastic Atahoc and Cribas formations generally occupy low topographic and structural positions, whilst the Maubisse Formation mostly forms mountain massifs, often resting on thrust planes, at high structural levels. Consequently the volcanic/carbonate successions of the Maubisse Formation have been commonly interpreted as allochthonous elements, overthrust onto the siliciclastic successions. In early interpretations, e.g. Audley-Charles (1968), Barber et al. (1977), the Maubisse Formation was interpreted as a unit of Asian origin which had been thrust over the autochthonous Australian clastic succession during arc±continent collision. As has already been mentioned, however, Wanner (1956) had earlier recognised that the two parallel successions contained essentially similar faunas. Stratigraphic interdigitation of the siliciclastic and carbonate/volcanic successions is now well established. In East Timor, to the west of Cribas, Grady and Berry (1977) described lithological units typical of the Maubisse Formation which are interlayered with units lithologically and faunally characteristic of the Cribas and Atahoc formations. In addition, in that area there are units with lithologically intermediate characteristics, such as green and green-brown, tuffaceous sandstones and siltstones. Red-brown siltstones and shales with occasional crinoid ossicles in the CribasAtahoc type section are very similar to red-brown calcareous siltstones and shales in the Maubisse Formation. The basalt horizon which in the type area is taken to mark the boundary between the Cribas and Atahoc formations is identical to some of the basaltic volcanic rocks in the Maubisse Formation. In the Maubisse region Hunter (1993) interpreted the contact between the Maubisse Formation and the structurally underlying Cribas Formation as a thrust contact (Fig. 4). However, the amount of slip and sense of shear along the contact is poorly constrained and it may represent minor internal slip within a sequence that is mostly stratigraphically continuous (Harris et al., 2000). Stratigraphic relationships have also been recorded between the Maubisse and the Atahoc/Cribas formations

by Sawyer et al. (1993) in West Timor. A conformable contact was mapped between the Maubisse Formation and the Atahoc Formation along a hillside above the Noil Laka river 5 km south of Kapan, although an absence of way-up criteria means that the stratigraphic order remains unclear. A stratigraphic contact between the Maubisse Formation and the Cribas Formation was recorded 6 km north of Kefamenanu. In this section at least three ¯ows of aphanitic pillow lava, containing blocks of crinoidal pink limestone and vesicular basalt, and representative of the Maubisse Formation, are succeeded by well bedded green and rust coloured shales with interbedded brown sandstone horizons typical of the Cribas Formation. The boundary between the Aileu Metamorphic Complex and the Maubisse Formation north of Maubisse (Fig. 4) was originally interpreted by Audley-Charles (1968) as a thrust, with the Aileu metamorphics structurally overlying the Maubisse block. Prasetyadi and Harris (1996) carried out four traverses across the contact between the Aileu Complex and other units near Hatolia, Letefoho, Aileu, and Laclo. Detailed studies of the contact in these areas showed that they were north-dipping thrust zones. On the other hand Barber and Audley-Charles (1976), Barber et al. (1977), Grady and Berry (1977) re-interpreted this contact as gradational in terms of both stratigraphy and metamorphic grade (Fig. 2). In the Letefoho and Aileu traverses Prasetyadi and Harris (1996) documented a compositional transition between the Cribas and Maubisse formations and slates of the Aileu Complex. Immediately to the north of Maubisse village there is a transition between Atomodesma- bearing shale and siltstone of the Cribas Formation, and slate and quartzite of the Aileu Complex. The limestone and calcareous shale unit of the Maubisse Formation also becomes increasingly clastic-rich northward. Red crinoidal limestone, identical to the much thicker units in the Maubisse region, is interlayered with slate and phyllite of the Aileu Complex. A similar transition is also found to the north of Hatolia (Prasetyadi and Harris, 1996) (Fig. 4). The various Permian formations of Timor clearly interdigitate, and it is reasonable to conclude that all the Permian successions accumulated in one heterogeneous depositional domain. The Maubisse Formation largely represents shallow marine environments deposited on topographic highs, formed as volcanic edi®ces capped by carbonate banks and more substantial carbonate platforms, whilst the Atahoc and Cribas formations represent deeper water deposits laid down in basins formed as faulted grabens (Fig. 9). The Aileu `Formation' is in part the metamorphic equivalent of the Maubisse, Atahoc and/or Cribas formations. 4. The Permo-Triassic boundary When the faunal richness of both the Permian and Triassic successions is considered (a second database compiled for the Triassic has so far recorded over 1300 species),

T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774

Timor should be an ideal place to accurately delineate the Permo-Triassic boundary. Unfortunately, a well-exposed Permo-Triassic boundary section has proved dif®cult to ®nd. In the faunally rich shelf carbonate successions the boundary, where it has so far been determined, is either an unconformity or a condensed sequence. Even in the less fossiliferous basinal successions, because of the absence of latest Permian±earliest Triassic fossils and the evidence for condensed sedimentary sequences, an apparent conformity cannot be con®rmed. In East Timor several potential Permo-Triassic boundary sections, not yet suf®ciently documented, can be identi®ed. The ®rst is the type section of the Cribas Formation, which apparently passes upward conformably into a Middle Triassic clastic/carbonate succession. Another potential Permo-Triassic boundary section, at Mount Lilu at the eastern end of the Hili Manu Range west of Manatuto (Fig. 4), was outlined by Berry et al. (1984). In the lower cliffs of Mount Lilu red ammonoid-rich limestones are overlain by red and grey shales and ®nally, at the mountain top, by white, well bedded limestones with chert lenses. Lithologically this section appears to mark a passage from the Permian Maubisse Formation into the Triassic Aitutu Formation. One spot sample from a red limestone near the transition from the red limestones into the overlying shales was dated as late Scythian by Berry et al. (1984) using conodonts. Elsewhere in the immediate area, spot samples include late Scythian ammonites in a red limestone, late Scythian ammonites and conodonts in a brown limestone, Anisian cephalopods in a red limestone, and upper Ladinian±Carnian dark grey conodont-bearing limestones (Nakazawa and Bando, 1968). In this area red limestones, typical of the Permian Maubisse Formation, are therefore found to extend upwards into the Early and Middle Triassic. The discovery of Atomodesma-bearing siliciclastics of the Cribas Formation beneath the Aitutu Formation in Maubisse village by Hunter (1993) provides another potential Permo-Triassic boundary section. The Triassic forms, Paragondolella, Epigondolella postera, Epigondolella quadrata, Metapolygnathus and Halobia, were identi®ed in the overlying limestones and siliciclastics of the Aitutu Formation (Hunter, 1993). In West Timor one of the better constrained stratigraphic sections across the Permo-Triassic boundary is found in the Oemofai river, in the Bisnain area of central West Timor (Barkham, 1993; Figs. 5 and 6). The Oemofai Member is underlain by a leached and dolomitised older Permian section in the ?Artinskian±Kungurian Bisnain Beds (Fig. 6). The dolomitisation probably represents subaerial exposure of the Early Permian section, either during the later part of the Artinskian±Kungurian regressive cycle, recognised elsewhere in the Bisnain area (Section 3.2), or in further regressive cycles during the Middle-Late Permian (see Section 7). The lowest 3 m of the Oemofai Member, a bivalve- (Atomodesma) and crinoid-bearing dense red micritic wacke/packstone, was interpreted by Barkham

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(1993) as a transgressive facies (Fig. 6). Upwards these pass into 9 m of grain-supported Atomodesma packstone, of Permian age. This is succeeded by 9 m of calcarenite with a pelagic ammonoid and shelly fauna. The ammonoid shells are leached, with the upper surfaces coated in ferromanganese minerals and are unidenti®able, but appear to be of Permian rather than Triassic aspect. The calcarenites were originally believed to contain Upper Permian±middle Scythian conodonts, but these have now been re-identi®ed as Norian (Upper Triassic) forms (see Section 6.5). The calcarenites are composed of prismatic shell fragments probably derived from reworking of the underlying shelly beds. There is a distinctive lack of any terrigenous clastic material. The upper beds of the calcarenite unit are typically burrowed, with mineralised in®lls of authigenic gypsum crystals replaced by calcite. These are succeeded by Norian deep water mudstones. The Permo-Triassic boundary in the Oemofai section is therefore either an unconformity, possibly cutting out the uppermost Permian and the Lower± Middle Triassic, or is represented by a condensed sequence spanning this period. No unconformity was recognised in this section in the ®eld. It will be necessary to make a careful bed-by-bed study of the Oemofai section to de®ne the position of the Permo-Triassic boundary more precisely. The Early Triassic of West Timor, as recognised by early workers (e.g. Welter, 1922), consists of limestones containing (downward) Anasibirites-, Owenites- and Meekoceraszone ammonites. These zones correspond to the upper part of the Lower Triassic, Scythian stage. At Noil Niti near Kapan, however, Permian crinoidal limestones reportedly pass into Meekoceras- zone limestones concordantly, without perceptible break or change in facies. The Lower Trias here contains the ammonites Pseudosagoceras multilobatum, Flemingites timorensis, Meekoceras indoaustralicum and M. timorense (Wanner, 1931b; Sherlock, 1947). Welter (1922) reported that between the uppermost Permian Productus and the lowest Triassic Meekoceras there is less than a centimetre of rock. Wanner (1931b), however, predicted that future work on the Scythian succession in Timor would yield a zonation for the Scythian ®ner than perhaps anywhere else in the world. Permo-Triassic boundary sections are also likely to be found in the Amarassi area of southwest Timor. The Amarassi fauna is the youngest of the classic Permian faunas of Timor (late Wuchiapingian?; see Section 5), and this same area also yields a substantial Triassic fauna, ®rmly dated at least as old as Anisian (von Arthaber, in Pakuckas, 1928). The ammonoid Episageceras noetlingi, Permian in age according to Haniel (1915), is associated with species of the Early Triassic Anasibirites horizon at Netu Kot (Smith, 1927). The Kasliu locality on the eastern edge of the Kekneno area may yield a further Permo-Triassic boundary section, as according to de Roever (1940), the Upper Permian (Wuchiapingian) succession described by Archbold and Bird (1989) is succeeded by the Lower Triassic Meekoceras ammonite zone.

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Fig. 10. Permian stratigraphic correlation chart, Timor and the Australian Northwest Shelf, also showing the various ages determined for the classic Permian faunas using different faunal groups, and a composite sea-level curve for the Permian of Timor and the Northwest Shelf. R ˆ regression; T ˆ transgression. The brick pattern represents dominant carbonate sedimentary environments and the grey shading clastic environments.

In summary, there are several localities in Timor where the Permo-Triassic boundary is likely to be present in a continuous, if usually condensed, stratigraphic section. The condensed sequences are interpeted as the result of strongly transgressive conditions in Timor at this time. High-standing sea levels across the Permo-Triassic boundary in Timor contrast with regressive conditions at the boundary implied by global eustatic sea level curves (e.g. Haq et al., 1987). In Timor the transgression was probably driven by local extensional tectonics, which were active through much of the Permian and into the Triassic (see Section 7). 5. The classic Permian fossil localities Several of the particularly rich Permian fossil localities in Timor have become well known internationally, even being accorded stage status on global stratigraphic correlation charts (e.g. Haq and van Eysinga, 1987). Haniel (1915) originally named the four main localities comprising, from the oldest in stratigraphical order: Somohole, Bitauni,

Basleo and Amarassi (Figs. 2 and 10). Additionally the Atsabe (Hatu Dame) locality from East Timor was placed between Somohole and Bitauni by Haniel (1915). Subsequently the Tae Wei locality (intermediate in age between Bitauni and Basleo: de Marez Oyens, 1938) and the Lidak locality (intermediate between Somohole and Bitauni, approximately equivalent to Atsabe/Hatu Dame: Simons, 1940) were added to this list from West Timor. The stratigraphic setting of these classic fossil localities is, however, not in general well understood, as the fossils were for the most part obtained either from soil or from eroded meÂlange, and frequently samples were purchased from local people. Consequently the stratigraphic relationships between the Timorese `stages' is not known in detail. In this section we brie¯y outline the composition of the various faunas and describe the local geology for some of the more important fossil localities, so that the context of the fossil collections may be better appreciated. In previous studies it has generally been assumed that the fauna from each locality represents a single relatively short time interval. As will become apparent, however, this is incorrect because most of the localities yield different faunal

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ages from different fossil groups. The various ages derived from the different faunal groups are plotted on Fig. 10. 5.1. Somohole (Fig. 2) The Somohole locality is apparently situated in the norhern part of the area mapped by Barkham (1993), although he was unable to relocate it. A mountain named Sonmahole (or Somnahole) is located about 9 km north of Bisnain village (Barkham, 1993, Fig. 3.1) (Fig. 5). According to Soejono Martodjojo (personal communication to W.M. Furnish and B.F. Glenister in Saunders, 1971) the Somohole locality is situated on the NW slope of the mountain, with the fossils preserved in a distinctive dark-weathering tuff. However, Barkham found no rocks of the age conventionally applied to the Somohole fauna (late Asselian±early Sakmarian: Grant, 1976) in that area. Furthermore, brief lithological descriptions of the Somohole locality by Grunau (1953) (we have not yet been able to trace the original descriptions) suggest that the Somohole locality forms either part of the Atahoc Formation or a transitional zone from the Atahoc Formation into the Maubisse Formation. Barkham found no Atahoc Formation in this area. In the Nekmelalat river to the west of Mount Sonmahole, the fauna is dated as Artinskian±Kungurian, and lithologies are typical of the Maubisse Formation. The fauna from Somohole in our database is relatively small (28 species), comprising sixteen ammonoid species (57%), seven crinoids (25%) and four brachiopods (15%). This fauna is comparable to the Lidak and Bitauni faunas (see below) in being dominated by ammonoids, but on the other hand the Somohole fauna contains a signi®cantly higher proportion of crinoid species than Bitauni. The age of the Somohole fauna based on ammonoids is late Asselian±early Sakmarian, most likely the latter (Grant, 1976; Glenister and Furnish, 1987). Echinoderms, in contrast, suggest a slightly younger, late Sakmarian±early Artinskian, age (Webster, 1998a). The transition from a deeper water succession containing a dominantly ammonoid fauna into a shallower water succession dominated by crinoids is consistent with a transition from the Atahoc Formation into the Maubisse Formation at the Somohole locality. 5.2. Hatu Dame (Atsabe) (Fig. 2) The fossil fauna collected from Hatu Dame near Atsabe is the richest Permian fauna so far discovered from East Timor. Based on cephalopods examined by Haniel (1915), this fauna was placed in a stratigraphic position intermediate between Somohole and Bitauni, and was subsequently identi®ed as approximately contemporaneous with the Lidak fauna collected by Simons (1940) in West Timor. Gerth (1950) suggested that the Hatu Dame fauna was Sakmarian in age, stratigraphically closer to Somohole than to Bitauni. Wanner (1956), however, argued for an Artinskian age, closer to the Bitauni fauna. The geology of Hatu Dame was described brie¯y by

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Wanner (1956), quoting Weber (unpublished). The locality is situated on the NW slope of the Ramelau Range, which in this area is predominantly formed of Triassic sandstones and claystones with basic eruptives, together with Permian limestones and red shales. Hatu Dame itself is a rocky summit (a `fatu' in Timor) 30±40 m high composed of a fossil-rich red crinoidal breccia and red shales. Our database contains 45 species from Hatu Dame, dominated by 18 coral species (40%), fourteen brachiopod species (31%) and eleven cephalopods (27%, including one nautiloid species). It may be signi®cant, however, that whilst all the cephalopods identi®ed are Early Permian types, eight species of brachiopod, four corals and one crinoid found at this locality have also been found in the supposedly Middle-Late Permian Basleo fauna, and all the other coral, brachiopod, crinoid and blastoid species have either been found only at Hatu Dame or at other undated localities in East Timor. This might suggest that the Hatu Dame locality includes both Lower and Middle-Upper Permian elements. Alternatively, the recognition of Basleo-type shallow marine faunal elements at Hatu Dame may support Webster's (1998a) suggestion (see below) that the Basleo fauna also contains a substantial Artinskian component. As the Hatu Dame succession is described as comprising crinoidal breccia and red shales (presumably interbedded), the latter interpretation seems most likely. 5.3. Lidak (Fig. 2) The Lidak fossil localities described by Simons (1940) are situated along the southern side of the Lidak mountains, 6±10 km SW of Atambua. They were described as small, isolated outcrops within the Sonnebait Series (elsewhere in West Timor equated with the Bobonaro MeÂlange Complex). The fossil-bearing rock is described as very similar in composition to that at Bitauni (see below), comprising dark red and brown limestone, typically crinoidal, often marly with much volcanic material. Also as at Bitauni, most of the fossils were collected from soil. In addition to ammonoids these localities yield brachiopods, bivalves, bryozoans and a few gastropods. Amygdaloidal pillow basalts also outcrop in this area. The Lidak fauna recorded in our database consists of only 17 species, comprising 10 ammonoids and one nautiloid (65% together), four brachiopods (24%) and two trilobites (12%). The fauna therefore again appears to be dominated by ammonoids as at Somohole and Bitauni, stratigraphically above and below Lidak. The composition of the Lidak fauna so far described, however, is likely to be severely skewed to the more obvious and striking elements of the fauna, and particularly in favour of the stratigraphically more useful ammonoids. Notably no crinoid species are recorded in our database from Lidak, despite the lithological description of crinoidal limestones. In addition, the bivalves, bryozoans and gastropods mentioned by Simons (1940) have also apparently never been described.

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Fig. 11. Simpli®ed geological map of the Basleo area, based on Wanner (1931b), de Marez Oyens (1940), de Waard (1955) and reconnaissance ®eldwork by TRC. Note that the area shown as Maubisse Formation indicates the dominant lithology in the meÂlange unit which also includes Triassic, ?Cretaceous and Palaeogene rocks. Base map from Bakosurtanal (1993).

5.4. Bitauni (Fig. 2) Bitauni village is located on the Kefamenanu±Atambua road, on the western edge of the area mapped by Barkham (1993) (Fig. 5). Two prominent hills with outcrop are surrounded by tilled ®elds, and to the south in the river Maubesi or Maubisse (the name unconnected with the Maubisse type locality) are outcrops of pillow lavas and intercalated tuffaceous limestones. The fossils from the classic Bitauni locality are found loose in the topsoil of the ®elds, and therefore their relationship to the local geology cannot be determined directly, although rock adhering to fossils consists of volcaniclastic mudstone similar to that in nearby outcrops. The outcrop areas also yield a less abundant fauna of Artinskian±Kungurian age, similar to that of the main fossil locality (see also Burck, 1923). The age of the Bitauni fauna has not yet been satisfactorily established. Smith (1927) interpreted the Bitauni fauna as comparable in age to the Leonard Formation of Texas, now approximately equated with the late Artinskian and Kungurian.. Waterhouse (1970) reported a Baigendzinian (late Artinskian) age based on ammonoids, but assigned an early Artinskian age based on brachiopods. Grant (1976) assigned a late Sterlitamakian (late Sakmarian) to Artinskian age range. NWA (Section 6.2) considers that the Bitauni brachiopod fauna correlates with those of the

Lower Ratburi limestone of Thailand and the Amb Formation of the Salt Range, Pakistan, indicating a Middle Permian, Roadian or even Wordian (Kazanian) age, considerably younger than the late Artinskian±early Kungurian age commonly assigned to the Bitauni fauna. Similarities with bivalves from Western Australia also suggest a Roadian age (NWA). The Bitauni fauna sensu stricto (a suite of ammonoids collected by Barkham from soil at Bitauni) was assigned a late Artinskian age by Barkham (1993), in agreement with the age determination based on ammonoids given by Waterhouse (1970). Samples from the Maubisse Formation in the wider Bisnain area to the east of Bitauni range from late Sakmarian to Late Permian (and younger), with the majority of Artinskian±Kungurian age (Barkham, 1993). The fauna identi®ed from Bitauni (101 species in our database) is considerably smaller than that at Basleo (see below), but is nevertheless substantial. At Bitauni ammonoids dominate, comprising 45% of identi®ed species, whilst our database records only four species of crinoid from Bitauni (4% of the fauna identi®ed). This contrasts strongly with Basleo which is dominated by crinoids (50% of the fauna by species), with ammonoids relatively less abundant (4% of species). Other important faunal elements at Bitauni are brachiopods (18 species, 18%), corals (15%) and gastropods (7%). For ammonoids the difference in the

