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Permian Brachiopoda from near KasliuVillage, West TimorN.W. Archbold a & P.R. Bird b ca Department of Geology , University of Melbourne , Parkville,Victoria, 3052, Australiab Department of Geology, Royal Holloway and Bedford NewCollege , University of London, Egham Hill , Egham, Surrey,TW20 0EX, United Kingdomc Scott, Pickford & Associates Ltd , 256 High Street, Croydon,Surrey, CRO INF, United KingdomPublished online: 27 Nov 2008.
To cite this article: N.W. Archbold & P.R. Bird (1989) Permian Brachiopoda from near KasliuVillage, West Timor, Alcheringa: An Australasian Journal of Palaeontology, 13:2, 103-123, DOI:10.1080/03115518908619045
To link to this article: http://dx.doi.org/10.1080/03115518908619045
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Permian Brachiopoda from near Kasliu Village, West Timor N. W. ARCHBOLD AND P. R. BIRD
ARCHBOLD, N. W., & BIRD, P. R., 1989:03:27. Permian Brachiopoda from near Kasliu Village, West Timor. Alcheringa 13, 103-123. ISSN 0311-5518. An assemblage of Permian Brachiopoda from the Maubisse Formation outcrops near
the village of Kasliu, West Timor, is described and figured. New taxa are Spiriferella orientensis sp. nov. and Eliva timorensis sp. nov. The assemblage is considered to be a correlative of the classic Basleo and Amarassi faunas of Timor. The assemblage was collected from outcrop and hence provides reliable data on the faunal association.
N. W. Arehbold, Department of Geology, University of Melbourne, Parkville, Victoria, 3052, Australia; P. R. Bird, Department of Geology, Royal Holloway and Bedford New College, University of London, Egham Hill, Egham, Surrey TW20 OEX, United Kingdom; present address: Scott, Pick ford & Associates Ltd, 256 High Street, Croydon, Surrey CRO 1NF, United Kingdom; received 29 September 1987.
Keywords: Brachiopoda, Permian, Timor, new taxa.
TIMOR became a classic South East Asian locality for Permian Brachiopoda through the systematic investigations of collections by Beyrich (1862, 1865), Martin (1881), Rothpletz (1891, 1892) and Broili (1916). Comparison of the Permian faunas of Timor with those from Western Australia, the Himalayas, southeast Asia and Irian Jaya is providing critical data for understanding the tectonic relationships of these regions during Permian times. In this paper we document new collections of Permian Brachiopoda from outcrops of the Maubisse Formation near the village of Kasliu, West Timor (Fig. 1). The collections were made by one of us (P.R.B.) during the course of mapping in the Kekneno Area of West Timor as part of an investiga- tion of the geology of Timor, organised jointly by the London University Consortium for Geological Research in Southeast Asia and the Geological Research and Development Centre, Bandung, Indonesia. In this paper P.R.B. is responsible for the details of the field geology (see Bird, 1987, for full details) and N.W.A. is responsible for the systematic descriptions, new taxa and the age determination.
0311/5518/89/010103-21 $3.00 © AAP
Considerable discussion and various tectonic models have sought to explain the high structural complexity of the geology of Timor. Major thrusting, melange develop- ment and faulting and deformation of the continental margin of Australia during collision with the Banda Arc have all been invoked to account for the geology (e.g. see Barber e t al . , 1977; Hamilton, 1979, ChamaIaun & Grady, 1978). The results of recent fieldwork in West Timor have emphasised the roles of thrust faulting (Rosidi et al. , 1979), wrench faulting and diapirism (Barber et al. , 1986). Age control by palaeontological dating is critical for con- straining structural and tectonic models, as demonstrated by Berry et al. (1984) who showed that a supposed stratigraphical inversion in the Manatuto region of East Timor does not exist, and on these grounds rejected the overthrust model (e.g. Barber et al. , 1977).
Many classical Permian brachiopod localities of Timor, e.g. Basleo and Bitauni, are not areas of outcrop, but are localities with boulders of Maubisse Formation within the Bobonaro Scaly Clay or in river beds. The best preserved fossils are often found in the top soil. Hence, the discovery of faunas from
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104 N. W. A R C H B O L D & P. R. BIRD A L C H E R I N G A
N • / E. Timor
W. Timor
5O I
100 km I
, • Wrench fault
f Thrust
J River
0 K Kasllu
tr- "-! Area of L ._1 Figure 2
Australian Margin
~ - ] Permo-Tr iass ic sediments
Overth rust Complex
~ ) Neogene L imestone
~ - ~ Permian Lst. + Volc.
Metamorphics
Ultrabasics
[ 1 Fig. 1. Location map of the Kekneno area in West Timor, showing the regional geology of Kekneno and the location of Fig. 2. Geological data from Rosidi et al. (1979) and Bird (1987).
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A L C H E R I N G A TIMORESE P E R M I A N B R A C H I O P O D S 105
distinct outcrop, even if in fault blocks, provides a reliable faunal association. Out- crops near Kasliu Village provide one such association.
All specimens are housed in the Museum of Victoria collections (NMVP prefix) and all figured specimens of new species other than holotypes are paratypes.
124 °10 ' 1 2 4 ° 12' 9 ° 3 0 , ~ / , ~ - ' - ,,, i ; -~ . i t S - - , , ~. 7 • - . . ' , . - ' ~ - ~ . s . [ , , " ~ , , < , - , ,', , ' - . . . . . . . , X . . . . . . . . . , , ~ , z , . , , - , , - . , . : ,
. . . . . , . . : ; , . , - . , . , - , , . : , , : , , - . . . . , , . , , . . . . . . . . . . ,,,_.,.., +~,-.-',.,,. i, , . . , - , _ , , - , . . . . . . 0 I I [ [ I ~ I ] . I . i . " P ' ~ ' O I , " X - - ~ I ~ I ~ - - ~ I I ~ ' % ~ . . . . / * ' ~ 1 / ~ 1 ~ Z u I 2 1 " . ~ . I ~ I , " ,
L, L ~'aZ_L~_ :~[ I I 1 1 "~ ' / 2-'," ";Y;;'-,'-%-_,.% " " "V. ' . " ' - ' " , ,~, ' . ' , ' : ' . ' . . '~ ' .
