Taxonomic review and evolutionary trends of Levipustulini and Absenticostini (Brachiopoda) from Argentina: Palaeobiogeographic and palaeoclimatic implicationsARTURO C. TABOADA & GUANG R. SHI

THE LEVIPUSTULINI in Argentina are represented by a great number of genera, including Bulahdelia Roberts (Taboada 1989), Levipustula Maxwell (Amos et al. 1963), Lanipustula Klets (Simanauskas 1996), Verchojania Abramov (Taboada 2008), Jakutoproductus Kaschirzev (Simanauskas & Archbold 2002; Taboada & Pagani 2010) and Piatnitzkya Taboada (1993). In this paper, we carry out a detailed taxonomic review of this tribe and discuss some of its key genera and species, such as Levipustula levis Maxwell and Lanipustula patagoniensis Simanauskas. In addition, we also report and document one new species (Lanipustula kletsi sp. nov.), as well as the first Southern Hemispheric record of Absenticosta Lazarev, 1991 in a related tribe Absenticostini. Absenticosta bruntoneileenae sp. nov. is suggested as a possible ancestral stock of the Patagonian Levipustulini

descendants. The development of similar phylogenetic lineages of Levipustulini in high latitude regions of both northern and southern hemispheres such as Siberia in Northeast Asia and Patagonia in southwestern Gondwana, could have been a consequence of parallel evolution. The progressive palaeobiogeographic isolation of Patagonia, coupled with its southward drift under cold palaeoclimatic conditions, during middle Carboniferous-early Permian is interpreted to have triggered the Levipustulini vicariance.

GEOLOGY AND BIOSTRATIGRAPHYArgentine material described here comes from two areas, the Uspallata-Iglesia Basin in western Argentina and the Tepuel-Genoa Basin in Patagonia; the two regions are currently separated by 1300 km (Fig. 1).

TABOADA, A.C. & SHI, G.R., 2011:07:29. Taxonomic review and evolutionary trends of Levipustulini and Absenticostini (Brachiopoda) from Argentina: palaeobiogeographic and palaeoclimatic implications. Memoirs of the Association of Australasian Palaeontologists 41, 87-114. ISSN 0810-8889.

The diagnosis and composition of the brachiopod Tribe Levipustulini Lazarev, 1985 is reviewed, leading to a detailed revision of the genera Levipustula Maxwell, 1951 and Lanipustula Klets, 1983, as well as a review of previous records of the species Levipustula levis Maxwell from Australia and Argentina. The presence of Lanipustula patagoniensis Simanauskas in Patagonia is confirmed with additional topotypic material described and illustrated. Based on this review, we reassign Levipustula levis from New South Wales, Australia to Lanipustula. Two new species, Lanipustula kletsi from the middle Pennsylvanian of Patagonia and the Absenticostinin Absenticosta bruntoneileenae from the latest Viséan of western Argentina, are proposed. Abstenticosta bruntoneileenae is suggested as a possible ancestral stock of the Patagonian Levipustulini through the lineage Lanipustula-Verchojania-Jakutoproductus-Piatnitzkya (Serpukhovian-middle Artinskian). The development of similar phylogenetic lineages of Levipustulini in high latitude regions of both northern and southern hemispheres (such as Siberia in Northeast Asia and Patagonia in southwestern Gondwana) is here interpreted as a consequence of parallel evolution. The progressive palaeobiogeographic isolation of Patagonia from mainland South America, coupled with its southward drift under cold palaeoclimatic conditions during middle Carboniferous-earliest Permian times, is proposed to have triggered the Levipustulini vicariance.

Arturo César Taboada (ataboada@unpata.edu.ar), CONICET-Laboratorio de Investigaciones en Evolución y Biodiversidad (LIEB), Facultad de Ciencias Naturales, Sede Esquel, Universidad Nacional de la Patagonia ‘San Juan Bosco’, Chubut, Argentina; Guang R. Shi (grshi@deakin.edu.au), School of Life and Environmental Sciences, Deakin University, Melbourne Campus at Burwood, 221 Burwood Highway, Burwood, Victoria 3125, Australia. Received 1 March 2011.

Keywords: Argentina, Brachiopoda, Levipustulini, late Palaeozoic, palaeobiogeography, parallel evolution.

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The Uspallata-Iglesia BasinThe Uspallata-Iglesia Basin (González 1985) was a narrow backarc area of deposition located between 28-34ºS (present-day latitudes), to the west of the ancient Protoprecordillera highland (Amos & Rolleri 1965). This basin has been traditionally subdivided into two different basins/subbasins, the Río Blanco to the north and the Calingasta-Uspallata to the south (Amos 1964). The Uspallata-Iglesia Basin presents a complicated stratigraphy due in part to the strong tectonism that affected most of the Late Palaeozoic deposits, now cropping out

discontinuously. Selected lithostratigraphic units for the entire Uspallata-Iglesia Basin and their lateral correlations based on key fossil content are shown in Figure 2, as are stratigraphic data on major glacial pulses.

In the Calingasta-Uspallata subbasin, Levipustulini and Absenticostini, among other brachiopods, have been recorded from the El Paso and Hoyada Verde members of the San Eduardo Formation (Mésigos 1953; nom. transl. González 1993; emend. Taboada 1997), in a deduced superpositional succession at the Barreal Hill (Fig. 3). Absenticosta bruntoneileenae sp.

Figure 1. Map of southern South America showing major palaeogeographic features and location of basins where Argentinean Levipustulini and Absenticostini were recorded (modified from Limarino & Spalleti 2006).

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nov. occurs together with the Rugosochonetes-Bulahdelia faunal assemblage from the El Paso Member, estimated to be late Viséan-earliest Serpukhovian in age (Taboada 1989, 2010). Levipustula levis and its faunal assemblage

were recorded in the Hoyada Verde Member and other lateral equivalents within the Calingasta-Uspallata subbasin, with an age not younger than Serpukhovian for its uppermost occurrence (Taboada 2010).

The Tepuel-Genoa BasinThe Tepuel-Genoa Basin is located in central-west Patagonia, between 43º-44º20´S and 69º30´-71ºW, flanked by the Somúncura and Deseado massifs to the northeast and southeast, respectively (Fig. 1). The basin infill is a nearly 6000 m thick, almost continuous sequence of clastic deposits ranging in age from early Carboniferous to early Permian (Suero 1948, 1953, 1958; Andreis et al. 1987; Archangelsky et al. 1996). The current upper Palaeozoic stratigraphic scheme of the basin, accepted by most Argentinean workers, was proposed by Page et al. (1984), with a tripartite succession of the Jaramillo, Pampa de Tepuel and Mojón de Hierro formations constituting the Tepuel Group, with its type section well exposed in the Tepuel Hill area. Other stratigraphic units are listed in the literature, but they are locally defined and generally regarded as lateral equivalents to parts of the Tepuel Group; these include the Las Salinas Formation (González 1972) in the Languiñeo area and Esquel and Valle Chico formations (Cucchi 1980) in the Esquel area. South of present-day 44ºS, the uppermost sections of the Upper Palaeozoic were reported by Suero (1953, 1958) and Ugarte (1966), and later recognised as a separate unit, named the Río Genoa Formation (Andreis & Cúneo 1989; Taboada & Pagani 2010). The stratigrafic column of the Tepuel Group with its main lithostratigraphic units and major glacial-related intervals and Levipustulini-bearing horizons, are shown in Figure 4. The stratigraphic and geographic location of the Lanipustula species described here from the Pampa de Tepuel and Las Salinas formations, are indicated in Figure 5.

SYSTEMATIC PALAEONTOLOGY

Order PRODUCTIDA Sarytcheva & Sokolskaya, 1959

Suborder PRODUCTIDINA Waagen, 1883Superfamily PRODUCTELLOIDEA Schuchert,

1929Family OVERTONIIDAE Muir-Wood & Cooper,

1960Subfamily PLICATIFERINAE Muir-Wood &

Cooper, 1960Tribe LEVIPUSTULINI Lazarev, 1985

Discussion. The Tribe Levipustulini Lazarev, 1985 (nom. transl. Brunton et al. 1995; ex

Figure 2. Time-space scheme showing selected lithostratigraphic units and location of glacial-related intervals of the Uspallata-Iglesia Basin. Stages and stratigraphic units of higher hierarchy are without scale, and the Permian timescale is based on Gradstein et al. (2004) and Gradstein & Ogg (2009) (note radiometric ages are not to scale). Biozones/faunas: M, Malimanian fauna; R-B, Rugosochonetes-Bulahdelia; L, Levipustula; M-M, Marginovatia-Maemia; T-S, Tivertonia-Streptorhynchus; C, Costa-tumulus. Black flag: barren intervals (Waterhouse, 2008). Other symbols include: ∆∆∆ - diamictites, and =O= - associated facies of glacial-related horizons. Cross-hatching delineates orogenic phases: Río Blanco orogenic phase (middle Viséan?), San Eduardo epeirogenic phase (middle Bashkirian) and San Rafael orogenic phase (late Early Artinskian) (modified from Taboada 2010).

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Levipustulinae Lazarev, 1985) is characterised by a compact set of genera comprising Levipustula Maxwell, 1951, Jakutoproductus Kaschirzev, 1959, Verchojania Abramov, 1970, Bulahdelia Roberts (in Roberts et al. 1976), Lanipustula Klets, 1983, Impiacus Lazarev & Suursuren (in Afanasjeva et al. 1988) (Lazarev 1985, 1990) and Piatnitzkya Taboada, 1993 (Brunton et al. 2000). All of these genera are currently ascribed to the Tribe by most authors. Nevertheless, the inclusion

of Jakutoproductus in the Levipustulinae was rejected by Shi (1994), who reassigned it to the subfamily Plicatiferinae Muir-Wood & Cooper, 1960 (see also Shi & Waterhouse 1996; Manankov 2004) because of its closer relationship to Plicatifera Chao, 1927 than to Levipustula. On the other hand, Jakutoproductus was retained in Levipustulini by subsequent authors (Brunton & Lazarev 1997; Brunton et al. 2000; Taboada & Pagani 2010; Simanauskas & Archbold 2002). The

Figure 3. A, Geographic location of main localities of the Calingasta-Uspallata and Río Blanco subbasins (modified from González 1985). B, Composite stratigraphic column of the San Eduardo Formation with the El Paso and Hoyada Verde members at Barreal Hill, showing location of faunal assemblages, Absenticosta bruntoneileenae sp. nov. and Levipustula levis occurrences, and glacial-related intervals recognised (modified from Taboada 1997).

Figure 4 (opposite). Time-space scheme showing the stratigrafic column of the Tepuel Group with its main lithostratigraphic units, recognised mayor glacial-related horizons throughout the sequence (based on Suero 1948; Taboada 2008; Taboada & Pagani 2010) and stratigraphic position of Patagonian Levipustulini records (B-H). The Carboniferous-Permian timescale is based on Gradstein et al. (2004) and Gradstein & Ogg (2009) (but note the radiometric ages are not drawn to scale). Symbols for diamictites and associated facies of glacial-related horizons are as in Figure 2. Symbols for brachiopod species: A, Absenticosta bruntoeileenae sp. nov. (FML-PI 1201-1). B, Lanipustula patagoniensis Simanauskas (MPEF-PI 370). C, Lanipustula kletsi sp. nov.(FML-PI 1702-2). D, Verchojania inacayali Taboada (FML-PI 3988). E, Verchojania archboldi Taboada (LIEB-PI 4). F, Jakutoproductus sabattiniae Taboada & Pagani (MPEF-PI 1807). G, Jakutoproductus australis Simanauskas & Archbold (DG-MLP 26170). H, Piatnitzkya borrelloi Taboada (FML-PI 766a) (all specimens in natural size; scale bar: 10mm).

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latter study, by Simanauskas & Archbold (2002), emphasised the close links in dorsal internal structures between genera such as Levipustula, Lanipustula, Verchojania, Jakutoproductus and Piatnitzkya. In addition, Simanauskas & Archbold (2002) have also indicated that the strong rugae observed in Jakutoproductus are only strongly developed in a minority of species, whereas in Plicatifera the strong regular rugae are also expressed on the shell interior, unlike those of Jakutoproductus. More recently, Jakutoproductus has been considered as a member of the Levipustulini lineage by Shi et al. (2008).