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number of species between Bitauni and Basleo is also signi®cant (45 and 23 respectively). Wanner (1956) recorded more than 800 individual specimens collected from Bitauni by the various sampling expeditions, whilst Wanner (1931a) recorded only 403 ammonoids from the geographically larger Basleo area. 5.5. Tae Wai The Tae Wei locality is situated about 5 km NE of Basleo (de Marez Oyens, 1938, 1940; Fig. 11). The rich ammonoid fauna reportedly weathers out of violet-brown tuffaceous marls similar to some of the layers at Bitauni. The faunal content of Tae Wei was outlined by de Marez Oyens (1938) and Gerth (1950), but has apparently never been described in detail (Glenister and Furnish, 1987). The Tae Wei fauna consists of 48 species in our database. The dominant element is ammonoids, with 23 species identi®ed (48% of species recognised at the locality). The other major elements of the fauna include gastropods (14 species, 29%) and crinoids (six, 13%). In the predominance of ammonoids the Tae Wei fauna is similar to the Bitauni fauna, rather than to the fauna of the geographically closer Basleo locality. It is also similar to Bitauni in terms of the ammonoid species present (de Marez Oyens, 1938). Of six ammonoid species that are recognised at Tae Wei and either at Bitauni or Basleo, all six are recognised at Bitauni, but only a single species (Adrianites cancellatus) has also de®nitely been identi®ed at Basleo. Thus although the Tae Wei fauna lies stratigraphically between Bitauni and Basleo, it more closely resembles Bitauni. Tae Wei is considered late Early Permian or Middle Permian in age, closer to the late Artinskian (?) Bitauni fauna than to the supposedly Middle or Late Permian (Wordian?) Basleo fauna (see Section 5.6). Gerth (1950) considered the Tae Wei ammonoids to be equivalent to the Sosio fauna of Sicily. Glenister and Furnish (1987) suggested that the Tae Wei fauna was either an admixture (human and/or tectonic!) of Baigendzhinian± Roadian and Wordian ammonoids, or a single fauna of intermediate (Wordian) age. However, this latter interpretation is not consistent with the rather older Artinskian±Kungurian age usually applied to the Bitauni fauna, and the Wordian age standardly applied to the less similar Basleo ammonoids. On the other hand if the Bitauni fauna is late Kungurian±early Roadian in age as interpreted by NWA from the brachiopod fauna, then the discrepancy would be less. 5.6. Basleo (Figs. 2 and 11) The Basleo locality is the faunally richest of the classic Timor sites. About half of the Permian species identi®ed from Timor have been recognised at Basleo, or from a small number of nearby localities. For the Basleo localities our database contains between 565 and 660 species, depending on the de®nition of the Basleo locality. For the stricter de®nition, 280 species (50%) are crinoids. The other main faunal elements are corals (17% of species identi®ed),

739

brachiopods (12%) and gastropods (8%), whilst only 23 ammonoid species have been identi®ed (4% of species at Basleo). This is in marked contrast with Bitauni (see above), where ammonoids are the dominant element in the fauna. Basleo is located 6 km ENE of Nikiniki, near the southern margin of the Neogene Central Basin (Figs. 2 and 11). The village of Basleo is situated on probable Triassic ¯ysch of the Babulu Formation, but in the river Noil Tobe, immediately to the north, is Bobonaro Complex meÂlange with abundant Permian fossils weathering into the soil, including crinoid stems, bryozoans, solitary corals and ammonoids. In addition shark's teeth are found in the soil, probably derived from the Early Cretaceous Nakfunu Formation which is also entrained into the meÂlange in this area. To the east, near the junction of Noil Tobe with the larger Noil Bunu river are solid outcrops of Maubisse Formation limestone (predominantly crinoidal limestone) and basaltic volcanics. The structure of the Basleo area appears chaotic at ®rst sight, typical of areas dominated by the Bobonaro MeÂlange Complex. However, reconnaissance geological mapping by de Waard (1955) suggests that, whilst the structure of the dominantly argillaceous sections is indeed chaotic, or at best dif®cult to determine because of extensive landsliding, more competent horizons such as the volcanic layers display a rather gentle style of folding along WNW±ESE axes (Fig. 11). No signi®cant geological ®eldwork has been carried out in this area since the 1950s, and considering its palaeontological importance, this area needs detailed geological mapping and systematic sampling from outcrop to better constrain the stratigraphic position and palaeoenvironmental signi®cance of the Basleo fauna. The age of the Basleo fauna is problematic. Based on ammonoids, this fauna was originally equated with the middle Productus Limestone of India by Haniel (1915), and with the Word Formation of Texas and the Sosio fauna of Sicily by Smith (1927). Wanner (1931a), however, considered the ammonoids somewhat younger than Sosio and the Wordian. Gerth (1950) assigned the Basleo ammonoids to a Basleo stage and a Timorites ammonoid zone, lower than the Cyclolobus zone of Amarassi, but higher than the Sosio stage (Tae Wei-equivalent) Waagenoceras zone. Based on brachiopods, Grant (1976) dated the Basleo fauna as early Kazanian. NWA considers the Basleo brachiopods slightly younger than Kazanian, probably early Wuchiapingian. In contrast, GDW considers many of the echinoderms to indicate a rather older, Artinskian age. For the bryozoans, Gilmour and Morozova (1999) have interpreted an U®mian (Roadian, early Middle Permian) age. Thus these four faunal groups from Basleo yield mutually exclusive ages. A key question is therefore whether or not all the various Basleo fossil localities are indeed contemporaneous. Wanner (1931a) considered all the fossil sites to be approximately the same age, based in particular on the dominant occurrence of the crinoid Timorocidaris sphaeracantha and

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the common occurrence of the blastoid Schizoblastus (now Deltoblastus) permicus at nearly all localities. One obvious solutionÐreworking of Artinskian echinoderms into a Middle-Late Permian basinÐappears to be untenable because of the excellent state of preservation of the echinoderm fauna. In the absence of suf®ciently high quality ®eld geological data linked to the palaeontological data, we cannot provide a de®nitive answer to this dilemma. On balance, however, we prefer the interpretation that there are several stratigraphic horizons represented in the Basleo area, ranging in age from Artinskian to early Wuchiapingian, with Artinskian/Early Permian faunas dominated by echinoderms, followed by a Roadian/early Middle Permian shallow marine fauna containing bryozoans; a Wordian/ later Middle Permian deeper marine fauna dominated by ammonoids; and an early Wuchiapingian/Late Permian shallower marine fauna dominated by brachiopods. 5.7. Ajer Mati (Fig. 2) This was the ®rst Permian locality identi®ed in Timor (Beyrich, 1862, 1865; Rothpletz, 1891, 1892; Wichmann, 1892), in a usually dry streambed (hence the name, `dead water') within present-day Kupang city. Wichmann (1892) indicated a coherent stratigraphic section along the Ajer Mati river, but the description given by Verbeek (1908) suggests that the outcrops are composed of meÂlange of the Bobonaro Complex. Lithologies encountered by Verbeek include crinoidal limestone, red calcareous and sandy clay-slates, sandy red and green limestones, serpentinite and diabase. No recent studies have been reported from this locality, and it is possible that it has been lost beneath urban development. 5.8. Kasliu The geology of the Kasliu locality, situated on the topographic shoulder between the Kekneno area to the west and the Mutis massif to the east (Fig. 7) has been described in detail by de Roever (1940) and Bird (1987). Archbold and Bird (1989) documented a brachiopod fauna, dated as Chhidruan (Wuchiapingian), younger than Kazanian, but older than Dzhul®an, collected from outcrop in this area. These authors correlate the Kasliu fauna with the brachiopod faunas of Basleo and Amarassi. The Kasliu locality is one of the few places in Timor where an uncondensed stratigraphic sequence is accurately dated as Upper Permian. 5.9. Amarassi (Fig. 2) As at Ajer Mati, Wichmann (1892) reported coherent Permian strata in the Kasimuti river south of Baun, whilst Verbeek (1908) encountered what would now be considered Bobonaro Complex meÂlange. Many of the fossils attributed to the Amarassi fauna were collected from isolated limestone blocks within river courses, rather than from outcrop. Burck (1923) provides a

detailed description of the individual blocks sampled from this locality during the 1916 expedition. In our database the Amarassi fauna consists of 152 species, larger in terms of species numbers than Bitauni, but considerably smaller than Basleo. It also differs from these other localities in being composite, with specimens collected from several localities in the extreme southwest of Timor, including the Ajer Mati locality and sites near Baun. The main elements of the Amarassi fauna include brachiopods (28% of the fauna identi®ed), bryozoans (18%), crinoids (16%), corals (13%) and cephalopods (9%, including two nautiloid species). Unlike the other classic Timor faunas the Amarassi fauna is not dominated by a single faunal group. The Amarassi fauna is the youngest of the classic Timor faunas. Haniel (1915) originally equated it with the upper Productus Limestone of India, and Gerth (1950) placed it in the Cyclolobus ammonoid zone (Chhidruan stage). More recently it has been dated as late Kazanian±early Dzhul®an by Grant (1976), and Chhidruan (Wuchiapingian), immediately post-Kazanian by Archbold and Bird (1989). Archbold now considers the Amarassi fauna to be probably late Wuchiapingian.

6. Palaeontology: Systematics and palaeogeographic signi®cance 6.1. Brachiopods (NWA) The Permian brachiopod assemblages of Timor have received very little modern study, apart from the investigations by Archbold and Bird (1989), Archbold and Barkham (1989) and Archbold (in Riding and Barkham, 1999). Nevertheless, it has long been recognised that in the classic fossil localities the `Bitauni' brachiopod fauna is of Kungurian±early U®mian age and that the `Basleo' and `Amarassi' brachiopod faunas are considerably younger (Waterhouse, 1976). These age estimates are understandable, because in the early 1970s less was understood about the middle Permian, and the Baigendzhinian, as originally proposed, included the Kungurian. Correlation of the Bitauni fauna with that of the Lower Ratburi Limestone of Thailand and the Amb Formation of the Salt Range, Pakistan indicates a Roadian or even Wordian (Kazanian) age in modern terms. Bivalve links with Australia also point to a Roadian age. Modern studies, describing discrete assemblages from individual localities, with at least some stratigraphy, are essential for resolving the full extent to which the Permian is represented in the geology of Timor. As presently understood, the Permian brachiopod faunas of Timor fall into three main age brackets best signi®ed by the locality names Bisnain, Bitauni and Basleo/Amarassi. The oldest assemblage is that known from Bisnain which is regarded as being late Sakmarian (Sterlitamakian) in age and may be contemporary with the Somohole ammonoid

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fauna or, more likely, just a little younger (Archbold and Barkham, 1989). The Bisnain assemblage includes representatives of the genera Arctitreta, Callytharrella, Stictozoster, Elivina, Neospirifer and Cyrtella that assist in constraining the age and are also signi®cant in terms of links to the contemporary faunas of Western Australia. Warm water genera include Martinia and Spirigerella. A mid-Permian fauna is indicated by brachiopods from Bitauni and correlatable localities such as Laktutus (Archbold, in Riding and Barkham, 1999), and some of the localities provided by Hamlet (1928). This fauna has links with the brachiopod fauna of the lower Ratburi Limestone of Thailand and is of Kungurian to U®mian age (i.e. extending into the Roadian) as discussed by Waterhouse (1981), Archbold (1999). The Bitauni fauna requires modern study but the presence of genera including Cimmeriella and Retimarginifera suggests links to Western Australia, and the Neospirifer record indicates a species of moderate size only, as compared with the large species known from many Late Permian faunas of the peripheral margin of Gondwana. The youngest Permian brachiopod assemblages from Timor are those from Basleo (including the Kasliu fauna, Archbold and Bird, 1989) and Amarassi. In modern terms, these two sub-assemblages are apparently early and late Wuchiapingian in age and respectively match the two Wuchiapingian faunas known from Western Australia (Archbold, 2000a,b). These Late Permian brachiopod assemblages from Timor include distinctive species of Chonetella, Waagenoconcha, Transennatia, Stenocisma, large Neospirifer, large Spiriferella, Martinia and Squamularia that are age-diagnostic and provide biogeographical links to both Western Australia and peripheral Gondwanan regions. The faunal provinciality of Permian brachiopods throughout Southeast Asia and Australia has been outlined by Shi and Archbold (1995, 1998), and Archbold and Shi (1996). In Sterlitamakian±Aktastinian (late Sakmarian±early Artinskian) times, Timor formed part of the Westralian province, together with sites in western Australia, western New Guinea and several stations in India and the Himalayas. At the same time an `Incipient Cimmerian' province including western Malaya and south Thailand (Sibumasu Terrane) was developing, in addition to the Cathaysian province of Southeast Asia. By late in the Early Permian (Baigendzhinian±early Kungurian), the Cimmerian province had divided into the Himalayan and Sibumasu subprovinces (Archbold and Shi, 1996). Timor formed part of the Himalayan subprovince, together with several faunas from northwest, central and south Tibet (Lhasa Block: Shi and Archbold, 1995). The Sibumasu subprovince includes faunas from Thailand, southern and northern Sumatra, and additionally western New Guinea, which together with Timor had formerly formed part of the Westralian province. By the Late Permian (Kazanian±Midian), only a southern Austrazean province and the more equatorial Cathaysian

741

province were recognised regionally. Faunas from Timor were not speci®ed in the study by Archbold and Shi (1996), because the Basleo and Amarassi faunas are slightly younger (Wuchiapingian). However, Archbold (1988) interpreted the Bitauni brachiopod fauna as showing links to that of Western Australia, and Grant (1976) correlated the Basleo brachiopods with the fauna from the Kalabagh Member of the Wargal Limestone in the Salt Range of Pakistan. Timor therefore continued to show af®nities in brachiopod faunas with Australia and parts of the Indian subcontinent during the Late Permian (Archbold, 2000b). 6.2. Bryozoans (PDT) The main source for information on Permian bryozoans from Timor is the monograph by Bassler (1929). Using the collections of J. Wanner, G.A.F. Molengraaff and H.A. Brouwer, Bassler described more than 70 species. However, he was apparently unaware of two earlier papers, by Beyrich (1865), in which three bryozoans are described and illustrated, two as new species, and Rothpletz (1892), in which a further two species are recorded, one in open nomenclature (Polypora sp.) and the second referred to a species (Fenestella virgosa Eichwald) originally described from the Carboniferous and Permian of Russia. Without recourse to the original material it is impossible to know whether any of the species discussed by Beyrich and Rothpletz are synonymous with Bassler's species: the illustrations and descriptions of both earlier authors are inadequate for precise taxonomic identi®cation. Ross (1978) and Sakagami (1985) both considered the bryozoans from the Permian of Timor to belong to a `Southern Tethys Region'. Although Gilmour and Morozova (1999) recognised a distinct Timor Province within the `Tropical Climatic Zone', their palaeogeographical map (based on Scotese and McKerrow, 1990) depicts Timor at a palaeolatitude greater than 308 and therefore outside the tropics. Uncertainties over the exact stratigraphical age of the bryozoans from the Timor Permian were noted by Ross (1979) who regarded the Bitauni Beds as Artinskian, the Basleo Beds as U®mian±Kazanian (Roadian±Wordian) and the Amarassi Beds as Kazanian (Wordian) (see Ross, 1979, Fig. 1). Gilmour and Morozova (1999), on the basis of bryozoan species shared between Timor and other provinces (China, Japan, South±Mongolia±Ussuria, Thailand± Malaysia), concluded that the bryozoan assemblage from the Basleo locality was of U®mian (Roadian) age and those from the Amarassi localities of Kazanian (Wordian) age. A massive turnover in bryozoan faunas occurred around the Palaeozoic±Mesozoic transition, but the extent to which this pivotal phase in bryozoan evolutionary history was due to a single end-Permian mass extinction is still unclear. According to Gilmour and Morozova (1999), extinction of bryozoans in the late Permian occurred gradually, such that

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Table 1 Nomenclature of stages and substages used in discussing the marine Permian ammonoid faunas of Timor (Section 6.3). Stages and substages identi®ed in Timor are indicated in bold

Late Permian

Stages

Substages

Tatarian

Dorashamian, Dzhul®an Chhidruan (including Amarassian) Wordian (including Capitanian)

Kazanian ( ˆ Guadalupian) Early Permian

Artinskian Sakmarian Asselian

Roadian (including Kungurian part and U®mian) Baigendzhinian (including Kungurian part) Aktastinian Sterlitamakian Tastubian

relatively few species were left by the latest Permian (see also Erwin, 1993). Furthermore, some of the major `Palaeozoic' groups which have been traditionally regarded as becoming extinct in the Permian are now known to have survived into the Triassic (Taylor and Larwood, 1988), albeit in low numbers. Prominent among the groups surviving into the Triassic are trepostomes. However, trepostomes are not especially well represented in the Permian of Timor, where they constitute only about 14% of bryozoan species. They are therefore outnumbered by both fenestrates (38%) and cystoporates (28% of species), the former not known with certainty from the Triassic and the latter very rare (SchaÈfer and Fois, 1987). The other suborders present in Timor are cryptostomes (18%) and ctenostomes (3%); no cyclostome species have been recorded. Without good data on the broader facies context of the Timorese bryozoans described by Bassler (1929) it is dif®cult to draw conclusions about palaeoenvironments. In general, however, modern and fossil bryozoans tend to live at shelf depths (,200 m) in environments where ambient water ¯ow is suf®cient to prevent anoxia and to replenish planktonic food resources. Hard or ®rm substrates for attachment are essential for most bryozoan species. For example, Bassler (1929, plate 227, Fig. 4) illustrated a colony of Fistulipora timorensis encrusting a brachiopod shell. Some Timorese bryozoans (e.g. Eridopora oculata, Bassler, 1929, plate 225, Fig. 5) encrusted crinoid columnals and it is possible that they colonised the stems of living crinoids that raised the colony to a higher tier among suspension feeders and possibly permitted survival in habitats where the sea-bed itself was anoxic. 6.3. Cephalopods (Ammonoids and Nautiloids) (HGO) The Permian ammonoid fauna of Timor is one of the best described and most diverse known (see Appendix A). The fauna has been well illustrated by Haniel (1915), Smith (1927) and collections are to be found in many museums, mainly of specimens derived from limestones, particularly from the Maubisse Formation. This skewed fauna indicates that sediments of Early Permian Sakmarian and Artinskian

(including Kungurian) age and Late Permian Kazanian and early Tatarian age occur in Timor. The scheme of stages and substages used for the ammonoids in this contribution (Table 1) does not conform exactly to the recommendations of the meeting of the Subcommission on Permian Stratigraphy in 1996 or to the scheme of Jin et al. (1997) derived from it. This is because Permian ammonoid faunas show a clear distinction between forms with a Late Carboniferous af®nity (Early Permian) and those which presage the Mesozoic faunas (Late Permian) so that it is dif®cult to de®ne the Middle Permian. Further research is necessary to correlate accurately the ammonoid faunas with the zonal schemes developed for the conodonts and fusulinids as given by Jin et al. (1997). Unfortunately, there have been problems in the precise dating of the Timor ammonoid faunas, and even in determining their relative ages. The classic ammonoid collections, as for other fossil groups, were often collected loose, and were not always precisely located geographically, let alone stratigraphically. For this reason it has been dif®cult to place the Timor faunas within a global framework of stages and substages. The Permian of Timor has, therefore, tended to be disregarded for correlation purposes (e.g. Glenister et al., 1993). In addition, the classi®cation of the Permian worldwide is itself the subject of continuing debate (Stepanov, 1973; Furnish, 1973; Archbold et al., 1993; Glenister et al., 1993). Recent studies in measured sections from the Permian of Timor, notably by Barkham (1986, 1993), Bird and Cook (1991), are of great value, as they have provided much more precise stratigraphical information than was available previously, and have permitted, in some instances, the matching of some of the earlier collections with the known stratigraphical succession. They also provide comparative data with faunas obtained, for example, in Western Australia (Archbold et al., 1993; Glenister et al., 1993). Barkham (1986) collected the following ammonoids from black shales of the Atahoc Formation ,10 m below pillow lavas in the Noil Bijau (his Locality 29) in the Bisnain area, West Timor (Figs. 5 and 8): Metalegoceras tschernyschewi