• . . - . - - .. . . - ~-# . , ;~ - ~ /~ ,~
. . . . . . . . . . t ~ . '~ ~ ' , ' ~ " ~ ~,
• ~~~.)~", . . . . . . . . . . . 0,~,~
' ~ ~i'l ~ " " \ " " . " I I~ r~,."~.'...'-'-'.,,'.l ' • • ".k" " . L~.X_._L__ - - J - - - - . L ~ ( , - ' , , , ~ . " , ' ; ~ ' , ( , , ~ 9 0 3 4
0 1 2 3 k m i., i . l i
Neogene Limestone ~ Metamorphic rocks
F ~ Permian Limestone ~ Ultrabasic rocks
& Volcanics • Fossil locality Permo-Triassic Clastics
Fig. 2. Map of the Kasliu area showing local geology and brachiopod sites,
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106 N. W. ARCHBOLD & P. R. BIRD ALCHERINGA
Stratigraphy The Maubisse Formation, originally defined in East Timor (Audley-Charles, 1968), is a series of Permian and Triassic carbonates and volcanics which contain rich subtropical faunas, although not true reef faunas as originally defined by Audley-Charles (cf. Hamilton, 1979, p. 123). Outcrops in the Kekneno area (Fig. 1) consist mainly of a dist inctive sequence of volcaniclast ic- bioclastic sands interbedded with pillow lavas and marls. This sequence, informally named herein the Kasliu Member of the Maubisse Formation, was first reported by Wanner (1913) and de Roever (1940). The Kasliu Member outcrops in the upper reaches of the Noil Witaeik, close to the village of Nuapin, and a poorly exposed sequence of pillow lavas and limestones crops out along the track from Kasliu village down to Noil Besasi. Thickness of this sequence is at least 200 metres. In the Kekneno area, outcrops of the Maubisse Formation are confined to the east of the Noil Tunsip wrench fault zone (Fig. 2). West of this fault zone, sediments of deep marine shelf to bathyal facies crop out over an area of more than 200 km 2. The sediments range from lowermost Permian to Upper Triassic. It is of note that the Maubisse Formation is not in stratigraphic contact with these sediments but rather is separated from them by a major structural feature - - the Noil Tunsip wrench fault.
Five main facies of the Maubisse Formation are recognised in outcrop as follows: Crinoidal dolomites, red marls and shales, bioclastic packstones and wackestones, volcaniclastic sands and pillow lavas. The brachiopods described herein from locality 337 were collected from outcrops of the volcaniclastic sands on the path from Kasliu to Noil Besasi. Those from locality 45 were collected from an isolated outcrop and boulders of the same facies as 337, 1 km SSE of Nuapin. The sands have the texture of grainstones and the volcanic clasts are well rounded with high sphericity. The rock consists of weathered iron rich basalt clasts and abundant bioclastic debris including brachiopod shells. A high energy environment is indicated but the significant number of articulated brachiopod specimens indicates
little evidence of transportation and hence probably rapid burial.
Age Comparison of the assemblage described herein with the classic brachiopod faunas of Timor indicates a close relationship with faunas described from Basleo and correlative localities by Broili (1916) and Hamlet (1928). Specimens described herein and referred to species such as Waagenoconcha waageni (Ro thp le tz ) , Transennatia tirnorensis (Hamlet), Neospirifer sp. nov. and Stenos- cisma timorense Hayasaka & Gan indicate substantial links with Basleo assemblages. Only Eliva timorensis sp. nov. represents a new generic record for these Basleo faunas. Grant (1976) noted that the Basleo fauna is a distinct reflection of the brachiopod fauna of the Kalabagh Member of the Wargal Limestone of the Salt Range, Pakistan, a view also expressed by Waterhouse (1976). The Kalabagh fauna is, in turn, close to that from the overlying Chhidru Formation. Both the Kalabagh and Chhidru faunas include the ammonoid Cyclolobus which is usually regarded as being younger than Kazanian and older than Dzhulf ian (Dickins, 1976; Waterhouse, 1976) although Grant (1970, 1976) favoured a Kazanian age. Cyclolobus occurs with the Amarassi fauna of Timor (Rothpletz, 1892) and has been reported from boulders of the Maubisse Formation in the Noil Tunsip Valley (probably derived from the Kasliu Area) by Dr H. G. Owen (personal communication to P.R.B., 1984).
The present assemblage is therefore con- sidered to be correlative with the Basleo and Amarassi faunas and hence is referred to the Chhidruan Stage above the Kazanian. The name Chhidruan is preferred to Punjabian (Stepanov, 1973) for reasons mentioned in Archbold (1982), although use of the name has difficulties because Chhidruan includes a fauna not from the Chhidru Formation (Waterhouse, 1976).
Systematic palaeontology Order P R O D U C T I D A Sarycheva & Sokolskaya 1959
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ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 107
S u b o r d e r P R O D U C T I D I N A W a a g e n 1883
Supe r f ami ly P R O D U C T A C E A Gray 1840
F a m i l y C H O N E T E L L I D A E Likharev 1960
Chone te l l a W a a g e n 1884
Type species. Chonetella 1884.
nasuta W a a g e n
Chone te l l a sp. (Fig. 3A-B).
cf. 1892 Chonetella nasuta Waagen; Rothpletz, p. 77, pl. 10, figs 12, 12a, 16.
cf. 1928 Chonetella nasuta Waagen; Hamlet, p. 13, pl. 3, figs 5-8.
