Intraspecific variation in Triassic Ophthalmidiid Foraminifera from Timor

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www.sciencedirect.com Revue de micropaléontologie 55 (2012) 39–52

Original article

Intraspecific variation in Triassic Ophthalmidiid Foraminifera from Timor Variation intraspécifique chez les Foraminifères Ophthalmidiidés du Trias du Timor David W. Haig ∗ , Eujay McCartain School of Earth and Environment (M004), The University of Western Australia, Stirling Highway, 6009 Crawley, Australia

Abstract Four ophthalmidiid species are described as free specimens extracted from mudstones and wackestones of Triassic age: Atsabella bandeiraensis nov. gen. nov. sp., Karaburunia atsabensis nov. sp., Ophthalmidium sp. cf. O. primitivum Ho and Spirophthalmidium grunaui nov. sp. Analysis of morphological variation found in large suites of specimens suggests that, as in modern miliolids, apertural characteristics, chamber shape and adult test size are features that vary within narrow limits and may be used to define species. In genera with milioline coiling in the post-embryonic stage, chamber arrangement may be highly variable. Post-embryonic coiling in K. atsabensis varies from quinqueloculine to almost spiroloculine and encompasses morphotypes that, in thin-section studies, have been attributed to a number of other genera. In Timor Leste, A. bandeiraensis, K. atsabensis and S. grunaui have been found with conodonts indicative of the Carnian, but the full local stratigraphic range of these species is uncertain. K. atsabensis occurs at another locality with conodonts suggestive of the Middle Triassic or less likely Carnian. Ophthalmidium sp. cf. O. primitivum has been found at one locality associated with A. bandeiraensis and K. atsabensis in a stratigraphic succession that suggests a correlation to the Carnian or Norian. The ophthalmidiids are found commonly associated with organic-cemented agglutinated and hyaline foraminifera and at some localities common to abundant ostracods and mollusc debris. They were most common in organic-rich carbonate mud of shallow-marine environments. © 2012 Elsevier Masson SAS. All rights reserved. Keywords: Foraminifera; Ophthalmidiida; Triassic; Timor

Résumé Nous décrivons quatre espèces de type « ophthalmidiid » à partir de spécimens prélevés dans des formations carbonatées (« mudstones » et « wackestones ») d’âge Triasique : Atsabella bandeiraensis nov. gen. nov. sp., Karaburunia atsabensis nov. sp., Ophthalmidium sp. cf. O. primitivum Ho et Spirophthalmidium grunaui nov. sp. L’analyse des variations morphologiques réalisées suggère que, comme dans le cas des espèces modernes de miliolidés, les caractéristiques de l’ouverture, la forme des loges et la taille du test adulte sont des caractéristiques qui varient dans des limites étroites et peuvent être utilisées pour définir les espèces. Dans les genres avec un enroulement de type milioliforme au stade post-embryonnaire, l’arrangement de la chambre peut être très variable. L’enroulement post-embryonnaire dans K. atsabensis varie du type « quinqueloculin » à proche de « spiroloculin ». Ces observations englobent des morphotypes, en analyse de lames minces, ont été auparavant attribués à un certain nombre d’autres genres. Au Timor Leste (Timor oriental), A. bandeiraensis, K. atsabensis et S. grunaui ont été observés en association avec des conodontes, attestant un âge carnien, mais l’attribution stratigraphique de ces espèces reste incertaine. Dans une autre localité du Timor oriental, Karaburunia atsabensis est observée en association avec des conodontes datés du Trias moyen. Ophthalmidium sp. cf. O. primitivum a été observé associé à A. bandeiraensis et K. atsabensis dans une succession stratigraphique que l’on attribue au Ladinien ou au Carnien. Les ophthalmidiidés se trouvent couramment associés à des formes de foraminifères agglutinés et hyalins et, dans certaines localités, en association avec un nombre moyen à abondant d’ostracodes et de débris de mollusques. Les « ophthalmidiids » sont fréquemment associés à des dépôts de carbonates fins (type « mudstone – wackestone ») et riches en matière organique, déposés en milieu marin peu profond. © 2012 Elsevier Masson SAS. Tous droits réservés. Mots clés : Foraminifères ; Ophthalmidiida ; Trias ; Timor

∗

Corresponding author. E-mail address: [email protected] (D.W. Haig).

0035-1598/$ – see front matter © 2012 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.revmic.2011.12.001