T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774

(Karpinsky), Metalegoceras sp., cf. Eoasianites beluense (Haniel), Peritrochia dieneri (Smith), Peritrochia gracilis (Smith), Stacheoceras arthaberi Smith, Stacheoceras tridens (Rothpletz), Neopronorites timorensis Haniel, Artinskia timorensis (Haniel). This stratigraphically well constrained faunal assemblage is ascribed to the Lower Permian, Sakmarian stage, Sterlitamakian substage. He also obtained Atsabites sp., indicating an Artinskian, possibly early Aktastinian age, from red calcarenites higher in the section. Bird (1987) collected the following ammonoids from the `Goniatite Beds' (250 m thick) in the Atahoc Formation, outcropping in the River Tunsip, near Nuapin, Kekneno area, West Timor (Figs. 7 and 8): near the base of the section Eoasianites beluense (Haniel), E. somoholoense (Haniel), Metalegoceras sp., Neopronorites timorensis Haniel, Neopronorites sp., Peritrochia timorense (Haniel), P. dieneri (Smith), cf. Agathiceras sp.; at 40 m Eoasianites beluense (Haniel); at 81 m (N.B. this is the same horizon as locality B of de Roever, 1940) Metalegoceras evolutum (Haniel) (some of large size), Neopronorites timorensis (Haniel), Peritrochia timorense (Haniel), P. dieneri (Smith) Stacheoceras arthaberi Smith; at 130 m Peritrochia timorense (Haniel) and a specimen of Eoasianites beluense (Haniel) was collected by Audley Charles (personal communication); at 160 mÐEoasianites sp. juv., Neopronorites timorensis (Haniel), Peritrochia timorense (Haniel), Agathiceras cf. A. sundaicum Haniel; at 197 mÐspecimens affected by dolomitization Metalegoceras cf. M.evolutum (Haniel), ?Eoasianites sp., Peritrochia dieneri (Smith); at 218 mÐMetalegoceras sundaicum (Haniel). These ammonoids indicate a Sakmarian (Sterlitamakian substage) Early Permian age, but a specimen of Jurasinites hanieli (Smith), obtained from an uncertain horizon near the base of this section may indicate an earlier Sakmarian (Tastubian) age for the basal beds of this succession. In addition Bird (1987) collected Eoasianites somoholense (Haniel), Neopronorites timorensis (Haniel), Peritrochia dieneri (Smith), P. gracilis (Smith), P. timorense (Haniel), Stacheoceras arthaberi Smith, from the Noil Nenbas (De Roever, in Simons, 1940); Peritrochia dieneri (Smith), P. timorense (Haniel) and loose, but possibly derived from this section, ?Metalegoceras evolutum (Haniel), Stacheoceras arthaberi Smith, from the upper part of the section in the Noil Bisane, and Peritrochia timorense (Haniel) in situ and P. dieneri loose, but probably derived from this section. All of these assemblages indicate an Early Permian, Sakmarian (Sterlitamakian) age. The following stratigraphically unconstrained ammonoid faunas have been reported from the classic Permian fossil localities of Timor (see Section 5): Somohole (Smith, 1927) (Fig. 5): Agathiceras sundaicum Haniel, Eoasianites beluense (Haniel), Eoasianites hanieli (Smith), Eoasianites melanesianum Smith, Peritrochia dieneri Smith, Metalegoceras somoholensis (Haniel), Metalegoceras evolutum (Haniel), Peritrochia timorense

743

(Haniel), Popanoceras boesei Smith, Stacheoceras dieneri (Smith), Sicanites papuanus (Smith), Neopronorites timorensis (Haniel). As reported above the geographical position of this locality (which has been used as a stage name) is uncertain (Section 5.1). The age of this fauna is essentially Early Permian Sakmarian (Sterlitamakian) but Sicanites papuanus in particular, suggests that later material of Artinskian (Baigendzhinian) age is also present. Hatu Dame (Atsabe) (Fig. 4) (Haniel, 1915; Wanner, 1956): Crimites oyensi (Haniel), Agathiceras sundaicum Haniel, Atsabites weberi Haniel, Metalegoceras sundaicum (Haniel), Peritrochia timorensis (Haniel), Popanoceras indoaustralicum Haniel, Propinacoceras transitorium Haniel, Stacheoceras dieneri (Smith), Neopronorites timorensis (Haniel), Thalassoceras dieneri Smith. Gerth (1950) considered this rich ammonoid fauna to be of Sakmarian age, whereas Wanner (1956) considered it to be Artinskian. A range of ages is indicated, from late Sakmarian (late Sterlitamakian) to early Artinskian. Lidak (de Roever, in Simons, 1940): Adrianites ex. aff. cancellatus globosa (Haniel), Agathiceras cf. A. sundaicum Haniel, Daraelites submeeki Haniel, Hyattoceras waageni Smith, Eothinites pseudomeneghinii (Haniel), Atsabites weberi Haniel, Peritrochia dieneri (Smith), Peritrochia gracile (Smith), Metalegoceras sundaicum (Haniel), Propinacoceras simile Haniel, Stenopronorites timorensis (Haniel), Waagenoceras lidacense de Roever. These specimens suggest an Artinskian (Baigendzhinian) age, but the presence of Waagenoceras suggests that later, Roadian, material is also present. They were obtained loose from the `Sonnebait Series' (Section 5.3) (Fig. 3) Bitauni (Barkham, 1986) (Fig. 5): Metalegoceras evolutum (Haniel), M. gigas (Smith), M. sundaicum (Haniel), M. tschernyschewi (Karpinsky), Stenpronorites timorensis (Haniel), cf. Propanoceras hanieli Smith, Adrianites sp., Agathiceras brouweri Smith, A. sundaicum Haniel, Artinskia timorensis (Haniel), Medlicottia sp., cf. Paragastrioceras sp., ?Ghazeloceras sp. nov., together with pseudorthoceratid nautiloids and `Endolobus' brouweri Haniel. The material suggests an Early Permian age range, from late Sakmarian (Sterlitamakian) to late Artinskian (including Kungurian) Roadian. These specimens were collected loose from soil between fatus of Maubisse Formation limestone near Bitauni (Section 5.4). Tae Wai (de Marez Oyens, 1938, 1940) (Fig. 11): Crimites cf. oyensi (Haniel), Adrianites cancellatus (Haniel), Agathiceras brouweri Haniel, Agathiceras cf. A. martini Haniel, Agathiceras cf. A. sundaicum Haniel, Artinskia simile (Haniel), Daraelites submeeki (Haniel), Epadrianites involutus Schindewolf, Hyattoceras sp., Peritrochia dieneri (Smith), Medlicottia sp., Metalegoceras sundaicum?, Stenopronorites cf. S. timorensis (Haniel), Popanoceras cf. indoaustralicum Haniel, Popanoceras sp., Propinacoceras sp., Rhiphaeites spp., Sicanites sp., Stacheoceras arthaberi Smith, Stacheoceras sp., Waagenoceras sp. The assemblage indicates a late Artinskian (Kungurian) Roadian age.

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Basleo (Haniel, 1915; Wanner, 1931a) (Fig. 11): Adrianites beyrichi Haniel, Adrianites cancellatus (Haniel), Adrianites rothpletzi (Haniel), Adrianites timorensis (Boehm), Adrianites wichmanni (Haniel), Agathiceras brouweri Smith, Cyclolobus persulcatus (Rothpletz), Episageceras nodosum (Wanner), [Gastrioceras] angulatum Haniel, Hyattoceras subgeinitzi Haniel, Medlicottia subprimas Haniel, Propinacoceras insulcatum Haniel, Propinacoceras simile Haniel, Stacheoceras tridens (Rothpletz), Sundaites levis Haniel, Timorites curvicostatus Haniel, Timorites hanieli Smith, Timorites striatus Haniel, Waagenoceras gemmellaroi Haniel, Waagenoceras intermedium Wanner, Xenaspis beedei Smith, Xenodiscus rotundus Haniel. This faunal list indicates a range of ages, from the late Artinskian (Kungurian) Roadian, into the Late Permian, including the Kazanian, Wordian and Tatarian (Chhidruan substage). Amarassi (Fig. 2): Adrianites timorensis (Boehm), Cyclolobus persulcatus (Rothpletz), Episageceras noetlingi (Haniel), Hyattoceras subgeinitz Haniel, Medlicottia subprimas Haniel, Parapopanoceras dyadicum Haniel, Stacheoceras tridens (Rothpletz), Sundaites levis Haniel, Timorites curvicostatus Haniel, Xenaspis sp., Xenodiscus rotundus Haniel. This fauna indicates only a Late Permian Tatarian age and probably Chhidruan. The name Amarassi, represents a fauna rather than a locality (Section 5.9), but it has been used as a substage name in the past (e.g. Furnish 1973). For determining the ages of the Permian stratigraphic units in Timor the sections measured and collected by Barkham in the River Bisnain area and by Bird in the Kekneno area are of particular importance (Figs. 5±8). The base of the Atahoc Formation in the Kekneno area is probably early Sakmarian (Tastubian), based on the occurrence of Jurasinites hanieli (Smith). There is no evidence of Asselian sediments on the basis of the ammonoid faunas. This is succeeded by a thick development of Sakmarian (Sterlitamakian) sediments, but there is no ammonoid evidence of the junction between late Sakmarian and early Artinskian. In the River Bisnain section, Barkham's (1986) locality 29, a late Sterlitamakian fauna in shales is succeeded by calcareous interbeds which, in the 10 m of sediment below the pillow lavas, have yielded a single Atsabites sp., probably of earliest Artinskian (Aktastinian) age. This suggests that the overlying volcanics are Artinskian. Thus, the Sterlitamakian in the Atahoc/Cribas basinal facies is now much better constrained than it has been hitherto. The remainder of the Lower Permian Artinskian (including the Kungurian) in the Atahoc and Cribas Formations is still not well con®rmed; ammonoid collections of Baigendzhinian and Roadian age having been obtained largely as soil samples with no stratigraphical control. The same applies to the shallower water Maubisse platform carbonate successions, where the range is from Sakmarian (Sterlitamakian) to late Artinskian (Roadian). Future work should be directed to elucidating these later

Early Permian successions in both the basin and platform successions. The Somohole fauna is essentially Sterlitamakian, but Sicanites papuanus (Smith) suggests a middle Artinskian (Baigendzhinian) presence in this area. Hatu Dame contains elements suggesting a latest Sterlitamakian age, but is essentially of early Artinskian age. The Bitauni area has yielded Sterlitamakian and late Artinskian (Roadian) species. Tae Wai has yielded a Roadian fauna. Basleo includes late Kungurian and Late Permian ammonoid faunas of Kazanian (Wordian) and Tatarian (Chhidruan) ages (Table 1). Upper Permian ammonoid faunas, apart from those at Basleo, are known from the Amarassi area. The earliest of these is of Tatarian (Chhidruan) age, but the presence of Dzhul®an (Lopingian) sediments in this area is also indicated. There is no certain evidence of the Dorashamian substage (Changsingian stage) at present; the latest Permian sediments and the earliest Triassic sediments appear to be absent in Timor (Section 4). Quite apart from the problems inherent in utilising the early ammonoid studies in Timor, there is considerable debate on the global superposition of the various provincial stages and substages within the Permian. The global stage and substage classi®cation of the marine Permian has been discussed recently by Archbold et al. (1993), Glenister et al. (1993), Jin et al. (1997) in which the problems of nomenclature are only too clear, and stem from the classic dilemma of geographically isolated sections, particularly in Russia, in which there is no ®rm evidence in the lithological succession for the preceding or succeeding substages (Stepanov 1973; Furnish, 1973). A simpli®ed classi®cation for the marine Permian, as used in this account, is given in Table 1. The collections made by Barkham (1993) and Bird and Cook (1991) from measured sections in Timor are of great value in this continuing debate. The Permian faunas of Western Australia have been described by Skwarko (1993) and their sparse ammonoid content by Glenister et al. (1993), who attempt a correlation with Permian successions elsewhere than Timor. Tastubian sediments with Jurasinites jacksoni (Etheridge) are developed in the Holmwood Shales in the Perth Basin, the upper part of which (Fossil Cliff Member) has yielded a Sterlitamakian fauna characterised by Metalegoceras. A similar fauna of Sterlitamakian age is present in the Poole Sandstone in the northern Canning Basin. Thus, there is good correspondence at genus level between the measured Western Australian sections and those of Timor. The relative paucity of ammonoids above the Sterlitamakian sediments make correlations with Timor impracticable. A possible direct correlation between Western Australia and the Bitauni area of Timor is provided by the occurrence of Metalegoceras kayi (Glenister, Windle and Furnish), which is very close to, if not conspeci®c with M. evolutum (Haniel), and not dissimilar to M. schucherti Miller and Furnish from the Capitanian of the United States.

T.R. Charlton et al. / Journal of Asian Earth Sciences 20 (2002) 719±774

6.4. Conodonts (RSN) In contrast to the large number of species of other Permian taxa recorded from Timor the conodont record is very sparse. Only two conodont taxonomic papers have so far been published (van den Boogaard, 1987; Nicoll and Metcalfe, 1998). Both of these papers are based on analysis of the matrix adhering to other fossils, or of samples collected for other purposes. The conodonts recorded by Berry et al. (1984) from the Maubisse Formation at Manatuto in East Timor proved to be of Triassic age. Van den Boogaard (1987) analysed material from the Maubisse Formation of West Timor held in the collections of the Geological Institute in the University of Amsterdam. The following conodont species were identi®ed: Diplognathus oertlii, Hindeodus sp., Neogondolella bisselli, Neospathodus sp. cf. N. praedivergens, Sweetocristatus sp., Sweetognathus sp. aff. S.whitei ( ˆ S. inornatus), Vjalovognathus shindyensis ( ˆ V. australis). This assemblage was identi®ed as of upper Artinskian age. Barkham (1993), as part of his study of the Maubisse Formation in West Timor, obtained conodonts from samples collected across the Permo-Triassic boundary in the Oemofai River (Fig. 5). Conodonts from what was thought to be the Permian part of the Oemofai Member were identi®ed as Neogondollela carinata (Clark) by L.Krystyn and considered to be indicative of the Late Permian. Nicoll has subsequently re-examined the sample in question (SB 98) and considers that only taxa of Late Triassic (Norian) age are present. Nicoll (AGSO Open ®le Report) and Nicoll and Metcalfe (1998) obtained Vjalovognathus australis Nicoll and Metcalfe and Hindeodus sp. from a sample (SB 18) collected by Barkham from the Khali Beds in the Maubisse Formation at a locality on the Bisnain River. Like the material of van den Boogaard (1987), this fauna was considered to be of Artinskian age. Conodont faunas of Kungurian age have been recovered from the Canning and Carnarvon basins in Australia (Nicoll and Metcalfe, 1998). There is clearly a need for a systematic sampling programme in Timor speci®cally directed at the study of Permian conodont biostratigraphy. Nicoll and Metcalfe (1998) recognised a north±south gradient in conodont species diversity in stratigraphic units of the same age in the Carnarvon and Canning Basins (Western Australia) and from Timor. Samples from the southernmost localities in the Carnarvon Basin produced the least diverse faunas and the fauna described by van den Boogaard (1987) had the greatest diversity. This diversity gradient was interpreted as re¯ecting the temperature gradient of the shallow water basins located on the Gondwanan margin of the Palaeo-Tethys (Early Permian) and Meso-Tethys (Late Permian) oceans. 6.5. Corals (JES) The Permian coral faunas of Timor are famous, not only for their richness and diversity, but also for the magni®cent

745

preservation of their skeletal carbonate. The total fauna consists of approximately 108 species of Rugosa and 25 species of Tabulata (Appendix A). These numbers are approximate only, because much of the systematic work is more than 50 years old, and descriptions of taxa lack data on population variation. This list of coral species utilises genus names as de®ned in Hill's (1981) revision of the Rugosa and Tabulata, and includes no subspecies. The faunal list contains 27 rugosan and four tabulate species that were proposed on the basis of only a single specimen; many of these will disappear when there is a systematic revision of the fauna. Gerth's (1921) monograph was the ®rst work to deal with the whole Permian coral fauna of Timor, and it forms the basis for all later studies, the majority of which have focused on the rugosans. Gerth (1921) authored 15 species and one genus of solitary rugosans, and one genus and three species of colonial corals. These colonial waagenophyllid corals were included by Minato and Kato (1965) in their revision of the Waagenophyllidae. Gerth's (1921) species, Ipciphyllum timoricum (Gerth), Lonsdaleiastraea vinassai Gerth and L. molengraaf® (Gerth) were ®tted into an evolutionary and temporal framework byMinato and Kato (1965, p. 58), who are the only modern authors to deal with these colonial forms. Gerth's (1921) work provides the best overall view of the tabulate corals and associated forms. This fauna has some taxa of rather generalised architecture, along with some forms unique to the Permian that are still not fully understood. Gerth (1921) authored six tabulate genera, as well as 18 species from this fauna, which are considerably different from those in Permian beds elsewhere in diversity, uniqueness of species, and their interaction with stalked echinoderms. Hehenwarter (1951) studied tabulate coral specimens but only from Basleo. A study by Koker (1924) of Permian corals from Timor proposed three new genera of Rugosa and discussed their skeletal microstructure and genesis. This aspect was later elaborated by Schindewolf (1942), and still later by Schouppe and Stacul (1955, 1959, 1966). The important monograph by Schindewolf (1942) on the Polycoeliidae and Plerophyllidae, is also partly based on specimens from Timor. Schindewolf (1940, 1942) proposed 16 new species of Timor rugosans, and in the 1942 monograph he also described the microstructure of the coral skeleton and relationships of coral groups. This has had a major effect on all subsequent discussion of rugose coral microstructure, diagenesis, and of relationships between the Rugosa and the Mesozoic and Cenozoic Scleractinia. Schouppe and Stacul (1955, 1959) published two major systematic works, the ®rst dealing with solitary corals having an axial columella. In this, as an introduction to the systematic part of their work, they discussed the construction of these coral skeletons at length; they then described 17 species, 10 of which were new. Their second monograph also dealt with solitary rugosans, those lacking the columella, introducing 27 new species and several new

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genera. These monographs mark the greatest single addition to our knowledge of the rugose corals of Timor. Schouppe and Stacul (1966) also published a major work on the structure of the Rugosa, based in large part on their studies of the Timor rugosans. The same rugosans provided Sorauf (1978, 1984) with material for scanning electron microscope study of skeletal microstructure and diagenesis, with electron microprobe (geochemical) analyses of the skeletal structures. This was based on the generally accepted premise that in the exceptionally well preserved Timor fauna the original calcite skeletal composition was retained, in contrast to Oekentorp's (1980, 1984, 1989) hypothesis that the original skeletons were aragonite, which were subsequently replaced by calcite during diagenesis. A valuable contribution by Fedorowski (1986) clari®ed a number of morphological and systematic questions arising from the work of Schouppe and Stacul (1955, 1959), made recommendations regarding their morphology and taxonomy and introduced several new generic names. A study by Niermann (1975) concentrated only on polycoeliid rugosans from Basleo, proposed 13 new species, and described a number of new subspecies. His descriptions were overly brief and poorly illustrated so that, although the species are suf®ciently well described and illustrated to validate his names, restudy of his types will be necessary before this work can be utilised. Important advances have recently been made in the understanding of some genera that had been questionably placed in the Tabulata, based on specimens from Timor. Plusquellec and Tourneur (1994) have shown that Trachypsammia is best regarded as an octocoral rather than as a tabulate, and Plusquellec et al. (1999) have discussed the paleobiology of the questionable tabulate coral Palaeacis. By far the greatest number of specimens and of species of solitary rugose corals from Timor have been collected from Basleo (Fig. 11). In the introduction to their monograph, Schouppe and Stacul (1955, p. 98) reported that they had studied approximately 12,000 specimens, collected in 1927; the overwhelming proportion of this material came from 30 collecting sites in the area of Basleo. The Polycoeliidae, which had not been studied by Schouppe and Stacul (1955), were studied by Niermann (1975) (13 new rugosan species). This same collection included the tabulate coral specimens studied by Hehenwarter (1951) (three new tabulate species). It is clear that the Basleo collections are by far the most diverse and abundant in both the Rugosa and Tabulata. Gerth (1921, pp. 124±129) discussed the geographic distribution of the faunas as they were then known, but this information is now taxonomically and geographically out of date. However, even taking into account only those species published by Gerth (1921), it is obvious that Basleo, Bitauni and their surroundings are the main source for Rugosa, and for all three species of colonial rugosans. Several other localities are worthy of mention for the occurrence of colonial rugosans, as reported by Gerth (1921, p. 126). Ipciphyllum timoricum (Gerth) occurs in

Amanuban at Oi Ekan (one specimen), at Kasliu (®ve specimens) and at Fatu Oinino (one specimen). Two other species of colonial corals were found, Lonsdaleiastraea vinassai Gerth (one specimen) at Poetain near Basleo, and Lonsdaleiastraea molengraf® (Gerth) (one specimen) at Noil Nunu Sono near Bitauni. Colonial rugose corals of Timor provide valuable evidence for palaeobiogeography. The distribution of Ipciphyllum timorensum, Lonsdaleiastraea vinassai and Lonsdaleiastraea molengraaf®i has been summarised on palaeobiogeographic maps of the waagenophyllid corals, and indicate that there was a connection from Timor to the shallow water areas of the Tethys (Minato and Kato, 1965, pp. 47, 152 and 156). Weyer (1979, p. 997) also noted that Ipciphyllum occurs in the Wuchiapingian of South China, although as a different species. Minato and Kato (1965) also noted in their phylogenetic diagrams that these species occur in strata of the Parafusulina and Neoschwagerina zones. Colonial Permian corals occur in limestones of the Maubisse Formation, as described above, associated with volcanics, suggesting that these corals lived in carbonate environments in small shelf areas, adjacent to active volcanoes (Fig. 9). Reconstructions of Permian coral palaeogeography distinguish two major realms, the Tethyan Realm, and the Cordillera±Arctic±Uralian Realm, separated by Pangea (Fedorowski, 1989). Numerous smaller epicratonic basins form distinctive biogeographic regions within these realms. The distribution of the colonial corals indicates clearly that Timor was connected to the major oceanic areas of Tethys that extended into present day Malaysia, China and Japan. Permian solitary corals belong to several faunal groups, generally referred to as `dissepimented' (called the `caniniidÐclisiophyllid fauna' by Hill, 1948) and `non-dissepimented', most commonly called the `Cyathaxonia fauna' (Hill, 1948; Kullman, 1966). These small, non-dissepimented solitary rugosans are found in deeper water shales, and some of these Permian corals have been regarded as nonzooxanthellate. Weyer (1979, 1997) described these corals from Russia, where they occurred in great abundance during the Kazanian, as forming part of a ªmonotonous ahermatypic Rugosa communityº. Fedorowski (1989, Fig. 1) showed that the greatest amount of diversity seen in the Kungurian and Kazanian corals of Tethys occurs within the Cyathaxonia fauna, and this also pertains to the Timor fauna. However, he also advised that the Cyathaxonia fauna should not be used as a paleoclimatic indicator (Fedorowski, 1989, p. 55), as species making up this fauna also occur with shelfdwelling species in shallow, warm water deposits, as well as in muddier and/or deeper and/or colder water deposits. Fedorowski (1989, p. 56) added that considerable adaptability is seen in Permian Cyathaxonia fauna genera, as in west Texas some occur in reefal environments. This also pertains to the abundant and diverse Tabulata from the Basleo localities. Palaeozoic tabulates, colonial and common reefdwelling corals, do not usually occur together with species