C o m m e n t s . A s i n g l e v e n t r a l v a l v e , NMVP60737, indica tes the presence o f this genus in the Kasl iu assemblage . The valve is s t rongly convex (width at ears 10.8 mm, es t imated length 7.8 ram) with dis t inct ears and s t rong radia l costae . No spine bases are present and the faint ret iculat ion o f the ventral o r n a m e n t o f Bibatiola G r a n t (1976, pl. 34, figs 24 and 33) is also absent . No ind ica t ion o f a no tched t rai l or V-shaped an te r io r extension is present but the valve an te r ior appea r s to be b roken . One of the specimens f rom Aje r Mat i , T imor , f igured by Rothple tz (1892, pl. 10, fig. 16) appea r s to vi r tual ly lack the no tched trai l as does one o f the Wesleoe
qlp B
H
¢
Fig. 3. A, B, Chonetella sp. NMVP60737, ventral valve in ventral view, x 2.5 and x 1. Waagenoconcha waageni (Rothpletz). C, D, NMVP60738, ventral valve in posteroventral and ventral views, x 1.2. E-l, Transennatia timorensis (Hamlet). E-G, NMVP60740, shell in dorsal, posteroventral and ventral views, x 1. H, I, NMVP60739, ventral valve in ventral and lateral views, x 1. J, K, Asperlinus sp. NMVPI20211, decorticated ventral valve in ventral view, x 1.2 and x 2.5.
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108 N. W. ARCHBOLD & P. R. BIRD ALCHERINGA
specimens of Hamlet (1928, pl. 3, fig. 7), therefore the development of the trail may be variable in the Timor species. The V-shaped trail appears to be strongly developed in Chonetella nasuta (see Waagen, 1884, pl. 81, figs 3-8; Muir-Wood, 1965, p. H460, fig. 323, la and lc and Grant, 1976, pl. 42, figs 1-17). Likharev's (1937, p. 88, pl. 13, figs 1-4) specimens, f rom the Nikitina Ravine, North Caucasus, virtually lack the V-shaped trail whereas his Chonetella triangularis f rom the same locality (1937, pl. 13, figs 5-7) is more triangular in outline and possesses a rounded, extended trail.
Locality. 337.
Family W A A G E N O C O N C H I D A E Muir- Wood & Cooper 1960
Waagenoeoneha Chao 1927
Type species. Productus humboldti d'Orbigny 1842
Waagenoconcha 3C-D)
1892
1916
cf. 1928
cf. 1928
1928
1978
waageni (Rothpletz) (Fig.
Productus waageni Rothpletz, p. 77, pl. 10, figs 19, 19a, 19b. Productus waageni Rothpletz; Broili, p. 14, pl. 4, figs 1-5. Productus humboldti Hamlet, p. 21, pl. 4, fig. 3 Productus humboldti var. irginae Hamlet, p. 23. Productuspurdoni Hamlet, p. 23, pl. 4, fig. 1; pl. 5, fig. 1. "Productus" waageni Rothpletz, Waterhouse, pl. 2, figs 2, 3.
C o m m e n t s . A single ven t ra l va lve , NMVP60738, (estimated width 53.5 mm, e s t ima ted length 50.5 m), s o m e w h a t decorticated anteriorly, indicates the presence of Waagenoconcha in the Kasliu assemblage. Short, sharp spine bases posteriorly and longer fine spine bases anteriorly compare well with those of shells attributed to W. waageni (see Waterhouse, 1978, pl. 2, fig. 2) from Basleo. Although decorticated in part, some strongly developed growth bands are present. As indicated by the above synonymy, probably only one species of Waagenoconcha is present in the Basleo assemblages of Timor and it is characterised by a distinct sulcus and distinct growth bands.
Broili 's (1916) specimens of W. waageni have been compared closely with W. imperfecta Prendergast (1943) f rom the correlative Hardman Formation, Canning Basin, Western Australia, by Prendergast (1943), Coleman (1957) and Muir-Wood & Cooper (1960). The two species are close in respect of shell outline, sulcus and prominent growth bands but spine bases are finer on the Western Australian species.
Locality. 337.
Family D I C T Y O C L O S T I D A E Stehli 1954
Transennatia Waterhouse 1975 = Gratiosina Grant 1976 = Asioproductus Chan (in Yang et al., 1977)
Type species. Productus gratiosus Waagen 1884
Transennatia timorensis (Hamlet). (Fig. 3E-I)
cf. 1865 Productussemireticulatus Beyrich, p. 82, pl. 2, figs 1, 2.
cf. 1892 Productus gratiosus Rothpletz, p. 76, pl. 10, figs 15, 15a-c.
1916 Productus gratiosus Rothpletz; Broili, p. 12, pl. 2, figs 4, 5, 7, 8, 11; non pl. 2, figs 12, 13 and ?, figs 9, 10.
1928 Productus gratiosus var. timorensis Hamlet, p. 20, pl. 2, figs 2-4.
1970 Spyridiophora? timorensis (Hamlet); Termier & Termier, p. 58, pl. 5, fig. 1, text figs 1-2.
Lectotype. Specimen figured by Hamlet (1928, pl. 2, fig. 3), selected by Waterhouse (1978, p. 119).
Comments. The Kasliu assemblage has yielded two specimens with the character is t ic ornament of Transennatia. The smaller of the two (NMVP60739; estimated width 24 mm, estimated length 18.5 mm) is an incomplete ventral valve with a distinct sulcus, reticulate ornament posteriorly and radial costae anteriorly which converge in the sulcus. It is of average size for the genus.
The la rger spec imen is a pa r t i a l ly decorticated shell of large size for the genus (NMVP60740; maximum width 30.7 ram, length 29.8 ram, width at hinge 25.7 mm, dorsal length 19.1 ram). Well defined convex ears are present as are a distinct ventral sulcus and low dorsal fold. Reticulate ornament is
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ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 109
developed posteriorly and radial costae anteriorly. Ventral costae are finer than those of Transennatia gratiosa (Waagen) as figured by Grant (1976, pl. 33, figs 19-26). Submature growth stages on NMVP60740 are typical of shell outlines of smaller Transennatia specimens and hence it seems likely that only
one species is present in the Basleo and correlative faunas of Timor, as indicated by the synonymy above. Hamle t ' s name t imorensis is avai lable , as no ted by W a t e r h o u s e (1978, p. 119) and the coincidental use of timorensis by Termier & Termier (1970) reinforces the name.
C IZ
!
Fig. 4. ?Reticulatia sp. A, B, NMVP60741, ventral valve in ventral and posteroventral views, x 1. C, NMVP60742, decorticated ventral valve in ventral view, x 1. D, NMVP60743, incomplete ventral valve in ventral view, x 1. E, NMVP60744, incomplete ventral valve in ventral view, x 1. F, NMVP60745, incomplete ventral valve in ventral view, x 1. G, NMVP60746, incomplete ventral valve in ventral view, x 1. H, NMVPI20210, ventral valve in ventral view, × 1.