According to Lazarev (1990), Brunton et al. (1995), Brunton & Lazarev, (1997) and Waterhouse (2001, 2002), the most important diagnostic features of most genera of Levipustulini appear to be the presence of spines on both valves tending to a quincunxial arrangement, with elongate or slightly swollen bases, but in Permian representatives the dorsal spines are usually absent. The distinctive spine pattern of the tribe is usually also matched by a subdued concentric ornament, mostly growth lines, weak rugae or lamellae, but again this concentric ornamentation may be lacking in some Permian genera. In addition, dorsal valves are commonly geniculate, the internal marginal structures are weak (i.e., low) or absent, and a shallow to moderate corpus cavity is present.

According to this characterisation, a similar set of genera can be easily recognised and assigned to the Tribe Levipustulini, here considered to comprise Bulahdelia, Impiacus, Lanipustula, Levipustula, Verchojania, Jakutoproductus and Piatnitzkya. The collective stratigraphic range for this tribe is from Late Viséan to Artinskian.

The Levipustulini was separated from the Plicatiferini by Brunton et al. (1995), Brunton & Lazarev (1997), Brunton et al. (2000), a view also accepted by Waterhouse (2001, 2002) and Simanauskas & Archbold (2002). Following Brunton et al. (1995), Levipustulini was placed within the Subfamily Plicatiferinae by a number of authors (Brunton & Lazarev 1997; Brunton et al. 2000; Simanauskas & Archbold 2002; Taboada 2008), but Waterhouse (2002), in an alternative classification of Productidina, assigned Levipustulini to the Subfamily Tubersulculinae (Family Avoniidae) in recognition of its well developed and uniform ventral spines. On the

other hand, Waterhouse’s (2002) scheme for suprafamilial position of Plicatiferinae was followed by Taboada (2008) and is also followed here.

Levipustula Maxwell, 1951

Type species. Levipustula levis Maxwell, 1951 from the Serpukhovian-Bashkirian? of Queensland, Australia.

Discussion. The type material of Levipustula levis came from the Neerkol beds of the Rockhampton district in Queensland, eastern Australia, and was described in detail by Maxwell (1951). With the exception of sample F11913b, our examination of the available type material included almost all specimens illustrated by Maxwell (1951), plus a few other specimens such as sample F11913a from the locality in Portion 200, Parish of Palen, and unlabelled samples without provenance, all of which are refigured here for comparison with Argentine material (Figs 2-5).

After Maxwell’s proposal of Levipustula, varied interpretations of the genus have been offered by different authors, based on a wide range of material from both eastern Australia and Argentina, resulting in a significant expansion of Maxwell’s original definition of the genus (Muir-Wood & Cooper 1960; Amos 1960; Campbell 1961; Böger & Fiebig 1963; Muir-Wood 1965; Kotljar & Popeko 1967; Winkler Prins 1968; Shi 1994; Shi & Waterhouse 1996; Brunton et al. 2000; Simanauskas & Archbold 2002). One diagnostic character not explicitly mentioned in the original description of Levipustula was the presence of dorsal spines, which could not be easily seen in the only dorsal external mould among the original specimens examined by Maxwell (1951). This dorsal exterior was illustrated as a plasticine cast by Maxwell (1951, pl. 2, fig. 4b). The lack of a mention of dorsal spines in the detailed description offered by Maxwell (1951) subsequently led some authors such as Muir-Wood & Cooper (1960), Muir-Wood (1965) and Waterhouse (1982), to believe that Levipustula either lacks dorsal spines or at most only has scarce dorsal spines. Brunton et al. (2000, fig. 299.3a) illustrated this dorsal valve external mould for the first time as part of the holotype specimen labelled F11900 (the

Figure 5. A, Simplified stratigraphic column of the Pampa de Tepuel Formation (lower section) as exposed at the western flank of the Tepuel Hill, showing location of fosiliferous beds, Lanipustula patagoniensis and glacial-related intervals (not to scale) recognised (adopted from Suero, 1948 and authors’ own observations). B, Simplified stratigraphic column of the Las Salinas Formation at the Languiñeo Hill, showing location of fosiliferous beds, Lanipustula patagoniensis, Lanipustula kletsi sp. nov. and glacial-related intervals (not to scale)(modified from González, 1972 and author’s own observations). C, Location map showing main localities of the Tepuel-Genoa Basin mentioned in the text.

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holotype also contains a ventral valve figured by Brunton et al. 2000, fig. 299.3b). It is from this dorsal valve external mould that Maxwell made the plasticine cast which he figured (Maxwell 1951, pl. II, fig. 4b). Our careful search of Maxwell’s collection indicates that the cast replica figured by Maxwell is actually missing from the collection, presumably lost. However, the holotype (specimen F11900) is present in the collection, which is an articulate internal mould with aspects of both dorsal and ventral valves. This specimen corresponds with Maxwell’s figures 4c and 4e of the plasticine cast. However, the external moulds of this specimen, figured by Maxwell (1951, figs. 4a and 4b) and Brunton et al. (2000, fig. 299.3a), are missing from the collection. Also of note is that the ventral valve external mould figured by Brunton et al. (2000, fig. 299.3b), according to our examination of the collection, actually does not belong to the holotype because its correct label is F11908, and it comes from a different locality to the true holotype specimen F11900 (see Maxwell 1951 for locality details).

The dorsal valve external mould figured by Brunton et al. (2000, fig. 299.3a) has indications of a few minute hollows near the hinge margin that probably correspond to small dorsal spines; but very likely dorsal spines are completely absent on the visceral disk. To clarify the nature of dorsal spines, we have also examined the latex cast of Levipustula levis from locality 145-5 of Ridgeland, northwest of Rockhampton, which were sent by Dr. John Roberts to Dr. Carlos González in 2000, now housed in the Palaeontological Institute of the Miguel Lillo Foundation (Tucumán). These latex casts, here figured (Fig. 4), include three well preserved dorsal valve exteriors, which do not exhibit any minute dorsal spines. In contrary, specimens assigned to L. levis by Campbell (1961) from Booral, New South Wales, about 1000 km south of the Rockhampton district, show densely arranged fine dorsal spines.

Therefore, these diagnostic dorsal spines, being absent in the type material of Levipustula, would exclude from the genus, all the specimens from NSW ascribed by Campbell (1961, 1962) to Levipustula. This also applies to the specimens from New South Wales listed as L. levis by Voisey (1940), as well as those illustrated by Muir-Wood & Cooper (1960) (see also Amos 1960, pl. 3, fig. 1), Muir-Wood (1965), McCarthy et al. (1974) and Brunton et al. (2000, only fig. 299.3c). The correct assignment of theses species to other genera will be discussed below.

According to the diagnosis and description by Maxwell (1951) and our own examination of

the type and complementary material mentioned above from Queensland, Levipustula is here diagnosed as follows: it has an oval to subcircular outline, a plano-convex to gently concavo-convex non-sulcate profile, sometimes with a short geniculation. The ornamentation of the ventral valve consists of irregular growth lines and spine bases often arranged in quincunx, with spines arising anteriorly and obliquely from the anterior end of elongate to slightly swollen ridge-like bases especially on the venter. The ventral cardinal margin has slender, long and curved spines. The dorsal valve contains prominent pits sometimes arranged in quincunx, growth lines more prominent than those of ventral valve, and occasionally weak rugae expressed more strongly anteriorly. Dorsal spines are very scarce or absent, fine and minute when present. The cardinal ridge and median septum are present in dorsal valve, but brachial impressions are indistinct or weakly impressed. The cardinal process is trilobed dorsally and the muscle scars are mainly smooth, and non-dendritic. The dorsal adductor scars are usually on raised subtriangular platforms.

Beside the type species, other material ascribed to Levipustula are from the Carboniferous of Siberia (Maxwell 1951) and Transbaikal (Kotljar & Popeko 1967), Europe (Campbell 1961; Böger & Fiebig 1963; Winkler Prins 1968; Rehòr & Rehòrova 1972; Korejwo 1986), Argentina (Amos 1960; Amos et al. 1963; Taboada 1997; Cisterna & Sterren 2008), Bolivia (Rocha-Campos et al. 1977), and questionably also from the early Permian of western Yunnan in southwest China (Shen et al. 2000). The Siberian Krotovia tolli Fredericks (1931) was closely compared with Levipustula levis by Maxwell (1951), but in our view the former appears to have a more densely arranged spines, different dorsal muscle scars, wide and strongly impressed brachial ridges and a quadrifid cardinal process; all these features would suggest that Krotovia tolli Fredericks is very different from Levipustula. Levipustula baicalensis (Maslennikov) (Kotljar & Popeko 1967) from Transbaikal was later assigned to Lanipustula Klets (1983). Levipustula breimeri Winkler Prins was cautiously compared with Tuberculatella Waterhouse by Waterhouse (1982), due to lack of sufficient comparative material. The other European species ascribed to Levipustula by Campbell (1961) and Böger & Fiebig (1963), such as L. rimberti (Waterlot) and L. piscariae (Waterlot), all exhibit closely spaced radial to quincunxially arranged elongate and swollen spine bases on the ventral valve and conspicuous fine dorsal spines (Demanet 1943), unlike Levipustula which has short and only slightly swollen spine-ridges on the ventral valve and either has very

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scarce, minute dorsal spines or lacks them. These European species are best included in Lanipustula Klets, especially so for Demanet’s specimens and probably also those figured by Rehór & Rehórova (1972) and Korejwo (1986). The single ventral exterior figured as Levipustula? sp. by Shen et al. (2000) does not contain enough information to warrant a detailed comparison. Previous reports of Levipustula from Argentina and Bolivia are discussed below.

Levipustula levis Maxwell, 1951 (Figs. 6-9)

1892 Productus brachythaerus Sowerby; Jack & Etheridge, p. 248.

1963 Levipustula levis Maxwell; Amos et al., p. 131, pl. I, figs 1-3.

1964 Levipustula levis Maxwell; Maxwell, p. 43, pl. 7, figs 17-20.

1964 Levipustula levis Maxwell; Hill & Woods, pl. C7, fig. 9.

1965 Levipustula levis Maxwell; McKellar, p. 8, pl. 2, figs. 8-11.

1977 Levipustula levis Maxwell; Rocha-Campos et al., p. 292-293, pl. 1, figs 1-5.

1997 Levipustula levis Maxwell; Taboada, pl. III, figs 13-14.

2000 Levipustula levis Maxwell; Brunton et al., p. 453, only figs 299.3a-3b.

2007 Levipustula levis Maxwell; Césari et al., p. 40, figs 3, 4.

2007 Levipustula levis Maxwell; González & Díaz Saravia, p. 56, fig. 4.

?2008 Levipustula levis?; Cisterna & Sterren, p. 142, figs 5a, 5b.

Comments. Maxwell’s original description was detailed. Although no specific diagnosis was provided for this species, Maxwell stated that diagnosis for the genus would be applicable for the species. Here we propose an emended diagnosis based on our examination of the type material and other published information mentioned above.

Emended diagnosis. Levipustula of transversely oval to subcircular outline and a plano-convex to gently concavo-convex non-sulcate profile, sometimes with a short geniculation. Hinge width slightly less than maximum width, which is located at about the posterior third of valve length. Although variable, shells are commonly up to 25 mm in width and 20 mm in length, with maximum width of 42 mm and maximum length of 35 mm (F11913a), and an average width/length ratio of 1.25. Spine density on ventral valve highly variable, between 3-5 per 5 mm across the venter at 10 mm from umbo, with slender,

curved halteroid spines along hinge line and on flanks, attaining 4-13 mm in length. Dorsal spines very scarce, minute and inconspicuous. Cardinal process bilobed ventrally, trifid dorsally, lacking alveolus. Dorsal anterior adductor scars smooth, usually on subtriangular raised platforms, posterior adductor scars not impressed. Ventral diductor scars are subrectangular in outline, striate radially and slightly raised.

Discussion. All the material examined shows great variation in some features, mainly in the density of ventral spines, length of spine-ridge bases and spine size. The quincunxial arrangement of spines is not always regular, nor is the convexity and size of individuals. The specimen F11913a, from Portion 200, Parish of Palen, County of Ward, not figured by Maxwell (1951), is a mould of a ventral valve exterior (Fig. 7H). This specimen appears to represent the largest (up to 40mm in width) specimen known for this species, with the lowest spine density (3 spines per 5 mm at 10mm from umbo). If this specimen is compared with the paratypes F11967 (Figs. 6H, 9H; also figured by Maxwell 1951, pl. 2, fig. 7) and F11909 (Figs. 6I, 9G; Maxwell 1951, pl. 2, fig. 5), both from the type locality (Portion 5V, Parish of Neerkol, County of Livingstone), the latter two specimens would be considered very small (26 mm and 28 mm maximum width, respectively), but have greater convexity and more densely distributed spines with 5 spines per 5 mm at 10 mm from umbo. Nevertheless, a ventral valve of another specimen from the type locality, labelled F13120 (Fig. 6G; Maxwell 1951, pl. 2, fig. 13), also has a lower spine density (3-4 spines per 5 mm at 10 mm from umbo) than the two paratypes, and looks intermediate in appearance among Maxwell’s original collection. This specimen therefore could be considered intermediate between morphological extremes, and hence highlights the great intraspecific variability of L. levis.