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of the Cyathaxonia fauna. In an interesting ecological occurrence in Timor, the tabulate corals Pseudofavosites stylifer, Aulohelia irregularis and Cladoconus magnus, are found coating or wrapped around crinoid columnals (Gerth, 1921), indicating that they lived in the water column, in relatively shallow but quiet seas, elevated from a (?)soft or turbid sea ¯oor. With a few exceptions, such as the colonial corals Ipciphyllum and Lonsdaleiastraea, the coral faunas do not shed much light on biostratigraphy in our present state of understanding. Minato and Kato (1965, p. 60) noted that the former is abundant within the Neoschwagerina zone, and that Lonsdaleiastraea vinassai Gerth and L. molengraaf® (Gerth) occur in this and the Parafusulina zones, with the latter the zone index coral for the Parafusulina zone, thus indicating a Kungurian to Kazanian (Wordian) age. However, their exact stratigraphic position in Timor is uncertain, partly because of the lack of systematic collecting and partly due to the occurrence of the corals in meÂlange. Adding to the imprecise biostratigraphic indications from solitary corals is the nature of the Timor fauna, which consists of species of small, non-dissepimented corals of the Cyathaxonia facies. These species tend to belong to simple, long-ranging genera, which lived in a wide range of environments, having different water temperatures, depths and clarity. They are dif®cult to classify into species and likewise it is dif®cult to generalise regarding their ecology. Fedorowski (1989, Fig. 1, p. 50) has summarised Permian worldwide coral diversity, with a preponderance of species (466) occurring in the Kungurian±Kazanian. The 290 species of the Cyathaxonia fauna and 106 species of massive corals show the highest diversity for any of the stages of the Permian. The rich Timor faunas were therefore part of widespread and diverse coral faunas which developed during the Kungurian±Kazanian in the late Middle and Late Permian. Fedorowski (1989) showed that much of this diverse fauna is composed of species of solitary Rugosa belonging to the Cyathaxonia facies, which occupied a deep shelf environment. Fedorowski (1989) also found that the Kungurian±Kazanian was the time of maximum diversity of massive colonial corals, which include the `Ipciphyllum-Lonsdaleiastrae fauna' of Timor. Ezaki (1991, 1994) found that simple colonial waagenophylids and the solitary non-dissepimented rugosans were abundant at the same time in both Iran and South China. The Permian rugosan corals of Timor, which seem to be a mixture of Kungurian and Kazanian rugosans, also occur within the same time period. The ¯ourishing tabulate faunas of Timor are largely endemic, although tabulates are present sparingly in other areas of Tethys. Hill (1957, p. 53) summarised the results of her study of the Timor coral fauna: ªConsidering the Timor corals as a whole, there is not enough speci®c differentiation between the faunas of these various localities for one to be able to accept without question that each represents a distinct

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stage.º Our understanding of the taxonomy and especially the ecology and biostratigraphy of these faunas has not increased much since that date. In order to allow the biostratigraphic use of this diverse and abundant coral fauna from the Permian of Timor, a major restudy of the species is needed, including their types, and stratigraphical relationships, in order to make full use of the temporal value of this rich coral fauna. Even the three species of colonial corals have problems of location and horizon of origin, and these are potentially the most biostratigraphically useful part of the fauna. 6.6. Echinoderms (GDW) The history of the collection and description of the Timor crinoids and blastoids was summarised by Webster (1998a) who compiled a list of 97 Timor localities from which echinoderms had been reported by various authors. In addition, Webster (1998a) noted the stratigraphic information, or lack thereof, as reported in the earlier literature, for each of nine localities: Somohole, Bitauni, Soefa, Nefotassi, Tae Wei, Dorf Sebot, Basleo, Amarassi, and Ajer Mati. He also pointed out that at some localities (i.e. Somohole and Bitauni), where a stratigraphic section was reported in the earlier literature, the crinoids may have come from a different lithologic unit from the ammonoids, but that nevertheless the stratigraphic age based on the ammonoids had been applied to the crinoids, and all the other fossils from the same locality. In addition, Webster noted that Burck (1923) had reported seven stratigraphic units from the Amarassi area, three of which could have yielded crinoids (one below and two above the ammonoid-bearing bed), yet all were considered to be the same stratigraphic age as that determined from the ammonoids. Unfortunately, in the crinoid descriptive literature (J. Wanner, 1910a,b, 1912, 1913, 1914±1929, 1916, 1920, 1923a,b, 1924, 1926, 1929a,b, 1930a,b, 1931a±d, 1932a,b, 1937, 1940a,b, 1941, 1942, 1949, 1950; de Marez Oyens, 1938, 1940) only the collection locality is given, not the precise stratigraphic horizon at that locality, preventing a determination of the true stratigraphic relationships among the various fossil groups. These practices, in combination with the indiscriminate collection and purchase of specimens by members of the Timor expeditions, as described above, leave considerable doubt about the true stratigraphic age and relationship of most of the echinoderm collections from Timor. Unquestionably, the Permian echinoderm species described from Timor are more diverse and numerically greater than the current total of all other known Permian echinoderm species worldwide (Bassler and Moodey, 1943). Species described after Bassler and Moodey's compilation are closing the gap, and as new Permian faunas continue to be described throughout the world, the Timor species will undoubtedly be numerically exceeded in the future (numerous publications are listed in Webster, 1973,

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1977, 1986, 1993, unpublished). However, at present the species recorded from Timor remain the largest known echinoderm fauna coming from a relatively small area for the Permian worldwide, and all other echinoderm faunas will continue to be compared with it. Echinoid species from Timor are few, based entirely on loose spines and interambulacral plates, and were all described by J. Wanner (1941). All taxa were reported from the Basleo `Beds' with `Cidaris' bitauniensis also reported from the Artinskian Bitauni Beds and Miocidaris permica also reported from the Artinskian Tae Wei Beds (J. Wanner, 1941). The generic assignment of most of the taxa is to some degree uncertain, because they are based on disarticulated plates. Permian echinoids, as compiled by Lambert and Thiery (1909±1925), Kier and Lawson (1978), are few and poorly known worldwide, and many are based on disarticulated corona plates and spines. This minimises the stratigraphic and correlative value of Permian echinoids worldwide. Blastoids attained their third greatest diversity in the Permian of Timor. The 12 ®ssiculate genera dominate the faunas, but the two spiraculate genera dominate numerically, with tens of thousands of thecae of Deltoblastus known from the oldest (Somohole) through youngest (Amarassi) Timor faunas (Waters, 1990; Webster, 1998a). Waters (1990) noted that most Timor genera are endemic, and that the non-endemics have their counterparts in Tethyan faunas reported from Artinskian strata of Australia and the southern Urals, and the Kazanian of Sicily. Webster and Sevastupolo (2001) reported Deltoblastus and Timoroblastus from late Sakmarian strata of Oman. Thus the Permian blastoids of Timor, as currently known, are restricted to the Tethyan realm. Crinoids dominate the echinoderm faunas of Timor and some species are known from hundreds and thousands of specimens, with over 90% of the crinoids coming from the vicinity of Basleo (J. Wanner, 1926). Webster (1998a) summarised the faunas for each of the nine localities that he discussed, recognising the preponderance of poteriocrinids over camerates and ¯exibles, as well as the crinoids over the blastoids, at most localities. Both the camerates and ¯exibles have remarkably high diversity in the Permian deposits of Timor compared with their much lower diversity in Late Carboniferous occurrences. Webster (1998a) also commented on the problems of determining the stratigraphic ages of the crinoids, as noted above, when comparing the faunas of the nine localities to one another. The Basleo faunas have a number of endemics, and clearly show the greatest diversity and abundance of all the Timor faunas. Few Timor taxa, at the species or genus level, are restricted to, or endemic in, only one of the nonBasleo localities. Crinoid genera and species are short-lived, as pointed out by Lane and Webster (1980). Thus the occurrence of the same crinoid species in the Basleo faunas, and at some of the localities dated as Artinskian based on the ammonoids, suggests that some of the Basleo faunas are

also of Artinskian age. This is supported by the recognition of 12 genera (eight species) of crinoids and three genera (one species) of blastoids common to the late Sakmarian± early Artinskian Callytharra Formation of Western Australia and the Basleo faunas (Webster, 1987; Webster and Jell, 1992, 1999). In addition, seven crinoid genera (one species) and one blastoid genus (one species) from post Callytharra Artinskian strata of Western Australia are common to the Basleo faunas (Webster and Jell, 1992, 1999). The systematics of a number of the echinoderms from Timor need revision. Breimer and Macurda (1972) discussed the ®ssiculate blastoids of Timor, and Macurda (1983) revised the ®ssiculates. Macurda (1983) also considered that the 17 species of Deltoblastus were probably variants of one or very few species. The numerous subspecies of Timoroblastus may also be variants of a single, or at most two or three, species. Several genera of the crinoids have ®ve or more species described by J. Wanner (1910a,b, 1912, 1913, 1914±1929, 1916, 1920, 1923a,b, 1924, 1926, 1929a,b, 1930a,b, 1931a±d, 1932a,b, 1937, 1940a,b, 1941, 1942, 1949, 1950) and again many of these species are considered variants of a single, or perhaps two or three, species. When proper analysis of these species is completed, the number of species known from Timor may be less than three-quarters of those currently recognized. At the generic level Webster and Lane (1987) considered that Timor species referred to Actinocrinites may represent a new genus. There may be a few other genera currently known from the Carboniferous, but unknown elsewhere in the Permian apart from Timor, that will be assigned to new genera. This may be especially true among the poteriocrinid cladids, several of which are known only from cups. If crowns of these taxa are found it could signi®cantly change their generic assignment. The palaeogeographic relationship of the Tethyan Permian echinoderm faunas was discussed by Webster (1998b). Permian Tethyan faunas are found from the southern Urals in the north, to Australia and New Zealand in the south, and from Timor and Thailand in the east, to India, Pakistan, Oman, Tunisia, and Sicily in the west. Equatorial faunas are those from the Urals, Timor, Thailand, Tunisia, and Sicily. Cooler water faunas, located south of 358S palaeolatitude, are those from Oman, Pakistan, India, Australia and New Zealand. Although poteriocrinid cladids dominate all faunas, there is a high percentage (68%) of endemics in the Tethyan faunas. Faunas ranged in age from early Sakmarian (Timor) to Wuchiapingian (Western Australia and Timor). Faunas from India, Oman, Australia and New Zealand, and some from Timor were living in clastic-rich or volcaniclastic environments, and all others in clay or carbonate-rich environments. Only the Tunisian fauna is associated with a reefal environment. Permian crinoid faunas from the western and northern margins of Pangaea are known from Bolivia, Mexico, several states of the southwestern part of the United States, Alaska, northern Canada, Germany, and the northern part of

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Russia. Most of these faunas are small, with less than ®ve taxa, and may have one genus in common with Timor. The Bolivian Artinskian fauna has six genera, two in common with Timor. The Artinskian fauna from southern Nevada (Lane and Webster, 1966; Webster and Lane, 1967) has seven genera in common with Timor. Both of these faunas have high percentages of poteriocrinid cladids with few camerates or ¯exibles. They apparently had little connection with the Tethyan faunas. Preservation of the Timor specimens is remarkable, in that the echinoderm stereom structure is exquisitely preserved and visible with relatively low magni®cation. However, crinoid and blastoid specimens have been transported short distances, as the proximal stem is rarely attached to the articulated cups or crowns, and holdfasts are not common in collections. Echinoids are known only from spines and interambulacral plates, suggesting disarticulation by scavengers or currents. Specimens were not reworked from older deposits and probably lived in an environment with high carbonate production, in close proximity to their site of deposition and burial. Webster and Jell (1992) suggested that the Timor faunas lived in a warm water environment favourable to higher carbonate productivity than the clastic-bearing carbonates and claystones of the Callytharra Formation of Western Australia. The warm water and high carbonate productivity are probably the key to the high diversity and abundance of the Timor faunas, especially those of the Basleo region. 6.7. Foraminifera (JEW) There are relatively few papers dealing directly with the Permian foraminifera of Timor, almost all deal exclusively with the large fusiform, microgranular-walled fusulines. Fusulines are very common in carbonate facies throughout the Middle and Late Carboniferous and Permian of the Tethyan Realm, and the evolution, stratigraphic ranges and palaeogeographical af®nities of the many fusuline genera are now very well known. They are very useful zonal fossils and there are numerous papers which link the Fusuline Zonation (®rst set up in Japan) with the stage terminology of SE Asia (see Kanmera et al., 1976). The earliest paper on fusulines of Timor is by Schubert (1915) in PalaÈontologie von Timor Volume 2. The material was collected during several expeditions to the island between 1909 and 1911 and of the four species of fusuline described and ®gured, three were new and were named after the leaders of the expeditions: Fusulina wanneri, F. molengraaf® and F. weberi. The paper also described a new species of smaller foraminifera, Geinitzina chapmani. The ®rst two-named fusuline species were described primarily from localities in the Benain River, West Timor, whilst F. weberi came from the slopes of Mt. Aubeon (Auveon), near Pualaca, East Timor. At the time Schubert (1915) believed these faunas to be Late Carboniferous in age. Thompson (1949) described material collected from

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several localities in West Timor during the Brouwer Expedition of 1937. Thompson (1949) revised Schubert's (1915) nomenclature and dating. Schubert's F. weberi was referred to the genus Palaeofusulina, F. wanneri was placed in Parafusulina, and F. molengraaf® referred to Schwagerina?. Thompson (1949) described part of Schubert's material, which Schubert had referred to F. granum-avenae of Roemer, as a new species (Schwagerina brouweri), the remainder being left as Schwagerina sp. These faunas were reassessed as probably of Early Permian age. Nogami (1963) studied collections of material made by geologists from the University of Kyoto in 1961, from HatoBuilico and Pualaca in East Timor. Four species were described: two, a Triticites and a Parafusuilina, were left in open nomenclature; the other two are Schwagerina nakazawae (to encompass F. granum-avenae of Schubert, 1915 (pars), and S. brouweri (pars) and Schwagerina sp., both of Thompson, 1949), and Schubert's F. weberi, which he placed correctly in Codonofusiella. These samples were now assigned to the `Lower' or `lowest' Permian. A more recent collection was made by Barkham (1993). He reported the following fusulines (identi®ed by O. Dawson): Monodiexodina sp., Schwagerina cf. pidingensis, Parafusulina gigantica, P. cf. japonica, Parafusulina sp. and Schwagerina sp., in carbonates from several localities in the Maubisse Formation of the Bisnain area of West Timor. The ®rst two species came from the Nekmelalat Member, the latter four from the Manufui Lavas. Monodiexodina wanneri is the most recent combination for Schubert's (1915) taxon, Fusulina wanneri, and it is indeed the typespecies of Monodiexodina; the specimens from Bisnain are very close. Both assemblages were considered to be of Artinskian±Kungurian (Early Permian) age. Other members the University of London Southeast Asia Research Group (Barkham, Barber and Charlton) have provided new material for the present study. Sample NK1 comes from the River Nekmalelat in the Bisnain area, and can be referred to the Nekmalelat Member of the Maubisse Formation. It contains relatively few fusulines, but these include Monodiexodina and a schwagerinid, with some smaller foraminifera belonging to Pachyphloia and/or Geinitzina. The rest (N.T. 2±5, 7, 8, TM 1028A, B, 1033, and TUK 14), also belonging to the Maubisse Formation, all come from the River (Noil) Tuke in West Timor. These specimens were collected from boulders in the river bed eroded from the hillsides of a steep tributary valley. In spite of a diligent search the fusulinid-rich rock has never been found in situ. Boulders in the river bed are packed with foraminifera, which are often arranged in current beds and their long axes are current-aligned. The fauna in the samples is practically a monotypic assemblage of Monodiexodina wanneri. Most of the fusulines faunas mentioned above are clearly of Early Permian age and belong to the Artinskian±Kungurian stages (Monodiexodina ranges down to the base of the Artinskian). On the other hand, Codonofusiella weberi, a

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very distinctive uncoiling form, also occurs in Timor, but in association with Schwagerina nakazawae and S. brouweri, but never with Monodiexodina wanneri. Our extensive knowledge of the stratigraphic ranges of fusuline genera (see, for example, Kanmera et al., 1976), shows that Codonofusiella ranges from the Neoschwagerina-Zone (which is later than the extinction of Monodiexodina in the Wordian), to the top of the Palaeofusulina-Zone (within the Changsingian). These Codonofusiella-limestones must, therefore be of Late Permian age. The palaeogeographical af®nities of the Permian marine faunas of Asia, based on the study of extensive datasets including fusuline foraminifera, have recently been much reviewed (see Shi et al., 1995; Archbold and Shi, 1996; Ueno, 2000). In the Permian, according to these authors, Timor lay between the Gondwana Realm and the Tethyan Ocean sensu stricto, in the southern transitional zone, also known as the Cimmerian Province. In terms of fusulines the transitional zone is characterised by low diversity faunas throughout the Permian, with a paucity of verbeekinid and neoschwagerinid fusulines, so common during Middle Permian (Bolorian/Kungurian±Midian) times in Tethys proper. Instead the transition zone is characterized by the genus Monodiexodina in the Early Permian (Artinskian or younger) and Codonofusiella in the Late Permian. Fusulines are also useful ecological indicators. By analogy with modern (but unrelated) fusiform alveoline foraminifera, they are thought to have harboured symbiotic algae, and would have lived in shallow, warm water, within the photosynthetic zone. Moreover, when found in situ, they are always in a carbonate platform environment, and never associated with clastic facies. This implies that Timor, although it was on the fringes of the tropical Tethyan domain, must have been from time to time `bathed' in warm-temperate waters, the climate probably becoming more sub-tropical as it drifted further north later in the Permian. Smaller (non-fusuline) foraminifera from the clastic facies of the Cribas Formation were reported by AudleyCharles (1968). These forms were identi®ed by D.J. Belford and comprise: Frondicularia sp. aff. woodwardii, Geintizina triangularis, Nodosaria sp. aff. postcarbonica and other unidenti®ed species of Cornuspira, Dentalina, Textularia, Trepeilopsis and the Lagenidae. The Cribas Formation is reliably dated on other evidence as of Kungurian to Late Permian age, possibly continuing in a condensed sequence into the Triassic. This assemblage of smaller foraminifera is compatible with an Early, rather than a Late Permian age. 6.8. Gastropods (NJM) The gastropods from the Permian of Timor are reasonably diverse, with a larger number of species than any of the Permian basins of mainland Australia. The majority of species have been described from Basleo. Along with the bellerophonts, the gastropod fauna resembles other Permian

gastropod faunas that are usually considered to have lived in the relatively warm, shallow marine waters of the lower latitudes. Most of the gastropods from Timor were collected at the classic fossil localities without satisfactory stratigraphic control. However, Straparollus was collected by Barkham (1993) from the Cribas Formation in the River Bisnain and from limestones of the Hoeniti Member of the Maubisse Formation in the River Hoeniti. Straparollus was also collected by Bird (1987) from bioclastic limestones of the Naibalan Member of the Atahoc Formation in the Noil Tunsip, together with Platyteichum and a pleurotomarid. Platyteichum has not previously been recognised outside Australia and New Zealand. Some at least locally signi®cant stratigraphic information can be obtained from in the reported gastropod faunas from the classic localities. For instance, in the genus Bayleia (primarily the genus Yvania of C. Wannner, 1942), the species B. bitauniensis has been recognised only at Bitauni; B. permica, B. reticulata and B. cf. salomonensis only at Tae Wei; and B. clathrata and B. conglobata only at Basleo. Wanner's species Yvania timorensis (?Ananias timorensis in Appendix B) has been reported from both Tae Wei and Basleo. Similarly Wanner (1942) reported Cyclonema bitauniensis and C. incertum from Bitauni, but C. basleoense, C. brouweri, C. molengraaf® and C. simile from Basleo. NJM, however, considers C. bitauniensis and C. brouweri to be synonymous (Appendix A). The majority of gastropod species from the Permian of Timor are based on a very small number of specimens, and often their characteristics have been rather inadequately illustrated, with the result that their proper identi®cation is often in doubt and we do not yet have a proper idea of the variation in characters of individual species. It therefore follows that very little weight should be put upon the number of species present in each of the faunas. It seems almost certain that Wanner separated more species of the genus Zygopleura than would be considered justi®ed today, his ®gures show specimens which can hardly be distinguished from specimens from the Lower Jurassic. There may similarly be more species of Timor vetigastropods than the material really justi®es. There is clearly room for further research into the taxonomy of these forms. The dif®culty in the taxonomic placement of many of the Palaeozoic gastropods is the inability of interpreting their soft part anatomy. We do not know whether a holostomatous Palaeozoic gastropod such as Cyclonema, Cyclites or Anomphalus had two gills or two kidneys or just one of each, and have so far failed to document enough other evidence to lead to their satisfactory classi®cation. It has not proven possible to discriminate between species of Platyceras as the morphology of their growth curves would have been dependent upon their position on a crinoid calyx with which they are widely associated (Bowsher, 1956). It is not known whether Platyceras are species speci®c to individual species of crinoids. The only limpet-like