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1. Introduction Ophthalmidiids became conspicuous during the Middle Triassic having evolved probably from non-septate cornuspiraceans (Zaninetti and Brönnimann, 1969). They are characterized by a proloculus followed by a planispirally coiled second chamber of variable length and then in latter whorls by variable coiling and chamber divisions, although the latter are often rudimentary and usually defined by kinking and narrowing of the chamber lumen. The Ophthalmidiida were probably the progenitors of the major evolutionary expansion within the Order Miliolida during the later Mesozoic and Cenozoic, following the Mississippian (early Carboniferous) appearance of the Class Miliolata (Vachard et al., 2010). In the Timor Triassic (Fig. 1), four genera have been identified. Three follow the classification of Loeblich and Tappan (1987): Karaburunia, Ophthalmidium and Spirophthalmidium, each with a single species, two of which are new. A fourth species is placed in a new genus Atsabella and the relationship of this to the widely recognized Gsollbergella is discussed. The Timorese species occur in very few of numerous samples that have yielded Triassic foraminiferal assemblages

from Timor Leste (Haig and McCartain, 2010) and are most abundant, as siliceous casts, in the residues of acid-digested Upper Triassic grey wackestone. Although the tests are replaced by silica, fine details of morphology are preserved. In Triassic literature, most records of the Ophthalmidiida are based on randomly orientated thin sections. For planispiral forms such as Ophthalmidium and Spirophthalmidium, equatorial sections provide a good representation of morphology although apertural details are often lost if the spiral plane is a little uneven. Sections of ophthalmidiids with milioline coiling (such as Karaburunia and the new Atsabella) are more difficult to interpret. A major aim of the present paper is to describe intraspecific morphological variation based on free specimens extracted from the rock in order to gain a better understanding of those features that limit species diagnosis. Comprehensive analyses of modern miliolids (e.g., Haig, 1988; Parker, 2009) show that apertural structure, chamber shape, wall-surface texture including ornamentation and ultrastructure and maximum test size are features that usually remain constant within a species. This paper investigates whether similar features define species among primitive Ophthalmidiida.

Fig. 1. Map of Timor showing extent of the Gondwana Megasequence (Permian to Middle Jurassic deposits of the East Gondwana interior rift system - see Fig. 2) and associated Australian Margin Megasequence (deposits that accumulated on passive margin after continental breakup). An enlarged map of Timor Leste shows outcrop areas of Triassic basinal facies. Approximate positions of studied samples (Table 1) are shown. Distribution of strata has been modified from Audley-Charles (1968) and Harris (2006). Megasequence nomenclature follows Keep and Haig (2010).

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Eastern Sulawesi (Late Triassic): Ophthalmidium spp. (11) Paraophthalmidium sp. (11) Sinto Ridge, Banda Sea (Late Triassic): Quinqueloculina sp. (10) [? = Karaburunia]

Seram (Late Triassic): Ophthalmidium spp. (12, 13) Paraophthalmidium sp. (13) Timor (Late Triassic): Gsollbergella spiroloculiformis (1) Karaburunia n. sp. (present study) Ophthalmidium n. sp. (present study) Spirophthalmidium n. sp. (present study)

Terranes now in S. E. Asia

Greater India

Australia

India

Antarctica 0

1000

Northern Carnarvon Basin (late Early to Early Middle Triassic): Gsollbergella spiroloculiformis (2) [?] ?Karaburunia rendeli (2) Ophthalmidium orbiculare (3) [Praeophthalmidium] Ophthalmidium sp. A (2) Ophthalmidium sp. B (2) Wombat Plateau/Northern Exmouth Plateau (Late Triassic): ?Ophthalmidium triadicum (4) [Spirophthalmidium] Ophthalmidium spp. (genus – sensu lato; 5, 6, 7).

km

Southern Africa

Kumaun and Spiti regions (Late Triassic): Ophthalmidium lucidum (8) Ophthalmidium triadicum (8, 9) [Spirophthalmidium] Ophthalmidium sp. (8)

East Gondwana interior rift system. Dashed lines indicate positions of rifts during final breakup (at 155 Ma in vicinity of Timor; 136 Ma off S. W. Australia). Rifting along Meso-Tethyan ocean boundary started during the Early Permian. East Gondwana

Ocean (Meso-Tethys on left) Fig. 2. Position of Timor within the East Gondwana interior rift system (modified from Harrowfield et al., 2005). Ages of continental breakup follow Heine and Müller (2005); age of initial rifting along Meso-Tethyan ocean boundary follows Metcalfe (2006). Foraminiferal records are from (1) Haig et al. (2007), (2) Apthorpe (2003), (3) Heath and Apthorpe (1986), (4) Quilty (1990), (5) Röhl et al. (1991), (6) Zaninetti et al. (1992), (7) Kristan-Tollmann and Gramann (1992), (8) Kristan-Tollmann (1984), (9) Bhargava and Gadhoke (1988), (10) Villeneuve et al. (1994), (11) Martini et al. (1997), (12) Al-Shaibani et al. (1983), (13) Martini et al. (2004).

2. Geological setting The Triassic of Timor (Fig. 1), reviewed by Charlton et al. (2009), was deposited in the East Gondwana Rift System (Haig and McCartain, 2010). Haig et al. (2007) illustrated specimens attributed to the ophthalmidiid genus Gsollbergella in thin section from the Carnian of Timor Leste. Other records of ophthalmidiids, usually in open species nomenclature and in thin section, from the East Gondwana Rift System include those from Eastern Sulawesi (Martini et al., 1997), Seram (Al-Shaibani et al., 1983; Martini et al., 2004) and Sinto Ridge (Villeneuve et al., 1994) to the north of Timor; and the Dampier Sub-basin of the Northern Carnarvon Basin (free specimens illustrated by Apthorpe, 2003; Heath and Apthorpe, 1986) and Wombat Plateau and the contiguous northern Exmouth Plateau (Quilty, 1990 - free specimens; Kristan-Tollmann and Gramann, 1992;

