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Alcheringa: An Australasian Journal of
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Permian Brachiopoda from near
Bisnain Village, West Timor
N.W. Archbold
a
& S.T. Barkham
b
a
Department of Geology , University of Melbourne , Parkville,
Victoria, 3052, Australia
b
Department of Geology, Royal Holloway and Bedford New
College , University of London, Egham , Hill, Egham, Surrey,
TW20 0EX, United Kingdom
Published online: 27 Nov 2008.
To cite this article: N.W. Archbold & S.T. Barkham (1989) Permian Brachiopoda from near Bisnain
Village, West Timor, Alcheringa: An Australasian Journal of Palaeontology, 13:2, 125-140, DOI:
10.1080/03115518908619046
To link to this article: http://dx.doi.org/10.1080/03115518908619046
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2. Permian Brachiopoda from near Bisnain Village,
West Timor
N. W. ARCHBOLD AND S. T. BARKHAM
ARCHBOLD,N. W., &BARKHAM,
S. T., 1989:03:27.PermianBrachiopodafromnear Bisnain
Village, West Timor. Alcheringa 13, 125-140. ISSN 0311-5518.
Collectionsof Permian Brachiopoda from the MaubisseFormation outcropsnear the
villageof Bisnain~WestTimor, are describedand figured.Newtaxa are Callytharrella khalii
sp. nov. and Elivina bisnaini sp. nov. The assemblageis consideredto be a correlativeof
the fauna of the Callytharra Formation of Western Australia and a Sterlitamakian (late
Sakmarian) age is indicated. The collectionscomefrom outcrop and henceprovidereliable
data on the faunal associations.
N. W. Archbold, Department of Geology, University of Melbourne, Parkville, Victoria,
3052, Australia; S. T. Barkham, Department of Geology, Royal Holloway and Bedford
New College, University of London, Egham Hill, Egham, Surrey, TW20 OEX, United
Kingdom; received 29 September 1987.
Keywords: Brachiopoda, Permian, Timor, new taxa, palaeoclimate.
PERMIAN brachiopods were first reported
from Timor in 1862 (Beyrich, 1862). Since
that report, collections have been systema-
tically described by Beyrich (1865), Martin
(1881), Rothpletz (1892), Broili (1916),
Hamlet (1928), Wanner & Sieverts (1935) and
Shimizu (1966). A few other species have been
described in other investigations but no new
collections have been described since
Shimizu's paper. The sequence of brachiopod
faunas from Timor has been reconstructed by
the study of the faunas themselves and
comparison with faunas in stratigraphic
sequence elsewhere (e.g. see Waterhouse &
Piyasin, 1970; Grant, 1976). Up until now,
two broad Permian brachiopod assemblages
have been recognised in Timor. These are the
late Early Permian Bitauni assemblage (the
age of which has been discussed by
Waterhouse, 1970, 1973, 1976, 1981, and
Grant, 1976) and a Late Permian Basleo-
Amarassi assemblage (age discussed by Grant,
1976, and Waterhouse, 1973, 1976). The new
assemblage described below and named by us
the Bisnain assemblage appears to be older
than either of the assemblages indicated
0311/5518/89/010125-16 $3.00 © AAP
above. The new assemblage is apparently
Sterlitamakian (late Sakmarian) in age by
direct comparison with elements of the
brachiopod fauna of the Sterlitamakian
Callytharra Formation of the Carnarvon
Basin, Western Australia (see Archbold, 1982,
for a summary of the age control of that
formation).
The new collections documented in this
study were made from outcrops of limestone,
attributed to the Maubisse Formation
(Audley-Charles, 1968), from near the village
of Bisnain (Fig. 1) along the Noil Hoeniti and
the Noil Khali. The collections were made by
one of us (S.T.B.) during the course of
mapping the Bisnain Area of West Timor as
part of an investigation 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 S.T.B. is responsible
for the details of the field geology and
N.W.A. is responsible for the systematic
descriptions and the age determination.
All specimens are housed in the Museum of
Victoria collections (NMVP prefix) and all
figured specimens of new species other than
holotypes are paratypes.
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3. 126 N. W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
i I
124"E 126"E
o 100
/
="""'
/
D .... .so./
Amarassi - -
1 I
Fig. 1. Locality map, island of Timor.
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 (cf. Hamilton, 1979,
p. 123). Outcrops in the Bisnain Area (Fig. 2)
consist of a distinctive sequence, approxi-
mately 1500 m thick, of fine to coarse
calcarenites with minor siltstones, shales and
calcareous shales. Four informal members are
outlined here, three of which are fossiliferous.
Maubisse Formation outcrops are separated
from younger units and Permian clastic units
with pillow lavas by fault contacts (Fig. 2).