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taxon in the fauna, `Patella cf. ottomana Kittl' of Wanner (1922) was±later considered by Wanner (1942) to be a craniate brachiopod; the identi®cation of this specimen might be worth further investigation. The absence of acteonid opisthobranchs, known from Carboniferous strata and occurring widely from the Triassic to the present day, probably relates to their great rarity and the poor sample size of the Timor gastropod faunas. 6.9. Mollusca and similar Phyla (NJM) The majority of benthonic Mollusca and mollusc-like phyla from the Permian of Timor are described by Wanner (1922, 1940, 1941, 1942) and Hamlet (1928). Information regarding the geographical distribution of these species is given in each of these papers. As for other fossil groups most specimens have been collected loose at the classic fossil localities. In West Timor the bivalve genus Atomodesma exarata was collected in situ by Barkham (1993) from the Cribas Formation in the River Bisnain and from the Maubisse Formation in the rivers Bafafor, Melelat and Maubisse. Atomodesma mytiloides was collected by Bird (1987) from bioclastic limestones of the Naibalan Member of the Atahoc Formation in the Noil Tunsip, Kekneno. Bird (1987) also reports that the Tutlap Member of the Cribas Formation is dominated by A. exarata to the exclusion of other forms. In these studies Atomodesma was identi®ed by J.M. Dickens who correlated the Tutlap Member with the West Australian Zone D2 of Kungurian age (Dickens, 1963). A. exarata has also been reported from Cribas, Aliambata and Loiquero in East Timor In general the the benthonic molluscs do not offer much input to solve stratigraphic problems, but the plicate species of Atomodesma may prove to be useful. For instance, A. multifurcata has so far been found only at localities in the Basleo area, and A(?). undulata only at Ajer Mati. A. exarata has notably not been recognised from the heavily sampled Basleo locality. It may be that this is stratigraphically signi®cant, perhaps indicating the absence of rocks of Kungurian age from Basleo. A. mytiloides, on the other hand occurs not only in the Kekneno area, but also at Bitauni and Basleo. This species may, therefore, have had a longer stratigraphic range. The hyolithid species Macrotheca timorensis has been described from the Susie Mepu locality near Basleo. The Hyolitha are still considered by many workers to be Mollusca, but they may be separated by their bilateral symmetrical, diverticulated gut, occasionally preserved by sediment in®ll (Runnegar et al., 1975). Hyolitha are quite rare in the Upper Palaeozoic and the genus Macrotheca can only be described as having a typical hyolithid shape. no `operculum' or shelled appendages are known. However, there have been no better suggestions for its sytematic placement. Apart from this, the only known hyolithid occurrence in Timor is from a fragment of a similar but perhaps distinct species collected at Bitauni (C. Wanner, 1941).

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The rostroconch Pseudoconcardium clipeoforme (Wanner) was reported from Tunium Eno in the Basleo area and ?Conocardium ornatum from Nefotassi 5 km north of the Somohole locality in West Timor. Rostroconchs super®cially resemble bivalves but, like the scaphopods, they did not develop a balanced system of elastic ligament and adductor musculature to control the opening and closing of their shell. They are usually considered to be the sister group of the bivalves. They are not known to have survived beyond the end of the Permian. The bivalve fauna of Timor differs from other Permian faunas in including a relatively small number of infaunal taxa. These are represented by shallow nestlers, including the Permophoridae and the ?Mytilidae, and shallow burrowing forms such as the trigonioids, crassatelloids and nuculanoids. The deeper burrowing, siphonate taxa of the Anomalodesmata and the elongated shells of the Orthonotida and Solemyoidea have not been discovered. Even among the shallow burrowers there are only a small number of specimens known. This may to some extent mirror sample size and possibly the dif®culty of preservation and collection of these usually thin-shelled taxa; but in addition, palaeoenvironmental factors are probably important. The extensive development of arenaceous clastics in the Australian Permian marine basins may have been more amenable for these deeper burrowers. In common with other deposits of Permian age a number of bivalve groups which otherwise might be expected in the Timor Permian fauna are absent. These include taxa normally attributed to the Lucinoida and the Megalodontoidea and similarly oyster-like, cemented taxa which are known only from the very latest Permian of Japan. Bellerophon timorensis umbilicata is recorded from Basleo and Bellerophon quadratus, B. timorensis gibber, B. galeatus expers from other localities in West Timor. The systematic placement of the bellerophont `snails' continues to be hotly disputed. Here the bilateral symmetry of both their early formed shells and their muscle attachment scars is taken as evidence that they did not undergo torsion. Torsion or at least partial torsion is taken to be the major synapomorphy of the Gastropoda. Some Bellerophontidae share shell microstructure type with some gastropods, but also with some bivalves. This could be interpreted as indicating that such a shell structure type is primitive for the three groups involved, and thus of little value in solving their relationships or, as preferred here, that the shell structure type, aragonitic crossed lamellar structure, has evolved separately on a number of different occasions during the history of the Mollusca. The Class Monoplacophora, which includes the bellerophontids, may prove to be a paraphylum, requiring further revision of the taxonomic position of this group. 6.10. Trilobites (RMO) Forming only a minor component of the Permian faunas of Timor (Wanner 1926, p. 34), trilobites were ®rst noted by

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Beyrich (1865), who described a new species, Phillipsia parvula on the basis of four small cranidia from Ajer Mati, and later Boehm (1908) ®gured an incomplete pygidium from Pualaca. Well preserved specimens collected during the expeditions of Wanner and others formed the subject of Tesch's (1923) contribution on trilobites to the PalaÈontologie von Timor series; most were attributed to his new taxon Proetus (Neoproetus) indicus, the material originating principally from the Bitauni and Basleo districts. Gheyselinck (1937) later redescribed Tesch's species, together with eight new ones, based on specimens collected from the Basleo district by a Dutch expedition in 1935. This remains the most comprehensive treatment of the Timor trilobites, although subsequently Hahn and Hahn (1970, 1972) reassigned some of these species, and these authors (1967, 1972) and Hahn et al. (1991) proposed new genera and subgenera based on Timor species. Hahn and Brauckmann (1975) redescribed and re®gured Beyrich's species, and Weller (1944), Owens (1983) ®gured specimens from Timor in their reviews of Permian trilobite genera. Elsewhere, trilobites have been mentioned brie¯y (e.g. Gageonnet and Lemoine 1958; Audley-Charles 1968), and Simons (1940) commented on the morphology of specimens from Lidak, to which he attributed an Artinskian age. The trilobites used as the basis for Tesch's (1923) and Gheyselinck's (1937) papers total approximately 100 specimens, some 80% of these being from the Basleo district; others undoubtedly exist in museum and private collections. Within the documented material seven species are represented, distributed among six genera, all members of the Phillipsiidae, one of three families of trilobites that survived into the Permian. The majority (about 70%) are accounted for by Neoproetus indicus (Tesch, 1923), which has been recorded from at least three of the principal Permian localities in Timor; ®ve additional species are known from the Basleo district, but only one or perhaps two from each of the remaining areas. Gheyselinck (1937) gave the source for most of the trilobites as the `Basleo Beds', and Hahn and Hahn (1970, 1972) and Owens (1983) assigned most Timor species to the Kazanian, the then commonly assumed age of this horizon. The collection of trilobites in situ from the Atahoc Formation in the River Bisnain section (Barkham, 1993), which is there dated as Artinskian, allows a more critical assessment of their stratigraphical distribution. Examination of these, now in the Natural History Museum, London, has con®rmed their identi®cation as Neoproetus indicus. It is assumed that other occurrences of this species are of a similar age, although many are apparently from clasts in the Bobonaro MeÂlange, or collected loose from the soil rather than from undisturbed sections. This is true, for example, of the occurrences in the Maubisse and Atahoc formations in the Bitauni district, and in the Maubisse Formation in the Basleo district (A.J. Barber, personal communication 2001). In the Basleo district, N. indicus has been collected from the Maubisse Formation at Noil Fatu, Mangan Tobe,

Tuninu, Kiumoko, Pantukak, Siloe Netu Kot, and Siloe Nifoe Koko, as well as from `Basleo', and `south central Timor'. At Tunino it is associated with Pseudophillipsia timorensis (Gheyselinck, 1937) and at Kiumoko and Siloe Netu Kot with Triproetus trigonoceps (Gheyselinck, 1937). Hildaphillipsia hildae (Gheyselinck, 1937) occurs with T. trigonoceps at Nipol Soempek. On the basis of these cooccurrences, it is probable that all these species, like N. indicus, are of Artinskian age, on the assumption that they originate from more or less the same horizon, and are not from two or more mixed ones. The two remaining species from this district, Triproetus gerthi (Gheyselinck, 1937) and Timoraspis breviceps (Gheyselinck, 1937) have been reported only from the vague location of `south central Timor' and could therefore be from any of the horizons within the Maubisse Formation that are apparently represented in the Basleo area (see above), with an age range from Artinskian to late Wuchiapingian. Outside the Basleo district, N. indicus occurs around Bitauni and Somohole, and possibly in the Lidak district of West Timor, and near Pualaca and in the Cribas Anticline in East Timor, in the presumed Artinskian parts of the Atahoc and Maubisse formations. The only con®rmed post-Artinskian trilobites from Timor are small cranidia identi®ed as Microphillipsia ? parvula (Beyrich, 1865) from Ajer Mati of possible late Wuchiapingian age. Elsewhere in the Amarassi district, Triproetus baungensis (Gheyselinck, 1937) from the Baun area occurs probably also at Basleo, if T. trigonoceps is a synonym. Since the occurrences of the latter may be of Artinskian age (see above), this is possibly also the age of T. baungensis. Indeterminate trilobites were listed by Tesch (1923) from Oesapikapitan in the Amarassi district. Of the seven species known from Timor, only one, N. indicus, has been identi®ed from elsewhere. A pygidium described by Kobayashi and Hamada (1979) from Kajibumi, near Kuala Lipis, Peninsular Malaysia, although distorted, is almost certainly referable to this species. All the genera known from Timor have been reported from other stations in the Tethyan realm, but represented by different species. Pseudophillipsia is especially widespread (see Owens and Hahn, 1993), and Neoproetus, Timoraspis and Hildaphillipsia are known from the Wordian of the Sosio Valley, Sicily. Triproetus has an almost cosmopolitan distribution in the Early Permian (Cisural Series), being widely distributed in Tethys (e.g. north Thailand, Oman), as well as in Svalbard, Alaska and west Texas. None of the genera from Timor is known from Australia, where Permian trilobites are represented in strata of Artinskian age from Western (Ditomopyge) and south-eastern Australia (Doublatia). With their forward-expanding glabellas, Neoproetus, Triproetus, Timoraspis and Pseudophillipsia probably had attached hypostomes; Fortey and Owens (1999, p. 434) inferred a predatory feeding habit for this type of morphology in such advanced Phillipsiidae. Hildaphillipsia, on the

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753

Fig. 12. Permian palaeostructure of the Timor±Australian Northwest shelf region, with seismic isochrons for the Permian of the Bonaparte Basin (McConachie et al., 1996). Contour interval is 0.1 s TWT, with shading at 1.0, 1.5 and 2.0 s. Structures in eastern and central Timor have been schematically restored to their locations prior to Neogene orogenesis.

other hand, is the last representative displaying the more primitive morphology in this family, in which the glabella tapers gently forwards, with the hypostome not rigidly attached, but presumed to have been ligamentally joined. There are arguments for this type of morphology being associated with particle feeding (Fortey and Owens, 1999). Cuticle thickness may provide some evidence of the zones particular trilobites inhabited on an onshore-offshore, or shallow-to-deep pro®le (Fortey and Wilmot, 1991), with, generally speaking, thicker cuticles on the inner shelf, and thinner ones on the outer shelf and slope. In the case of the Timor trilobites, the cuticle of Neoproetus indicus is comparatively thick and robust, whilst in such species as Hildaphillipsia hildae and Timoraspis breviceps it is much thinner. Comparison of these with the examples given by Fortey and Wilmot (1991) suggest that Neoproetus indicus possibly inhabited a shallow or inshore environment, as represented by the Maubisse Formation, and parts of the Atahoc Formation. Other species associated with it, but with thinner cuticles, although presumably occurring in water of a similar depth, may have inhabited more protected ecological niches (Fortey and Wilmot 1991, p. 149). 7. Palaeogeographic interpretation The stratigraphic record of Timor commences with the

Atahoc Formation in the late Asselian or early Sakmarian (Bird, 1987). It is likely that the basinal Atahoc Formation slightly predates the carbonate-volcanic Maubisse Formation, which is dated reliably only from the late Sakmarian (Barkham, 1993). The Sakmarian section in the Maubisse Formation at Bisnain (the Hoeniti Member), does not include volcanics or volcaniclastics; nor does the Sakmarian section of the Atahoc Formation in the Bisnain and Kekneno areas, nor the Atahoc type section in East Timor, except at the very top (Audley-Charles, 1968). However, the classic Somohole locality, with a late Asselian±Sakmarian ammonoid fauna (Grant, 1976) but a late Sakmarian±early Artinskian echinoderm fauna (Webster, 1998a,b), reportedly contains volcanic tuffs (e.g. Wanner, 1923a). Unfortunately this locality has not been revisited during more recent work. We have not yet found satisfactory stratigraphic descriptions of the Somohole succession, but Grunau (1953) published a schematic stratigraphic column showing shales interbedded with limestones. Based on the apparent passage from a late Asselian±early Sakmarian ammonoid-dominated fauna into a late Sakmarian±early Artinskian echinoderm-dominated fauna, it seems likely that the Somohole succession marks an upward passage from the Atahoc Formation into the basal Maubisse Formation. This passage may re¯ect the development of shallow marine environments due to volcanic shoaling and/or eustatic regression. Bird (1987) suggested that the Sakmarian Atahoc

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Fig. 13. Schematic palaeogeographies for the Permian of Timor.

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Formation of the Kekneno area was deposited in a narrow, approximately E±W trending graben, with sedimentary sourcing from both the northern and southern rift ¯anks. The Atahoc Formation of the Bisnain area was probably also deposited in a graben with an E±W basinal elongation, based on the north to south basinward distribution of facies described in Section 3.2, and assuming that the present thrust structure in this area re¯ects a degree of control by pre-thrusting, basement extension faults (Barkham, 1993). Similarly Neogene thrust structures in the Aileu, Cribas and Loiquero areas of northern East Timor most likely suggest inversion of an E±W elongated basin or basins, which developed originally during the Early Permian. The Aileu sedimentary basin, now represented by the Aileu Complex, partly of Permian age, appears to have had a northern siliciclastic sedimentary source at some time (Barber and Audley-Charles, 1976; Prasetyadi and Harris, 1996). The Maubisse Formation in East Timor lies to the south of this north coast Atahoc/Aileu basin. The E±W graben of the Kekneno area may have formed a westward extension of the north coast grabens in East Timor, but it seems less likely that the Bisnain area formed part of the same E±W rift, given its more southerly location and the occurrence of the Maubisse Formation to the north of the Atahoc Formation in the Bisnain area. A regional palaeotectonic map for the Permian is shown in Fig. 12, and a schematic palaeogeographic map for the Sakmarian of Timor in Fig. 13a. The Kekneno area is interpreted as occupying a location at the junction of the South Banda Graben (the present-day Aileu, Cribas and Loiquero basinal areas in East Timor) and the older NW±SE trending Bonaparte Graben (see Section 8.3). It is suggested that the Bisnain, Basleo and Laktutus successions were deposited in, or on the margins of, a smaller graben subparallel to the South Banda Graben, but developed within the former structural high on the eastern ¯ank of the Bonaparte Graben. The oldest identi®ed Permian volcanism outside the Somohole area occurs in the Artinskian of the Bisnain area (Banfafor Member, Manufui Lavas and Nekmelalat Member of the Maubisse Formation, and also in the less reliably dated upper Atahoc Formation: Barkham, 1993). Volcanism in the Maubisse Formation was also dated as Artinskian±Kungurian at Laktutus (Barkham, 1993), but older Permian rocks are not exposed in that area. In the Cribas Anticline the ®rst volcanic horizon occurs at the top of the poorly dated, but probably Artinskian, Atahoc Formation. Hence the Artinskian of Timor seems to mark a transition from extension with only limited volcanism, to rifting with major volcanism (Fig. 13b). The geochemistry of the volcanism is compatible with a rift setting, being tholeiitic to mildly alkaline in character, with a trace element geochemistry suggesting extrusion in a rift-valley or ocean-¯oor setting (Berry and Jenner, 1982; Barkham, 1993). The Artinskian±Kungurian succession of the Bisnain area additionally records a major phase of marine regres-

755

sion. This is indicated by general shallowing of the marine Artinskian±Kungurian successions of the Maubisse Formation, by the subsequent development of palaeosols and probable subaerial karsti®cation, and by the development of the Khali Beds as a major regressive wedge at the edge of the Bisnain carbonate/volcanic platform. In the basinal area to the south, the boundary between the poorly dated upper Atahoc Formation and the Kungurian lower Cribas Formation is marked by a limestone interval, and this may be a distal facies equivalent of the regressive Khali Beds. Similar limestones form the boundary beds between the Atahoc and Cribas formations in the Kekneno area, suggesting that regression also affected the Kekneno Basin at this time. Evidence for contemporaneous structural extension, together with the occurrence of volcanic beds in the Atahoc-Cribas boundary beds at both Bisnain and Cribas, suggests that this regression was very likely tectonically driven rather than purely eustatic. This Artinskian±Kungurian regressive phase may also help to explain the well preserved Artinskian crinoid fauna at Basleo (Webster, 1998a). Outcrops examined by TRC in the Basleo area suggest that Basleo was primarily a basinal depositional area, part of the interpreted E±W Basleo Graben in Fig. 13. During the Artinskian, sea levels may have fallen suf®ciently in the Basleo segment of the graben for crinoids to ¯ourish in situ in a relatively shallow marine environment. Alternatively, extension in the Basleo Graben may have been restricted to an area north of Basleo until the Artinskian, but block faulting may have extended further south at that time, downfaulting the Basleo area and allowing the accumulation of the crinoidal limestones. The Tuke Beds, now found some 9 km south of Basleo, are also dated as Artinskian (Thompson, 1949), and these may represent a laterally equivalent nearshore facies (Barkham, 1993) which accumulated on the southern margin of the Basleo Graben. Substantial volcanism continued into the ?Middle and Late Permian. In many localities there appears to be an upward transition within the Maubisse Formation from predominant limestone into a predominantly volcanic succession (e.g. the Maubisse type area; at Pualaca; and at Kasliu). In the Kasliu area this volcanism is dated as Late Permian (Archbold and Bird, 1989), whilst a Late Permian age is suspected for the upper 500 m volcanic pile at Maubisse (Audley-Charles, 1968). Locally volcanism continued into the Early Triassic in East Timor (Grady and Berry, 1977), but in West Timor Triassic volcanism appears to be limited to tuffaceous interbeds (e.g. Wanner, 1931b). Following the Artinskian±Kungurian regression, the ?Middle-Late Permian of Timor seems to be characterised overall by a major phase of marine transgression (Fig. 13c). In the Bisnain area, the Cribas Formation above the basal limestones is a deep marine, poorly oxygenated basinal succession (Barkham, 1993), and similar conditions probably also prevailed in the Kekneno Basin (Bird, 1987).