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110 N. W. ARCHBOLD & P. R. BIRD ALCHERINGA
Large col lect ions are requi red to con f i rm the on togene t ic range o f the species and H a m l e t ' s syn typic series consists o f smal l specimens, but the i l lust ra t ions o f H a m l e t ' s Productus in f la tus (see Hamle t , 1928, pl. 2, figs 7, 8) a p p e a r to indicate tha t large Transennatia is present in Basleo corre la t ive locali t ies.
Locality. 45.
Reticulatia M u i r - W o o d & C o o p e r 1960
Type species. Productus heucoensis King 1931
?Reticulatia sp. (Fig. 4A-H)
cf. 1916 Productusspiralis Broili, p. 11, pl. 3, figs 3-5, non cet.
cf. 1928 Productuschitichunensis Hamlet, p. 17, pl. 3, figs 1, 2.
1940 Productussemireticulatus Martin; de Roever, p. 130.
Comments. The Kasl iu assemblage includes seven d ic tyoc los t id specimens (NMVP60741- 60746, 120210) the generic ident i ty o f which is not cer ta in because detai ls o f the ears , an te r ior cos tae and the dorsa l valve are lacking. The p ronounced ret iculate o r n a m e n t on the visceral disc and the relatively fine even costae an te r io r ly suggest Reticulatia ra ther than Costiferina M u i r - W o o d & Cooper (1960) .a l though the la t te r cannot be ruled out . The closely related Callytharrella A r c h b o l d (1985) is charac te r i sed by the presence o f p r o m i n e n t costa te ears and a lack o f over lapp ing growth lamel lae on the an te r io r o f the ventra l valve, features which canno t be de te rmined on the present mater ia l . Compar i sons with relatively finely cos ta te d ic tyoclos t ids f rom Basleo and corre la t ive local i t ies are indica ted in the synonymy but whether d i f ferent species are present can only be determined by ontogenetic studies based on large col lect ions.
Localities. 45 and 337.
Fami ly L I N O P R O D U C T I D A E Steh| i 1954
S u b f a m i l y L I N O P R O D U C T I N A E Stehli 1954
Asperlinus W a t e r h o u s e & Piyas in 1970
Type species. Productus asperulus W a a g e n 1884
Asperlinus sp. (Fig. 3J-K)
cf. 1892 Productus asperulus Rothpletz, p. 76, pl. 10, fig. 14.
cf. 1928 productus asperulus Rothpletz; Hamlet, p. 27, pl. 5, figs 2, 4, 5.
Comments. A single, small ventra l valve in ternal m o u l d with minor a moun t s o f adheren t shell indicates the presence o f Asperlinus. Cos tae are dist inct , rad ia l , sharp and wavy and increase by branching . Traces o f re t icula te o rnamen t occur pos te r io r ly . Dimens ions o f NMVP120211 are as fol lows: m a x i m u m width 9.5 mm, es t imated hinge width 7.8 mm, valve length 7.6 mm. Mater ia l is insuff ic ient for de te rmining the charac te r o f the T i m o r species.
Locality. 45.
Order R H Y N C H O N E L L I D A Kuhn 1949
S u p e r f a m i l y S T E N O S C I S M A T A C E A Oehler t 1887
Family S T E N O S C I S M A T I D A E Oehlert 1887
Subfami ly S T E N O S C I S M A T I N A E Oehler t 1887
Stenoscisma C o n r a d 1839
Type species. Terebratula schlottheimii yon Buch 1835
Stenoscisma timorense ( H a y a s a k a & Gan) (Fig. 5A-Z , AA-BB)
1865 Camarophoria crumena Beyrich, p. 73, pl. 1, figs 11, 12.
Fig. 5. Stenoscisma timorense (Hayasaka & Gan). A, B, NMVPI20212, shell in dorsal and ventral view, x 1.2. C-F, NMVP120213, shell in posterior, ventral, anterior and dorsal views, x 1.2. G, H, NMVPI20214, juvenile shell in dorsal and ventral views, x 1.3. I, NMVP120215, juvenile ventral valve in ventral view, x 1.3. J, NMVP120216, ventral valve in ventral view, × 1.2, note weakly developed costae. K-M, NMVP120217, shell in dorsal, posterodorsal and ventral views, × 1. N, NMVP120218, shell in ventral view, x 1. O, P, NMVPI20219, shell in ventral and dorsal views, x 1.2. Q-S, NMVP120220, shell in dorsal, ventral and anterior views, x 1.2. T-V, NMVP120221, shell in dorsal, ventral and anterior views, x 1, x 1.2 and x 1.2. W-Y, NMVPI20222, shell in dorsal, ventral and anterior views, x 1.2, x 1 and × 1. Z, AA, BB, NMVP120223, shell in ventral, dorsal and posterior views, × 1.
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ALCHERINGA T I M O R E S E P E R M I A N B R A C H I O P O D S 111
A
K
I..
E
M
U
D
F
R
U V
S
T
Y
W A
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112 N.W. ARCHBOLD & P. R. BIRD ALCHERINGA
1892
1916
1928
1940
1940
1965
cf. 1963
cf. 1976
cf. 1978
cf. 1979
cf. 1982
cf. 1985
Camarophoria pinguis Rothpletz, p. 48, pl. 10, figs 3, 7, 9. Camarophoriapurdoni Broili, p. 55, pl. 11, figs 7-18. Camarophoria purdoni Broili; Hamlet, p. 57, pl. 9, figs 5-8. Camarophoria timorensis Hayasaka & Gan, p. 129, pl. 9(3), figs 1-7. Camarophoria purdoni Broili; de Roever, p. 130. Stenoscisma purdoni (Broili); Grant, p. 149, pl. 20, figs 1-4. Camarophoriapurdoni Broili; Ustritskiy, p. 21, pL 8, figs 1-6. Stenoscisma tirnorense Chang & Ching, p. 193, pl. 11, figs 20-27. Stenoscisma timorense Likharev & Kotlyar, p. 17, fig. 8. Stenoscisma timorense Kochirkevich, p. 52, pl. 12, figs 1-6. Stenoscisma timorensis Zhan & Wu, pl. 4, figs 17-18. Stenoscisma cf. timorense Liu & Waterhouse, p. 22, pl. 4, fig. 13-14, pl. 5, figs 1-4.