The stratigraphically lowest member is the
1030 m thick Khali Member characterised by
medium to predominantly coarse calcarenites
with minor shale, chert, and silty horizons and
abundant crinoidal, bryozoan and less
common brachiopod bioclastic detritus. The
base is a fault breccia and the top, with a fault
breccia, is indicated by an abrupt change to
the uniform, fine, structureless, saccharoidal
calcarenite with no fossils of the 210 m thick
Bisnain Member. With abrupt change the
Bisnain Member passes vertically into the
Oemofai Member of 150 m of fine to
predominantly coarse calcarenites rich, at
some levels, in crinoidal debris but very poor
in retrievable brachiopods (only one
incomplete specimen referred herein to
Callytharrella khalii and one specimen of
Elivina bisnaini have been collected). The
Hoeniti Member, with a faulted basal contact,
is a coarse calcarenite, 115 m thick, with
significant intervals of interbedded calcareous
shales and richly fossiliferous in crinoidal,
bryozoan and brachiopod detritus. Further
details of the sequence are provided by
Barkham (1986).
Localities. All localities are indicated on
Figure 2; their stratigraphical details are as
follows: Locality 1,215 m above base of Khali
Member; Locality 3, 256 m above base of
Khali Member; Localities 17 & 30, 225 m
above base of Khali Member; Locality 175,
332 m above base of Khali Member; Locality
95, base of Oemofai Member; Locality 103,
85 m above base of Oemofai Member;
Locality 133, 6 m above base of Hoeniti
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4. ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 127
i
.*2_-
o
1-
158
157iS~:~,
151
t50-/~ :',(:':"
141~ ,.'2:,
) 95 ~ / ~ ; , .
'
~
'
:
"
:
"
~
'
~
t
:
'
~
-
~
2
~ /~ ' 17!30
124"45'E
j k.
Fig. 2. Geological
SKETCH MAP OF THE GEOLOGY
OF TH BISNAIN AREA
5 0 5
Kilometres I I 1
/ ( 1 ......... 7 +v !~
map of the Bisnain Area showing Permian brachiopod localities.
AAA
Pleistocene carbonates
Triassic carbonates]
Triassic elastics J
Hoeniti Member
Oemofai Member
Bisnain Member
Khali Member
Cribas Formation
Atahoc Formation
Thrust boundary
Lithological boundary
Location numbers
(Aitutu Formation )
(Permian carbonates)
(Maubisse Formation)
Member; Locality 141, 59 m above base of
Hoeniti Member; Locality 150, 92 m above
base of Hoeniti Member; Locality 151, 96 m
above base of Hoeniti Member; Locality 153,
101 m above base of Hoeniti Member;
Locality 157, 112 m above base of Hoeniti
Member; Locality 158, 115 m above base of
Hoeniti Member.
Age
Despite the thickness of the sections involved,
the brachiopod faunas available for study are
reasonably uniform in composition. Notably,
the species Elivina bisnaini sp. nov. occurs
throughout the section and Callytharrella
khalii sp. nov. also demonstrates a consider-
able range. For this reason the faunas are
treated as a single assemblage. However, all
locality occurrences are listed under the
descriptions and specimens are figured from
each locality in case futur6 collections should
indicate the potential for zonation.
Comparison of the assemblage described
herein with the classic brachiopod faunas of
Timor indicates some generic links with the
relatively poorly localised and understood
Bitauni assemblages. Generic links may
include Callytharrella (see Productus semi-
reticulatus of Broili, 1916, as discussed by
Archbold, 1985, p. 24) and Elivina (see
Shimizu, 1966, pl. 17, figs 1-13, so-called
Spiriferella rajah, a possible Elivina). How-
ever, the Bitauni assemblages are poorly
understood and are poorly localised being
from soil horizons and isolated boulders so
that mixing of ages may have occurred. While
Bitauni ammonoids indicate a late Artinskian
age (including Kungurian in the sense of
Ruzhentsev, 1956, and Glenister & Furnish,
1961) not all Bitauni brachiopods are from
ammonoid localities.
Comparison of the Bisnain assemblage with
brachiopod faunas of the Western Australian
intracratonic basins revealed considerable
links with the fauna of the Callytharra
Formation. Elivina bisnaini sp. nov. is close
to E. hoskingae Ar~hbold & Thomas (1985)
and Callytharrella khalii sp. nov. recalls C.
callytharrensis (Prendergast, as revised by
Archbold, 1985). Of considerable interest is
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5. 128 N. W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
the first reported occurrence in Timor of
Punctocyrtella. Large Punctocyrtella species
of the nagmargensis type with a distinctive
furrow in the dorsal valve are found through-
out peripheral Gondwanan regions in
sequences of Tastubian and Sterlitamakian
age as summarised by Archbold & Thomas
(1986) and Archbold & Gupta (1986). In
Western Australia, Punctocyrtella australis
(Thomas, 1971)occurs in the Tastubian Lyons
Group fauna and ranges up into the
Sterlitamakian Callytharra Formation.