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During the Middle Permian (Roadian) the Basleo area initially accumulated relatively shallow marine sediments containing bryozoans (Gilmour and Morozova, 1999), succeeded by a dominantly ammonoid fauna of approximately Wordian age. The Roadian bryozoan fauna may therefore record the commencement of transgression following the Artinskian±Kungurian regression. The Middle and Late Permian may be separated by a short phase of regression, which would explain the apparent passage in the Basleo fauna from predominant ammonoids in the Middle Permian to a more mixed fauna (dominated by brachiopods?) in the Late Permian (early Wuchiapingian). The Kasliu succession, located on a structural high immediately east of the Kekneno Basin, was probably also deposited during relatively high sea levels in the early Late Permian (Wuchiapingian). In the main basinal areas, such as the Kekneno and Cribas basins, the Upper Permian has not been de®nitively identi®ed from fossil evidence, although there is apparent conformity between the Permian and Triassic successions, and it is likely that the MiddleLate Permian is represented by a condensed sequence (Bird and Cook, 1991). The Amarassi fauna is interpreted to have developed during the ®nal Permian phase of transgression, which apparently commenced in the late Wuchiapingian (Fig. 13c). Despite being composite, including fossils from several localities across southwest Timor, there has been no suggestion in the earlier literature that the Amarassi fauna is anything other than Late Permian in age (unlike the other three classic fossil localites, all of which have yielded different apparent ages from different fossil groups). However, a caveat must be placed even on this, as the Amarassi trilobites could be as old as Artinskian, if some apparently comparable trilobites from Basleo are also Artinskian in age (see Section 6.9). A possible explanation for the general concurrence in ages in the Amarassi fauna is that the extreme southwest of Timor remained emergent throughout most of the Permian, being submerged below sea level only in the Late Permian. The Amarassi succession was deposited on the western ¯ank of the Bonaparte Graben, at the northern end of a persistent structural high, including the Ashmore Platform and Scott Plateau regions of the modern NW Australian continental margin. This area may have remained generally emergent throughout much of the Permian (Bradshaw et al., 1988). The latest Permian (Changhsingian) has not been de®nitively recognised anywhere in Timor. By this time the entire Timor region may have subsided below sea level, and consequently the Permo-Triassic boundary in most, if not all, parts of Timor is represented by a condensed stratigraphic sequence. Because globally the Permo-Triassic boundary is interpreted as a time of relative regression, it appears that in Timor tectonic movements continued to play a greater role than eustasy in controlling local sea levels. Transgressive conditions probably continued well into the Triassic in Timor.

8. Regional correlations 8.1. Stratigraphic relationship to the Northwest Shelf Mory (1988) divided the Permian succession of the Northwest Shelf between two groups, the Late Carboniferous±Early Permian Kulshill Group, and the Middle Permian±Early Triassic Kinmore Group (Fig. 10). The oldest formation in the Kulshill Group, the Kuryippi Formation, predates the oldest recognised sedimentary rocks in Timor, and is not considered further here. The Treachery Shale, which overlies the Kuryippi Formation, consists of carbonaceous shale and tillite deposited during the ®nal stages of Carbo-Permian glaciation. It is dated as early Asselian, and therefore probably also predates the Timor succession, which did not begin to accumulate until late Asselian or early Sakmarian times. The Treachery Shale is a transgressive unit, deposited during rising sea levels associated with deglaciation. The commencement of Permian sedimentation in Timor may in part be connected with this transgression, combined with contemporaneous extension which initiated the Timor sedimentary basins at this time. However, the Timor succession would appear to be essentially post-glacial, as no dropstones have been recorded in the Atahoc Formation. The succeeding Keyling Formation (late Asselian± Sakmarian) is a thick ¯uviodeltaic siliciclastic succession deposited, according to Mory (1988), as a result of erosion in the Australian hinterland related to post-glacial rebound. The age range of this formation corresponds with the ammonoid-rich lower Atahoc Formation, ammonoids from the Somohole fossil locality and of the Bitauni ammonoid fauna. Sea levels on the outer Australian margin were apparently relatively high at this time, leading to the deposition of open marine sedimentary successions in Timor and on the outer Northwest Shelf. The ®ne-grained lower Atahoc Formation is a distal facies equivalent of the ¯uviodeltaic Keyling Formation. The Fossil Head Formation at the top of the Kulshill Group consists of siltstone and sandstone with minor limestone. It is dated as Artinskian±Kungurian, and is therefore equivalent in age to the Maubisse Formation at Bisnain (including part of the Bitauni fauna), and the upper Atahoc Formation throughout Timor. The widespread distribution of the Fossil Head Formation across the Northwest Shelf suggests the continuation of relatively high sea levels on the Australian margin, in contrast to tectonically induced regression in Timor at this time. The stratigraphy of the Hyland Bay Subgroup (the Hyland Bay Formation of Mory, 1988), which represents the Middle-Upper Permian elements of the Permo-Triassic Kinmore Group, has recently been reviewed by Gorter (1998). This subgroup consists (upwards) of the Pearce, Cape Hay, Dombey, Tern and Penguin Formations. Two regionally extensive limestone intervals are recognised in both the Pearce and Dombey formations,

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while the remainder of the succession is predominantly siliciclastic. The lower part of the Pearce Formation is not well dated, and could be as old as Kungurian, whilst the upper part of the formation is dated as U®mian (Roadian, early Middle Permian). The formation has a widespread distribution on the Northwest Shelf, typically two limestone units separated by a shaly clastic succession. To the southeast the formation becomes increasingly dominated by clastics, and onshore in northwestern Australia it probably passes into a sandstone succession. The Pearce Formation limestones represent transgressive intervals, with the intervening clastics corresponding to a short-lived phase of regression. The base of the Pearce Formation is sharply unconformable on the underlying Kulshill Group. The succeeding Cape Hay Formation is essentially Midian/Kazanian (Capitanian±Wordian) in age, although it may extend stratigraphically somewhat upwards and downwards (Gorter, 1998). In the southern Bonaparte Basin the formation is characterised by several cycles of ¯uvial, tidal and barrier-bar delta clastic sediments. A prominent regional seismic marker horizon in the lower part of the formation is designated the `Onset of Major Progradation' (Gorter, 1998), and the formation is essentially a regressive cycle. This regressive phase on the NW Shelf may correspond to the regression inferred from the Basleo sequence in Timor, corresponding to the ?Wordian succession dated by ammonoids, separating the Roadian succession dated by bryozoans, and the Wuchiapingian succession dated by brachiopods. The Dombey Formation comprises two limestone intervals separated by a clastic succession. The lower limestone unconformably overlies the Cape Hay Formation, and consists of bryozoan biomicrite, calcilutite, micaceous shale and siltstone. The succeeding clastic interval contains both shales and sandstones, whilst the upper limestone consists of marls passing upward into ®nely crystalline limestones with interbeds of shale. The limestone intervals thicken to the north and west towards Timor. In the distally located Sahul Shoals-1 well on the Ashmore Platform (Fig. 1), these limestones contain a brachiopod fauna compositionally very similar to the Basleo and Amarassi faunas (Archbold, 1988). As with the limestones in the Pearce Formation, Gorter (1998) interpreted the two Dombey limestone horizons as representing periods of rapid transgression, whilst the intervening clastics represent a minor regressive phase. The Dombey Formation is dated as Wuchiapingian, and it is therefore very tempting to correlate the lower Dombey limestone with the early Wuchiapingian Basleo brachiopod fauna, and the upper Dombey limestone with the late Wuchiapingian Amarassi succession. The ®nal two Upper Permian formations recognised by Gorter (1998) are the Tern and Penguin formations, which are both dated to the Changhsingian, or latest Permian. The Tern Formation comprises about 100 m of sandstone and shale in the southern Bonaparte Basin, but this thins out to

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the north, west and east. The Penguin Formation, composed dominantly of claystones with minor siltstones and sandstones, is also thickest in the southern Bonaparte Basin, thinning northwards into distal claystones. These two formations record several cycles of transgression and regression, but these are most marked at the southern end of the Bonaparte Basin, well onto the Australian continental margin, whilst more distal parts of the shelf accumulated thin claystones. The transgressive±regressive cycles are therefore low (third?) order cycles within a higher (second?) order sea level highstand. This is consistent with the continuing transgressive conditions identi®ed at this time in Timor. In summary, the Permian stratigraphy of Timor can be correlated fairly closely with the Australian Northwest Shelf in terms of sequence stratigraphy (timing of transgressive and regressive phases) allowing for contemporaneous tectonism in Timor, but lithostratigraphic correlation is not close, except perhaps for the distally located exploration wells such as Sahul Shoals-1. The Permian of the inner Northwest Shelf is dominated by relatively proximal clastic successions including several important sandstone intervals. The clastic successions of Timor are predominantly shaly, consistent with a distal position for Timor on the northwest Australian margin during the Permian. On the other hand, the shallow marine successions of Timor are dominated by basic volcanics and shelf carbonates. Volcanics are rare, if present at all, in the Permian of the Northwest Shelf, and carbonate sequences were only developed occasionally, particularly during transgressive phases. Presumably the shallow marine and emergent areas of Timor formed isolated bathymetric/topographic highs on the outer margin of the Australian continental block, and were therefore largely protected from clastic input from the present-day Northwest Shelf region. During transgressive phases, the carbonate environments expanded southward from Timor onto the Australian margin in the absence of signi®cant clastic input. At the same time, and particularly during regressive phases, the Timor basinal grabens received distal clastic input from the emergent Australian continent throughout much of the Permian, channelled particularly along the Bonaparte Graben (Fig. 12). 8.2. Palaeomagnetism Three palaeomagnetic studies have been reported from the Permian of Timor: 1. Chamalaun (1977a) determined a palaeolatitude of 348S from the Cribas Formation in its type locality near Cribas village, East Timor. The Cribas Formation was interpreted by Chamalaun as Late Permian in age, following Audley-Charles (1968), but the age range of the formation is now established as Kungurian±Late Permian, and probably continuing into the Triassic (Bird, 1987; Barkham, 1993).

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Fig. 14. Palaeotectonic setting of the Timor region through the Permian (Charlton, 2001). The palaeolatitudes shown may correspond approximately to the position of Timor at the beginning of the Permian and at the beginning of the middle Permian in current terminology (Jin et al., 1997), based on the poorly constrained Australian polar wandering curve (Klootwijk, 1996) and palaeoenvironmental data from Timor (e.g. Riding and Barkham, 1999).

2. Chamalaun (1977b) reported a palaeolatitude of 268 from the Maubisse Formation in its type area in East Timor, and showed that at the a 95 con®dence level the Maubisse and Cribas palaeomagnetic data sets are indistinguishable. Blundell (1979) pointed out, however, that this uncertainty still permitted a separation of 2000 km between the two sites. The age of the magnetisation is not well constrained, as the Maubisse Formation at Maubisse apparently extends from the Early Permian (from at least the late Tastubian±Sterlitamakian: Hasibuan, 1994) to the Late Permian (Audley-Charles, 1968), and near Manatuto, into the Triassic (Berry et al., 1984). 3. In West Timor Wensink and Hartosukohardjo (1990) determined a palaeolatitude of 398 from the Maubisse Formation. This was a composite of two localities, one 7 km east of Kefamananu with a mean palaeolatitude of 37.78, and a second near the village of Suanae 20 km SW of Kefamenanu with a palaeolatitude of 43.28. Again no detailed determination of the site ages was given by the authors. Palaeoenvironmental indicators suggest that in the late Sakmarian Timor lay at temperate latitudes (the Hoeniti Member at Bisnain), but by the latest Artinskian/earliest Kungurian Timor had moved to sub-tropical latitudes (the Nannait Member at Laktutus). By the Tatarian/Late Permian Timor appears to have reached near tropical latitudes (Riding and Barkham, 1999). During the Asselian Timor may have been located at high temperate to sub-polar latitudes, if it was ®xed essentially to northwestern Australia, which was experiencing glaciation at that time.

Through the Permian, therefore, Timor appears to have moved over a considerable range of latitudes, from near polar to at least sub-tropical. The palaeomagnetic results obtained so far are consistent with the palaeoenvironmental data, but do not provide any signi®cant control, as they are not adequately constrained by age. There is clearly scope for further detailed palaeomagnetic study of the Permian of Timor in palaeontologically well-dated material, linked rigorously to the stratigraphy. 8.3. Permian regional tectonic setting The regional tectonic setting of Timor in the Permian, as interpreted by Charlton (2001), is shown in Fig. 14. The Bonaparte Graben passing NW±SE through the Timor region is an older Palaeozoic feature, probably an aulacogen formed during Devonian rifting (Lee and Gunn, 1988; Metcalfe, 1996). The formation of the E± W trending South Banda Graben through northern Timor in the Early Permian was associated with the separation of the Sibumasu Terrane of SE Asia from northern Gondwanaland (Metcalfe, 1996). In Fig. 14 it is suggested that the oceanic rift that developed between Sibumasu and northern Gondwanaland passed eastwards in the Timor region into an area of intracontinental extension, comparable to the northern end of the present-day Red Sea rift. Extension continued within Gondwanan eastern Indonesia through the Permian and into the Triassic, but did not develop to the stage of full oceanic rifting until a later phase of continental breakup in the Late Jurassic±Early Cretaceous.

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Acknowledgements Past and present members of the University of London Southeast Asian Research Group are deeply indebted to successive Directors of the Geological Research and Development Centre, Bandung for their continued administrative and logistic support for the research programme in Timor and to the many geologists from GRDC who participated enthusiastically in the ®eldwork, including Soebardjio Tjokrosapoetro, Tohap Simandjuntak, Suharsono, Ichsan Umar, Kusnama Brata, Kustomo Hasan, Said Aziz, Syarif Hidayat and Herman Sugilar. We are also indebted to the people of West Timor who acted as guides, companions and ®eld support teams. Dr Ron Berry (Tasmania) has made useful comments on an earlier version of the text. Shuji Niko (Hiroshima) updated the orthocone nomenclature. Appendix A. Listing of species identi®ed in the Permian of Timor Ammonoids (HGO) Adrianites beyrichi (Haniel) Adrianites cancellatus forma globosa (Haniel) Adrianites (Crimites) oyensi (Haniel) Adrianites rothpletzi (Haniel) Adrianites timorensis (Boehm) Adrianites timorensis (Boehm) var. involuta Haniel Adrianites wichmanni (Haniel) Agathiceras brouweri Smith Agathiceras globosum (Haniel) Agathiceras involuta Haniel Agathiceras martini Haniel Agathiceras sundaicum Haniel Arcestes megaphyllus Beyrich Artinskia simile (Haniel) Artinskia timorensis (Haniel) Atsabites sp. Cyclolobus persulcatus (Rothpletz) Daraelites submeeki Haniel Endolobus (Solenocheilus) brouweri Haniel Eoasianites (Somoholites) beluensis (Haniel) Eoasianites hanieli (Smith) Eoasianites somoholense Haniel Epadrianites involutus Schindewolf Episageceras nodosum Wanner Episageceras noetlingi Haniel Eumedlicottia subprimas Haniel Gastrioceras angulatum (Haniel) Gastrioceras beluense Haniel Gastrioceras hanieli Smith Gastrioceras melanesianum Smith Gastrioceras somoholense Haniel Ghazeloceras? sp. Hyattoceras subgeinitzi Haniel

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Hyattoceras waageni Smith Jurasinites hanieli Smith Lecanites pseudo-meneghinii (Haniel) Lecanites weberi (Haniel) Marathonites dieneri Smith Marathonites gracilis Smith Medlicottia (Artinskia) artiensis Gruenewaldt var. timorensis Haniel Medlicottia magnotuberculata Tschernyschew Medlicottia orbignyana Vernuil Medlicottia sp. det. aff. orbignyana Vernuil Metalegoceras evolutum (Haniel) Metalegoceras gigas (Smith) Metalegoceras somoholense (Haniel) Metalegoceras sundaicum (Haniel) Metalegoceras tschernyschewi (Karpinsky) Neopronorites timorensis Haniel Neostacheoceras hanieli Schindewolf Neostacheoceras tridens (Rothpletz) Paragastrioceras? sp. Paralegoceras australe Smith Paralegoceras wanneri Smith Parapopanoceras dyadicum Haniel Parapronorites timorensis (Haniel) Peritrochia dieneri Smith Peritrochia gracilis (Smith) Peritrochia timorense (Haniel) Perrinites brouweri Smith Perrinites subcumminsi (Haniel) Popanoceras boesei Smith Popanoceras clausum (Gemmellaro) Popanoceras indo-australicum Haniel Pronorites cf. postcarbonarius (Karpinsky) Pronorites uralensis Karpinsky var. timorensis Haniel Propinacoceras sp. det. aff. af®ne Gemmellaro Propinacoceras insulcatum Haniel Propinacoceras simile Haniel Propinacoceras transitorium Haniel Prostacheoceras dieneri (Smith) Pseudoschistoceras sp. Rhiphaeites spp. (2) Sicanites sp. det. aff. mojsisovicsi Gemmellaro Sicanites papuanus Smith Somoholites deroeveri Saunders Stacheoceras arthaberi Smith Stacheoceras timorense (Haniel) Stacheoceras wanneri (Haniel) Stenopronorites timorensis (Haniel) Sundaites levis Haniel Tainoceratid Thalassoceras dieneri Smith Timorites curvicostatus Haniel Timorites hanieli Smith Timorites striatus Hanniel Waagenoceras gemmellaroi Haniel Waagenoceras intermedium Wanner

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Waagenoceras lidacense de Roever Xenaspis beedei Smith Xenaspis sp. Wanner Xenodiscus rotundus Haniel Bellerophonts (NJM) Bellerophon galeatus C. Wanner Bellerophon galeatus var. expers C. Wanner Bellerophon quadratus C. Wanner Bellerophon timorensis C. Wanner Bellerophon timorensis var. gibber C. Wanner Bellerophon timorensis var. umbilicata C. Wanner Bellerophon sp. 1±2 C. Wanner Bucania lyelli Gemmellaro Bucanopsis kattaensis (Waagen) Bucanopsis kattaensis Waagen var. carinata C. Wanner Bucanopsis timorensis (Hamlet) Euphemites cf. carbonarius (Cox) Euphemites sundaicus C. Wanner Patellilabia timorense Hamlet Bivalves (NJM) Atomodesma exarata Beyrich Atomodesma exarata Beyrich var. elongata C. Wanner Atomodesma exarata Beyrich var. lata C. Wanner Atomodesma multifurcata C. Wanner Atomodesma Evenia mytiloides Beyrich Atomodesma Evenia timorensis C. Wanner Atomodesma? undulata Rothpletz Atomodesma variabilis (? ˆ multifurcata) C. Wanner ?Avicula sp. Hamlet ?Aviculopecten brouweri Hamlet ?Aviculopecten sundaensis Hamlet Carbonarca? inaequivalvis C. Wanner Cardiomorpha? sp. Cypricardinia mytiliformis C. Wanner Edmondia sp. Euchondria weiensis C. Wanner Eurydesma sp. Grammysia? sp. Palaeolima cf. retifera Shumard Lithodomus timorensis C. Wanner Merismopteria cf. macroptera (Morris) Myalina? sp. indet. Myoconcha timorensis C. Wanner Mysidioptera? sp. Nuculana oyensi C. Wanner ?Cyrtorostra atavum Waagen Parallelodon subtilistriatus C. Wanner Parallelodon sundaicum Hamlet Pecten sp. Hamlet Pinna sp. Hamlet Pleurophorus? dubius C. Wanner Pleurotomariid sp. indet.