Lectotype. Kochirkevich (1979, p. 52) made the following designation: 'example figured by Broili (1916) on plate 11, fig. 10 and Hayasaka & Gan (1940) on plate 3, fig. 2'. Unfortunately, the Broili specimen was not refigured on Hayasaka & Gan's pl. 9(3), fig. 2 which is, in fact, a less typical specimen of the species. Rather, Broili's figure was repeated on page 128 of Hayasaka & Gan's text as their text fig. 2. In the interests of nomenclatural stability, Broili's specimen (Broili, pl. 11, fig. 10) is confirmed as the lectotype herein.
Comments. S. timorense is by far the most abundant species in the Kasliu assemblage with some 41 specimens present of which 12 (NMVP120212-120223) are illustrated herein. As indicated by Hayasaka & Gan (1940) the species is highly variable in terms of the development of costae. They recorded variations from 1 to 6 costae in the sulcus; the present population varies from 3 to 6. Present material affords no new information on the species so no further description is provided here.
As indicated by the comparisons listed in the synonymy, the species, or specimens comparable to it, has been recorded from the Himalayas, Tibet, northwest China, Inner Mongolia and the Soviet Far East. Whether
or not these reports represent true S. timorense may be debated but with such a variable species as S. timorense it is not unexpected that comparable specimens will occur elsewhere. Only by investigating large collections will the variability of these other reports be ascertained and the species relation- ships be confirmed.
Localities. 45 and 337.
Order SPIRIFERIDA Waagen 1883
Suborder SPIRIFERIDINA Waagen 1883
Superfamily SPIRIFERACEA King 1846
Family SPIRIFERIDAE King 1846
Subfamily NEOSPIRIFERINAE Waterhouse 1968
Neospirifer Frederiks 1924
Type species. Spirifer fasciger yon Keyserling 1846
Neospirifer sp. (Fig. 6A)
cf. 1916 SpiriferfascigerBroili, p. 34, pl. 7, figs 1, 2, 3. ? 1940 Spiriferfasciger de Roever, p. 130.
Comments. A single large, incomplete ventral valve (NMVP120224; estimated maximum width 128 mm, valve length 58 mm +, ventral interarea height 11 mm) indicates the presence of a new distinctive species of Neospirifer comparable to that recorded from Basleo and correlative localities by Broili (1916, pl. 7, figs 1-3). The large size of the valve, gentle plicae, fine costae and wide sulcus recall species such as N. grandis Archbold & Thomas (1986) and related species as discussed in that study. Unlike those species, however, the Timor species is characterised by a relatively low ventral interarea, particularly considering the size of the shells involved. The small internal mould figured by Broili (1916, pl. 6, fig. 11) may be a juvenile of the species. Other Neospirifer specimens from Bitauni (Broili, 1916, pl. 6, figs 12-14) are a distinct species as is N. timorensis (Martin, 1881) as discussed by Archbold & Thomas (1986, p. 136). The lectotype of N. timorensis (Martin) is figured herein.
Locality. 337.
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A L C H E R I N G A TIMORESE P E R M I A N B R A C H I O P O D S 113
Subfamily SPIRIFERELLINAE Waterhouse 1968
Spiriferella Chernyshev 1902
Type species. Spirifer saranae de Verneuil 1845
Spiriferella orientensis sp. nov. (Figs 7A-N, 8A-H)
1940 Spirifer spec. de Roever, p. 130.
I-Iolotype. NMVP 120225, an incomplete shell with portion of the dorsal valve missing.
I::
Fig. 6. Neospirifer sp. A, NMVPI20224, ventral valve in ventral view, × 1. B-E, Neospirifer timorensis (Martin).B.-E, Rijksmuseum Leiden 12040, shell in dorsal, ventral, posterior and anterior views, × 1.
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114 N . W . ARCHBOLD & P. R. BIRD ALCHERINGA
Material. NMVPI20225-120237, a col lec t ion o f two m a t u r e shells with po r t i on o f dorsa l valves a t t ached and eleven i so la ted ventra l valves ranging f rom juveni le to ma tu re specimens.
Measurements. See Table 1.
Specimen Maximum Hinge Ventral Number width width length
NMVP 120225" 40.9 31.0 54.5 NMVP 120226 52.6 32.0 - - NMVP 120227 45.9 30.0e 60.6 NMVP120228 46.0e 36.0e - - NMVPI20229 38.0e 30.0e - - NMVPI20230 35.8 25.6 37.8 NMVP120231 30.5 20.5 34.0+ NMVP 120232 29.2 21.2 28.0 NMVP 120233 25.4 20.0e 23.4 NMVP 120234 22.0e 15.3 20.2 NMVP120235 19.0 + - - 22.0 NMVP120236 18.0+ 14.5 18.5 NMVP 120237 16.8 11.9 14.8
Table I. Measurements of Spiriferella orientensis sp. nov. in mm. e = estimate. * = holotype
Diagnosis. Large, e longate species with sharp costae on both valves increasing in number by branching . Hinge na r row and sulcus b r o a d l y V-shaped.
Description. Large biconvex species with elongate outl ine. Max imum width at matur i ty an te r io r o f mid length . Hinge na r row with no ears.
Ventral valve s t rongly convex with u m b o strongly over tu rned . Ventral in terarea poor ly known but re la t ively high, na r row and concave. De l thy r ium not known. Vent ra l sulcus b r o a d , mode ra t e ly deep, V-shaped in cross-sect ion and sharply d e m a r c a t e d f rom lateral f lanks. Distinct median costae present . Sulcal bound ing cos tae high, sharp , fo rm dist inct p l ica t ions ; b ranch ing cos tae arise at abou t 2 cm f rom u m b o . Sulcal f lanks car ry one costa in i t ia l ly which branches a b o u t 1 to
1.5 cm f rom u m b o . All sulcal cos tae sharp . La te ra l s lopes o f valve car ry b r o a d , r o u n d e d costae which b ranch between I and 1.5 cm of umbo . Vent ra l valve an te r ior thin, pos te r io r heavily th ickened. Ventral in ter ior unknown. Dorsa l u m b o smal l , sharp . Dorsa l in te ra rea low. Valve thin. Dorsal fas t ig ium na r row, sharp , with p o o r l y def ined , na r row, shal low median groove. Flanks of fastigium each carry one cos ta which branches f rom med ian cos ta (fold) at 0.75 cm app rox ima te ly . La te ra l costae sha rp , f o rm pl ica t ions , b ranch at 1.5 to 2.0 cm f rom umbo . Dorsa l in te r io r unknown .