Other genera present in the Bisnain assem-
blage are compatible with a Sterlitamakian
age, although the material is not always
adequate for specific comparison. Of interest
is the record of Stictozoster sp., a genus often
of Kungurian-Ufimian age (Archbold, 1981;
Waterhouse, 1981)but also recorded from the
Sterlitamakian of Western Australia
(Archbold, 1984). Faunas from the
Sterlitamakian-Aktastinian of peninsula
Thailand (Waterhouse, 1981) include a
possible representative of Callytharrella (in
the form of Stereochia koyaoensis ) as well
as small Streptorhynchus and coarsely ribbed
Stenoscisma which suggest comparison with
the Bisnain assemblage.
On the basis of the comparisons of the
Bisnain material at the species level and the
apparent significant links with the
Sterlitamakian Callytharra Formation fauna
of Western Australia, it is considered most
likely that the Bisnain assemblage is of
Sterlitamakian age. The Bisnain faunas are of
significance because they come from an
outcropping sequence and hence provide data
on reliable faunal associations unlike most
previous Timor collections.
Palaeoclimate
The Permian sedimentary record of the
intracratonic basins of Western Australia
commenced with the well established
sequences of tillitic and glacio-marine deposits
indicative of cold water temperatures
(Teichert, 1950). Cold seas persisted until the
end of the Tastubian (Dickins, 1978;
Archbold, 1982) after which a marked
amelioration of water temperature and
climate is indicated for Western Australian
Sterlitamakian units such as the Callytharra
Formation (Teichert, 1950; Dickins, 1978).
This amelioration is particularly indicated by
the occurrence of warm water Tethyan
brachiopods within the faunas (Teichert,
1950; see also Archbold, 1984 for specific
examples) and hence a temperate climate is
indicated for units such as the Callytharra
Formation.
The similarity of brachiopod species of the
Bisnain brachiopod assemblages with
elements of the Callytharra assemblage points
to a similar water temperature regime for the
two faunas. Dominant elements in the Bisnain
assemblages such as Elivina and Callytharrella
are particularly noteworthy. Additional
elements such as the rare martiniid and
?Spirigerella may be taken to indicate warmer
temperatures but the distinctive presence of
Punctocyrtella would appear to indicate a
temperate water regime as for other
correlative peripheral Gondwanan faunas
(Archbold & Gupta, 1986). This temperate
climate for the Sterlitamakian interval of the
Maubisse Limestone of Timor is in contrast
to the subtropical and tropical climates usually
attributed to the younger Permian intervals of
the same unit (Audley-Charles, 1968) and is
consistent with the model that the Maubisse
Limestones were deposited originally on the
northern margin of Gondwana (cf. Hamilton,
1979, p. 125).
Systematic palaeontology
Order STROPHOMENIDA Opik 1934
Suborder ORTHOTETIDINA Waagen 1884
Superfamily DERBYIACEA Stehli 1954
Family STREPTORHYNCHIDAE Stehli
1954
Arctitreta Whitfield 1908
Type species. Arctitreta pearyi Whitfield 1908
Arctitreta? sp. (Fig. 3A-C)
Comments. A single juvenile shell,
NMVPI20251, indicates the presence of a
streptorhynchid in the Bisnain assemblage.
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6. ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 129
The shell is flattened, maximum width
10.7 mm, length 10.0 mm, with small ears and
distinct radial costellae on both valves,
approximately 9 per 5 cm at the valve
anteriors. The costellae are flat and uneven,
and increase by intercalation and subdivision.
The ventral umbo is slightly hooked. The shell
interior is unknown.
The single specimen recalls juveniles stages
of growth on Arctitreta plicatilis (Hosking) as
redescribed by Thomas (1958, pl. 1l, fig. lc).
Costellae on the Timor shell fall within the
size range of those of A. plicatilis.
Streptorhynchids have been recorded from
both the Bitauni and Basleo faunas of Timor
by Broili (1916, pl. 1, figs 1-5) but the figured
specimen from Bitauni is large and was
relatively transverse during juvenile growth
stages, while the Basleo Streptorhynchus
pseudopelagonatus possesses a strongly
hooked ventral umbo, very fine costellae and
a dorsal sulcus.
Locality. 151, with Elivina bisnaini sp. nov.
Order PRODUCTIDA Sarycheva & Sokol-
skaya 1959
Suborder PRODUCTIDINA Waagen 1883
Superfamily PRODUCTELLACEA
Schuchert & Le Vene 1929
?Family PRODUCTELLIDAE Schuchert &
Le Vene 1929
Stictozoster Grant 1976
Type species. Stictozoster leptus Grant 1976
Stictozoster sp. (Fig. 3D-E)
Comments. A single ventral valve,
NMVP120252, demonstrates the presence of
Stictozoster. The distinctly convex valve,
maximum width 22.9 mm, hinge width
15.0 mm, valve length 18.5 ram, valve
thickness approximately 9.00 mm, is typical
of the genus. The external ornament of fine
spine bases in concentric rows is well
developed, about 6-7 spine bases per 5 mm at
10 mm from the umbo.