Oriocrassatella plana (Golowkinsky) Sanguinolites? bitauniensis C. Wanner Schizodus? cf. curtus Meek & Worthen Blastoids (GDW) Angioblastus depressus Wanner Angioblastus variabilis Wanner Anthoblastus brouweri Wanner Anthoblastus stelliformis Wanner Calycoblastus tricavatus Wanner Ceratoblastus nanus Wanner Deltoblastus batheri (Wanner) Deltoblastus crassus (Jansen) Deltoblastus delta (Bather) Deltoblastus delta elongatus (Wanner) Deltoblastus delta subglobosus (Wanner) Deltoblastus jonkeri (Wanner) Deltoblastus magni®cus (Wanner) Deltoblastus molengraaf® (Wanner) Deltoblastus molengraaf® sebotensis (Wanner) Deltoblastus permicus (Wanner) Deltoblastus permicus ellipticus (Wanner) Deltoblastus pseudodelta (Wanner) Deltoblastus somoholensis (Wanner) Deltoblastus timorensis (Boehm) Deltoblastus timorensis globosus (Wanner) Deltoblastus verbeeki (Wanner) Deltoblastus verbeeki (Wanner) elongatus (Jansen) Dipteroblastus permicus Wanner Indoblastus granulatus Wanner Indoblastus nuciformis Wanner Indoblastus weberi (Wanner) Nannoblastus cuspidatus Wanner Nannoblastus pyramidatus Wanner Neoschisma timorense Wanner Neoschisma verucosum Wanner Notoblastus oyensi Wanner Orbitremites malaianus Wanner Orbitremites malaianus constrictus Wanner Pteroblastus brevialatus Wanner Pteroblastus brevialatus depressus Wanner Pteroblastus decemcostis Wanner Pteroblastus ferrugineus Wanner Pteroblastus gracilis Wanner Rhopaloblastus timoricus Wanner Sphaeroschisma somoholense Wanner Thaumatoblastus longiramus Wanner Thaumatoblastus longispinus Wanner Timoroblastus coronatus Wanner Timoroblastus coronatus altus Wanner Timoroblastus coronatus basiplanus Wanner Timoroblastus coronatus carinatus Wanner Timoroblastus coronatus constrictus Wanner Timoroblastus coronatus depressus Wanner Timoroblastus coronatus elongatus Wanner

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Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus Timoroblastus

coronatus erectus Wanner coronatus expansus Wanner coronatus informis Wanner coronatus ingens Wanner coronatus levis Wanner coronatus longipedatus Wanner coronatus pentagonalis Wanner coronatus semiconstrictus Wanner coronatus tesselatus Wanner coronatus ungulatus Wanner weiensis Wanner

Brachiopods (NWA) Arctitreta? sp. Asperlinus sp. `Athyris' globulina Waagen Athyris cf. semiconcava Waagen Athyris timorensis Aulosteges dalhousi Davidson Aulosteges tibeticus Diener Callispirina cf. wimani Callytharrella khalii Archbold and Barkham Camarophoria antisella Broili Camarophoria crassa Hamlet Camarophoria globosa Tscernyschew Camarophoria nucula Schwellwien Camarophoria pinguis Waagen Camarophoria tenui-striata Hamlet Cardiocrania sp. Chonetella nasuta Waagen Chonetes dubia Hamlet Chonetes jonkeri Hamlet Chonetes molengraaf® Broili Chonetes rothpletzi Broili Cleiothyridina royssii (LeÂveilleÂ) var. subexpansa Waagen Cleiothyridina subexpansa (Waagen) Composita cf. C. advena Derbyia grandis Waagen Dictyoclostus gratiosus (Waagen) Dictyoclostus gratiosus (Waagen) var. timorensis (Hamlet) Dictyoclostus semireticulatus (Martin) Dictyoclostus spiralis (Waagen) Dielasma biplex Waagen Dielasma burcki Hamlet Dielasma elongatum Schlotheim var. orientalis Grabau Dielasma?itaitubense Derby Dielasma nummulus Waagen Dielasma sundaensis Hamlet Dielasma truncatum Waagen Eliva timorensis Archbold and Bird Elivina bisnaini Archbold and Barkham Elivina cf. brachythyride Enteletes derbyi Waagen var. demissa Schwellwien

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Eumetria cf. grandicosta (Davidson) Globiella foordi (Etheridge) Hamletella altus (Hamlet) Krotovia? opuntia (Waagen) Leptodus catenata Wanner Leptodus nobilis Waagen Linoproductus cora (D'Orbigny) Lythonia sp. Marginifera transversa Waagen Marginifera typica Waagen Martinia carinthiaca Schwellwien Martinia corculum Kutorga Martinia elongata Waagen Martinia glabra (Martin) Martinia nucula Rothpletz Martinia simensis var. substricta Tschernyschew Martinia timorensis Hamlet Meekella striato-costata Cox Neophrycodothyris indica (Waagen) Neospirifer mooshakhailensis (Davidson) Notothyris leuchsi Hamlet Notothyris cf. mediterranea Gemmellaro Notothyris minuta Waagen Notothyris nucleolus Kutorga Notothyris triplicata Diener Oldhaminella sp. Orbicoelia sp. Orthis indicaeformis Hamlet Orthis? molengraaf® Hamlet Orthothetes crenistria var. senilis Phillips Orthothetes (Streptorhynchus) semiplanus Waagen Paralyttonia sp. Phricodothyris sp. Plicatifera minor (Schwellwien) Poikilosakos variabilis Productella? patula Girty Productus abichi Waagen Productus asperulus Waagen Productus cancriniformis Tschernyschew Productus chitichunensis Diener Productus (Pustula?) elegans M'Coy Productus humboldti D'Orbigny Productus humboldti D'Orbigny var. irginae Stuckenberg Productus opuntia Waagen Productus punctatus Martin Productus purdoni Davidson Productus semireticulatus Martin Productus transversalis Tscernyschew Productus waageni Rothpletz Productus (Marginifera) wanneri Broili Punctocyrtella sp. Reticularia broilii Hammlet Retzia (Eumetria) grandicosta Davidson (Waagen) Retzia (Hustedia) indica Waagen Retzia (Hustedia) indica Waagen var. tenui-striata

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Hamlet Retzia (Hustedia) radialis Phill. var. grandicosta Davidson Rhipidomella cora D'Orbigny Rhynchonella hanieli Broili Rhynchonella (Uncinulus) jabiensis Waagen Rhynchonella (Uncinulus) timorensis Beyrich Rhynchonella wichmanni Rothpletz `Rhynchonella' cf. wynnei Waagen Richthofenia? lawrenciana de Koninck Flem. Rugaria molengraaf® Broili Schizophoria? indicaeformis (Hamlet) Schuchertella beyrichi Rothpletz Spirifer lyra Kutorga `Spirifer' oldhamianus Waagen Spirifer simaanensis Hamlet Spirifer basleoensis Hayasaka and Hosono Spirifer tibetanus (Diener) Spiriferella orientensis Archbold and Bird Spiriferella rajah (Salter) Spiriferina cristata Schlotheim Spiriferina (Mentzlia) nefotassiensis Hamlet Spiriferina wimani Hamlet Spiriferinellina sp. cf. insculpta (Schlotheim) Spirigera subtilita (Hall) Spirigera timorensis Rothpletz Spirigerella grandis Waagen Spirigerella timorensis Squamularia grandis Squamularia lineata (Martin) Squamularia? indica (Waagen) Stenoscisma gigantea (Diener) Stenoscisma globosa (Tschernyschew) Stenoscisma purdoni (Davidson) Stenoscisma (Camarophoria) purdoni (Davidson) var. gigantea Diener Stenoscisma timorense (Hayasaka and Gan) Stictozoster sp. Streptorhynchus beyrichi Rothpletz Streptorhynchus cf. crenistria Phill. Streptorhynchus interplicatus Rothpletz Streptorhynchus pectiniformis Davidson Streptorhynchus pseudo-pelargonautus Broili Strophalosia indica Waagen Terebratula (Dielasma) elongata Schlotheim Terebratula himalayensis Davidson var. sparsiplicata Waagen Transennatia timorensis (Hamlet) Waagenoconcha waageni (Rothpletz) Bryozoans (PDT) Acanthocladia acuticosta Bassler Acanthocladia rectifurcata Bassler Acanthocladia regularis Bassler Clausotrypa conferta Bassler

Clausotrypa minor Bassler Clausotrypa separata Bassler Dyscritella adnascens Bassler Dyscritella spinulosa Bassler Eridopora parasitica (Waagen and Wentzel) Eridopora oculata Bassler Fenestella [s.l.] aspratilis Bassler Fenestella [s.l.] basleoensis Bassler Fenestella kukaensis Bassler Fenestella [s.l.] latericrescens Bassler Fenestella [s.l.] orientalis Bassler Fenestella [s.l.] parviuscula Bassler Fenestella [s.l.] pulchradorsalis Bassler Fenestella [s.l.] regina Bassler Fenestella virgosa Eichwald Fistulipora basleoensis Bassler Fistulipora crassilabiata Bassler Fistulipora crebriseptata Bassler Fistulipora labratula Bassler Fistulipora lunatifera Bassler Fistulipora? mackloti (Beyrich) Fistulipora milleporacea Bassler Fistulipora molengraaf® Bassler Fistulipora muÈlleri Beyrich Fistulipora octonaria Bassler Fistulipora parallela Bassler Fistulipora simillima Bassler Fistulipora timorensis Bassler Fistulipora wanneri Bassler Fistulocladia typicalis Bassler Fistulotrypa ramosa Bassler Goniocladia indica Waagen and Pichler Goniocladia laxa (Bassler) Goniocladia timorensis Bassler Hexagonella turgida Bassler Hinganella laeviuscula (Bassler) Neorhombopora gratiosa (Bassler) Neorhombopora orientalis (Bassler) Neorhombopora pulchra (Bassler) Penniretepora crassicaulis (Bassler) Penniretepora ¯exicaulis (Bassler) Penniretepora pulchella (Bassler) Penniretepora scalaris (Bassler) Phyllopora? robusta Bassler Polypora brouweri Bassler Polypora consanguinea Bassler Polypora macrops Bassler Polypora magnidiscus Bassler Polypora timorensis Bassler Polypora tripliseriata Bassler Rhabdomeson consimile Bassler Rhabdomeson grande Bassler Septoptera orientalis Bassler Sphragiopora crateriformis Bassler Stenopora parvulipora (Bassler) Stenopora spicata (Bassler)

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Streblascopora fasciculata (Bassler) Streblascopora germana (Bassler) Streblotrypa minutula Bassler Streblotrypa? ignota Bassler Tabulipora tenuinervis Bassler Thamniscus cf.dubius (Schlotheim) Thamniscus humilis Bassler Thamniscus megastoma Bassler Ulrichotrypa magnopora Bassler Ulrichotrypa permiana Bassler Ulrichotrypa ramulosa Bassler Ulrichotrypella wanneri (Bassler) Ulrichotrypa zonata Bassler Voigtella? timorensis (Bassler) ?Voigtella sp. Conodonts (RSN) Diplognathus oertlii Neogondolella carinata (Clark) Hindeodus sp. Neogondolella bisselli, Neospathodus cf. praedivergens, Sweetocristatus sp. Sweetognathus cf. behnkeni Sweetognathus aff. whitei Vjalovognathus australis Nicoll and Metcalfe Vjalovognathus shindyensis. Taxonomic comments: Sweetognathus aff. whitei ˆ S. inornatus; Vjalovognathus shindyensis ˆ V. australis Corals (rugose) (JES) ?Amplexocarinia abichi (Waagen and Wentzel) ?Amplexocarinia beyrichi (Gerth) Amplexocarinia bitauniensis Schouppe and Stacul Amplexocarinia composita Schouppe and Stacul Amplexocarinia geyeri Heritsch Amplexocarinia heritschi Schouppe and Stacul Amplexocarinia naliensis (Gerth) Amplexocarinia subtilis Schouppe and Stacul Asserculinia prima Schouppe and Stacul Basleophyllum indicum (Koker) Basleophyllum pachyderma (Koker) Basleophyllum solidum Schouppe and Stacul Calophyllum angustum (Rothpletz) Calophyllum clausum (Nierman) Calophyllum multiseptatum (Koker) Calophyllum pennatum (Nierman) Calophyllum rarum (Nierman) Duplocarinia micron (Schouppe and Stacul) Duplocarinia tenueseptatum (Schouppe and Stacul) Duplocarinia timorica Fedorowski Duplophyllum calyculatum (Koker) Duplophyllum schindewol® Schouppe and Stacul

763

Duplophyllum wanneri Schouppe and Stacul Duplophyllum zaphrentoides Koker ? Duplophyllum permicum (Schouppe and Stacul) Endamplexus dentatus Koker Endamplexus kokeri Schouppe and Stacul Endamplexus makros Schouppe and Stacul Endamplexus raripusolosus Schouppe and Stacul Endothecium apertum Koker Endothecium decipiens Koker Euryphyllum breviseptatum Schouppe and Stacul Euryphyllum cainodon (Koker) Euryphyllum coniculiforme Schouppe and Stacul Euryphyllum hilli Schouppe and Stacul Euryphyllum robustum (Koker) ?Falsiamplexus jonkeri (Koker) Ipsiphyllum timoricum (Gerth) Kabakovitchiella arcuata (Schouppe and Stacul) Kabakovitchiella duplex (Schouppe and Stacul) Kabakovitchiella thomasi (Schouppe and Stacul) Lonsdaleiastraea molengraaf® (Gerth) Lonsdaleiastraea vinassai Gerth Lophophyllidium gracile Schouppe and Stacul Lophophyllidium parvum Schouppe and Stacul Lophophyllidium (Lophbillidium) elongatum Wang Lophophyllidium (Lophbillidium) martini Schouppe and Stacul Lophophyllidium (Lophbillidium) spinosum (Martin) Paralleynia amphibolos Schouppe and Stacul Paralleynia leptoseptata Schouppe and Stacul Paralleynia soshkinae Schouppe and Stacul Pentamplexus simulator Schindewolf Pentamplexis subtilis Nierman Pentaphyllum elegans Nierman Pentaphyllum frons Nierman Pentaphyllum parallelum Nierman Pentaphyllum praetercrescens Nierman Pentaphyllum subcylindicum Schindewolf Pleramplexus dissimilis Schindewolf Pleramplexus grandis Nierman Pleramplexus similis Schindewolf Plerophyllum aequabile Schindewolf Plerophyllum bitaunense Koker Plerophyllum gerthi Schindewolf Plerophylum radiciforme Gerth Plerophyllum tenue Koker Plerophyllum weberi (Gerth) Productiophyllum brouweri (Schouppe and Stacul) Productiophyllum incertum (Koker) Productiophyllum pusillum (Schouppe and Stacul) Prosmilia compressa Koker Prosmilia cyathophylloides (Gerth) Sochkineophyllum laxa Nierman Spineria diplochone (Koker) Spineria ultima (Koker) Spineria uniformis Schouppe and Stacul Tachylasma beyrichi (Rothpletz)

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Tachylasma densum Hill Tachylasma gracile Schindewolf Tachylasma isoseptatum (Koker) Tachylasma makrodeuterum Nierman Tachylasma mediacompactum Nierman Tachylasma paululum Nierman Tachylasma ponderosum Schindewolf Tachylasma superans Schindewolf Tachylasma timorense (Gerth) Tachylasma variabile Schindewolf Tachylasma (Prionophyllum) crassiseptum (Schindewolf) Timorphyllum breviseptatum Schouppe and Stacul Timorphyllum complicatum Wang Timorphyllum wanneri Gerth U®mia formosum Schindewolf U®mia isophyllum Schindewolf U®mia kobayashii Schindewolf. U®mia multitabulatum Schindewolf U®mia persymmetricum Schindewolf. Verbeekiella arboricalis Schouppe and Stacul Verbeekiella australis (Beyrich) Verbeekiella gerthi Schouppe and Stacul Verbeekiella rothpletzi (Gerth) Wannerophyllum asteroides Schouppe and Stacul Wannerophyllum cristatum (Gerth) Wannerophyllum elongatum Schouppe and Stacul Wannerophyllum excedens Schouppe and Stacul Wannerophyllum minus Schouppe and Stacul Wannerophyllum timoricum Schouppe and Stacul Wannerophyllum torquatum (Rothpletz) Wannerophyllum tubulosum (Gerth) Corals (tabulate) (JES) Aulohelia irregularis Gerth Aulohelia laevis Gerth Aulopora timorica Gerth Aulopora minima Hehenwarter Cladochonus crassus (McCoy) Cladochonus magnus Gerth Cladochonus beecheri (Grabau) `Dictyopora' incrustans Gerth ?Favosites permicus Gerth ?Favosites relicta Gerth Gertholites curvata (Waagen and Wentzel) Gertholites jabiensis (Waagen and Wentzel) Gertholites monstrosa (Gerth) Gertholites lobata (Gerth) Heterocoenites variabilis Gerth, Heterocoenites crassus Gerth Michelinia indica Waagen and Wentzl Michelinia bursifera Hehenwarter Palaeacis regularis Gerth Palaeacis tubifer Gerth Pseudofavosites stylifer Gerth

Trachypsammia dendroides Gerth Trachypsammia monoseptata Hehenwarter Schizophorites dubiosus Gerth Stylonites porosus Gerth Crinoids (GDW) Abrachiocrinus conicus Wanner Abrachiocrinus timoricus Wanner Acariaiocrinus angulosus Wanner Acariaiocrinus clavulus Wanner Actinocrinites brevispinus Wanner Actinocrinites brouweri Wanner Actinocrinites carinatus Wanner Actinocrinites dilatatus Wanner Actinocrinites exornatus Wanner Actinocrinites permicus Wanner Actinocrinites spinaetectus Wanner Actinocrinites timoricus Wanner Allosycocrinus? grandis Wanner Allosycocrinus? medius Wanner Allosycocrinus pusillus Wanner Ancistrocrinus depressus Wanner Ancistrocrinus vermistriatus Wanner Apographiocrinus quinquelobus (Wanner) Apographiocrinus rugosus (Wanner) Apographiocrinus verbeeki pumilus (Wanner) Apographiocrinus verbeeki vermistriatus Wanner Asymmetrocrinus poteriocrinoides Wanner Atremacrinus calyculus Wanner Basleocrinus conicus Wanner Basleocrinus obliquus Wanner Basleocrinus pocillum Wanner Basleocrinus pusillus Wanner Basleocrinus striategranulatus Wanner Basleocrinus turbinatus Wanner Benthocrinus cryptobasalis Wanner Bolbocrinus curvatus Oyens Bolbocrinus hieroglyphicus Wanner Bolbocrinus hieroglyphicus exornatus Wanner Bolbocrinus hieroglyphicus tenuisculptus Wanner Bolbocrinus hieroglyphicus tuberculatus Wanner Bolbocrinus irregularis Wanner Bolbocrinus pusillus Wanner Bolbocrinus rex Wanner Bolbocrinus turbinatus Wanner Bolbocrinus waldthauseniae Wanner Bolbocrinus waldthauseniae basleoensis Wanner Cadocrinus amarassicus Wanner Cadocrinus variabilis canaliculatus Wanner Cadocrinus variabilis Wanner Calycocrinus amarassicus Wanner Calycocrinus crassus Wanner Calycocrinus curvatus Wanner Calycocrinus curvatus conicus Wanner Calycocrinus curvatus coronatus Wanner

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Calycocrinus curvatus depressus Wanner Calycocrinus curvatus elongatus Wanner Calycocrinus curvatus informis Wanner Calycocrinus curvatus labrosus Wanner Calycocrinus curvatus subcoronatus Wanner Calycocrinus curvatus subturbinatus Wanner Calycocrinus curvatus turbinatus Wanner Calycocrinus erectus Wanner Calycocrinus granulatus Wanner Calycocrinus granulatus altior Wanner Calycocrinus kupangensis Wanner Calycocrinus labiatus Oyens Calycocrinus major Wanner Calycocrinus malaianus Wanner Calycocrinus millericrinoides Wanner Calycocrinus nuciformis Wanner Calycocrinus patella Wanner Calycocrinus perplexus Wanner Calycocrinus piriformis Wanner Calycocrinus poculum Wanner Calycocrinus similis Wanner Calycocrinus spinosus Wanner Calycocrinus tuberculatus Wanner Calycocrinus venemai Wanner Calycocrinus venemai angulatus Wanner Calycocrinus venemai Wanner planus Oyens Camptocrinus indoaustralicus Wanner Ceratocrinus exornatus Wanner Ceratocrinus gracilis Wanner Cibolocrinus spinosus Wanner Cibolocrinus timorensis Wanner Coenocystis angulosus Wanner Coenocystis perforatus Wanner Coenocystis somoholensis Wanner Contignatocrinus contignatus (Wanner) Cranocrinus timoricus Wanner Cydonocrinus turbinatus Wanner Cydonocrinus turbinatus minor Wanner Calceolispongia aculeatus (Wanner) Calceolispongia bicornutus (Wanner) Calceolispongia bifurcus (Wanner) Calceolispongia cornutus (Wanner) Calceolispongia elegans (Wanner) Calceolispongia horridus (Wanner) Calceolispongia mammeatus (Wanner) Delocrinus crassus (Wanner) Delocrinus? malaianus Wanner Depaocrinus ottowi Wanner Dichostreblocrinus timorensis Oyens Embryocrinus hanieli Wanner Eutelecrinus mangostanus Wanner Eutelecrinus piriformis Wanner Eutelecrinus poculiformis Wanner Eutelecrinus subglobosus Wanner Graphiocrinus amplior Wanner Graphiocrinus beyrichi Wanner