Outside Australia, L. levis has been described or figured from the upper Palaeozoic of Argentina (Amos 1960; Amos et al. 1963; Cúneo & Sabattini 1987; Taboada 1997; Cisterna & Sterren 2008) and Bolivia (Rocha-Campos et al.1977). In Argentina, L. levis was first reported by Amos (1960) according to specimens from Patagonia (Chubut province) and later from western Argentina (Precordillera, San Juan province) (Amos et al. 1963). Patagonian specimens were sent by Amos to Australian experts, who agreed with Amos’ taxonomic determination (Campbell 1961; Maxwell 1964). Unfortunately, to date the exact localities of the Argentine material examined by these two Australian palaeontologists remain unclear. More recently, the Patagonian specimens

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were reviewed by Simanauskas (1996), who referred most of the material illustrated by Amos (1960) to a new species in a different genus; Lanipustula patagoniensis Simanauskas, 1996. Recently, Taboada (2008) also examined this material and excluded specimens MLP 5086c and MLP 5086d of Amos (1960, pl. 2, figs. 10-11; Amos 1979, p. 78, fig. a) from the synonymy of L. patagoniensis; instead assigning them, and also material from the Pampa de Tepuel Formation at La Carlota Post locality, to another new species in a different genus, Verchojania inacayali Taboada. 2008. Verchojania inacayali is here considered also to include labelled specimens MLP 5086a and 5086b of Amos (1960, pl. 3, figs. 2, 5).

Amos (in Amos et al. 1963: p. 131, pl. I, figs.

1-4) illustrated, as Levipustula levis, specimens from La Capilla Formation at Las Cambachas, western Argentina, and this was accompanied by a brief description. He indicated that those specimens were almost identical to Patagonian “L. levis”, but were smaller. The western Argentine Levipustula levis material was also stated to display more erect ventral spines with a somewhat irregular distribution pattern. Material collected by us from the western Argentine locality show signs of recrystallisation that would have obscured its ornamentation details, including spine arrangement. Besides, Amos (in Amos et al. 1963) reported the occurrence of L. levis in the Hoyada Verde and Leoncito localities, both only a few kilometers to the

Figure 6. Levipustula levis Maxwell, 1951, from the type locality: Portion 5V. Parish of Neerkol, County of Livingstone, Queensland. A-B, D-E, holotype, F11900, composite internal mould in dorsal view, composite internal mould in antero-ventral view, latex cast dorsal valve interior, composite internal mould in postero-ventral view. C, F, F11901-a, dorsal valve internal mould, latex cast dorsal valve interior. G, F13120, latex cast ventral valve exterior. H, paratype F11967, latex cast ventral valve exterior. I, paratype F11909, latex cast ventral valve exterior. J, paratype, F11901-b, latex cast ventral valve exterior. (all figures x 1.5; scale bar: 10mm).

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Figure 7. Levipustula levis Maxwell, 1951. A, F, F11908, external ventral mould, latex cast ventral valve exterior, from Portion II, Parish of Baywulla, County of Yarrol, Queensland. B-D, F, F11906, composite internal mould in ventral and dorsal views, latex cast dorsal valve interior, from Portion 2162, Parish of Neerkol, County of Livingstone, Queensland. E, G, latex cast ventral valve exteriors of unlabeled specimens of the type material. H, F11913a, latex cast of ventral valve exterior, from Portion 200. Parish of Palen, County of Ward, Queensland. I, F11914, latex cast dorsal valve interior, from Portion 200, Parish of Palen, County of Ward, Queensland. J, F11915, latex cast dorsal valve interior, from Portion 198. Parish of Palen, County of Ward, Queensland. K, latex cast dorsal valve interior of an unlabeled specimen of the type material (all figures x 1.5; scale bar: 10mm).

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south, and from where comparable additional material has been discovered (Taboada 1997; Césari et al. 2007; Cisterna & Sterren 2008). This material and also other material from the La Capilla Formation at the Villa Corral locality (González & Taboada 1987) are here illustrated. Our examination of all these materials from both western Argentina and Patagonia reveals a complementary characterisation for the Argentine Levipustula levis, as follows: Shell of gently concavo-convex profile with a suboval to subcircular outline, maximum width and length of 22 mm and 16 mm, respectively, with an average width/length ratio of 1.25. Hinge width less than maximum width, which is located in the posterior third of valve length. Ventral valve with fine, mostly regular concentric growth lines, and spines in a regular quincunxial arrangement, with 5 spines in 5 mm at 10 mm from the umbo. Spine-ridge bases elongate, slightly swollen, of 2/3 mm

maximum length. Dorsal valve with small dimples or pits, concentric growth lines, and few sparse minute spines located posteriorly near the hinge line. Scarce, poorly preserved interiors show radially striate diductor scars and slightly raised subtriangular to suboval platforms, where anterior adductors are attached. Cardinal process bilobed ventrally, trifid dorsally. As Amos (in Amos et al. 1963) and more recently, Taboada (2006), have indicated, western Argentine specimens are comparatively small, on average, compared to the type specimens of Levipustula levis from eastern Australia, but they are otherwise externally almost indistinguishable.

Rocha-Campos et al. (1977) have illustrated L. levis from the upper part of the Taiguati Formation (Macharetí Group) at Parapeti River in the southern subandean area of Bolivia. The Bolivian material appears comparable in shell size to individuals from western Argentina than to the

Figure 8. Levipustula levis Maxwell from the locality 145-5, Ridgeland, northwest of Rockhampton, Queensland. A, FML-PI 3558-1, latex cast ventral valve exterior. B, FML-PI 3558-2, latex cast ventral valve exterior. C, FML-PI 3558-3, latex cast dorsal valve exterior. D, FML-PI 3559-2, latex cast dorsal valve exterior. E, FML-PI 3560, latex cast dorsal valve exterior. F, FML-PI 3561, latex cast dorsal valve interior. G, FML-PI 3559-1, latex cast dorsal valve interior (all figures x 1.5; scale bar: 10mm).

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Australian type material of L. levis.

Lanipustula Klets, 1983

Type species. Pustula baicalensis Maslennikov, 1960, from the Gutay Formation, upper part of lower to middle Carboniferous of the Transbaikal region, Russia.

Discussion. Lanipustula was proposed to differentiate and accommodate a set of species previously variably assigned to several different genera, such as Avonia Thomas, Pustula Thomas and Levipustula Maxwell, all widely recorded in the lower to middle Carboniferous of the Transbaikal and Khabarovsk regions of eastern Russia (Maslennikov 1960; Kotljar & Popeko 1967; Zavodowsky & Stepanov 1970; Klets 1983, 2005a, 2005b). Although accepted by most authors (Abramov & Grigorjeva 1983, 1986; Astafieva-Urbajtis et al. 1983; Pavlova & Manankov 1983; Lazarev 1985, 1990, 1991), the validity of Lanipustula has been questioned by Roberts (in Roberts et al. 1993, 1995) who

considered its minor morphological differences to not be of generic significance. More recently, Lanipustula was considered valid by Simanauskas (1996), Brunton & Lazarev (1997), Brunton et al. (2000), Simanauskas & Archbold (2002) and Waterhouse (2002), with additional species recorded from the Carboniferous of Verkhoyansk (Abramov & Grigorjeva 1983, 1986), the lower Carboniferous (upper Viséan) to the lower Permian (Asselian) of Mongolia (Astafieva-Urbajtis et al. 1983; Lazarev 1991; Manankov 2004), as well as from the lower Pennsylvanian of Argentina (Simanauskas 1996). As mentioned above, Lanipustula is also possibly represented in Europe, notably by L. piscariae (Waterlot), L. rimberti (Waterlot) and, in particular, species figured by Demanet (1943).

Klets (1983) has emphasised the distinction between Lanipustula and Levipustula in the position of the cardinal ridges and the shape and outline of anterior adductor scars. In Klets’ view, the cardinal ridges do not diverge from the cardinal margin in Lanipustula, its anterior adductor scars are distinctively pyriform and posterior adductor

Figure 9. Levipustula levis Maxwell from Argentina. A, FML-PI 1267, partially decorticate ventral valve exterior. B-C, FML-PI 1268, partially decorticate articulate specimen in ventral and dorsal views. D-E, FML-PI 3824, latex cast ventral valve exterior, impression ventral valve exterior. F, FML-PI 3240-1, mould dorsal valve exterior. G, paratype F11909, latex cast ventral valve exterior. H, paratype F11967, latex cast ventral valve exterior, from the type locality for comparison. I, FML-PI 3558-3, latex cast dorsal valve exterior from locality 145-5 of Ridgeland for comparison (all figures x 1.5; scale bar: 10mm).

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scars poorly defined or not visible, and the anterior ridges strongly elevated. All these characters used to distinguish Lanipustula from Levipustula were only considered intraspecific by Taboada (2006). For example, the ‘invisible posterior adductor scars’ considered by Klets to be characteristic of Lanipustula has also been confirmed for some species of Levipustula. It is important to note that the presence or absence of posterior adductor scars could be an artifact of preservation and therefore it should not be taken as a character for a generic distinction. The usually raised subtriangular platforms could be where both the anterior and posterior adductors were located, but discernable only with exceptional preservation. The cardinal ridges are usually also parallel to the hinge line in Levipustula but occasionally show a slight divergence from the hinge (Fig. 8F). In addition, considerable similarity of the elevated anterior edge of the anterior adductor scar platforms exists in the genera. Notwithstanding these minor, ‘intraspecific’ differences, Lanipustula and Levipustula are significantly different in the abundance of dorsal spines, as was pointed out by Brunton et al. (2000), and also in the development of concentric ornamentation in that Lanipustula has strong rugae and/or lamination on both valves.

According to the description and illustration by Klets (1983), Lanipustula has a subcircular to transversely subelliptical outline and a gently concavo-convex nonsulcate profile, a short geniculation, sometimes with a barely discernible fold. Ornamentation of the ventral valve comprises concentric rugae and growth lines with short swollen spine bases and oblique spines quincunxially arranged. The dorsal valve exterior is ornamented with concentric rugae, stronger laterally and on the auricles. Abundant, minute hollow spines are present in concentric rows anterolaterally. The ventral diductor scars are typically subtriangular in outline and striate. The cardinal process is bilobate internally, trifid externally and the cardinal ridges are parallel to the hinge line. The median septum is grooved

posteriorly or has an alveolus. The adductor scars are located on raised subtriangular platforms and are distinctly separated. The brachial ridges are weakly expressed.

Complementary topotypic material of Lanipustula patagoniensis Simanauskas is here described and discussed, and another new Patagonian species of Lanipustula, a possible descendant of the former, is proposed. In addition, a new species of Absenticosta from western Argentina is described and suggested as the probable ancestor of the Patagonian Levipustulini lineage Lanipustula-Verchojania-Jakutoproductus-Piatnitzkya.

Lanipustula patagoniensis Simanauskas, 1996 (Fig. 10)

1960 Levipustula levis Maxwell; Amos, pl. III, only figs. 1, 3.

1996 Lanipustula patagoniensis; Simanauskas, p. 303-304, figs. 3A-3E.

Comments. Simanauskas (1996) has given a detailed description of the species, and an expanded diagnosis is provided below, based on that of Simanauskas (1996) and our observations of abundant topotypic and other material.

Material. Silicone moulds of holotype MLP 26196, paratypes MLP 26200a, 26200b, 26200c and specimen MLP 5088a, and one hundred external and internal moulds of both valves, FML-PI 890-893, 1693-1/1693-4, 1694, 1694-1/1694-17, 1701-1/1701-31 and MEF-PI 2111, 2127, 2320, 2388, 2391, 2574-2575, 2578 from the “bed with Productus and Fenestella” of Suero (1948) and bed “Ft-13” of Freytes (1971).