Röhl et al., 1991; Zaninetti et al., 1992) to the south of Timor (Fig. 2). 3. Methods and materials The studied ophthalmidiids are free specimens that come from seven samples (Table 1) including the washed residues of mudstone and the acid-digested residues of wackestone. Standard micropalaeontological processing techniques were used for the disaggregation of samples. Friable mudstone was gently crushed, boiled in water with added detergent and the clay dispersant sodium hexametaphosphate and then wet sieved through 2 mm, 150 ␮m and 63 ␮m meshes. For indurated limestone, 330 g portions of 1–3 cm rock pieces were placed in 5 L of water, 500 mL of glacial acetic acid and 2 L of buffer solution (prepared by filtering spent solution from previous processing). Samples

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Table 1 Location, stratigraphic setting, depositional environment and age of samples containing free specimens of ophthalmidiids from Timor. Samples

UWA No.

◦S

◦E

A7012A

144199

–8.91461

125.42158

A dark grey wackestone collected from low in a displaced block of a unit comprising thin-bedded indurated dark grey wackestone interbedded with thin slightly less indurated grey marl and some beds of peloidal packstone and grainstone (Brisbout, 2010, unpubl.). The wackestone beds differ from those of other sections described here in lacking a typical basinal fauna (e.g., radiolarians, calcispheres, bivalve-filaments). Foraminifera (including abundant organic-cemented agglutinated types), ostracods, brachiopods, crinoidal debris and sponge spicules are present. This unit is underlain by a tidal-deltaic mudstone-sandstone facies that overlies a shallow-marine skeletal, peloidal, oolitic limestone; and is overlain by a megalodon limestone (Brisbout, 2010, unpubl.). This association of facies suggests the thin-bedded wackestone unit may represent deposition in a shallow normal-marine to metahaline lagoonal setting. Palynomorph assemblages from 30 m below this section indicate an age no older than Carnian. Foraminifera from the basal limestone unit in the overall succession (observed in thin section), about 100 m below the sampled level, include Triadodiscus eomesozoicus, Trocholina ventroplana, Endotriadella wirizi, Endoteba ex. gr. E. obturata, Planiinvoluta carinata, Duotaxis inflata, D. metula, Siphovalvulina alpina, Siphovalvulina sp., Ophthalmidium sp. suggestive of a Ladinian to Carnian age. Rare thin sections of possible Triasina have been observed in a bed at the top of the thin-bedded wackestone unit suggesting a Norian or Rhaetian age for this level. The age of sample A7012A is probably Carnian or Norian

C659a

143763

–8.68860

126.01370

From one of two thick (50–100 cm) beds of dark grey wackestone interbedded with weakly laminated to massive grey silty mudstone. These strata are very similar to those discussed for the M625a, b succession and a similar “basinal” depositional setting and depositional processes are interpreted. One conodont element fragment was recovered from the sample. The fragment represents the anterior third of a P element, which displays a high blade and a platform that gently tapers to the anterior point with no obvious step. These characteristics suggest a probable Middle Triassic age for this element with the possibility of a Carnian age

M625a, b

143445

–8.52230

126.00370

These mudstone samples were taken from an outcrop dominated by mudstone with rare thin to medium beds of grey wackestone and bivalve-filament packstone. Horizons rich in halobiid bivalves are present. Calcareous mudstone-wackestone beds are moderately bioturbated and include radiolaria, calcispheres and bivalve-filaments. The succession suggests a “basinal” depositional setting below storm-wave base but within neritic water depths. Fluid mud density currents, distal tails of coarser density currents or hemipelagic sedimentation from fluvio-deltaic derived hypopycnal plumes may have deposited the mudstone. Carbonate sediment was sourced from shallow water and deposited through either hemipelagic settling from carbonate platform derived hyperpycnal currents or hypopycnal plumes. The limestone beds were deposited during periods with limited deposition of siliciclastic sediment and bivalve-filament packstone suggests concentration of these by sediment winnowing by either density currents or possible water currents. The age is uncertain

M644a

143908

–8.61360

126.10170

From a wackestone that forms part of a succession dominated by thin to thick wackestone beds and rare packstone-grainstone beds interbedded with thin beds of mudstone. The limestone strata are very similar to those described for samples M625a, b and similar processes are likely responsible for the deposition of this succession. Among many possibilities, a more distal position relative to a fluvio-deltaic source and a more proximal position to a carbonate factory may explain the higher limestone to mudstone ratio of this succession compared to those of samples C569a, M625a, b, S6101a and S6103a. Conodont elements recovered from this sample are attributable to Carniepigondolella aff. zoae-samueli and Metapolygnathus lindae (tentatively identified by M. Orchard, 2008) indicating the late Carnian