Stictozoster is known from the late Early
Permian of Thailand (Grant, 1976) and Irian
Jaya (Archbold, 1981). Both accounts record
species with ventral valves close to the Timor
specimens but with ventral spines a little more
narrowly spaced, about 9 per 5 mm on the
Thai species and ll per 5 mm on the Irian
Jayan shell at 10 mm from the ventral umbo.
Stictozoster has also been reported, with a
query, from the Sterlitamakian faunas of
Thailand (Waterhouse, 1981, pl. 8, fig. 2) and
the Callytharra Formation of Western
Australia (Archbold, 1984). It should be
noted, however, that the Western Australian
species possesses considerably coarser spine
bases than the present Timor shell and other
species referred to the genus. Ufimian and
Kazanian species of the Arctic also possess
fine spines as outlined by Archbold (1981,
p. 9) and Waterhouse (1981, p. 74).
Locality. 175. No other brachiopods present.
Superfamily PRODUCTACEA Gray 1840
Family DICTYOCLOSTIDAE Stehli 1954
Callytharrella Archbold 1985
Type species. Dictyoclostus callytharrensis
Prendergast 1943.
Cailytharreila khalii sp. nov. (Figs 3F-M,
4A-J)
Holotype. NMVP120253, an incomplete
ventral valve from locality 3.
Material. NMVP 120253-120264, a collection
of crushed shells and ventral valves from the
following localities: NMVP120253-120255, a
crushed shell and two ventral valves from
locality 3; NMVP120256-120257, two crushed
ventral valves from localities 17 and 30;
NMVP120258, an incomplete ventral valve
from locality 95; NMVP120259-120260, two
ventral valves from locality 133;
NMVP120261, a crushed shell from locality
141; NMVP120262-120264, 3 ventral valves
from locality 157.
Size ranges. Most specimens are unsuitable for
measurement but the largest specimen
NMVP 120259 yields the following: maximum
width 66 mm, length 61 mm +.
Diagnosis. Large for genus. Ornamentation
fine on trail. Fasciculae weak on trail. Ventral
sulcus shallow to distinct.
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7. 130 N.W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
Description. Large of genus; outline
transverse, widest at hinge at maturity. Ears
distinct, thickened, tips of ears squared with
quadrate outline. Ventral umbo low, pointed.
Ventral sulcus arises between 2 and 3 cm from
umbo, varies between shallow and moderately
deep anteriorly. Visceral disc strongly
reticulate. Rugae fine; reticulate ornament
persists for approximately 4 cm from umbo.
Costae fine, continue anteriorly along trail;
costa with rounded crests, bifurcate or
trifurcate at anterior of spine bases. Costae
number some 10 per cm at 5 cm from umbo
and are normally less than 1.0 mm wide on
anterior trail. Weak fasciculae developed in
trail and costae converge in trail.
Ventral spines scattered, also row on ears,
scattered over visceral disc with occasional
coarser spine on trail, up to 1.7 mm thick at
base. Dorsal exterior poorly known although
reticulate ornament extends over visceral disc,
ears appear costate and spines are absent.
Interior unknown.
Discussion. The species is named because of
its distinctive combination of ornament and
sulcal characters. The large size of the species
and its large ears, although imperfectly
known, indicate a species of Callytharrella
allied to C. callytharrensis which is distin-
guished by its coarser ornament and generally
more pronounced sulcus.
Stereochia koyaoensis Waterhouse (1981,
especially pl. 16, figs 3, 5 and 11) is a
distinctive sulcate form with distinct ears.
Ventral ornament is fine, as is the new species
but the sulcus of the Sterlitamakian Thai
species is deeper than that of C. khalii. The
trail of S. koyaoensis is also short when
compared with that of C. khalii.
Dictyoclostids recorded from Bitauni by
Broili (1916, pl. 2, figs 15, 16) appear to
possess a short trail and short quadrate ears
and hence are less likely to be representatives
of Callytharrella. The large specimen from
Sonnebikoe (Broili, 1916, pl. 2, fig. 14) recalls
Callytharrella in size but details of the ears are
lacking.
Localities. 3, 17 + 30, 95, 133, 141, 157. See
appendix for faunal associations.
Order SPIRIFERIDA Waagen 1884
Suborder SPIRIFERIDINA Waagen 1884
Superfamily SYRINGOTHYRIDACEA
Frederiks 1926
Family SYRINGOTHYRIDIDAE Frederiks
1926
Subfamily PERMOSYRINXINAE Water-
house 1986
Punctocyrtella Plodowski 1968
Type species. Punctocyrtella
Plodowski 1968.
spinosa
Punctocyrteila sp. (Fig. 5A-D)
Comments. A single, worn dorsal valve
(NMVP120265) indicates the presence of this
significant genus in the Bisnain assemblage.