M i c r o - o r n a m e n t o f radia l capi l lae , fine concentric growth lines and minute, ill def ined pustules on cross-over points .
Discussion. Spiriferella orientensis sp. nov. appears to belong to the S. rajah (Salter, 1865) g roup o f species but possesses dis t inct ive enough features such as a shor t hinge and sharp cos tae for a new species to be es tabl ished. The large Basleo suite a t t r ibu ted to S. rajah by Broili (1916, pl. 4, fig. 19; pl. 5, figs 1-11; pl. 5, figs 1-6) includes n u m e r o u s ma tu re large specimens charac te r i sed by a wide aur icu la te hinge. They m a y well be m a t u r e s p e c i m e n s o f S. in terpl icatus (Rothple tz , 1892) as indicated by W a t e r h o u s e (1966). Cos tae , especial ly those o f the sulcus and dorsa l valve, o f the Basleo suite are not as sharp as those o f the present mate r ia l , (see also W a t e r h o u s e , 1978, pl. 3, figs 14-15). H i m a l a y a n S. rajah, as i l lus t ra ted by G u p t a & W a t e r h o u s e (1979, pl. 1, figs 10-14; pl. 2, figs 1-10) is a h ighly var iable species, at t imes with a shor t hinge and e longate out l ine but with lower rounde d costae and less sharp dorsa l fold. Nepalese S. rajah (Wate rhouse , 1978, pl. 4, figs 1-7; pl. 14, figs 1-13) have more r o u n d e d and lower cos tae and a shal lower sulcus than S. orientensis sp. nov. as do specimens f rom the Chinese H i m a l a y a s (Chang & Ching, 1976, pl. 17, figs 3-12).
Fig. 7. A-N, Spiriferella orientensis sp. nov. A-C, holotype, NMVP120225, shell in dorsal, posteroventral and ventral views, x 1. D-G, NMVP120226, shell in dorsal, ventral, posteroventral and lateral views, x 1. H, NMVPI20232. ventral valve in ventral view, x 1. I, NMVP120231, ventral valve in ventral view, x 1. J, NMVP120233, ventral valve in ventral view, x 1. K, NMVP120237, juvenile ventral valve in ventral view, x 1. L, NMVPI20236, ventral valve in ventral vie'w, x 1. M, NMVPI20234, ventral valve in ventral view, x 1. N, NMVP120235, ventral valve in ventral view, x 1. O, Spiriferella sp. Rijksmuseum Leiden 12041, ventral valve in ventral view, × 1.
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A L C H E R I N G A TIMORESE PERMIAN BRACHIOPODS 115
oLO,
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116 N.W. ARCHBOLD & P. R. BIRD ALCHERINGA
Spiriferella rajah shares some features with Spiriferella keilhavii (von Buch, 1847) as discussed by Waterhouse & Waddington (1982) who referred some Northern Hemis- phere reports of S. rajah to von Buch's species (Waterhouse & Waddington, 1982, p. 28). North Chinese and Inner Mongolian reports of S. rajah (summarised by Lee et al., 1985, p. 123, pl. 2, figs 3, 4, 6, 9, 12) also comprise specimens with rounded costae and hence are distinct from the new species. The magnificent suite of specimens from the Soviet Far East described by Frederiks (1916, p. 80, pl. 5, figs 1-9) and attributed to S. rajah includes elongate shells with relatively high ventral in terareas but costae are finer, more numerous and more rounded than those of S. orientensis sp. nov. Other Soviet Far East specimens attributed to S. rajah and S. grandis Kotlyar (in Likharev & Kotlyar, 1978, pl. 18, figs 6-8) are comparable with the suite described by Frederiks.
S. rajah has been repor ted f rom a "Bi tauni" fauna of east Timor by Shimizu (1966, pl. 3, figs 1-11)and a "Bi tauni" fauna from Letti (Broili, 1915, pl. 21, figs 11, 17-18) but all specimens are small with well rounded fine costae and hence are not close to S. orientensis sp. nov. Broili's (1915, pl. 21, figs 17-18) specimens have such fine costae that comparison with Eliva is possible.
Spiriferella nepalensis Legrand-Blain (1977) from the late Early Permian of Nepal is an elongate species with a relatively narrow hinge which may be ancestral to S. orientensis sp. nov. Dorsal costae of the Nepalese species are relatively low and rounded unlike those of S. orientensis.
The second specimen attributed to Spirifer timorensis by Martin (1881, pl. 2, fig. 8) is, in fact, a representative of Spiriferella with a relatively wide hinge with ears and fine costae. A plaster replica is figured herein (Fig. 70) .
Localities. 45 and 337.
Eliva Frederiks 1924
Type species. Spirifer lyra Kutorga 1844
Eliva timorensis sp. nov. (Fig. 9A-N)
Holotype. N M V P 1 2 0 2 3 8 , a c o m p l e t e decorticated shell.
Material. NMVP120238-120243, a complete shell, an incomplete shell and four ventral valves.
Measurements. See Table 2.
Specimen Maximum Hinge Ventral Dorsal Number width width length length
NMVP120238* 18.0 11.0 23.1 18.9 NMVP 120239 17.0 11.0e 22.5e 19.5 NMVP120240 26.0e -- 27.9 -- NMVP120241 19.8 -- 21.0+ -- NMVP120242 16.8 10.0e 20.0 -- NMVP120243 14.2 -- 15.0 --
Table 2. Measruements of Eliva timorensis sp. nov. in ram. e = estimate. * = holotype.
Diagnosis. Moderately biconvex elongate species with fine costellae, a narrow, relatively deep sulcus and low but clearly demarcated fastigium.