Diagnosis. Large Lanipustula, subcircular to transversely subrectangular in outline with gently concave dorsal valve and a convex ventral valve, with its strongest convexity in the posterior third of valve and flattened anteriorly, non-sulcate. Well

Figure 10. Lanipustula patagoniensis Simanauskas, 1996. A, MLP 26196, silicone mould of the interior of an articulate specimen (holotype). B, MLP 26200a, silicone mould of the interior of an articulate specimen (paratype). C, MLP 26200b, silicone mould of the exterior of an articulate specimen (paratype). D, FML-PI 1701-11, internal mould of ventral valve in antero-ventral view. E, MLP 26200c, silicone mould of dorsal valve interior (paratype). F, MLP 5088a, silicone mould of dorsal valve interior. G, FML-PI 1701-10, silicone mould of the exterior of an articulate specimen in dorsal view. H, MEF-PI 2368, internal mould of an articulate specimen in dorsal view. I, MEF-PI 370, silicone mould of ventral valve exterior. J, FML-PI 1693-1a, latex mould of ventral valve exterior. K, MEF-PI s/nº, internal mould of ventral valve. L, MEF-PI 2368, internal mould of an articulate specimen in dorsal view. M, MEF-PI 2574-1, silicone mould of ventral valve exterior in antero-ventral view. N, LIEB-PI 147 internal mould of ventral valve in antero-ventral view. O, R, FML-PI 891, latex mould of cardinal process (x 2) and external mould of dorsal valve. P-Q, FML-PI 890a, internal mould of dorsal valve and its latex mould. S-T, FML-PI 1693-2, external mould of dorsal valve and its latex mould. U, FML-PI 1693-1, external mould of dorsal valve. V, FML-PI 1694-3b, external mould of dorsal valve. W, FML-PI 1694-3a, external mould of dorsal valve (all figures natural size, otherwise as indicated; scale bar: 10mm).

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developed ventral trail, dorsal valve with short but well marked geniculation. Hinge width less than maximum width, which is located at midlength of the valve. Shell size in mature specimens commonly between 25-35 mm in width and 20-25 mm in length, with largest sizes up to 45 mm and a width/length ratio of 1.20-1.48. Ventral valve with weak rugae and growth lines, slightly swollen ridge-like spine bases in quincunxial arrangement (3-4 per 5 mm at 10 mm from umbo) with fine subprostrate spines more oblique anteriorly. Ventral trail with shorter spine bases and longer oblique to recumbent spines (up to 5 mm in length) variably arranged, from quincunxial to broadly concentric, and only concentric growth lines anteriorly. A row of 4-5 suberect spines are present along hinge line. A thin myophragm separates large subtriangular diductor scars, striate longitudinally and flabellate anteriorly. Dorsal valve more rugose than ventral valve, with small dimples, a row of minute, erect circular spines near the hinge margin and variably abundant (5-9 per 5 mm at 10 mm from hinge), suberect spines often in concentric rows over the disc. Dorsal trail with concentric growth lines, weaker rugae and scarce spines. Dorsal anterior adductor scars smooth, usually on subtriangular raised platforms and posterior adductor scars smooth, slightly impressed. Cardinal ridges usually parallel to hinge line or slightly divergent. Brachial ridges rarely impressed, set at approximately 45º to median septum. Alveolus present in juvenile specimens but variably closed at maturity or absent in gerontic stage. Cardinal process is bilobed ventrally and trifid dorsally.

Discussion. This Patagonian species exhibits diagnostic features of Lanipustula, comprising abundant external dorsal spines, rugose-lamellose concentric bands with growth lines on both valves and quincunxial arrangement of swollen ridge-like spine bases with oblique spines anteriorly on ventral valve. Internally, Patagonian material shows typical, but variable, anteriorly raised subtriangular adductor scar platforms and slightly impressed posterior adductor scars, besides other features. Nevertheless, L. patagoniensis is the largest of any species in the genus (width up to 40 mm) and has a well developed ventral trail and geniculation, unlike younger representatives of the genus. These characteristics of L. patagoniensis closely resemble younger representatives of Impiacus Lazarev, particularly the type species I. dzhinsetuensis Lazarev & Suursuren, 1988 (see also Lazarev 1991; Brunton et al. 2000) from the upper Viséan of the Gobi Altai, Mongolia. Lazarev (1991) has proposed a phylogenetic lineage from Absenticosta through Impiacus to

Lanipustula. As will be discussed below, this possible phylogenetic link of L. patagoniensis to Lazarev’s proposed Absenticosta-Impiacus-Lanipustula lineage is significant and could be evidence of a comparable or parallel evolutionary trend, albeit delayed, for the development and diversification of the Tribe Levipustulini in Argentina.

The synonymy of Lanipustula patagoniensis, according to Simanauskas (1996), has included all material from Patagonia previously assigned to Levipustula levis Maxwell by Amos (1960), with the exception of one specimen MLP 5088a (Amos 1960, pl. III, only figs. 1, 3). But this specimen, in our view, should also belong here. Several other specimens figured by Amos (1960, pl. 2, figs. 10-11 and pl. 3, figs. 2 and 5) have already been transferred to Verchojania inacayali Taboada (2008). Still one other specimen, figured by Amos (1960, pl. III, figs. 4a, 4b), is here reassigned to Lanipustula kletsi sp. nov., described below.

Material figured by Campbell (1961) as Levipustula levis Maxwell from the Booral Formation of New South Wales, Australia, appears to be conspecific with or at least very close to L. patagoniensis. Likewise, Lanipustula would also embrace other specimens from New South Wales, such as those figured by Muir-Wood & Cooper (1960, pl. 50, figs. 12-15), Amos (1960, pl. 3, fig. 1), Muir-Wood (1965, figs. 342.5a-b), McCarthy et al. (1974, figs. 4G-H) and Brunton et al. (2000, fig. 299.3c).

To date, L. patagoniensis is known from two different horizons within the Pampa de Tepuel Formation. One of these was reported by Suero (1948) as “beds with Fenestella and Productus”, which was wrongly equated by Simanauskas (1996, fig. 1) with the bed Ft1-9a of Freytes (1971) as the latter is located nearly 1000 m stratigraphically below the former and only bears scarce fossil remains according to Freytes (1971). The second L. patagoniensis-bearing horizon was indicated by Simanauskas (1996) as bed Ft1-13 of Freytes (1971), which was also incorrectly positioned by Simanauskas (1996) as being 1000 m stratigraphically above the “beds with Fenestella and Productus” (Simanauskas 1996, fig. 1). In fact, detailed field work by the senior author has revealed that the two L. patagoniensis-bearing horizons from the type section of the Pampa de Tepuel Formation, i.e., the “beds with Fenestella and Productus” and “Ft1-13”, are stratigraphic equivalents exposed at two different localities separated by a 1.3 km long creek on the western flank of Tepuel Hill, one north of the creek at S43º41’41”W70º44’32.5’’ at 1167 m above sea level, the other south of the creek at S43º42’18.1’’W70º44’09.8’’ at 1213 m

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above sea level. As a result of this revelation, the correct stratigraphic distribution of L. patagoniensis and hence the L. patagoniensis assemblage (i.e., formerly Levipustula levis Zone) has been restricted to a narrow horizon of a few metres thick in the middle third of the Pampa de Tepuel Formation. Nevertheless, recent fieldwork carried out by Dr. Alejandra Pagani and one of the authors (ACT) has revealed a new record of Lanipustula patagoniensis stratigraphically 450 m above (S43º41’22.7’’, W70º43’37.3’’) the “beds with Fenestella and Productus” of Suero (1948). Stratigraphic correlation indicates this horizon would also be present in the lowermost part of the Las Salinas Formation (Member LS-1 of González 1972) in the Languiñeo Hill (near Olivera Post), where fragmentary specimens of L. patagoniensis have been collected (labeled FML-PI nº 1703, Palaeontological Institute, Miguel Lillo Foundation, Tucumán, Argentina).

Lanipustula kletsi sp. nov. ( Fig. 11)

1960 Levipustula levis Maxwell; Amos, pl. 3, only figs. 4a, 4b.

Material. Holotype FML-PI 1702-9, paratypes FML-PI 1702-1, 1702-2, 3837, 3839. Other material, FML-PI 1702-3/1702-22, 3840, LIEB-PI 294-295, from the Las Salinas Formation, LS9 member (González 1972), Languiñeo Hill, around S43º05’37,1’’, W70º13’00,5’’ and S43º05’05.0’’, W70º12’14.7’’.

Etymology. In honour and memory of the late Russian geologist and palaeontologist Dr. Alexandr Klets.

Diagnosis. Medium size Lanipustula, subelliptical to subcircular to variably transverse outline with planar to gently concave dorsal valve with a short geniculation. Maximum width located at about posterior third of valve. Ventral valve with moderate to strong rugae and concentric growth lines, short swollen spine bases and fine oblique spines in quincunxial arrangement (4-5 per 5 mm at 5 mm from umbo). Dorsal valve with generally weak rugae, but becoming stronger anterolaterally, with small dimples and suberect spines (6-7 per 5 mm at 10 mm from hinge), in roughly quincunxial to concentric arrangement.

Description. Medium-sized Lanipustula , subelliptical, subcircular to variably transverse in outline with planar to gently concave dorsal valve with short geniculation. Ventral valve slightly convex with its strongest convexity in the posterior third of valve and flattened anteriorly.

Hinge width less than maximum width, which is located at the posterior third of valve. Shell size in mature specimens commonly between 18-20 mm in width and 13-17 mm in length, with largest size up to 25 mm and a width/length ratio between 1.00-1.54. Ventral valve with moderate to strong rugae and concentric growth lines, short swollen spine bases and fine oblique spines in quincunxial arrangement (4-5 per 5 mm across the venter at 5 mm from umbo). A row of 4-5 suberect spines emerge at 45º from hinge line and reach up to 6 mm in length; slender and slightly curved spines on flanks and trail, up to 8 mm long. A thin myophragm separates striate, longitudinally subtriangular diductor scars extending 1/3 of valve length. Dorsal valve with weak rugae, but comparatively stronger laterally and anteriorly, with small dimples and suberect spines (6-7 per 5 mm at 10 mm from hinge), in roughly quincunxial to concentric arrangement. Dorsal internal features variably expressed. Median septum varies from a thin median ridge extending 1/3 valve length posteriorly arising from short buttress plates surrounding an alveolus (Fig. 7K-L, FML-PI 1702-5), to a breviseptum that reaches 1/2 valve length, arising from a massive posterior platform without alveolus (Fig. 11P, FML-PI 1702-7). One specimen (Fig. 11T, FML-PI 1702-21) shows an intermediate stage, with a breviseptum from a posterior platform with a shallow alveolus developed. Dorsal anterior and posterior adductor scars subelliptical in outline, smooth, located on variably raised subtriangular platforms. Cardinal ridges parallel to hinge line, extending laterally as a marginal rim (Fig. 11R, FML-PI 3837). Cardinal process bilobed ventrally and trifid dorsally. Brachial ridges usually well developed (Fig. 11N-O, FML-PI 1702-9), diverging approximately at 55º to median septum, smooth on inner surface, bounded by a low ridge. Endospines densely distributed and minute, disposed toward the anterior margin of the valve.

Discussion. Lanipustula kletsi sp. nov. has all the diagnostic characters of the genus plus the well preserved posterior adductor scars, which were thought absent by Klets (1983) when he diagnosed the taxon based on the Russian material. Previously, Lazarev (1991) has also reported posterior adductor scars in Lanipustula toalensis Lazarev.

Lanipustula kletsi sp. nov. closely resembles the type species L. baicalensis (Maslennikov) (Maslennikov 1960; Kotljar & Popeko 1967; Grigorjeva & Manankov 1970; Zavodowsky & Stepanov 1970; Klets 1983, 2005b; Astafieva-Urbajtis et al. 1983; Pavlova & Manankov 1983; Manankov 2004), from which it can

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Figure 11. Lanipustula kletsi sp. nov. A-B, Paratype FML-PI 1702-1, internal mould of ventral valve and latex mould of ventral valve exterior. C, Paratype FML-PI 1702-2, latex mould of ventral valve exterior. D, FML-PI 1702-8, mould ventral valve exterior. E, FML-PI 1702-18, internal mould of ventral valve. F, LIEB-PI 295, external mould of ventral valve. G-H, FML-PI 1702-17, external mould of dorsal valve (continued opposite)

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be distinguished in the weaker expression of the subtriangular adductor platform and the more uniform size of the spines around the auricles (unlike the Russian species whose auricular spines are 2-3 times thicker than those distributed elsewhere). In addition, L. baicalensis (Maslenikov) lacks dorsal spines along the cardinal margin and auricles whereas these spines are clearly present in L. kletsi sp. nov. Other species comparable with L. kletsi sp. nov. are L. natalinensis Abramov & Grigorjeva (1983) and L. tyraensis Abramov & Grigorjeva (1986), from the upper and lower Carboniferous of the Verkhoyansk region in eastern Russia, respectively. These two species have a greater density of ventral spines and a globose profile when compared with L. kletsi sp. nov. In addition, L. toalensis Lazarev (1991) from the upper Viséan of northern Mongolia, is almost twice as large, has a very low profile, longer spine bases and almost prostrate ventral spines, unlike the gentle concavo-convex profile, short swollen spine bases and oblique ventral spines of the Patagonian species.