S6101a

144519

–8.83349

125.65385

A grey wackestone sample from a section that passes from weakly calcareous mudstone with rare thin to medium grey wackestone beds into an interval dominated by the latter and thin beds of weakly calcareous mudstone. These strata are very similar to those described for samples M625a, b and similar depositional processes and setting are interpreted for this succession. The anterior two thirds of a single conodont P element was recovered from this sample. This fragment displays a high free blade with a platform initiated halfway down the fragment with a pronounced step and uplifted platform margins. These characteristics suggest an age no older than Carnian. On the aboral surface there is no indication of flaring of the keel and the presence of the basal pit. This suggests this element belongs to one of several genera restricted to the Carnian

S6103a

144531

–8.83517

125.66085

Similar to sample C659a this grey wackestone sample was taken from a thick grey wackestone bed within a mudstone-dominated succession. These strata are very similar to those described for sample M625a, b and were likely deposited in a similar setting by similar processes. Age is uncertain

Stratigraphic setting, depositional environment and age

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were left to digest in the acetic acid solution for 4 to 5 days and then wet sieved with the 150 ␮m and 63 ␮m size fractions retained. Residues were picked using a very fine sable-hair brush under a reflecting light stereomicroscope. All illustrated specimens have been lodged in the Commonwealth Palaeontological Collection of Geoscience Australia, Canberra. The rock samples and processed residues are housed in the collections of the Earth Science Museum at the School of Earth and Environment at the University of Western Australia. 4. Timor ophthalmidiid species The classification follows Loeblich and Tappan (1987, 1992). Morphological terminology follows Hottinger (2006). Order MILIOLIDA Lankester, 1885 Superfamily CORNUSPIRACEA Schultze, 1854 Family OPHTHALMIDIIDAE Wiesner, 1920 Genus Atsabella nov. gen. Type species: Atsabella bandeiraensis nov. sp.; by monotypy; Triassic, Timor. Etymology: The genus is named after the Atsabe Subdistrict, Timor Leste. Diagnosis: Test almost ovoid in lateral longitudinal view with broadly rounded base and broadly rounded to broadly acute apertural end; moderately compressed in facial (apertural) view with almost flat or gently convex sides and broadly rounded margin; initial chamber arrangement obscure; later chambers defined by constricted foramina and rapid expansion of the subsequent chamber lumen, added in variable milioline arrangement with final chambers almost spiroloculine; wall imperforate, uniformly thin except for slight thickenings at aperture and foramina; aperture a narrow arch of variable height with slightly thickened rim, without base or any toothplate structure, maybe slightly asymmetric. Remarks: It is difficult to compare the free specimens on which Atsabella is based to “Agathamminoides gsollbergensis Zaninetti, 1969” the originally designated type species of the monotypic “Agathamminoides Zaninetti, 1969”. Zaninetti’s species was based on thin sections with the holotype (Zaninetti, 1969: fig. 1B; a “longitudinal” section) showing a broadly ovoid outline similar to that in Atsabella, but with apparently less confined foramina between final chambers (although seen only in one dimension in A. gsollbergensis). A transverse section included among the type material (Zaninetti, 1969: fig. 1C) of A. gsollbergensis is distinctly triangular with regular quinqueloculine arrangement of whorls in contrast to a more irregular milioline arrangement tending spiroloculine in Atsabella. Zaninetti (1976) placed “Agathamminoides gsollbergensis Zaninetti, 1969” into synonymy with “Agathamminoides spiroloculiformis (Oraveczné Scheffer, 1968)” assuming the later was the senior subjective synonym. Agathammina spiroloculiformis was described as a new species by Oraveczné Scheffer in the “Annual Report of the Hungarian Geological Institute 1968 which was published in January 1971 not 1968 (see Oraveczné Scheffer, 1971). Because Agathamminoides Zaninetti, 1969 was preoccupied by Agathamminoides Vangerow, 1964, Zaninetti (1979) established a new name