Description. Average to large species, moderately biconvex with elongate outline. Maximum width at maturity anterior of mid- length. Hinge narrow. Ears absent.
Ventral valve moderately convex with umbo sharply pointed. Ventral interarea poorly known but appears to be relatively low, narrow and concave. Delthyrium not known. Ventral sulcus narrow, relatively deep, V- shaped in cross-sect ion and dist inctly demarcated f rom lateral flanks. Thin, sharp, median costa present in sulcus. Ventral costae fine, well rounded. Costae arise at umbo. Sulcal bounding costae branch within 0.5 cm of umbo as do lateral flank costae. Individual initial costae give rise by branching to fascicles of three or five costae. Fascicles weakly expressed because of lack of lateral plications. Ventral valve anterior thin, posterior with minor thickening. Ventral interarea poorly known but dental plates and short adminicula apparently present.
Dorsal umbo small, sharp. Dorsal interarea low. Valve thin. Dorsal fastigium low, broadens anteriorly with low fold at anterior commissure. Fastigium clearly demarcated by shallow depression on each side. Fastigium and lateral flanks covered with fine, low, well rounded branching costae. Costae branch within a few m m of umbo. Dorsal interarea unknown.
Micro-ornament of radial capillae.
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A L C H E R I N G A T I M O R E S E P E R M I A N B R A C H I O P O D S 117
Discussion. The distinctive outline and ornament of E. timorensis sp. nov. leave little doubt as to its generic position. Cooper & Grant (1976, p. 2238) have shown that the genus can be recognised by those features rather than details of the dental plates and adminicula.
E. timorensis sp. nov. differs from E. lyra (Kutorga, 1844, pl. 9, fig. 7; syntypic series
refigured by Chernyshev, 1902, pl. 8, figs 4-5) by its narrower deeper sulcus and its demarcated fastigium. E. lyra is also a more robust species than E. timorensis sp. nov. E. lyra is characteristic of the Asselian of the Urals (Kalashnikov, 1980) and Frederiks (1932) described additional specimens under a variety of subspecific names from the Asselian of the Pechora Basin, north Urals.
it Fig. 8. Spiriferella orientensis sp. nov. A-E, NMVPI20227, ventral valve in posteroventral and ventral views, x 1.2 and portions of surface enlarged, x 8. F, NMVP120230, ventral valve in ventral view, x 1. G, NMVP120229, ventral valve in ventral view, x 1.2. H, NMVP120228, ventral valve in ventral view, x 1.2.
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118 N. W . A R C H B O L D & P. R. B I R D A L C H E R I N G A
E. lyra has also been recorded from the Early Permian of the Tian Shan (Keidel, 1906,
pl. 13, fig. 3), the Early Permian of Laos (Mansuy, 1913, pl. 5, fig. 10a-10g), and the
r ! ) F:
i v
~" i~ I̧̧ : i ! ~ ~
0
i V
I y
Fig. 9. A-N, Eliva timorensis sp. nov. A, K, NMVPI20240, ventral valve in ventral and posterior views, x 1.2 and x 1.5. B-D, NMVP120238, holotype, shell in ventral, lateral and dorsal views, x 1.3. E, NMVP120241, ventral valve in ventral view, x 1.2. F, NMVP120243, ventral valve in ventral view, x 1. G, H, L, NMVPI2039, shell in ventral, lateral and dorsal views, x 1.2. I, J, NMVPI20242, ventral valve in ventral and posteroventral views, x 1.1. M, N, NMVP120210A, incomplete dorsal valve in dorsal view, x 1.3 and portion of surface enlarged, x 8. O-R, Phricodothyris sp. O-R, NMVPI20250, natural cast of shell in dorsal, ventral, posterior and anterior views, x 1.3. S-U, Cleiothyridina spp. S, T, NMVP120245, decorticated shell in ventral and dorsal views, x 1. U, NMVPI20244, decorticated dorsal valve in dorsal view, x 1.2. V-Y, ?Spirigerella sp. V-X, NMVPI20246, shell in ventral, dorsal and lateral views, x 1.2. Y, NMVPI20247, ventral valve in ventral view, x 1.
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A L C H E R I N G A T I M O R E S E P E R M I A N B R A C H I O P O D S 119
Early Permian of the Karateke Mountains (Vadasz, 1911, pl. 1, fig. 13a-b), however these forms possess relatively simple costae with little evidence of fasciculation and hence are not close morphologically to E. timorensis sp. nov. The tiny (juvenile?) E. lyra recorded by Vinassa de Regny & Gortani (1905, pl. 14, fig. 28) from the Carnic Alps possesses an elongate outline and very fine fasciculate costae but its small size precludes,closer comparison with E. timorensis sp. nov.
Texan species described as E. inflata and E. shumardi by Cooper & Grant (1976) have a narrower, deeper sulcus like that of E. timorensis sp. nov. but they tend to be wider species with sharper costae and less clearly demarcated fastigia. Their outlines are also less oval than that of E. timorensis. The Texan species are late Kazanian in age.
E. lyra has been recorded from Basleo and Noil Simaan by Hamlet (1928, p. 38, pl. 6, figs 5, 6). Only specimens from Noil Simaan were illustrated and they have wider sulci and coarser costae than the material at hand and are unlikely to be conspecific. The age of the Noil Simaan locality is not clear; the species occurs with the enigmatic "Spiri fer" simaanensis Hamlet. Broili (1915, pl. 21, figs 17, 18) figured two specimens from Letti that have ventral sulci and costae close to those of E. timorensis. He referred them to Spirifer rajah but an assignment to Eliva appears more probable. The Letti faunas are generally con- sidered to be of "Bi tauni" type (Waterhouse, 1976) but more than one horizon may be present.
Locality. 45.
Superfamily RETICULARIACEA Waagen 1883
Family ELYTHIDAE Frederiks 1924
Subfamily ELYTHINAE Frederiks 1924
Phricodothyris George 1932
Phricodothyris sp. (Fig. 90-R)
? 1940 Squarnularia lineata: de Roever, p. 130.