The older Serpukhovian L. patagoniensis Simanauskas (1996), as described above, differs from L. kletsi sp. nov. mainly in its stronger concavo-convex profile, larger size, weaker ventral rugae, stronger dorsal rugae, longer swollen spine bases, lesser density of ventral spines, but greater density of dorsal spines and higher dorsal subtriangular adductor scar.

Tribe ABSENTICOSTINI Waterhouse, 2002

Absenticosta Lazarev, 1991

Type species. Absenticosta uldzejtuensis Suursuren & Lazarev, 1991 (in Lazarev 1991), from the upper? Viséan Urkhiyn-Khundiy Formation, Khangay Highland, Mongolia.

Comments. The diagnosis and discussion provided by Lazarev (1991) indicates that Absenticosta has a concavo-convex visceral disc, a distinct trail and concentric rugae, sometimes interrupted and undulose. Bases of spines are tubercular, usually without elongate bases. Spines on the ventral valve are randomly scattered, but are absent at the cardinal angles. Spines on the dorsal valve are straight and confined primarily to the geniculation. The cardinal ridge deviates

slightly from the cardinal margin and continues, with thickening, to the anterior margin of the visceral disc.

Besides the type species, from the upper Viséan of Mongolia and the Transbaikal region (Plicatifera plicatilis in Kotljar & Popeko 1967), the genus also includes an unnamed species from the upper Viséan of Mount Bus-Ula, Mongolia (Lazarev 1991). The genus is also possibly present in northeastern Russia (Ganelin in Lazarev 1991). Concentric ornamentation in Absenticosta resembles Plicatifera Chao (1927) but this character in the latter genus is regular, unlike the highly irregular (in strength, undulation and bifurcation patterns) rugae of Absenticosta (Lazarev 1991). Absenticosta differs from the closely related genus Impiacus Lazarev & Suursuren (1988), mainly in its more sharply defined concentric sculpture and absence of spines at the cardinal angles (Lazarev 1991).

Absenticosta bruntoneileenae sp. nov. (Fig. 12)

? 2001 Tuberculatella peregrina (Reed); Siman-auskas & Cisterna, p. 217-220, figs. 5a-5g.

Material. Fifteen specimens of external and internal moulds of ventral and dorsal valves. Holotype FML-PI 1201-1, Paratypes 1201-2/1201-4, 1204-4. Other material 1201-8, 1204-1, 1204-3, 1204-5/1204-6, 1204-9, from the El Paso Member of the San Eduardo Formation (latest Viséan), Elloy creek (Barreal hill) around S31º40’42.9’’, W69º25’16.7’’.

Etymology. Dedicated to Mrs. Eileen Brunton, devoted wife of the outstanding British brachiopodologist, the late Dr. Howard Brunton.

Diagnosis. Medium-sized to large Absenticosta, subelliptical in outline with gently concave to evenly convex profile and a short trail. Hinge width slightly less than maximum width. Shell size commonly between 15-22 mm in width and 11-15 mm in length with a width/length ratio from 1.31 to 1.47. Short, fine subprostrate to oblique spines scattered over ventral valve with a density of 2-3 in 5 mm at 5 mm from umbo. Scarce (up to two), suberect spines located on umbonal slopes toward cardinal margin and only one stout spine at each cardinal angle. Conspicuous subcircular shallow dimples over dorsal valve, along with

and its silicone mould. I, Paratype FML-PI 3839, external mould of dorsal valve. J, FML-PI 3840, external mould of dorsal valve. K-L, FML-PI 1702-5, internal mould of dorsal valve and its silicone mould. M, LIEB-PI 294, internal mould of dorsal valve. N-O, Holotype FML-PI 1702-9, internal mould of dorsal valve and its latex mould. P, FML-PI 1702-7, interior of dorsal valve. Q, FML-PI 1702-20, internal mould of dorsal valve. R, Paratype FML-PI 3837, internal mould of dorsal valve. S, FML-PI 1702-14, external mould of dorsal valve. T, FML-PI 1702-21, internal mould of dorsal valve (all figures x 1.5; scale bar: 10mm).

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fine, erect, minute spines in roughly concentric arrangement, 5 in 5 mm on dorsal visceral disc.

Description. Medium to large size Absenticosta, regularly transverse subelliptical outline, gently concave to evenly convex profile with short trail. Ventral valve evenly convex, flattened anteriorly toward the trail. Hinge width slightly less than maximum width, which is located at the posterior third of valve length. Shell size commonly between 15-22 mm in width and 11-15 mm in length with a width/length ratio from 1.31 to 1.47. Ventral valve with low concentric and irregular rugae, variably undulating and occasionally discontinuous. Growth lines inconspicuous. Short, fine subprostrate to oblique spines scattered over the valve with a density of 2-3 in 5 mm at 5 mm from umbo. Scarce (up to two), suberect spines located on umbonal slopes near cardinal margin and only one stout spine at cardinal angle (left auricle of specimen FML-PI 1201-1). A narrow, shallow ventral sulcus possibly present (FML-PI 1201-1).Ventral interior with thin, long myophragm extending 1/3 of valve length, which separates large, deeply radially striate, subtriangular diductor scars extending 2/3 of valve length. Dorsal valve gently concave, abruptly and shortly geniculate. Ornamentation of low concentric rugae, expressed better posteriorly, generally weaker than those of ventral valve. Concentric growth lines better developed

anteriorly. Conspicuous subcircular shallow dimples over dorsal valve; dorsal spines minute, erect, arranged in roughly concentric pattern, 5 in 5 mm on disc. Partially decorticate dorsal valve interiors reproduce external ornamentation and exhibit radial rows of fine pustules. Cardinal process bilobed internally. Other characters not preserved.

Discussion. The shell outline, profile and ornamentation of the Argentinean specimens suggest assignment to Absenticosta Lazarev (1991). As pointed out by Lazarev (1991), the genus usually lacks spines on the auricles, but larger and younger representatives could exhibit a single spine on each auricle, as indeed seen in our new Patagonian species. The presence of auricular spines suggests a similarity to Impiacus Lazarev & Suursuren, 1988, but the low concentric ornamentation and spine pattern of our new species indicate a clear difference from this genus.

Absenticosta bruntoneileenae sp. nov. can be distinguished from the type species A. uldzejtuensis Suursuren & Lazarev (in Lazarev 1991) (Lazarev & Suursuren 1992; Brunton et al. 2000) by its having low and fewer concentric rugae over the ventral valve and weaker and more irregularly developed concentric ornamentation on the dorsal valve.

Absenticosta bruntoneileenae sp. nov. constitutes the first record of the genus in the

Figure 12. Absenticosta bruntoneileenae sp. nov. A-C, Holotype FML-PI 1201-1, exterior of ventral valve in ventral, posterior and lateral views. D, Paratype FML-PI 1201-2, internal mould of ventral valve. E, Paratype FML-PI 1201-4, partially decorticate dorsal valve interior. F, FML-PI 1201-6, external mould of dorsal valve. G, Paratype FML-PI 1204-3, fragmentary articulate specimen in dorsal view. H, FML-PI 1201-7, partially decorticate dorsal valve interior. I, FML-PI 1201-3, dorsal valve external mould of a juvenile specimen. J, Paratype FML-PI 1204-4, internal mould of ventral valve. K, FML-PI 1201-5, partially decorticate ventral valve exterior (all figures x 1.5; scale bar: 10mm.)

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Southern Hemisphere and documents another brachiopod with antitropical distribution (see below).

PALAEOBIOGEOGRAPHIC ORIGIN AND EVOLUTION OF THE PATAGONIAN LEVIPUSTULINI LINEAGEThe Levipustulini have long been recognised from northeast Asia (Kolyma, Omolon, Verkhoyansk, Okhotsk and Mongolian regions) where they are commonly found in association with low-diversity temperate brachiopod faunas ranging in age from Carboniferous to early Permian (Klets 1983, 2005a, b; Ganelin & Durante 2002; Klets & Kislyakov 2000; Klets et al. 2006; Ganelin & Biakov 2006; Ganelin 2010). As in Gondwana, late Palaeozoic marine faunas of northeast Asia lack warm-water fossil elements, except those from long-travelled accreted oceanic terranes (Ganelin & Tschernjak 1996; Shi 2006). This similarity in late Palaeozoic marine faunal composition and distribution patterns between the high latitude regions of both hemispheres has led to the recognition of biogeographic bipolarity (or antitropicality) by a number of authors (e.g., Ustritskiy 1971; Shi et al. 1995; Shi & Grunt 2000). This distinctive late Palaeozoic biogeographic phenomenon has usually been taken to imply bipolar cooling patterns at high latitudes (Waterhouse 1967, 1969, 1976; Dickins 1985) and has also been referred to as a potential correlation interval between high latitudes of both hemispheres (Roberts et al. 1976; Archbold 2005, Shi & Shen 2005; Shi 2006; Taboada et al. 2005; Taboada 2008; Taboada & Pagani 2010).

The Tribe Levipustulini commonly exhibits antitropical distribution but two separate, morphologically similar lineages from this tribe are shared only between northeast Asia and Patagonia (Tepuel-Genoa Basin). The parallel lineages are both composed of Lanipustula, Verchojania and Jakutoproductus, with Absenticosta here interpreted as the possible common ancestor for both lineages. The two lineages are believed to have diverged in the early Carboniferous, followed by parallel evolution in the high latitude regions of both hemispheres under comparable cool climate conditions during the Late Palaeozoic (Shi et al. 2008). In this scenario, the development of these two related lineages could be regarded as a consequence of parallel evolution in vastly separate geographic locations but with similar or comparable climatic/ecological conditions. This is an alternative interpretation to the ‘trans-oceanic dispersal’ model that has so far been advocated as a mechanism of initiating and sustaining late Palaeozoic biogeographic bipolarity (see Shi & Grunt 2000, reference therein provided). If

parallel evolution was indeed the mechanism for initiating the late Palaeozoic bipolarity between Patagonia and northeast Asia, it must have commenced during the late Viséan at a time when the Earth was experiencing a global cooling phase allowing the transoceanic and global dispersal of some cool-water brachiopod genera (e.g. Absenticosta, among others) from lower to higher latitudes. Ancestral (Viséan) Absenticosta was restricted to the southern shelf waters of Siberia, then located in the low to middle latitudes of the Northern Hemisphere during the middle and late Palaeozoic (Cocks & Torsvik 2007). It is likely that Absenticosta might have migrated westward from southern Siberia through the Austro-Panthalassic-Rheic corridor (Taboada & Shi 2009) southward to southwestern South America (western Argentina). This possible presence of an early Carboniferous (Viséan in particular) trans-Pangean-type Austro-Panthalassic-Rheic corridor as a migration pathway and connection between the Palaeotethys and eastern Panthalassa (or southwestern Gondwana in this case) has also been suggested to explain the temporal and spatial distribution patterns of several other Carboniferous brachiopod genera such as Yagonia (Taboada & Shi 2009) and Aseptella (Cisterna & Simanauskas 1999; Martinez Chacón & Winkler Prins 1999). In this model, Absenticosta bruntoneileenae sp. nov., recorded in the El Paso Member (latest Viséan) from western Argentina, either represents a direct immigrant from Siberia, or a descendant of a yet undiscovered ancestral taxon that had evolved through allopatric speciation. Lazarev (1991) has discussed a phylogenetic lineage from Absenticosta through Impiacus to Lanipustula in the Northern Hemisphere. The morphological resemblance of Absenticosta bruntoneileenae sp. nov. to earlier Impiacus and, to a lesser extent, L. patagoniensis, is here interpreted as evidence of a comparable evolutionary trend in Patagonia. Temporally compared to the evolution of the lineage in northeast Asia, the evolution of the parallel Patagonian lineage was almost synchronous with its northern counterparts. As illustrated in Figure 4, the Patagonian Levipustulini lineage of Lanipustula (Serpukhovian) through Verchojania (Moscovian-Asselian) to Jakutoproductus (early Artinskian) commenced in the latest Viséan and terminated in the early Artinskian. The lineage is characterised by having its ventral spines becoming progressively more quincunxially arranged, coupled with a progressively weaker and more variable concentric ornamentation pattern of growth lines and rugae (synapormorphy). In addition, ventral spines are often oblique or subprostrate with elongate or slightly swollen

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bases throughout the lineage, but dorsal spines appear to have evolved from being abundant for early Carboniferous taxa to either scarce or completely lost for Permian representatives. Many of these evolutionary features are manifest in the Levipustulini genus Piatnitzkya, the youngest (middle Artinskian) in this lineage in Patagonia. This genus, located stratigraphically at the upper end of the proposed Patagonian Levipustulini phylogenetic lineage, demonstrates evidence for many of the suggested evolutionary trends; for example, it has no dorsal spines, nor concentric ornamentation. In the Northern Hemisphere (northeast Asia), Jakutoproductus appears to represent the youngest genus in the Levipustulini phylogenetic lineage, where the youngest Jakutoproductus tends to have a larger shell size, more conspicuous ornament, a deeper angular sulcus and a more marked geniculation and trail (Shi 1994), besides a quadrifid cardinal process (Manankov 2004).