The valve is large, estimated width 80 + mm,
length 39 ram, with a pronounced fastigium
and fold. The fastigium carries a prominent
median furrow. Lateral flanks of the
fastigium are smooth and the lateral flanks of
the valve carry coarse, simple costae, at least
12 on each flank. The shell surface is worn
and the shell substance punctate.
This distinctive dorsal valve is closely
related to Punctocyrtella nagmargensis and its
allies known from a wide range of peripheral
Gondwanan regions including the Pamirs
(Grunt & Dmitriev, 1973), Afghanistan
(Plodowski, 1970; Termier et al., 1974), Tibet
(Hu, 1983; Jin, 1985), Kashmir (Bion, 1928)
and Western Australia (Thomas, 1971).
Punctocyrtella, with a distinct dorsal furrow,
is characteristic of these Tastubian-Sterlita-
Fig. 3. A-C, Arctitretasp. NMVPI20251,juvenileshell in ventral, dorsal and ventral views, x 2.5, x 2.5 and x
I. D, E, Stictozostersp. NMVP120252,ventral valvein posteriorand ventral views, x 1.5. F-M, Callytharrella
khaliisp. nov. F-H, NMVP120253,holotype,ventral valvein posterior, ventraland anterior views, x 1.2, x 1,
x 1.3. 1, J, NMVP120254,ventral valvein ventral and posteriorviews, x 1.2 and x 1.4. K, L, NMVP120262,
ventral valvein posterior and ventral views, x 1. M, NMVP120255,crushed shell in ventral view, z 1.
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8. ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 131
Q
H
I_
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9. 132 N.W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
makian Gondwanan faunas and hence the
new record from Timor is significant.
Younger Punctocyrtella may have lost the
dorsal furrow as suggested by Waterhouse
(1983), and may have a lower ventral interarea
(Waterhouse, 1987).
Locality. 133, with Callytharrella khalii sp.
nov.
Superfamily SPIRIFERACEA King 1846
Family SPIRIFERIDAE King 1846
Subfamily NEOSPIRIFERINAE Waterhouse
1968
Neospirifer Frederiks 1924
Type species. Spirifer fasciger von Keyserling
1846
Neospirifer sp. (Fig. 5E)
Comments. A single incomplete ventral valve
(NMVP120266) possesses the characteristic
fine costae in fasciculae of Neospirifer. The
specimen is relatively small (estimated width
48 mm), no growth lines are preserved and
therefore no ontogenetic observations can be
made on the specimen. Referral of the
specimen to any named species or lineage of
Neospirifer is not possible.
Locality. 141, with Callytharrella khalii sp.
nov.
Subfamily SPIRIFERELLINAE Waterhouse
1968
Elivina Frederiks 1924
Type species. Spirifer tibetana Diener 1897
Elivina bisnaini sp. nov. (Fig. 6A-Z, AA-LL)
Holotype. NMVP120270, a ventral valve
from locality 3.
Material. NMVP 120267-120295, a collection
of specimens from the following localities:
NMVP120267, a ventral valve from locality
1; NMVP120268-120278, 10 ventral valves
and an incomplete shell from locality 3;
NMVP 120279-120285, 6 ventral valves and 1
incomplete shell from localities 17 and 30;
NMVP120286, a ventral valve from locality
103, NMVP120287-120289, 3 ventral valves
from locality 150; NMVP120290, a ventral
valve from locality 151; NMVP120291, a
ventral valve from locality 153;
NMVP120292-120293, 2 ventral valves from
locality 157; NMVP10294-120295, 2 ventral
valves from locality 158.
Size ranges. Hinge width, 5.9-28.0 mm;
maximum width, 8.5-36.6 mm; ventral valve
length, 10.4-42.5 mm; ventral interarea
length, 2.5-7.0 mm (largest specimen not
measurable).
Diagnosis. Medium to large for genus.
Elongate outline. Costae simple or branching
producing fascicles of 3 or 4 costae at sub-
maturity and maturity. Umbonal shoulders
accentuate triangular posterior outline.
Description. Medium to large for genus,
biconvex, convexity of dorsal valve not well
known because of imperfect material. Outline
elongate with maximum width anterior of
mid-length. Hinge narrow.
Ventral valve strongly convex. Ventral
umbo pointed, overhangs narrow interarea
which is convex and bisected by narrow,
triangular delthyrium. Delthyrium filled with
massive, flattened apical callosity at maturity.