Comments. A single natural cast of a small shell (NMVP120250) with preserved con-
centric lamellae and indistinct spine bases demonstrates the presence of the Elythidae in the assemblage. The specimen (maximum width 22.3 mm, ventral length 20.9 mm, dorsal length 18.4 mm), wider than long, is feebly uniplicate and so recalls smaller elythids from Basleo, figured by Broili (1916, pl. 7, fig. 7; pl. 8, fig. 12) that are also wider than long. Other elythids figured by Broili (1916) from Basleo are more elongate and hence may
i i
Fig. 10. Dielasmatids indet. A-C, NMVP120248, decorticated shell in dorsal, ventral and lateral views, x 1. D-F, NMVP 120249, decorticated shell in ventral, dorsal and lateral views, x 1.
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120 N. W. ARCHBOLD & P. R. BIRD ALCHERINGA
represent a separate genus. The present specimen is distinct from Early Permian elythids from Western Australia (Archbold & Thomas , 1984) referred to Spirelytha Frederiks (1924).
Locality. Close to locality 337 but strati- graphic relationship to 337 unknown.
Order ATHYRIDIDA Dagys 1974
Subo rde r A T H Y R I D I D I N A B o u c o t , Johnson & Staton 1964
Superfamily ATHYRIDACEA Mccoy 1844
Family ATHYRIDIDAE McCoy 1844
Subfamily ATHYRIDINAE McCoy 1844
Cleiothyridina Buckman 1906
Type species. Atrypa pectinifera Sowerby 1840.
Cleiothyridina spp. (Fig. 9S-U)
Comments. Two specimens in the Kasliu assemblage indicate the presence of two species of Cleiothyridina or its allies. One specimen (NMVP 120244) is an isolated dorsal valve (maximum width 28.5 ram, valve length 22.5 mm) showing no trace of a dorsal fold and hence is probably a true representative of Cleiothyridina. It recalls submature growth stages of several specimens illustrated by Broili (1916, pl. 10, figs 8, 9, 12). The second specimen (NMVP120245) an incomplete, decorticated shell (maximum width estimated 34 ram, ventral length 27.1 mm, dorsal length 26.4 mm) has a distinct ventral sulcus developed anteriorly and a low broad dorsal fold anteriorly. It probably represents a submature individual of the forms with pronounced sulci figured from Basleo by Broili (1916, pl. 10, figs 10, 11, 13) and which may be referable to Pinegathyris Grunt (1980). Cleiothyridina penta Prendergast (1935, pl. 2, figs 13-15) is a related form from the correlative Hardman Formation of Western Australia.
Locality. 45.
Family SPIRIGERELLIDAE Grunt 1980
Spirigerella Waagen 1883 Type species. Spirigerella derbyi Waagen 1883
?Spirigerella sp. (Fig. 9V-Y)
Comments. Two small specimens, one decorticated shell (NMVPI20246) and one ventral valve (NMVPI20247), probably represent juveniles of Spirigerella or an allied genus such as Composita. The specimens are elongate and possess a low dorsal fold and shallow ventral sulcus anteriorly. The ventral sulcus arises some 0.5 cm from the ventral umbo and is initially little more than a median flattening. The specimens may be juveniles of Basleo Spirigera timorensis Rothpletz as figured by Broili (1916, pl. 9, figs 7-14; pl. 10, figs 1-2) but are inadequate for precise deter- mination. Measurements of the two specimens are as follows: NMVP120247, ventral maximum width 11.8 mm, ventral length 12.8 mm; NMVPI20246, maximum width 13.3 mm, ventral length 15.3 mm, dorsal length 13.8 mm.
Locality. 45.
Order TEREBRATULIDA Waagen 1883
Suborder TEREBRATULIDINA Waagen 1883
Superfamily TEREBRATULACEA Waagen 1883
Family DIELASMATIDAE Schuchert 1913
Dielasmatids indet. (Fig. 10A-F)
Comments. Two large terebratulid shells (NMVPI20248-120249) tha t are bo th decorticated and damaged indicate the presence of large terebratulids in the assemblage. Only Hamlet (1928, pl. 10, fig. 9) has ever figured a large terebratulid from Timor and her specimen is more elongate with a higher dorsal fold than the present specimens. The material is insufficient for generic determination but gross shell form may indicate a non-plicate species of Gilledia or Fletcherithyris.
Localities. 45 and 337.
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A L C H E R I N G A T I M O R E S E P E R M I A N B R A C H I O P O D S 121
Acknowledgements P . R . B . is g r a t e fu l to I c san U m a r , K u s n a m a Bra ta and T o m H a s s a n o f G . R . D . C . B a n d u n g a n d T o n y Ba rbe r , T i m C h a r l t o n a n d Sa rah C o o k (Roya l H o l l o w a y a n d B e d f o r d N e w Co l l ege , U n i v e r s i t y o f L o n d o n ) fo r the i r ass i s tance d u r i n g f i e l d w o r k . N . W . A . t h a n k s Dr C. F. W i n k l e r P r i n s ( R i j k s m u s e u m v a n G e o l o g i e en M i n e r a l o g i e , L e i d e n ) f o r i n f o r m a t i o n a n d p las t e r cas ts o f M a r t i n ' s (1881) T i m o r m a t e r i a l . N . W . A . ' s w o r k is s u p p o r t e d by the A u s t r a l i a n Resea rch G r a n t s S c h e m e . Isabel M u n r o t y p e d the m a n u s c r i p t .
R E F E R E N C E S ARCHBOLD, N. W., 1982. Classification and evolution of
the brachiopod family Rugosochonetidae Muir- Wood 1962. Proceedings o f the Royal Society o f Victoria 94, 1-9.
ARCHBOLD, N. W., 1985. Studies on Western Australian Permian brachiopods. 5. The family Dictyoclostidae Stehli 1954. Proceedings o f the Royal Society o f Victoria 97, 19-30.
ARCHBOLD, N. W., & T~OMAS, G. A., 1984. Permian Elythidae (Brachiopoda) from Western Australia. AIcheringa 8, 311-326.
ARCHBOLD, N. W., & THOMAS, G. A., 1986. Neospirifer and Trigonotreta (Spiriferida, Brachiopoda) from the Permian of Western Australia. Alcheringa 10, 125-161.
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