LATE PALAEOZOIC PALAEOGEOGRAPHY OF PATAGONIA Patagonia has been considered by some workers as a suspect terrane accreted to Gondwana in late Palaeozoic times (Ramos 1984, 2008). Alternative interpretations also exist, suggesting that Patagonia had always remained as an integral part of South America since the early Palaeozoic (Dalla Salda et al. 1990, 1994). Recently,

Pankhurst et al. (2006) have suggested that northern Patagonia (North Patagonian Massif) was already part of Gondwana in Ordovician times. In contrast, southern Patagonia (Deseado Massif), perhaps together with parts of West Antarctica (see also Hervé et al. 2006; Ramos 2008), was continental crust to the south of the major plate boundary (Fig. 13), and seems to have belonged to a parautochthonous entity that started to collide with the northern Patagonian block in Late Palaeozoic times (Pankhurst et al. 2006; Von Gosen 2008; Balhburg et al. 2009). Pankhurst et al. (2006) have also suggested that the southern Patagonian continental block, on which the foreland mid Carboniferous–early Permian Tepuel-Genoa Basin was formed before, during and after the collision, would not have been separated very far from the southern South America margin during the Palaeozoic drifting. If we accept that the present-day Patagonia represents two amalgamated blocks as suggested by Pankhurst et al. (2006), their respective late Palaeozoic floras and faunas would have followed different biogeographical and evolutionary pathways, the significance of which would depend on the degree of their mutual geographical and climatic separation throughout the late Palaeozoic. To date, very limited published information is available in regards to this aspect of Patagonia’s late Palaeozoic history.

There have been more and longer late Palaeozoic

Figure 13. Schematic reconstruction of SW Gondwana showing Late Palaeozoic plate configurations (simplified from Pankurst et al. 2006). The North Patagonian Somuncurá Massif had formed part of continental South America since Ordovician (or at least Devonian) times, separated from the Southern Patagonian Deseado Massif (or Deseado terrane) to the south. The question marks indicate the current uncertainty about the true extent of Southern Patagonian Deseado Massif. The Antarctic Peninsula would consist of Late Carboniferous additions, but is shown as at the present day for the purpose of easy identification. FI=Falkland/Malvinas Islands plateau, EWM=Ellsworth-Whitmore mountains crustal block.

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glacial episodes recognised in Patagonia than in western Argentina (southwestern margin of Gondwana) (Taboada 2008, 2010; Taboada & Pagani 2010). This would suggest that Patagonia was located closer to the palaeopole than western Argentina throughout the late Palaeozoic (Taboada et al. 2005; Taboada & Pagani 2010; Taboada & Shi 2010). This scenario agrees with a possible local block rotation of Patagonia (Rapalini 2005), allowing a displacement between its northern boundary and Gondwana up to about 1000 km in the middle to late Carboniferous (Ramos 2008; Rapalini et al. 2010). As a consequence, strong faunal contrasts were established between western Argentina and Patagonia since middle Carboniferous times when the late Bashkirian-early Moscovian Marginovatia-Maemia fauna appeared in western Argentina (Uspallata-Iglesia Basin), north of Patagonia. This fauna was linked to a climatic warming event in western Argentina (Taboada 1997, 2006). During this relative warming event, a number of brachiopod genera and species, notably including Marginovatia, Maemia, Reticularia and Neochonetes granulifer (Owen), seem to have migrated to western Argentina by taking advantage of the available trans-Pangean-type Austro-Panthalassic-Rheic corridor. However, the deglaciation event and subsequent global warming in the Bashkirian may have sufficiently warmed up the trans-Pangean-type Austro-Panthalassic-Rheic corridor to the extent that it now served as a thermal barrier for the biotic interchange between the Northern Hemisphere and southwestern Gondwana where Patagonia was located in a subpolar setting (probably up to 70ºS) experiencing a cold climatic regime. This warming event and the interpreted rise of a thermal barrier across western and northwestern Argentina (and northern Chile, Peru and Boliva further north) is here considered as the trigger for the initiation of biogeographical differentiation between western Argentina and Patagonia in the Bashkirian. As a result, Pennsylvanian-early Permian (Artinskian) cold-water adapted faunal assemblages among others, were developed in Patagonia, allowing the evolution of the postulated Absenticosta-Lanipustula-Verchojania-Jakutoproductus-Piatnitzkya lineage in Patagonia, accompanied by increasing faunal endemism and palaeobiogeographic isolation.

The weak glacial record since Bashkirian times in western Argentina contrasts with Australian evidence of widespread glaciations until Permian times. This asymmetrical climatic record reflects the clockwise rotation of Gondwana suggested for the Carboniferous and consequent drifting of western Argentina to lower latitudes unlike

eastern Australia which drifted toward the south palaeopole. Patagonia, located in an intermediate palaeogeographic setting between western Argentina and Australia during this time, was also rotated and displaced across latitudes, but it must have been shifted to lower latitudes later and more slowly than western Argentina. Patagonia would have been separated from western Argentina and progressively drifted toward southern higher latitudes, thus more or less following the plate motion of Australia in this regard. Consequently, we suggest the isolation of Patagonia throughout the late Palaeozoic, coupled with its southward drift toward a colder climate over this period, could have triggered the parallel evolution of the Levipustulini lineage between Patagonia in southwestern Gondwana and Siberia in northeast Asia.

A thalassocratic regime during the late Asselian-Sakmarian and a global climatic amelioration since the late Sakmarian, could explain the fast recovery of migration pathways for the biotic interchange among Patagonia and other Gondwanan areas, particularly Western Australia and the Cimmerian regions, now stretching from the Middle East to Southeast Asia. This biotic interchange pathway probably occurred through a narrow oceanic connection or an epicontinental seaway incursion as is shown by the relative high diversity of the Cimmeriella fauna from Patagonia (Taboada & Pagani 2010). This scenario is in agreement with the palaeogeographic reconstruction of the Tepuel-Genoa Basin at the western border of the Deseado Massif constituting a discrete terrane detached from the northern Somuncurá Massif, which by this time would have been amalgamated with the southwestern margin of South America (Pankhurst et al. 2006). On the other hand, the Patagonian-Western Australia-Cimmerian palaeobiogeographical connection would be more difficult to explain, if Patagonia had been a single block made of the amalgamated Somuncurá and Deseado Massifs, as was postulated by Ramos (2008). In Ramos’ reconstruction model, a direct marine pathway for the biotic interchange between the Tepuel-Genoa Basin and the easternmost mentioned regions would have been blocked. In our view, the collision of the Patagonian Deseado Massif against mainland South America began after early Permian times.

ACKNOWLEDGEMENTSThis paper is dedicated to the memories of two very close colleagues: the late Professors Neil W. Archbold and late Dr. Alexandr Klets. The former supported ACT to visit Australia in 2004 to examine the type material of Levipustula levis.

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We thank Dr. Alexandr Klets for his provision of comparative fossil material from Siberia and his constructive discussions about Lanipustula. Valuable comments and suggestions by two anonymous reviewers improved the final version of the manuscript. Gratitude is expressed to Mr. Mario Campaña for drawings. Material studied from Argentina was collected during several fieldtrips with staff and logistic support of the Laboratory on Evolution and Biodiversity (LIEB-UNPSJB), the Miguel Lillo Foundation and/or the Egidio Feruglio Palaeontological Museum. The National Research Council of Argentina (CONICET) and the LIEB have provided laboratory facilities. GRS thanks the Australian Research Council for financial support which enabled ACT to visit Australia in 2008 and 2010.

REFERENCESAbrAmov, b.S., 1970. Biostratigrafiia kamennougolnykh

otlozhenii Sette-Dabana (Iuzhnoe Verkhoiane). Nauka, Moskva, 178 p.

Ab r A m o v, b.S. & Gr i G o r j e wA, A.D . , 1983. Biostratigrafiia i brakhiopody srednego i verkhnego karbona Verkhoiania. Paleontologicheskii Institut, Trudy 200, 168 p.

Ab r A m o v, b.S. & Gr i G o r j e wA, A.D . , 1986. Biostratigrafiia i brakhiopody nizhnego karbona Verkhoiania. Nauka, Moskva, 192 p.

AfAnASjevA, G.A., boGoSlovSkAiA, m.f., lAzArev, S.S., PAvlovA, e.e. & SuurSuren, Sh., 1988. Novye taksony vizenskikh brakhiopod i ammonoidei iz Gobiiskogo Altaia. Sovmestnaya Sovetsko-Mongolskaya Paleontologicheskaya Ekspeditsiya, Trudy 33, 51-58.

AmoS, A.j., 1960. Algunos Chonetacea y Productacea del Carbonifero Inferior y Superior del Sistema de Tepuel, Provincia de Chubut. Revista de la Asociación Geológica Argentina 15, 81-107.

AmoS, A.j., 1964. A review of the marine Carboniferous stratigraphy of Argentina. 22nd International Geological Congress, New Delhi, Proceedings 9(9), 53-72.

AmoS, A.j., 1979. Guia Paleontologica Argentina. Parte I: Paleozoico. Faunas Carbónicas y Pérmicas. Consejo Nacional de Investigaciones Cientificas y Técnicas (Buenos Aires), 154 p.

AmoS, A.j. & rolleri, e.o., 1965. El Carbónico marino en el Valle Calingasta-Uspallata (San Juan y Mendoza). Boletín de Informaciones Petroleras (Buenos Aires) 368, 1-23.

AmoS, A.j., bAlDiS, b. & CSAky, A., 1963. La fauna del Carbonífero medio de la Formación La Capilla y sus relaciones geológicas. Ameghiniana 3(4), 123-132.

AnDreiS, r. & Cúneo, r., 1989. Late Paleozoic high constructive deltaic sequences from Northwestern Patagonia, Argentine Republic. Journal of South American Earth Sciences 2, 10-24.

ArChAnGelSky, S., AzCuy, C.l., CéSAri, S.n., González, C.r., hüniCken, m.A., mAzzoni, A. & SAbAttini, n., 1996. Correlación y edad de las biozonas. 203-226 in Archangelsky, S. (ed.), El Sistema Pérmico en la República Argentina y en la República Oriental del Uruguay. Academia Nacional de Ciencias, Córdoba.

ArChbolD, n.w., 2005. The Permian Time Scale: correlation of the marine Permian of Greater Gondwana. 47 in Pankhurst, R.J. & Veiga, G.D. (eds), Gondwana 12: Geological and Biological Heritage of Gondwana, Abstracts. Academia Nacional de Ciencias, Córdoba.

AStAfievA-urbAjtiS, k.A., DurAnte, m.v., mAnAnkov, i.n. & PAvlovA, e.e., 1983. Kharakteristika kompleksov organicheskikh ostatkov. Sovmestnaya Sovetsko-Mongolskaya Paleontologicheskaya Ekspeditsiya, Trudy 21, 13-24.

bAhlburG, h., vervoort, j.D., AnDrew, Du frAne, S.A., boCk, b., AuGuStSSon, C. & reimAnn, C., 2009. Timing of crust formation and recycling in accretionary orogens: Insights learned from the western margin of South America. Earth-Science Reviews 97, 215–241.

böGer, h. & fiebiG, h., 1963. Die fauna des Westdeutschen Oberkarbons 2. Die articulaten brachiopoden des Westdeutschen Oberkarbons. Palaeontographica, Abt.A 122(4-6), 111-165.

brunton, C.h.C. & lAzArev, S.S., 1997. Evolution and classification of the Productellidae (Productidae), Upper Paleozoic Brachiopods. Journal of Paleontology 71(3), 381-394.

brunton, C.h.C., lAzArev, S.S. & GrAnt, r.e., 1995. A Review and New Classification of the Brachiopod Order Productida. Palaeontology 38(4), 915-936.

brunton, C.h.C., lAzArev, S.S., GrAnt, r.e. & jin yu-GAn, 2000. Productidina. 424-609 in Kaesler, L.R. (ed.), Treatise on Invertebrate Paleontology, Part H, Brachiopoda Revised 3. Geological Society of America, Boulder and University of Kansas Press, Lawrence.