43

“Gsollbergella Zaninetti, 1979” for her 1969 genus but cited the type species as “Gsollbergella spiroloculiformis (Oraveczné Scheffer, 1968)”. Gsollbergella spiroloculiformis was based on orientated thin sections of free specimens (Oraveczné Scheffer, 1971: p. 91, 102, 103, pl. 2, figs. 1–5). All of the illustrated specimens are longitudinal sections. In comparison to Atsabella, G. spiroloculiformis has chambers that are of more uniform width throughout length and a more fusiform test. The shape of the aperture in G. spiroloculiformis was not described by Oraveczné Scheffer and is unclear in the figures. However, the paratype figured in Oraveczné Scheffer (1971: pl. 2, fig. 2) appears to have a long neck on the foramen beneath the final aperture, reminiscent of the genus Karaburunia Langer, 1968. Atsabella bandeiraensis nov. sp. Plate 1, Figs. 1–13 Holotype: CPC41334, from A7012a, shallow normal marine to metahaline lagoonal facies, probably Carnian or Norian (Table 1). Other material: 38 specimens from the type locality, preserved as silicified casts; and a few specimens, preserved as original calcite, from several other localities (Table 2); from lagoonal and basinal facies; stratigraphic range uncertain, possible Middle Triassic, Carnian, possible Norian (Table 1). Etymology: The species is named after Bandeira Gorge in the Atsabe Subdistrict, Timor Leste, where the type locality occurs. Diagnosis: See diagnosis of genus (defined by monotypy). Description: The following characters are in addition to those used to define the genus. Variation in the longitudinal profile is illustrated by comparing the broadly ovoid holotype (Plate 1, Fig. 1) with the more fusiform paratype illustrated in Plate 1, Fig. 7. The test is small (Fig. 3) with maximum length ranging from about 0.19 mm to about 0.48 mm and maximum width from about 0.13 mm to about 0.33 mm. The holotype lies within the mid-range of length and toward the smaller end of the width range (Fig. 3). Sutures between whorls and between chambers within the whorl are usually flush. The external surface is smooth. The narrow arched aperture located above the peripheral margin of the test is a constant feature of all Timor specimens, although the height of the aperture varies (e.g., compare Plate 1, Fig. 4 with Plate 1, Fig. 6). Remarks: Specimens from the Locker Shale (upper Triplexisporites playfordii Zone; Spathian to Lower Anisian) of the Northern Carnarvon Basin, Western Australia identified by Apthorpe (2003: p. 11, pl. 2, figs. 8–16) as Gsollbergella spiroloculiformis probably belong to Atsabe bandeiraensis. The stratigraphic range of A. bandeiraensis may therefore be from at least the Spathian or Lower Anisian to the Carnian or Norian. Genus Karaburunia Langer, 1968 Type species: Karaburunia rendeli Langer, 1968; Middle Triassic (Anisian), Turkey. Karaburunia atsabensis nov. sp. Plate 1, Figs. 14–18; Plate 2, Figs. 1–32 Holotype: CPC41343, from A7012a, shallow normal marine to metahaline lagoonal facies, probably Carnian or Norian (Table 1).

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Plate 1. Figs. 1–13. Atsabella bandeiraensis nov. sp.; from A7012A, Carnian or Norian. Figs. 1 and 2. Holotype, longitudinal and facial (apertural) secondary-electron images, CPC41334. Figs. 3 and 4. Longitudinal and facial (apertural) secondary-electron images, CPC41335. Figs. 5 and 6. Longitudinal and facial (apertural) secondary-electron images, CPC41336. Figs. 7 and 8. Longitudinal and facial (apertural) secondary-electron images, CPC41337. Fig. 9. Longitudinal secondaryelectron image showing thin wall, CPC41338. Fig. 10. Longitudinal view under glycerine in transmitted light, CPC41339. Fig. 11. Longitudinal view under glycerine

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Table 2 Distribution of ophthalmidiids in studied samples. Samples

A7012a

C659a

M625a

M625b

M644a

S6101a

S6103a

Preparation type Palynology Conodont age

ar

ar C-F Middle Tr

wr

wr

ar

ar

ar

Late Carn.

Carnian

Miliolida species Atsabella bandeiraensis nov. sp. Karaburunia atsabensis nov. sp. Ophthalmidium sp. cf. O. primitivum Ho, 1976 Spirophthalmidium grunaui nov. sp. Other biogenic groups Conodonts Fish/shark teeth Foraminifera – hyaline types Foraminifera - organic-cemented agglutinates Mollusc debris Echinoderm debris Ostracods Radiolaria Sponge spicules

C A R

C

C

R R

C C R

R A

R

R

R R

C R

A

C

A C A

C C C

R R C A

R C R

R A

R C R

C

A: adundant; C: common; R: rare; ar: acetic acid digestion residue; wr: residue of rock boiled with detergent in water and washed over 150 ␮m and 63 ␮m sieves; C: S. quadrifidus Zone; D: S. speciosus Zone; F: A. reducta Zone; zones follow Helby et al. (1987); identified by J. Backhouse.

maximum width (mm)

0.35

0.30

0.25

0.20

0.15

assemblage from A7012a assemblage from M625a

holotype 0.10 0.10

0.15

0.20

0.25

0.30

0.35

0.40

0.45

0.50

0.55

maximum length (mm) Fig. 3. Maximum width versus maximum length measurements for Atsabella bandeiraensis nov. sp.

Other material: Over 100 specimens from the type locality, preserved as silicified casts; and a few specimens from several other localities (Table 2); from lagoonal and basinal facies; stratigraphic range uncertain, possible Middle Triassic, Carnian, possible Norian (Table 1). Etymology: The species is named after the Atsabe Subdistrict in which the type locality occurs. Diagnosis: Species of Karaburunia with small fusiform test (< 0. 5 mm in maximum length); post-embryonic stage with variable milioline coiling ranging from quinqueloculine

with a triangular cross-section to almost spiroloculine with an ovoid cross-section; chamber lumen of almost constant width throughout length; flush sutures; thick wall; prominent narrow cylindrical apertural neck of variable length, with narrow everted lip around aperture. Description: Test free, small (< 0.5 mm long); fusiform outline with variable length to width ratio (Fig. 4); peripheral margin broadly rounded; cross-sectional shape varying from almost triangular (Plate 2, Fig. 30) to elongate ovoid (Plate 2, Fig. 4); coiling milioline, varying from almost quinqueloculine to almost

Plate 1 (Continued). in transmitted light, CPC41340. Fig. 12. Longitudinal view under glycerine in transmitted light, CPC41341. Fig. 13. Longitudinal view under glycerine in transmitted light, CPC41342. Figs. 14–18. Karaburunia atsabensis nov. sp. From A7012A, Carnian or Norian. Figs. 14 and 15. Holotype; longitudinal and facial (apertural) secondary-electron images, CPC41343. Fig. 16. Longitudinal view under glycerine in transmitted light, CPC41344. Fig. 17. Longitudinal view under glycerine in transmitted light, CPC41345. Fig. 18. Longitudinal view under glycerine in transmitted light, CPC41346. Scale bars = 0.1 mm.