Dental plates and adminicula buried in
massive valve thickening. Teeth short, stout,
pointed, Ventral muscle field elongate,
pointed anteriorly. Adductor scars narrow,
long, clearly differentiated from diductors on
each side. Valve interior pitted, at times with
weakly developed plicae reflecting external
ornamentation. Sulcus broad, U-shaped in
cross-section, with distinct median sulcal
costa. Sulcus arises at umbo. Costae arise at
umbo, sulcal bounding costae may bifurcate
Fig. 4. Callytharrella khaliisp, nov. A, NMVPI20256, ventral valve in ventral view, × 1. B, NMVPI20257, ventral
valve in ventral view, × 1.2. C, NMVPI20259, ventral valve in ventral view, × 1. D, NMVP120260, ventral valve
in ventral view, x I. E, NMVP120258, ventral valve in ventral view, × 1. F, NMVPI20261, crushed shell in dorsal
view, × 1. G, H, NMVPI20263, ventral valve in ventral and anteroventral views, x 1. I, J, NMVP120264, ventral
~alve in ventral and anterovenlral views, × 1.
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10. ALCHERINGA T1MORESE PERMIAN BRACH1OPODS 133
A
D
C
iz
O
F (3
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11. 134 N. W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
Fig. 5. A-D, Punctocyrtella sp. dorsal valve in dorsal, posterior, anterior and posterodorsal views, × 1. E, Neospirifer
sp. E, ventral valve in ventral view, x 1.
within 0.6 cm of umbo or remain simple and
bifurcate at greater distance anteriorly. Costae
branch asymmetrically and may form sulcal
bounding fascicles of 3 or 4 costae by valve
anterior. Costae well rounded with consider-
able variation within species in terms of
relative size, bifurcation distances and
appearance of fascicles as shown in Figure 6.
Valve anterior not thickened, posterior with
massive fibrous thickening.
Dorsal valve poorly known, apparently
convex, thin, with distinct fastigium.
Fastigium with distinct median groove.
Lateral flanks of valve with costae, simple on
available material.
Micro-ornament consists of distinct radial
capillae, increasing in number by branching
and intercalation, crossed by fine growth
lamellae. Minute pustules occur on the cross-
over sites of capillae and lamellae.
Fig. 6. Elivina bisnaini sp. nov. A, NMVP120267, ventral valve in ventral view, x 1. B, NMVP120268, ventral
valve in ventral view, x 1. C, NMVP120269, ventral valve in ventral view, x 1. D, E, NMVP120270, ventral valve
in ventral and dorsal views, x 1. F, G, NMVP120271, ventral valve in ventral and dorsal views, x 1. H, NMVP120272,
ventral valve in ventral view, x 1. I, J, NMVP120273, ventral valve in ventral and dorsal views, x 1. K, NMVP120274,
ventral valve in ventral view, x 1. L, M, NMVPI20275, ventral valve in ventral and dorsal views, x 1.2. N,
NMVPI20276, juvenile ventral valve in ventral view, x 1. O, P, NMVP120278, incomplete shell in ventral and
posterior views, x 1 and x 1.3. Q, NMVP120277, ventral valve in ventral view, x 1. R, NMVP120279, ventral
valve in ventral view, x 1. S, NMVP120280, ventral valve interior view, x 1. T, NMVP120281, ventral valve in
ventral view, x 1. U, NMVP120282, aberrant ventral valve in ventral view, x I. V, NMVP120283, gerontic ventral
valve in ventral view, x 1. W, Y, Z, NMVPI20285, incomplete shell in ventral and dorsal views, x 1, portion of
ventral surface enlarged, x 7. X, NMVP120284, ventral valve in ventral view, x 1. AA, NMVP120286, ventral
valve in ventral view, x 1.2. BB, NMVPI20287, ventral valve in ventral view, x 1. CC, NMVP120293, ventral
valve in ventral view, x 1. DD, EE, NMVP120289, ventral valve in ventral and dorsal views, x 1. FF, NMVP120288,
ventral valve in ventral view, × 1. GG, HH, NMVPI20290, ventral valve in ventral view, × 1 and × 2.5. II,
NMVP120291, ventral valve in ventral view, × 1. J J, NMVPI20294, ventral valve in ventral view, × 1. KK,
NMV120292, ventral valve in ventral view, × 1. LL, NMVP120295, ventral valve in ventral view, × 1.
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12. ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 135
,Q
v ~
QF GG
"E F ..Id
J CC
nD
KK
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13. 136 N. W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
Discussion. Elivina bisnaini sp. nov. is a
variable species of the genus and recalls
individuals of the equally variable E.
hoskingae Archbold & Thomas (1985) from
the Sterlitamakian Callytharra Formation and
Fossil Cliff Member of Western Australia.
Individuals of E. bisnaini can be closely
similar to individuals of E. hoskingae (e.g.
compare Fig. 6R with fig. 31 in Archbold &
Thomas, 1985, p. 45). Nevertheless, ventral
costae of the present species tend to bifurcate
earlier than those of most specimens of E.
hoskingae. Umbonal shoulders of E.
hoskingae are also often more rounded than
those of E. bisnaini sp. nov. The two species
appear to be closely related and both, because
of the variability displayed, indicate the need
for caution when assigning individual
specimens to species. A third species that
shares features with E. bisnaini sp. nov. and
E. hoskingae is E. tenuisulcatus (Merla) from
the Early Permian of the Karakorum. Merla's
account of the species (1934, p. 273, pl. 26,
figs 14-21) includes specimens of comparable
size with the above species and with relatively
coarse ventral costae. Costae branch a little
later in ontogeny than typical E. bisnaini sp.
nov.