CAmPbell, k.S.w., 1961. Carboniferous Fossils from the Kuttung Rocks of New South Wales. Palaeontology 4(3), 428-474.

CAmPbell, k.S.w., 1962. Marine Fossils from the Carboniferous Glacial Rocks of New South Wales. Journal of Paleontology 36(1), 38-52.

CéSAri, S.n., Gutiérrez, P.r., SAbAttini, n., ArChAnGelSky, A., AzCuy, C.l., CArrizo, h.A., CiSternA, G., Cúneo, r.n., DíAz SArAviA, P., González, C.r., Di PASquo, m., PAGAni, m.A., leCh, r., Sterren, A., tAboADA, A.C. & verGel, m.m., 2007. Paleozoico Superior de Argentina: un registro fosilífero integral en el Gondwana occidental. Asociación Paleontológica Argentina, Publicación Especial 11, Ameghiniana 50º aniversario, 35-54.

ChAo, y., 1927. Productidae of China, Part I: Producti. Palaeontologia Sinica, Series B 5(2), 1-192.

AAP Memoir 41 (2011) 111

CiSternA, G. A. & SimAnAuSkAS, t., 1999. Aseptella (Brachiopoda) en el Paleozoico Tardío de Argentina. Revista Española de Paleontología 14(1), 117-122.

CiSternA, G.A. & Sterren, A.f., 2008. Late Carboniferous Levipustula fauna in the Leoncito Formation, San Juan province, Argentine Precordillera: biostratigraphical and palaeoclimatological implications. Proceedings of the Royal Society of Victoria 120(1), 137-147.

CoCkS, l.r.m. & torSvik, t.h., 2007. Siberia, the wandering northern terrane, and its changing geography through the Palaeozoic. Earth-Science Reviews 82, 29–74.

CuCChi, r.j., 1980. La Formación Esquel: nueva interpretación estratigráfica. Revista de la Asociación Geológica Argentina 35(2), 167-173.

Cúneo, r. & SAbAttini, n., 1987. Flora y Fauna de la base de la Formación Río Genoa en la localidad de Ferraroti, Pérmico inferior de Chubut, Argentina. Memorias del 4º Congreso Latinoamericano de Paleontología 1, 283-298.

DAllA SAlDA, l., CinGolAni, C., vArelA, r., 1990. The origin of Patagonia. Comunicaciones, Una revista de geología andina 41, 55–61.

DAllA SAlDA, l., vArelA, r., CinGolAni, C.A., ArAGón, e., 1994. The Río Chico Paleozoic crystalline complex and the evolution of Northern Patagonia. Journal South American Earth Sciences 7, 377–386.

DemAnet, f., 1943. Les horizons marins du Westphalien de la Belgique et leurs faunes. Musee Royal d’Histoire Naturelle de Belgique, Memoires 101, 1-166.

DiCkinS, j.m., 1985. Late Paleozoic glaciation. BMR Journal of Australian Geology & Geophysics 9, 163-169.

freDeriCkS, G.n., 1931. Verkhnepaleozoiskaia fauna Kharaulakhskikh gor. Akademiia Nauk SSSR, Otdelenie Matematicheskikh i Estestvennykh Nauk, Izvestiia 2, 199- 228.

freyteS, e., 1971. Informe geológico preliminar sobre la Sierra de Tepuel (Deptos. Languiñeo y Tehuelches, prov. de Chubut). Informe YPF. Buenos Aires (unpublished).

GAnelin, v.G., 2010. Brachiopods in the Late Palaeozoic benthic biota of northeast Asia. Geological Society of Australia, Abstracts 95, 43-44.

GAnelin, v.G. & biAkov, A.S., 2006. The Permian biostratigraphy of the Kolyma-Omolon region, Northeast Asia. Journal of Asian Earth Sciences 26, 225-234.

GAnelin, v.G. & DurAnte, m.v., 2002. Biostratigraphy of the Carboniferous of Angaraland. Newsletter on Carboniferous Stratigraphy 20, 23-26.

GAnelin, v.G. & tSChernjAk, G.e., 1996. Marine basins of Northeast Asia. 207-234 in The Carboniferous of the World, b.III. Instituto Tecnológico GeoMinero de España.

González, C.r., 1972. La Formación Las Salinas, Paleozoico superior de Chubut (República Argentina). Parte I. Estratigrafía, facies y ambientes de sedimentación. Revista de la Asociación Geológica Argentina 27(1), 95-115.

González, C.r., 1985. Esquema bioestratigráfico del Paleozoico Superior marino de la Cuenca Uspallata-Iglesia, República Argentina. Acta Geológica Lilloana 16(2), 231-244.

González, C.r., 1993. Late Paleozoic faunal succession in Argentina. 12º Congres International de la Stratigraphie et Geologie du Carbonifere et Permien, Comptes Rendus 1, 537-550.

González, C.r. & DíAz SArAviA, P., 2007. Faunas y paleoclimatología del Paleozoico Superior de Argentina: revisión crítica. Acta Geológica Lilloana 20(1), 41-72.

González, C.r. & tAboADA, A.C., 1987. Nueva localidad fosilífera del Carbónico marino en la provincia de San Juan. 10º Congreso Geológico Argentino, Acta 3, 103-105.

GrADStein, f.m. & oGG, j.G., 2009. The Geologic Time scale. 26-34 in Hedges, S.B. & Kumar, S. (eds), The Timetree of Life. Oxford University Press, Oxford.

GrADStein, f.m., oGG, j.G., Smith, A.G., bleeker, w. & lourenS, l.j., 2004. A new Geologic Time Scale, with special reference to Precambrian and Neogene. Episodes 27(2), 83-100.

GriGorjewA, A.D. & mAnAnkov, i.n., 1970. Produktidy. 76-80 in Obut, A.M. (ed.), Karbon Omolonskogo I Iugo-Zapadnoi Chasti Kolymskogo Massivov. Institut Geologii i Geofiziki, Trudy 60.

hervé, f., miller, h. & PimPirev, Ch., 2006. Patagonia-Antarctica connections before Gondwana break-up. 217-228 in Fütterer, D.K., Damaske, D.K., Miller, H. & Tessensohn, F. (eds), Antarctica. Springer-Verlag, Berlin.

hill, D. & wooDS, j.t., 1964. Carboniferous Fossils of Queensland. Queensland Palaeontographical Society, Brisbane, 1-32.

jACk, r.l. & etheriDGe, r., 1892. The geology and palaeontology of Queensland and New Guinea. Publication of the Geological Survey of Queensland 42, 1-756.

kASChirtzev, A.S., 1959. Polevoi Atlas Fauni Permskikh Otlozhenii Severo-Vostoka SSSR. Akademiya Nauk SSSR, Leningrad, 85 p.

kletS, A.G., 1983. A New Carboniferous Productid Genus. Paleontological Journal 1, 70-75.

kletS, A.G., 2005a. Benchmarks for correlation of Carboniferous and Permian marine deposits in Siberia and Far East. Russian Geology and Geophysics 46(3), 256-272.

kletS, A.G., 2005b. Upper Paleozoic of Marginal Seas of Angarida. Institute of Petroleum Geology, Novosibirsk, 240 p.

kletS, A.G. & kiSlyAkov, S.G., 2000. Regional Stratigraphic Divisions of the Carboniferous in the

AAP Memoir 41 (2011)112

North of Khabarovsk Territory. Geology of Pacific Ocean 16, 31-40.

kletS, AG., buDnikov, i.v., kutyGin, r.v., biAkov, A.S. & Grinenko v.S., 2006. The Permian of the Verkhoyansk-Okhotsk region, NE Russia. Journal of Asian Earth Sciences 26, 258-268.

korejwo, k., 1986. Biostratigraphy of the Carboniferous Deposits of the Swidnik Blocks (Lublin Coal Basin). Acta Geologica Polonica 36(4), 335-346.

kotlyAr, G.v. & PoPeko, l.i., 1967. Biostratigrafiia, mshanki i brakhiopody verkhnego paleozoiia Zabaikalia. Zapiski Zabaikalskogo Filiala Geograficheskogo Obshchestva SSSR, Trudy Otdeleniia Geologii 28(5), 1-257.

lAzArev, S.S., 1985. Brakhiopody Echinoconchidae i Buxtoniidae. Paleontologicheskii Zhurnal 1, 64-74.

lAzArev, S.S., 1990. Evoliutsiia i sistema produktid. Paleontologicheskii Institut, Trudy 242, 1-171.

lAzArev, S.S., 1991. The phylogenetic series of Carboniferous brachiopods Absenticosta-Impiacus-Lanipustula from Mongolia. Paleontological Journal 25(4), 65-78.

lAzArev, S.S. & SuurSuren, Sh., 1988. Otriad Product ida . 53-55 in Afanasjeva, G.A. , Bogoslovskaia, M.F., Lazarev, S.S., Pavlova, E.E. & Suursuren, Sh., Novye taksony vizenskikh brakhiopod i ammonoidei iz Gobiiskogo Altaia. Sovmestnaya Sovetsko-Mongolskaya Paleontologicheskaya Ekspeditsiya, Trudy 33.

lAzArev, S.S. & SuurSuren, Sh., 1992. New productids (Brachiopoda) in the Carboniferous of Mongolia. 61-69 in Grunt, T.A. (ed.), New taxa of fossil invertebrates of Mongolia. Sovmestnaya Mongolskaya Paleontologicheskaya Ekspeditsiya, Trudy 41.

limArino, C.o. & SPAlletti, l.A., 2006. Paleogeography of the upper Paleozoic basins of southern South America: An overview. Journal of South American Earth Sciences 22, 134-155.

mAnAnkov, i.n., 2004. New species of Early Permian brachiopods and biostratigraphy of the boreal basin of Mongolia. Paleontological Journal 38(4), 366-372.

mArtínez ChACón, m.l. & winkler PrinS, C.f., 1999. Distribución paleobiogeográfica de Aseptella (Productida): conexión entre los braquiópodos carboníferos de España y Argentina. Trabajos de Geología, Universidad de Oviedo 21, 221-227.

mASlennikov, D.f., 1960. Novye vidy permskikh ekhinokonkhid Zabaikalia. 341-343 in Markowskii, B.P. (ed.), Novye Vidy Drevnikh Rastenii I Bespozvonochnykh SSSR 1. Nedra, Moskva.

mAxwell, w.G.h., 1951. Upper Devonian and Middle Carboniferous Brachiopods of Queensland. University of Queensland, Department of Geology, Papers 3(14), 1-27.

mAxwell, w.G.h., 1964. The Geology of the Yarrol Region. Part I. Biostratigraphy. University of

Queensland, Department of Geology, Papers 5(9),1-79.

mCCArthy, b., runneGAr, b. & kleemAn, j.D., 1974. Late Carboniferous-Early Permian Sequence North of Drake, N.S.W. Dates, Deposition and deformation of Beenleigh Block. Queensland Government Mining Journal 75, 1-7.

mCkellAr, r.G., 1965. An Upper Carboniferous Brachiopod Fauna from the Monto District, Queensland. Geological Survey of Queensland, Publication 328, 1-15.

méSiGoS, m., 1953. El Paleozoico superior de Barreal y su continuación austral. Sierra de Barreal. Provincia de San Juan. Revista de la Asociación Geológica Argentina 8(2), 65-109.

muir-wooD, h.m., 1965. Suborder Productidina. 439-510 in Moore, R.C. (ed.), Treatise on Invertebrate Paleontology. Part H. Brachiopoda. Geological Society of America, New York and University of Kansas Press, Lawrence.

muir-wooD, h.m. & CooPer, G.A., 1960. Morphology, classification and life habits of the Productoidea (Brachiopoda). Memoir of the Geological Society of America 81, 1-447.

PAGe, r.f., limArino, C.o., lóPez GAmunDi, o. & PAGe, S., 1984. Estratigrafía del Grupo Tepuel en su perfil tipo y en la región de El Molle, provincia del Chubut. 9º Congreso Geológico Argentino, Actas 1, 619-632.