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Plate 2. Figs. 1–32. Karaburunia atsabensis nov. sp. showing variability in test profiles. Secondary-electron images. From A7012A, Carnian or Norian. Figs. 1 and 2. Longitudinal and facial (apertural) secondary-electron images, CPC41347. Figs. 3 and 4. Longitudinal and facial (apertural) secondary-electron images, CPC41348. Figs. 5 and 6. Longitudinal and facial (apertural) images, CPC41349. Figs. 7 and 8. Longitudinal and facial (apertural) images, CPC41350. Figs. 9 and 10. Longitudinal and facial (apertural) images, CPC41351. Figs. 11 and 12. Longitudinal and facial (apertural) images, CPC41352. Figs. 13 and 14. Longitudinal and facial (apertural) images, CPC41353. Figs. 15 and 16. Longitudinal and facial (apertural) images, CPC41354. Figs. 17 and 18. Longitudinal and facial (apertural) images, CPC41355. Figs. 19 and 20. Longitudinal and facial (apertural) images, CPC41356. Figs. 21 and 22. Longitudinal and facial (apertural) images, CPC41357. Figs. 23 and 24. Longitudinal and facial (apertural) images, CPC41358. Figs. 25 and 26. Longitudinal and facial (apertural) images, CPC41359. Figs. 27 and 28. Longitudinal and facial (apertural) images, CPC41360. Figs. 29 and 30. Longitudinal and facial (apertural) images, CPC41361. Figs. 31 and 32. Longitudinal and facial (apertural) images, CPC41362. Scale bar = 0.1 mm.

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0.30

maximum width (mm)

holotype of Quinqueloculina nucleiformis Kristan-Tollman

holotype of Karaburunia rendeli Langer

0.25

range of Palaeomiliolina tibetica Ho

0.20 holotype of Sigmoilina? triadica Langer 0.15 holotype of Karaburunia atsabiensis nov. sp.

0.10 0.20

0.25

0.30

0.35

0.40

0.45

0.50

0.55

range of Palaeomiliolina tenuis Ho 0.60

0.65

0.70

maximum length (mm) Fig. 4. Maximum width versus maximum length measurements for Karaburunia atsabensis nov. sp. Assemblage from A7012A. Dimensions of the holotypes of other species that may belong within Karaburunia are shown for comparison.

spiroloculine; early whorl obscure but initial chamber arrangement more irregular than later chambers (Plate 1, Figs. 16–18) that in most specimens are added at an angle of about 180◦ ; chamber lumen a narrow tube that changes little in width along length; sutures flush; wall thick with smooth surface although surface ultrastructure not certain because of preservation; aperture at end of conspicuous narrow cylindrical apertural neck of variable length, surrounded by a narrow everted lip. Comparisons: In terms of test length versus width measurements, the holotypes of Quinqueloculina nucleiformis Kristan-Tollmann (1964: p. 61, pl. 9, figs. 9–11), Karaburunia rendeli Langer (1968: p. 592, pl. 1, figs. 6–8) and Sigmoilina? triadica Langer (1968, p. 592, pl. 1, figs. 9–11) fall well outside of the range of the type assemblage of K. atsabensis (Fig. 4). In terms of size, Palaeomiliolina tenuis Ho (in Ho et al., 1976: p. 24, pl. 24, figs. 29–31c) and P. tibetica Ho (in Ho et al., 1976: p. 24, pl. 24, figs. 28a and b, 32a–33b) are mostly larger than K. atsabensis and have more incised sutures than the Timor species. The holotype of Spiroophthalmidium fusiformis Trifonova (refigured by Trifonova, 1993: pl. 10, fig. 13) and the range of test dimensions for this species recorded by Trifonova (1993: p. 54) lie within the equivalent range for K. atsabensis. Although Trifonova (1993: pl. 10, figs. 13–16) included a broad range of test shapes in S. fusiformis (which she transferred to Ophthalmidium), similar to that displayed in K. atsabensis, the apertural neck seems much more variable in width and length than in the Timor species. Genus Ophthalmidium Kübler and Zwingli, 1870 Type species: Oculina liasica Kübler and Zwingli, 1866; Lower Jurassic (Toarcian), Switzerland. Ophthalmidium sp. cf. O. primitivum Ho in Ho et al., 1976 Plate 3, Figs. 1–4 cf. 1976. Ophthalmidium primitivum Ho in Ho et al., pl. 23, 24, pl. 24, figs. 21, 22; Triassic (Upper Tulong Formation), Tibet.