Spiriferella rajah and Spiriferella sp. nov.
of Shimizu (1966, pl. 17, figs 1-13) from a
Bitauni correlated locality in East Timor,
appear to be a species of Elivina with ventral
costae coarser than typical E. bisnaini. The
dorsal fastigium of Shimizu's material is
comparable to that known for E. bisnaini but
details of the ventral interior are not well
known for the East Timor material. A
possible Elivina from Letti (Broili, 1915,
pl. 21, fig. 11) possesses ventral costae that
exhibit only minor bifurcations and hence
fascicles of costae are not evident from the
figure. Specimens attributed to Spirifer lyra
and Spirifer tibetanus by Hamlet (1928, pl. 6,
figs 5, 6; pl. 7, figs 1, 2) from Noil Simaan
and Wesleoe, Timor respectively also indicate
species of Elivina but the former specimens
are shells with fine costae and strongly
rounded umbonal shoulders while the latter
specimens have coarse, often simple costae.
Hence up to four additional species of Elivina
may be present in the Timor and Letti islands
but none appear to be comparable with E.
bisnaini. Whether or not the classic species
Spirifer kupangensis Beyrich (1865, p. 78, pl.
1, figs 6a-c), type specimen refigured by Frech
(1902, p. 589), represents a species of Elivina
is arguable but it is not close morphologically
to either E. bisnaini or E. hoskingae in view
of its coarse ventral costae and strongly
biconvex shell.
E. tschernyschewi Waterhouse &
Waddington (1982) from the Sakmarian of
the Urals (see Chernyshev, 1902, pl. 7, figs
2-6) is a species of rounded to elongate outline
with ventral sulcal costae forming fascicles of
3 to 4 costae. The Soviet species can possess
a relatively deep sulcus which is V-sided.
Waterhouse & Waddington (1982, p. 33)
designated as holotype the specimen of
Spiriferella tibetana not Diener, Chernyshev
1902 illustrated by Chernyshev (1902) on plate
17, figure 3. Presumably plate 7, figure 3 was
the intention of those authors and so their
original designation is treated by us as being
a lapsus calami.
The Spirifer tibetanus of Mansuy (I913, pl.
3, fig. 11; pl. 4, fig. la-lc) from Khamkeut,
Laos, of Early Permian age, is a somewhat
more trigonal species that E. bisnaini sp. nov.
but its costae recall those of some specimens
of the Timor species.
E. occidentalis (Schellwien, 1900, pl. 11,
figs 10-13) from the Sakmarian of the Carnic
Alps, appears to be characterised by finer
ventral costae than are normal for E. bisnaini
sp. nov. as is also shown by the specimen
figured by Heritsch (1938, pl. 8, figs 9-I1) and
referred to Elivina tibetanus. Heritsch (1931,
p. 27, pl. 2, figs 79, 80) figured a further shell
(as Spirifer lyra) from the Carnic Alps that
also appear to belong to Schellwien's species
in view of its fine costae.
Localities. 1, 3, 17 and 30, 103, 150, 151,153,
157, 158. See appendix for faunal associations
of each locality.
Superfamily RETICULARIACEA Waagen
1883
Family ELYTHIDAE Frederiks 1924
Elythid indet. (Fig. 7A-B).
Comments. A single, worn incomplete shell
(NMVP120296) maximum width 16.9 mm,
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14. ALCHERINGA TIMORESE PERMIAN BRACHIOPODS 137
B
G
1 |
D
Fig. 7. A, B, elythid indet. NMVP120296, incomplete shell in dorsal and ventral views, x 1. C, D, martiniid indet.
NMVP120297, ventral valve in ventral view and view of ventral interarea, × 1. E-G, Stenoscisma? sp. E, F,
NMVPI20298, crushed shell in ventral and dorsal views, x 1.2. G, NMVPI20299, dorsal valve in dorsal view, ×
1. H-J, Cleiothyridinasp. H, I, NMVP120300,crushed shellin dorsal and posterodorsal views, x 1.2.J, NMVPI20301,
dorsal valve in dorsal view, x 1. K, Spirigerellasp. NMVPI20302, shell in ventral view, x 1.2.
may represent an elythid of the Phricodothyris
type as discussed by Archbold & Thomas
(1984). It is figured for the sake of complete-
ness of the fauna.
Locality. 17.
Superfamily MARTINIACEA Waagen 1883
Family MARTINIDAE Waagen 1883
Subfamily MARTINIINAE Waagen 1883
Martiniid indet. (Fig. 7C-D)
Comments. A single large incomplete ventral
valve (NMVP120297) represents a martiniid
of uncertain affinity. The specimen is figured
for completeness. Large martiniids are known
from Basleo and Port Timor (Broili, 1916, pl.