Graphiocrinus beyrichi nustoiensis Wanner Graphiocrinus declivis Wanner Graphiocrinus? depressus Wanner Graphiocrinus? depressus labiosus Wanner Graphiocrinus? excavatissimus Wanner Graphiocrinus? excavatissimus ornatus Wanner Graphiocrinus ovoides Wanner Graphiocrinus pumilis Wanner Graphiocrinus punctatus Wanner Graphiocrinus rotundatus Wanner Graphiocrinus scrobiculatus Wanner Graphiocrinus subamplior Wanner Graphiocrinus verbeeki Wanner Hemistreptacron carinatum Wanner Hemistreptacron carinatum ornatum Wanner Hypocrinus schneideri Beyrich Indocrinus crassus Wanner Indocrinus elegans Wanner Indocrinus nodosus Wanner Indocrinus turgidus Wanner Isocatillocrinus indicus Wanner Jonkerocrinus? conicus Oyens Jonkerocrinus spinosus Wanner Lopadiocrinus angustecavatus Wanner Lopadiocrinus brouweri Wanner Lopadiocrinus brouweri vermistriatus Wanner Lopadiocrinus granulatus Wanner Lopadiocrinus granulatus labiosus Wanner Lopadiocrinus tuberculatus Wanner Loxocrinus booni Oyens Loxocrinus dilatatus Wanner Loxocrinus globulus Wanner Malaiocrinus crassitesta Wanner Malaiocrinus pusillus Wanner Malaiocrinus sundaicus Wanner Metaeutelecrinus fritillus (Wanner) Metallagecrinus acutus (Wanner) Metallagecrinus dux (Wanner) Metallagecrinus excavatus (Wanner) Metallagecrinus indoaustralicus (Wanner) Metallagecrinus in¯atus (Wanner) Metallagecrinus ornatus (Wanner) Metallagecrinus procerus (Wanner) Metallagecrinus quinquebrachiatus (Wanner) Metallagecrinus quinquelobus (Wanner) Metasycocrinus piriformis (Rothpletz) Mollocrinus ornatissimus Wanner Mollocrinus paucituberculatus Wanner Mollocrinus poculum Wanner Monobrachiocrinus ®ciformis Wanner Monobrachiocrinus ®ciformis carinatus Wanner Monobrachiocrinus ®ciformis elongatus Wanner Monobrachiocrinus ®ciformis granulatus Wanner Monobrachiocrinus waitzi Oyens Neocatillocrinus incissus Wanner Neodichocrinus nanus Wanner

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Neolageniocrinus timorensis Wanner Neoplatycrinus dilatatus Wanner Neoplatycrinus major Wanner Neoplatycrinus somoholensis Wanner Neoplatycrinus transitorius Wanner Neozeacrinus peramplus Wanner Neozeacrinus? springeri Wanner Nereocrinus antiquus Wanner Nereocrinus granulatus (Wanner) Notiocrinus ekadensis Wanner Notiocrinus timoricus Wanner Oklahomacrinus expansus (Wanner) Palaeoholopus pretiosus Wanner Parabursacrinus compressus Wanner Parabursacrinus conus Wanner Parabursacrinus? gracilis Wanner Parabursacrinus magni®cus Wanner Parabursacrinus magni®cus granulatus Wanner Parabursacrinus nefotassiensis Wanner Parabursacrinus procerus Wanner Parabursacrinus pyramidatus Wanner Parabursacrinus pyramidatus granulatus Wanner Paracatillocrinus granulatus Wanner Paracatillocrinus spinosus Wanner Paradoxocrinus patella Wanner Paraeutelecrinus elongatus (Wanner) Paraeutelecrinus erectus (Wanner) Paraeutelecrinus subwelteri Wanner Pareutelecrinus welteri Wanner Paraplasocrinus transitorius (Wanner) Parastachyocrinus granulatus (Wanner) Parastachyocrinus in¯atus Wanner Parastachyocrinus obliquus (Wanner) Parastachyocrinus malaianus Wanner Parastachyocrinus malaianus ornatus Wanner Parasycocrinus fastigate-pileatus Oyens Pargraphiocrinus exornatus Wanner Parindocrinus oyensi Wanner Peripterocrinus gracilis Wanner Permiocrinus immaturus Wanner Permobrachypus adhaerens Wanner Petrocrinus beyrichi Wanner Petrocrinus boschi Oyens Petrocrinus kukaensis Wanner Pilidiocrinus permicus Wanner Plagiocrinus infratectus Wanner Plagiocrinus jaekeli Wanner Plagiocrinus torycrinoides Wanner Platycrinites cf. rugosus Miller Platycrinites wachsmuthi Wanner Platycrinites wachsmuthi apnaÈensis Wanner Platycrinites wachsmuthi frequentior Wanner Platycrinites wrighti Oyens Plesiocrinus piriformis Wanner Pleurocrinus depressus Wanner Pleurocrinus globosus Wanner

Pleurocrinus goldfussi Wanner Pleurocrinus pusillus Wanner Pleurocrinus spectabilis Wanner Poteriocrinus malaianus Wanner Proapsidocrinus cuspidatus Wanner Proapsidocrinus permicus Wanner Prochoidiocrinus nodusus Wanner Prolobocrinus? gracilis (Wanner) Prolobocrinus permicus Wanner Prolobocrinus striatus Wanner Prophyllocrinus dentatus Wanner Pumilindocrinus angulosus (Wanner) Pumilindocrinus pumilus (Wanner) Roemerocrinus gracilis Wanner Roemerocrinus gracilis granulatus Wanner Roemerocrinus scrobiculatus Wanner Roemerocrinus turbinatus Wanner Rimosindocrinus brevijugatus (Wanner) Rimosindocrinus pachycephalus (Wanner) Rimosindocrinus rimosus (Wanner) Rumphiocrinus singularis Wanner Spaniocrinus validus Wanner Spheniscocrinus spinosus Wanner Stachyocrinus zea Wanner Stomiocrinus minimus Wanner Stomiocrinus piriformis Wanner Stomiocrinus subglobosus Wanner Strongylocrinus molengraaf® Wanner Stuartwellercrinus pusillus Wanner Stuartwelleri jonkeri (Wanner) Stuartwelleri minimus (Wanner) Stuartwelleri propinquus (Wanner) Sundacrinus elongatus Wanner Sundacrinus granulatus Wanner Sundacrinus triangulus Wanner Sundacrinus vastus Wanner Synbathocrinus campanulatus Wanner Synbathocrinus campanulatus elongatus Wanner Synbathocrinus campanulatus in¯atus Wanner Synbathocrinus constrictus Wanner Synbathocrinus constrictus sinuosus Wanner Syntomocrinus sundaicus Wanner Synyphocrinus indicus Wanner Synyphocrinus trautscholdi Wanner Synyphocrinus weidneri Wanner Tapinocrinus timoricus (Wanner) Tapinocrinus spinosus (Wanner) Tenagocrinus sulcatus (Wanner) Teratocrinus? bulbosus Wanner Teratocrinus spathulifer Wanner Teratocrinus? triangulatus Wanner Thetidicrinus piriformis Wanner Timorocidaris baculiformis Wanner Timorocidaris clavaeformis Wanner Timorocidaris fungiformis Wanner Timorocidaris fusiformis Wanner

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Timorocidaris cf. fusiformis Wanner Timorocidaris pistilliformis Wanner Timorocidaris sphaeracantha Wanner Timorechinus alatus (Wanner) Timorechinus alatus major (Wanner) Timorechinus decurtatus (Wanner) Timorechinus lacertosus (Wanner) Timorechinus maximus (Wanner) Timorechinus mirabilis (Wanner) Timorechinus mirabilis vermistriatus (Wanner) Timorechinus multicostatus (Wanner) Timorechinus sp. indet. aff. multicostatus (Wanner) Timorechinus nefotassiensis (Wanner) Timorechinus ovatus (Wanner) Timorechinus pentagonalis (Wanner) Timorechinus pentagonalis nodosus (Wanner) Timorechinus proboscideus (Wanner) Timorechinus spinosus (Wanner) Timorechinus spinosus amarassicus (Wanner) Timorechinus spinosus incissus (Wanner) Trimerocrinus minimus Wanner Trimerocrinus pumilus Wanner Trimerocrinus pumilus pentagonus Wanner Trimerocrinus ventriosus Wanner Ulocrinus? conoideus Wanner Ulocrinus? indicus Wanner Wannerocrinus glans Oyens Wrightocrinus jakovlevi (Wanner) Xenocatillocrinus wrighti Wanner Echinoids (GDW) Archaeocidaridae gen. and sp. indet. Cidaris bitauniensis Wanner Cidaris jonkeri Wanner Miocidaris permica Wanner Neoschisma verrucosum Wanner Permocidaris? timorensis Wanner Radiolus radiatus-tubulatus Radiolus sp. 1±4 Wanner Spiraculata sp. Foraminifera (JEW) Archaeodiscus sp. Cornuspira sp. Dentalina sp. Frondicularia sp. Frondicularia sp. aff. woodwardii Howchin Genitzina triangularis Chapman and Hall Lagenidae Nodosaria sp. aff. postcarbonica Spandel Tetraxis conica Ehrenberg Textularia sp.

Trepilopsis sp. Fusulinids (JEW) Codonofusiella weberi (Schubert) Monodiexodina wanneri Parafusulina cf. japonica Parafusulina gigantica Schwagerina brouweri Thompson Schwagerina cf. pindingensis Schwagerina nakazawae Nogami Schwagerina? molengraaf® (Schubert) Triticites sp. Gastropods (NJM) Agnesia acuminata C. Wanner Agnesia? conformis C. Wanner Ananias timorensis Anomphalus? sundaicus C. Wanner `Cyclonema' basleoense C. Wanner `Cyclonema' bitauniensis C. Wanner `Cyclonema' brouweri C. Wanner `Cyclonema' incertum C. Wanner `Cyclonema' molengraaf® (C. Wanner) `Cyclonema' molengraaf® var. rugosa C. Wanner `Cyclonema' simile C. Wanner Euomphalus cf. catilliformis de Koninck Euomphalus crotalostomiformis C. Wanner Euomphalus sinistrorsus C. Wanner Euomphalus sundaicus C. Wanner Loxonema? sp. Macrocheilus cf. brancoi Gemmellaro Naticopsis cf. mediterranea Gemmellaro Naticopsis ovalis C. Wanner Naticopsis cf. ovalis C. Wanner Naticopsis praealta C. Wanner Naticopsis retusa C. Wanner Naticopsis sp. 1 C. Wanner Omphalotrochus gerthi C. Wanner Pagodina arctiformis C. Wanner Pagodina? incerta C. Wanner Pagodina rugosa C. Wanner Pagodina typus C. Wanner Patella cf. ottomana Kittlinger Patellostium timorense Hamlet Phymatifer nodocarinatus (C. Wanner) Platyceras varians abundans C. Wanner Platyceras varians latum C. Wanner Platyceras varians pretiosum C. Wanner Platyceras varians sundaicum C. Wanner Platyceras varians tortum C. Wanner Platyceras varians varians C. Wanner Platypleurotomaria planiapicata C. Wanner Platyteichum sp. Pleurotomaria cf. carinifera Girty

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Pleurotomaria? clathrata C. Wanner Pleurotomaria? minima C. Wanner Pleurotomaria? sequens Waagen Pleurotomaria subheterospira C. Wanner Pleurotomaria sp. 1±5 C. Wanner Schizosoma pusilliforme (C. Wanner) Schizosoma pusilliforme var. weiensis C. Wanner Soleniscus? sp. cf. chemnithiaeformis (Gemmellaro) Soleniscus ventriculosus C. Wanner Spiroraphella tunsiehensis C. Wanner Straparollus? discoides C. Wanner Straparollus? oyensi C. Wanner cf. Straparollus permianus King Trachydomia tuberosa C. Wanner Turbonellina concinna C. Wanner Turbonellina concinna var. nodulosa C. Wanner Turbonellina dacqueÂi Hamlet Turbonellina grippi Hamlet Turbonellina nitida C. Wanner Turbonellina cf. novosselovkensis Yakowlew Umbonellina? spiralis C. Wanner Worthenia punctata C. Wanner Yvania bitauniensis C. Wanner Yvania conglobata C. Wanner Yvania permica C. Wanner Yvania reticulata C. Wanner Yvania cf. salomonensis (Gemmellaro) Yvania? wanneri (Hamlet) Zygopleura angulata C. Wanner Zygopleura crassa C. Wanner Zygopleura dubia C. Wanner Zygopleura dubia var. geniculata C. Wanner Zygopleura ignorata Trautschold Zygopleura nitida C. Wanner Zygopleura simplex C. Wanner Zygopleura simplex var. latior C. Wanner Hyolithid (NJM) Macrotheca timorensis C.Wanner Nautiloids (Shuji Niko) Atahococeras timorense Niko, Nishida and Nakazawa Bactrites sp. (Orthoceras sp. indet. Nr. 2, Haniel, 1915) Bactrites ? sp. (Orthoceras sp. indet. Nr. 3, Haniel, 1915) Bitaunioceras bitauniense (Haniel, 1915) Discites (Domatoceras) arthaberi Haniel Domatoceras sp. Lopingoceras maubesiense (Haniel) Mooreoceras sp. Nautilus (Aganides) bitauniensis Haniel Nautilus molengraaf® Haniel Nautilus wanneri Haniel Neorthoceras verbeeki (Haniel) Neorthoceras ? sp. (Orthoceras sp. indet. Nr. 1, Haniel,

1915) ªOrthocerasº welteri Haniel, 1915 ªOrthocerasº sp. (Orthoceras sp. indet. Nr. 4, Haniel, 1915) ªOrthocerasº sp.(Orthoceras sp. indet., Martin, 1887) Pleuronautilus dyadicus Haniel Rhiphaeonautilus sp. ?Tainonautilus sp. Temnocheilus sp. indet. Rostroconchs (NJM) Conocardium clipeoforme C. Wanner Conocardium ornatum Hamlet Sponges Aulacospongia? parvula Gerth Aulacospongia bulbosa Gerth Aulacospongia hanieli Gerth Caryospongia? dyadica Gerth Hindia permica Gerth Hindia permica var. bitaoeniensis Gerth Hindia wanneri Gerth Mastophyma globosa Gerth Mastophyma jonkeri Gerth Palaeodesma tubulosa Gerth Palaeojerea molengraaf® Gerth Palaeophyma cucumeriformis Gerth Palaeophyma piriformis Gerth Phacellopygma campana Gerth Phacellopygma praemorsa Gerth Pycnospongia timorensis Gerth Timorella permica Gerth Virgula? malayica Gerth Spores Entylissa cymbatus Limitisporites cf. moersensis Krauselisporites sp. Veryhachium sp. Radiolarian Rhapolodictum sp. Trilobites (RMO) Hildaphillipsia hildae (Gheyselinck, 1937) Microphillipsia? parvula (Beyrich, 1865) Neoproetus indicus (Tesch, 1923) Pseudophillipsia timorensis (Gheyselinck, 1937) Timoraspis breviceps (Gheyselinck, 1937) Triproetus baungensis (Gheyselinck, 1937) [probably includes T. trigonoceps (Gheyselinck, 1937)] Triproetus gerthi (Gheyselinck, 1937) (probably includes

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T. brevicauda (Gheyselinck, 1937), and T. teschi (Gheyselinck, 1937)) Problematic Myzostoma? sp. Shamovella obscura (Maslov) Appendix B. Bibliography of Timor Permian palaeontology The following are the principal references to the palaeontology of Timor (i.e. those featuring primary descriptions of the specimens or containing substantial faunal lists): Ammonoids: Haniel (1915), Smith (1927), Wanner (1932a), de Marez Oyens (1938), Gerth (1950), and Niko et al. (2000). Bivalves: Wanner (1922), Hamlet (1928), Wanner (1940). Brachiopods: Broili (1916), Hamlet (1928), Wanner and Sieverts (1935), Hayasaka and Gan (1940), Hayasaka and Hasono (1951), Hayasaka (1953), Shimizu (1966), Archbold and Bird (1989), Archbold and Barkham (1989), Hasibuan (1994), Kato et al. (1999). Blastoids: Wanner (1924, 1931d, 1932b, 1940b), Webster (1998a,b). Bryozoa: Bassler (1929). Conodonts: van den Boogaard (1987). Corals: Penecke (1908), Gerth (1921, 1922), Koker (1924), Gerth (1926, 1927b), Hehenwarter (1951), Schouppe and Stacul (1955, 1959), Fedorowski (1986). Crinoids: Wanner (1910b, 1912, 1916, 1920, 1923a,b, 1929a,b, 1930a,b, 1931c, 1937, 1940a), de Marez Oyens (1940), Wanner (1942, 1949, 1950), Webster (1998a). Echinoids: J. Wanner (1941). Fusulinids: Schubert (1915), Thompson (1949), Nogami (1963). Gastropods: C. Wanner (1922), Hamlet (1928), C. Wanner (1942). Hyolithid: C. Wanner (1941). Ostracods: Grundel and Kozur (1975), Bless (1987). Sponges: Gerth (1909, 1927a). Trilobites: Tesch (1923), Gheyselinck (1937), Hahn and Hahn (1970, 1972), Hahn and Brauckmann (1975), Owens (1983). General: Beyrich (1862, 1865), Martin (1881), Rothpletz (1891, 1892), Boehm (1908), Wanner (1926, 1931a), Simons (1940), Wanner (1956), Audley-Charles (1968), Bird (1987), Barkham (1993) Detailed references are given in the Reference List. References Archbold, N.W., 1988. Permian Brachiopoda and Bivalvia from Sahul Shoals No. 1, Ashmore Block, Northwestern Australia. Proceedings of the Royal Society of Victoria 100, 33±38.

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Archbold, N.W., 1999. Additional records of Permian brachiopods from near Rat Buri, Thailand. Proceedings of the Royal Society of Victoria 111 (1), 71±86. Archbold, N.W., 2000a. In: Burov, B.V., Esaulova, N.K., Gubareva, V.S. (Eds.). The Australian Permian: Correlations with the Russian Permian Sequences. Proceedings of the International Symposium Upper Permian Stratotypes of the Volga Region, GEOS, Moscow, pp. 53±70. Archbold, N.W., 2000b. Palaeobiogeography of the Australasian Permian. Association of Australian Palaeontologists Memoir 23, 287±310. Archbold, N.W., Barkham, S.T., 1989. Permian Brachiopoda from near Bisnain village, West Timor. Alcheringa 13, 125±140. Archbold, N.W., Bird, P.R., 1989. Permian Brachiopoda from near Kasliu village, West Timor. Alcheringa 13, 103±123. Archbold, N.W., Shi, G.R., 1996. Western Paci®c Permian marine invertebrate palaeobiogeography. Australian Journal of Earth Sciences 43, 635±641. Archbold, N.W., Dickins, J.M., Thomas, G.A., 1993. Correlation and age of Permian marine faunas in Western Australia. In: Skwarko, S.K. (Ed.). Palaeontology of the Permian of Western Australia. Bulletin of the Geological Survey of Western Australia, 136. pp. 11±18. Audley-Charles, M.G., 1968. The Geology of Portuguese Timor. Memoir of the Geological Society of London, 4. Audley-Charles, M.G., Harris, R.A., 1990. Allochthonous terranes of the Southwest Paci®c and Indonesia. Philosophical Transactions of the Royal Society A31, 571±587. Bachri, S., Situmorang, R.L., 1994. Geological Map of the Dili Quadrangle 2406±2407, scale 1:250,000. Geological Research and Development Centre, Bandung. Bakosurtanal (Badan Koordinasi Survey dan Pemetaan Nasional), 1993. Peta Rapabumi Indonesia, 1:25,000, Sheet 2406-113, Oinlasi. Barber, A.J., Audley-Charles, M.G., 1976. The signi®cance of the metamorphic rocks of Timor in the development of the Banda Arc, eastern Indonesia. Tectonophysics 30, 119±128. Barber, A.J., Audley-Charles, M.G., Carter, D.J., 1977. Thrust tectonics on Timor. Journal of the Geological Society of Australia 24, 51±62. Barber, A.J., Tjokrosapoetro, S., Charlton, T.R., 1986. Mud volcanoes, shale diapirs, wrench faults and melanges in accretionary complexes, Eastern Indonesia. American Association of Petroleum Geologists Bulletin 70 (11), 1729±1741. Barkham, S.T., 1986. Preliminary report on ®eld work in central West Timor, October±December 1985. London University Consortium for Geological Research in Southeast Asia, Report 43 (unpublished). Barkham, S.T., 1993. The Structure and Stratigraphy of the Permo-Triassic Carbonate Formations of West Timor, Indonesia. Unpublished PhD Thesis, University of London. Bassler, R.S., 1929. The Permian Bryozoa of Timor. PalaÈontologie von Timor 16 (28), 37±90. Bassler, R.S., Moodey, M.W., 1943. Bibliographic and faunal index of Paleozoic pelmatozoan echinoderms. Geological Society of America Special Paper 45, 734. van Bemmelen, R.W., 1949. The Geology of Indonesia. Government Printing Of®ce, The Hague. Berry, R.F., Grady, A.E., 1981. Deformation and metamorphism of the Aileu Formation, north coast, East Timor, and its tectonic signi®cance. Journal of Structural Geology 3, 143±167. Berry, R.F., Jenner, G.A., 1982. Basalt geochemistry as a test of tectonic models of Timor. Journal of the Geological Society of London 139, 593±604. Berry, R.F., Burrett, C., Banks, M., 1984. New Triassic faunas from East Timor and their tectonic signi®cance. Geologica et Palaeontologica 18, S127±S137. Beyrich, E., 1862. Untitled report commencing: Uber Gebirgesarten und Versteinerungen, welche von dem Artze Dr Schneider in der Gegend von Koepang auf der Insel Timor gesammelt wurden. Zeitschrift der Deutschen Geologische Gesellschaft 14, 537. Beyrich, E., 1865. Uber eine Kohlenkalk-Fauna von Timor. Abhandlungen der Koniglichen Akademie der Wissenschaften zu Berlin 1864, 59±98.

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