Material: 10 specimens. Remarks: This species has a small test (Fig. 5) with almost circular equatorial outline and about six narrow tubular chambers in post-embryonic stage, each between one-half-whorl and one-whorl in length. Chamber divisions are marked by a slight kink and constriction in the tube. The aperture is at the end of a short neck that arises, in the plane of coiling, abruptly at right angles from the equatorial margin. In Timor specimens the aperture is semicircular and has a narrow everted lip. Ho (in Ho et al., 1976) did not describe or illustrate the shape of the aperture in O. primitivum. The species has a greater number of whorls with narrower chamber lumens than in the holotype of O. liasicum (following the reillustration and redescription of the type specimen by Wood, 1946: p. 461, pl. 29, fig. a, pl. 30, fig. 1). It also has a distinct apertural neck that is not apparent in the type specimen of O. liasicum. O. orbiculare Burbach (1886: p. 499, pl. 5, figs. 3–6) from the Early Jurassic of Germany does not have the abrupt 90◦ swing of the short apertural neck away from the equatorial contour. O. exiguum Koehn-Zaninetti (1969: p. 64–67, pl. 6, fig. D, fig. 14A–G) has a more fusiform equatorial outline than present in O. primitivum. Praeophthalmidium tricki Langer (1968: p. 591, pl. 1, figs. 1–5) has a similar equatorial profile and apertural neck to O. primitivum but differs in thick wall layers covering the central part of the test forming a lenticular axial profile. Genus Spirophthalmidium Cushman, 1927 Type species: Spiroloculina acutimargo Brady, 1884; Holocene, Atlantic and Pacific Oceans. Spirophthalmidium grunaui nov. sp. Plate 3, Figs. 5–13 Holotype: CPC41366 from M644a, shallow-marine basinal carbonate-rich mud facies, late Carnian (Table 1). Other material: 30 topotypes.

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Plate 3. Figs. 1–4. Ophthalmidium sp. cf. O. primitivum Ho, 1976. From A7012A, Carnian or Norian. Secondary electron images. Fig. 1. Equatorial view under glycerine in transmitted light, CPC41363. Fig. 2. Peripheral view showing aperture, secondary electron image, CPC41363. Fig. 3. Equatorial view under glycerine in transmitted light, CPC41364. Fig. 4. Equatorial view under glycerine in transmitted light, CPC41365. Figs. 5–13. Spirophthalmidium grunaui nov. sp., from M644a, late Carnian. Fig. 5. Holotype, equatorial (lateral) view under glycerine in transmitted light, CPC41366. Fig. 6. Holotype, peripheral view, secondary-electron image,

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maximum width (mm)

0.35

0.3

0.25

holotype 0.2 0.2

0.25

0.3

0.35

0.4

0.45

0.5

maximum length (mm) Fig. 5. Maximum width versus maximum length measurements for Ophthalmidium sp. cf. O. primitivum. Assemblage from A7012A. Dimensions of the holotype (Ho et al., 1976: pl. 24, fig. 21) of O. primitivum are shown for comparison.

Etymology: The species is named in honour of Hans R. Grunau who wrote some of the pioneering papers on the geology of Timor Leste. Diagnosis: A medium size species of Spirophthalmidium, very compressed, with narrow chambers which decrease very gradually in width along length toward aperture; periphery truncate with rounded edges; about five whorls developed with about eight chambers in post-embryonic stage; sutures flush; wall smooth; aperture a very small opening at end of long neck, without lip. Description: Test free, of medium size (< 0.9 mm in maximum length, Fig. 6), very compressed with narrow truncated margin with rounded corners (Plate 3, Fig. 6); equatorial outline usually elongate fusiform with narrowly drawn out apertural end and an often constricted opposite end although irregular shaped tests are present (e.g., Plate 3, Fig. 12); post-embryonic chamber arrangement spiroloculine with chamber divisions marked by a slightly constricted kink in the narrow chamber tube; about eight chambers developed in post-embryonic stage; chamber lumen increases very gradually in maximum width with growth but each chamber narrows gradually towards the aperture; sutures flush; wall thick, with smooth surface (although ultrastructure uncertain due to preservation); aperture a very small opening at end of the chamber tube that extends past previous chamber outline as a very long neck lacking a lip. Comparisons: Spirophthalmidium grunaui has more whorls and narrower chambers than S. triadicum Kristan (1957: p. 290, pl. 25, figs. 2a–4) from the Austrian Rhaetian. It differs from S. macfadyeni (Wood and Barnard, 1946: p. 92, 101, pl. 9, figs. a–g) from the Early Jurassic of England in having narrower chambers that do not show such rapid decrease in diameter along chamber length. A similar distinction is made with