PAnkhurSt, r.j., rAPelA, C.w. & tASSone, A., 2006. Gondwanide continental collision and the origin of Patagonia. Earth-Science Reviews 76, 235–257.

PAvlovA, e.e. & mAnAnkov, i.n., 1983. Brakhiopody. 33-51 in Afanasjeva, G.A. (ed.), Biostratigrafiia, fauna i flora karbona khrebta Deng-Nuru v iuzhnoi Mongolii. Sovmestnaya Sovetsko-Mongolskaya Paleontologicheskaya Ekspeditsiya, Trudy 21.

rAmoS, v.A., 1984. Patagonia, un continente paleozoico a la deriva? 9º Congreso Geológico Argentino, Actas 2, 311-325.

rAmoS, v.A., 2008. Patagonia: A paleozoic continent adrift? Journal of South American Earth Sciences 26, 235-251.

rAPAlini, A.e., 2005. The accretionary history of southern South America from the latest Proterozoic to the Late Paleozoic: some paleomagnetic constraints. 305-328 in Vaughan, A.P.M., Leat, P.T. & Pankhurst, R.J. (eds.), Terrane Processes at the Margins of Gondwana. Geological Society, London, Special Publications 246.

rAPAlini, A.e., lóPez De luChi, m., mArtínez DoPiCo, C., linC klinGer, f., Giménez, m. & mArtínez, P., 2010. Did Patagonia collide with Gondwana in the Late Paleozoic? Some insights from a multidisciplinary study of magmatic units of the North Patagonian Massif. Geologica Acta 8(4), 349-371.

rehór, f. & rehórovA, m., 1972. Makrofauna

AAP Memoir 41 (2011) 113

uhlonosneho karbonu ceskoslovenske casti hornoslezske panve. Ostravske Muzeum-Sustova Paleontologicka sbirka. Preci venujeme 100, 1-133.

robertS, j., ClAoué-lonG, j. & joneS, P. j., 1995. Australian Early Carboniferous time. 23-40 in Berggren, W.A., Kent, D.V., Aubry, M.P. & Hardenbol, J. (eds), Geochronology, Time Scales and Global Stratigraphic Correlation. SEPM Special Publication 54.

robertS, j., hunt, j.w. & thomPSon, D.m., 1976. Late Carboniferous Marine Invertebrate Zones of Eastern Australia. Alcheringa 1, 197-225.

robertS, j., joneS, j. & jenkinS, t.b.h., 1993. Revised correlations for Carboniferous marine invertebrate zones of eastern Australia. Alcheringa 17, 353-376.

roChA-CAmPoS, A.C., CArvAlho, r.G. & AmoS, A.j., 1977. A Carboniferous (Gondwana) Fauna from Subandean Bolivia. Revista Brasileira de Geociencias 7(4), 287-303.

SArytChevA, t.G. & SokolSkAjA, A.n., 1959. O klassifikatsin lozhnoporistykh brakhiopod. Akademiia Nauk SSSR, Doklady 125(1), 181-184.

SChuChert, C. 1929. Classification of brachiopod genera, fossils and Recent. 10-25 in Schuchert, C. & Levene, C.M. (eds.), Fossilium Catalogus I: Animalia 42, Brachiopoda. W. Junk, Berlin.

Shen, S., Shi, G. r. & zhu, k., 2000. Early Permian brachiopods of Gondwana affinity from the Dingjazhai Formation of the Baoshan Block, Western Yunnan, China. Rivista Italiana di Paleontologia e Stratigrafia 106(3), 263-282.

Shi, G.r., 1994. The Late Palaeozoic brachiopod genus Jakutoproductus Kashirtsev 1959 and the Jakutoproductus verchoyanicus Zone, northern Yukon Territory, Canada. Alcheringa 18, 103-120.

Shi, G.r., 2006. The marine Permian of East and Northeast Asia: an overview of biostratigraphy, palaeobiogeography and palaeogeographical implications. Journal of Asian Earth Sciences 26, 175-206.

Shi G.r. & Grunt t.A., 2000;. Permian Gondwana Boreal antitropicality with special reference to brachiopod faunas. Palaeogeography, Palaeoclimatology, Palaeoecology 155, 239-263.

Shi, G.r. & Shen, S.z., 2005. Permian marine biotas of East and NE Asia and Gondwana links: implications for Permian global marine biogeography and northern Eurasian-Gondwana correlations. 333 in Pankhurst, R.J. & Veiga, G.D. (eds), Gondwana 12: Geological and Biological Heritage of Gondwana, Abstracts. Academia Nacional de Ciencias, Córdoba.

Shi, G.r. & wAterhouSe, j.b., 1996. Lower Permian Brachiopods and Molluscs from the Upper Jungle Creek Formation, Northern Yukon Territory, Canada. Geological Survey of Canada, Bulletin 424, 1-241.

Shi, G.r., ArChbolD, n.w. & zhAn, l.P., 1995.

Distribution and characteristics of mid-Permian (Late Artinskian-Ufimian) mixed/transitional marine faunas in the Asian region and their palaeogeographical implications. Palaeogeography, Palaeoclimatology, Palaeoecology 114, 241-271.

Shi, G.r., tAboADA, A.C. & kletS, A.G., 2008. Gondwana-Eurasian Permian biogeographic links: implications for Permian global marine biogeography, inter-continental correlations and climate change. 186-187 in Xiao, W.J., Zhai, M.G., Li, X.H. & Liu, F. (eds), Gondwana 13: Programs and Abstracts. Institute of Geology and Geophysics, Dali.

SimAnAuSkAS, t., 1996. Una nueva especie de Lanipustula (Productoidea, Brachiopoda) del Paleozoico superior de Patagonia, Argentina. Ameghiniana 33(3), 301-305.

SimAnAuSkAS, t. & ArChbolD, n.w., 2002. Early Permian Jakutoproductus (Productida: Brachiopoda) from Patagonia, Argentina. Alcheringa 26, 465-474.

SimAnAuSkAS, t. & CiSternA, G., 2001. Braquiópodos articulados de la Formación El Paso, Paleozoico superior, Precordillera Argentina. Revista Española de Paleontología 16(2), 209-222.

Suero, T., 1948. Descubrimiento del Paleozoico superior en la zona extraandina del Chubut. Boletín de Informaciones Petroleras (Buenos Aires) 287, 31-48.

Suero, T., 1953. Las sucesiones sedimentarias suprapaleozoicas de la zona extraandina del Chubut. Revista de la Asociación Geológica Argentina 8(1), 37-63.

Suero, T., 1958. Datos geológicos sobre el Paleozoico superior en la zona de Nueva Lubecka y alrededores (Chubut extraandino, prov. de Chubut). Revista Museo de La Plata (N.S.) 5, Geología 30, 1-28.

tAboADA, A.C., 1989. La fauna de la Formación El Paso, Carbonífero inferior de la Precordillera sanjuanina. Acta Geológica Lilloana 17(1), 113-129.

tAboADA, A.C., 1993. Un nuevo Linoproductidae (Brachiopoda) del Pérmico temprano de Patagonia. 12º Congres International de la Stratigraphie et Geologie du Carbonifere et Permien, Comptes Rendus 1, 589-598.

tAboADA, A.C., 1997. Bioestratigrafía del Paleozoico superior marino del Valle de Calingasta-Uspallata, provincias de San Juan y Mendoza. Ameghiniana 34(2), 215-246.

tAboADA , A.C., 2006. Levipustula Maxwell, 1951 y Lanipustula Klets, 1983 (Brachiopoda, Levipustulini) en Argentina: revisión preliminar. 9º CongresoArgentino de Paleontología y Bioestratigrafía, Resúmenes, p. 193.

tAboADA, A.C., 2008. First record of the Late Paleozoic brachiopod Verchojania in Patagonia, Argentina. Proceedings of the Royal Society of Victoria 120(1), 305-319.

AAP Memoir 41 (2011)114

tAboADA, A.C., 2010. Mississippian-Early Permian brachiopods from western Argentina: Tools for middle- to high-latitude correlation, paleobiogeographic and paleoclimatic reconstruction. Palaeogeography, Palaeoclimatology, Palaeocology 298, 152-173.

tAboADA, A.C. & PAGAni, m.A., 2010. The coupled occurrence of Cimmeriella-Jakutoproductus (Brachiopoda: Productidina) in Patagonia: implications for Early Permian high to middle paleolatitudinal correlations and paleoclimatic reconstruction. Geologica Acta 8(4), 519-540.

tAboADA, A.C. & Shi, G.r., 2009. Yagonia Roberts (Brachiopoda: Chonetidina) from the Malimán Formation, Lower Carboniferous of western Argentina: paleobiogeographical implications. Alcheringa 33, 223-235.

tAboADA, A.C. & Shi, G.r., 2010. Taxonomic review and evolutionary trends of Levipustulini and Absenticostini (Brachiopoda) from Argentina: palaeobiogeographic and palaeocl imatic implications. Geological Society of Australia Abstracts 95, 115.

tAboADA, A.C., ArChbolD, n.w., González, C.r. & SAbAttini, n., 2005. The Late Carboniferous-Early Permian Tepuel fauna of Patagonia: updated brachiopods records. 349 in Pankhurst, R.J. & Veiga, G.D. (eds), Gondwana 12: Geological and Biological Heritage of Gondwana, Abstracts. Academia Nacional de Ciencias, Córdoba.

uGArte, F.R., 1966. El Suprapaleozoico del Río Genoa (Chubut) e inmediaciones. Acta Geológica Lilloana 7, 379-406.

uStritSky, V.I., 1971. Biostratigrafiia verkhnego paleozoia Arktiki. (Biostratigraphy of the Upper Paleozoic in the Arctic.). Nauchno-Issledovatel’skii Institut Geologii Arktiki, Trudy 164, 1-279.

uStritSky, V.I., 1974. O bipoliarnosti faun pozdnego paleozoia. Paleontologicheskii Zhurnal 2, 33- 37.

vAi, G.G., 2003. Development of the palaeogeography of Pangaea from Late Carboniferous to Early Permian. Palaeogeography, Palaeoclimatology, Palaeoecology 196, 125-155.

von GoSen, W., 2008. Stages of Late Palaeozoic deformation and intrusive activity in the western

part of the North Patagonian Massif (southern Argentina) and their geotectonic implications. Geological Magazine 146, 48–71.

voiSey, A.H., 1940. The upper Palaeozoic rocks in the country between Manning and Karuah Rivers, New South Wales. Proceedings of the Linnean Society of New South Wales 65, 192-210.

wAAGen, W., 1883. Salt Range Fossils. I. Productus-Limestone Fossils. Geological Survey of India, Memoirs, Palaeontologia Indica, series 13, 4(2), 391-546.

wAterhouSe, J.B., 1967. Cool water faunas from the Permian of the Canadian Arctic. Nature 216, 47-49.

wAterhouSe, J.B., 1969. World correlations of New Zealand Permian Stages. New Zealand Journal of Geology and Geophys 12, 713-737.

wAterhouSe, J.B., 1976. World correlations for Permian marine faunas. Papers from the Department of Geology, University of Queensland 7(2), 232 p.

wAterhouSe, J.B., 1982. New Carboniferous Brachiopod Genera from Huai Bun Nak, North-East Thailand. Palaeontologische Zeitschrift 56, 39-52.

wAterhouSe, J.B., 2001. Late Paleozoic Brachiopoda and Mollusca from Wairaki Downs, New Zealand. With notes on Scyphozoa and Triassic ammonoids and new classifications of Linoproductoidea (Brachiopoda) and Pectinida (Bivalvia). Earthwise 3, 1-195.

wAterhouSe, J.B., 2002. Classification within Productidina and Strophalosiidina. Earthwise 5, 1-62.

wAterhouSe, J.B., 2008. Golden Spikes and Black Flags – macro-invertebrate faunal Zones for the Permian of East Australia. Proceedings of the Royal Society of Victoria 120, 345-372.

winkler PrinS, C.F., 1968. Carboniferous Productidina and Chonetidina of the Cantabrian Mountains (NW Spain): Systematics, Stratigraphy and Palaeoecology. Leidse Geologische Mededelingen 43, 41-126.

zAvoDowSky, v.m. & StePAnov, D.l., 1970. Tip Brachiopoda. 70-182 in Kulikov, M.V. (ed.), Polevoi Atlas Permskoi Fauny i Flory Severo-Vostoka SSSR. Magadanskoe Knizhnoe Isdatelstvo, Magadan.