8, figs 17-21; pl. 9, fig. 1) and from Bitauni
equivalents on Letti (Broili, 1915, pl. 21, figs
1 and 3) and hence a wide time range is
indicated.
Locality. 17, with Elivina bisnaini sp. nov.,
Callytharrella khalii sp. nov. and Cleio-
thyridina sp.
Order RHYNCHONELLIDA Kiihn 1949
Superfamily STENOSCISMATACEA
Oehlert 1887
Family STENOSCISMATIDAE Oehlert 1887
Stenoscisma Conrad 1839
Type species. Terebratula schlottheimii yon
Buch 1835
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15. 138 N. W. ARCHBOLD & S. T. BARKHAM ALCHERINGA
Stenoscisma? sp. (Fig. 7E-G)
Comments. Two poorly preserved specimens
(NMVP120298-120299) are indicative of
stenoscismatids in the assemblage. Despite
being incomplete, the most noticeable aspect
of the specimens is the very coarse plications
in the ventral sulcus and on the dorsal
fastigium and lateral flanks of the shell. This
feature and the general shell outline suggest
a species allied to the Camarophoria globosa
of Hamlet (1928) from Bitauni (Hamlet, pl.
9, fig. 4), Hamlet's specimen in larger and has
coarser plicae than true Stenoscisma? globosa
(Chernyschev, 1902, pl. 46, figs 2, 3) from the
Sakmarian of the Urals although the two may
well be allied when better known. Also
probably allied to the Timor material is the
Gerassimovia bactriana Grunt (in Grunt &
Dmitriev, 1973, pl. 9, figs 3-5), a large form
with coarse plicae from the Aktastinian of the
Pamirs.
Localities. 3 and 17 with Elivina bisnaini sp.
nov. and Callytharrella khalii sp. nov.
Order ATHYRIDIDA Dagys 1974
Superfamily ATHYRIDACEA McCoy 1844
Family ATHYRIDIDAE McCoy 1844
Cleiothyridina Buckman 1906
Type species. Atrypa pectinifera Sowerby
1840
Cleiothyridina sp. (Fig. 7H-J)
Comments. Cleiothyridina is represented by
2 specimens (NMVP120300-120301) one of
which has been compressed and hence appears
more transverse than would have originally
been the case. The general form of the
specimens is consistent with that of small
species such as Cleiothyridina ailakensis
(Reed) as recorded by Grunt & Dmitriev
(1973) from the Aktastinian of the Pamirs or
young C. seriata Grant (1976, pl. 54) from the
late Early Permian of Thailand. C.
baracoodensis (Etheridge, 1903) from the
Sterlitamakian Callytharra Formation of
Western Australia is also comparable, at its
submature stages of growth, with the present
material but further comparison is not
warranted given the limited nature of the
Timor material.
Loaclities. 17 and 157 with Elivina bisnaini
and Callytharrella khalii sp. nov.
Family SPIRIGERELLIDAE Grunt 1980
Spirigerella Waagen 1883
Type species. Spirigerella derbyi Waagen 1883
Spirigerella sp. (Fig. 7K)
Comments. A single crushed shell indicates
the presence of Spirigerella or an ally in the
Bisnain assemblage. The specimen
(NMVP120302) is relatively large, maximum
width 28.5 mm, length 33.2 mm, and hence
is comparable with S. timorensis (Rothpletz,
1892) as figured by Broili (1916, pl. 9, figs
7-14) from Basleo. Hence this is an element
of the assemblage that may, on the surface,
indicate a younger age but the species has also
been recorded from Bitauni (Hamlet, 1928,
p. 54) and hence the genus may range
throughout much of the Permian of Timor.
Locality. 3 with Elivina bisnaini sp. nov. and
Callytharrella khalii sp. nov.
Acknowledgements
S.T.B. is grateful to Dr A. J. Barber for
assistance and advice. N.W.A.'s work is
supported by the Australian Research Grants
Scheme. We thank Isabel Munro for typing
the manuscript.
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1
3
17 & 30
175
95
103
133
141
150
151
153
157
158
Elivina bisnaini sp. nov.
Callytharrella khafii sp. nov., Elivina bisnaini sp. nov., Stenoscisma? sp.
Callytharrella khalii sp. nov., Elivina bisnaini sp. nov., elythid indet., martiniid
indet., Stenoscisma? sp., Spirigerella? sp.
Stictozoster sp.
Callytharrella khalii sp. nov.
Elivina bisnaini sp. nov.
Callytharrella khalii sp. nov., Punctocyrtella sp.
Callytharrella khalii sp. nov., Neospirifer sp.
Elivina bisnaini sp. nov.
Arctitreta? sp., Elivina bisnaini sp. nov.
Elivina bisnaini sp. nov.
Callytharrella khalii sp. nov., Elivina bisnaini sp. nov.
Elivina bisnaini sp. nov.
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