S. northamptonensis Wood and Barnard (1946: p. 88, 95, 103, pl. 5, figs. a–n, pl. 6, figs. a–k, pl. 7, figs a–o, pl. 8, figs. a–i) also from the English Early Jurassic. 5. Discussion The large suites of free specimens available for this study show great variation in chamber arrangement within the interpreted species with, for example, tests assigned to K. atsabensis ranging from almost spiroloculine to quinqueloculine. In previous studies this range of variation in Triassic ophthalmidiids has been classified among several genera. Chamber shape, including the internal shape of the chamber lumen, apertural shape and test size are stable features within the species recognized here, following a pattern observed in modern miliolids by Haig (1988) and Parker (2009). The importance of wall-surface ultrastructure for taxonomic differentiation could not be assessed because of the preservation of the studied material. All of the studied species lack ornament. Several problems exist in placing the Timor fauna in the context of Triassic ophthalmidiid evolution and in making biogeographic evaluations both regionally and globally. Most ophthalmidiids recorded elsewhere from Triassic deposits have been identified from thin sections and categorised within narrow morphological limits based mainly on particular chamber arrangements, chamber shapes as seen in section and size. However, the three-dimensional shape of chambers and the shape of the aperture are difficult to interpret from thin sections. It is therefore difficult to compare species established through the study of thin sections with those recognized here. The lack of precise age determinations for most of the Timor occurrences is another problem for the comparison of faunas.

Plate 3 (Continued) holotype, CPC41366. Fig. 7. Equatorial (lateral) view under glycerine in transmitted light, CPC41367. Fig. 8. Equatorial (lateral) view under glycerine in transmitted light, CPC41368. Fig. 9. Equatorial (lateral) view under glycerine in transmitted light, CPC41369. Fig. 10. Equatorial (lateral) view under glycerine in transmitted light, CPC41370. Fig. 11. Equatorial (lateral) view under glycerine in transmitted light, CPC41371. Fig. 12. Equatorial (lateral) view under glycerine in transmitted light, CPC41372. Fig. 13. Equatorial (lateral) view under glycerine in transmitted light, CPC41373. Scale bars = 0.1 mm.

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maximum width (mm)

0.6

0.5

0.4 holotype

0.3

0.2 0.4

0.5

0.6

0.7

0.8

0.9

maximum length (mm) Fig. 6. Maximum width versus maximum length measurements for Spirophthalmidium grunaui nov. sp. Assemblage from M644a.

None of the ophthalmidiid species identified in Timor has a stratigraphic range that can be assigned unequivocally to a narrow stratigraphic interval. Several of the Timor samples containing ophthalmidiids have yielded rare conodonts (Table 2), the major index fossil group for the Triassic, but other samples are barren of these microfossils. Palynomorphs have been extracted from one of the samples and are tentatively tied to the northwest Australian spore-pollen zonation (Table 2) that includes broad zones indirectly correlated to the international stages (Helby et al., 1987). The conodont evidence presented in Table 1 suggests that A. bandeiraensis, K. atsabensis and S. grunaui are present in the Carnian of Timor. Based on palynological and foraminiferal evidence in underlying and overlying units (Table 1), a Carnian or Norian age is suggested for the occurrence of A. bandeiraensis, K. atsabensis and O. sp. cf. O. primitivum in sample A7012a. The ophthalmidiids seem to have a wide facies distribution in muddy limestones in Timor. Atsabella bandeiraensis, K. atsabensis and O. sp. cf. O. primitivum appear to be most abundant in the normal-marine to metahaline lagoonal facies interpreted for sample A7012a (Table 1) with A. bandeiraensis and K. atsabensis also common to rare in neritic basinal carbonate muds. Spirophthalmidium grunaui has only been found in the latter facies. 6. Conclusions Opthalmidiids form a small part of the Upper Triassic foraminiferal fauna found in basinal and lagoonal muds in Timor. Assemblages of free specimens extracted from the muds provide ideal material for study of intraspecific variation. Four species have been described, each belonging to a different genus: Atsabella nov. gen., Karaburunia, Ophthalmidium and Spirophthalmidium. Chamber arrangement is particularly variable and in Karaburunia the milioline coiling ranges from

almost spiroloculine to quinqueloculine. Stable features for taxonomic discrimination at species level are chamber shape, including the internal shape of the chamber lumen, apertural shape and test size. Acknowledgements Our work has been supported by funding from Eni Australia and Eni Timor Leste, the University of Western Australia and Geoscience Australia. We thank Myra Keep for leading the University of Western Australia geological research efforts in Timor Leste and Alfredo Pires the Secretaria de Estado dos Recursos Naturis for Timor Leste, Francisco da Costa Monteiro and his staff of SERN, Gualdino da Silva and his staff at the Timor Leste Authoridade Nacional do Petróleo and Noberta Soares da Costa and her staff of the Timor Leste Direcc¸ão Nacional de Geologia e Recursos Minerais for their support of our work. The study would not have been possible without the field assistance provided by many of our Timorese friends. Mike Orchard and Bob Nicoll are thanked for advising us on the conodont age determinations and we are grateful to John Backhouse who contributed the palynological zonation. We thank Thomas Ilhe and Julien Bourget for assistance with translation of the abstract. Roberto Rettori, Taniel Danelian and an anonymous reviewer are thanked for making valuable comments on the original manuscript. The electron microscopy was undertaken by D. Haig using facilities at the UWA Centre for Microscopy, Characterisation and Analysis. References Al-Shaibani, S.K., Carter, D.J., Zaninetti, L., 1983. Geological and micropaleontological investigations in the Upper Triassic (Asinepe Limestone) of Seram, Outer Banda Arc, Indonesia. Archives des Sciences, Geneva 36, 297–313.

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